Evolution of tail fork depth in genus Hirundo

A classic example of a sexually selected trait, the deep fork tail of the barn swallow Hirundo rustica is now claimed to have evolved and be maintained mainly via aerodynamic advantage rather than sexually selected advantage. However, this aerodynamic advantage hypothesis does not clarify which flight habits select for/against deep fork tails, causing diversity of tail fork depth in hirundines. Here, by focusing on the genus Hirundo, we investigated whether the large variation in tail fork depth could be explained by the differential flight habits. Using a phylogenetic comparative approach, we found that migrant species had deeper fork tails, but less colorful plumage, than the other species, indicating that migration favors a specific trait, deep fork tails. At the same time, tail fork depth but not plumage coloration decreased with increasing bill size – a proxy of prey size, suggesting that foraging on larger prey items favors shallower fork tails. Variation of tail fork depth in the genus Hirundo may be explained by differential flight habits, even without assuming sexual selection.     

Divergent rectrix moult: the implications and conditions of moult sequence

Wing and tail morphology strongly affect flight performance which may consequently decline during feather moult due to the creation of feather gaps in the flight‐surface. Hence, the size and shape of moult‐related gaps may directly affect flight capacity. Here, I examined the divergent rectrix moult sequence compared to the more common distal moult sequence. In the divergent moult, the focus of rectrix moult is shifted from the tail centre (R₁; rectrices numbered distally from mid‐tail outward) to another rectrix (R₂ or R₃), and then rectrices are moulted bidirectionally, towards the tail centre and outwards. The result of this moult sequence is the splitting of the tail gap into multiple smaller gaps. Using a large moult database including 5669 individuals of 47 Western Palaearctic passerine species, I found evidence of divergent moult sequence for only seven species. Using comparative and experimental approaches, I found that the divergent rectrix sequence is correlated with higher moult speed and lower aerodynamic cost. Furthermore, the divergent rectrix sequence is more common among adults than juveniles. This work focused on the feather moult sequence – a seldom studied aspect of the avian life‐history. I propose that moult‐related aerodynamic costs may be an important evolutionary factor not only in moult speed, but also in moult sequence.     

Limits on the evolution of tail ornamentation in birds

In birds, elongated tails are one of the most common and most studied ornaments. However, the tail also has an aerodynamic function, playing a role in turning and slow flight. If a tail is to function as an ornament, then there will be an inevitable conflict between the aerodynamic role and the signaling role. Aerodynamic theory has developed to the point where it is possible to predict the aerodynamic and mechanical consequences of ornamental tails of different sizes and shapes. Tail elongation will influence many different mechanical and aerodynamic parameters. For at least some and possibly all of these parameters, there will be limits that are placed by the bird's anatomy, morphology, or physiology on the extent to which the effect of tail elongation could be tolerated. For example, if a particular tail morphology meant that the power required to fly exceeded the power available from the flight muscles, then the bird would obviously be unable to fly with such a tail. To examine whether these considerations could limit the development of ornamental tails, the effect of long tails of different shapes was examined on three parameters: static balance, lift-to-drag ratio, and the cost of flight. All three of these parameters were found to limit the range of possible tail morphologies that could be developed by birds. These limits were most acute for small birds, which may not be able to operate with a long tail of any shape. Slightly larger birds would be able to develop elongated streamers and forked tails but not pintails or wedge-shaped tails. Medium to large birds are less constrained and could develop a much wider range of tails than small birds, but there will always be limits to the sizes of tail ornament that could be developed. Thus the physical consequences of ornamentation on ecology and behavior are likely to be responsible for some of the patterns of ornamentation observed in nature.     

Great tits do not compensate over time for a radio‐tag‐induced reduction in escape‐flight performance

The use of biologging and tracking devices is widespread in avian behavioral and ecological studies. Carrying these devices rarely has major behavioral or fitness effects in the wild, yet it may still impact animals in more subtle ways, such as during high power demanding escape maneuvers. Here, we tested whether or not great tits (Parus major) carrying a backpack radio‐tag changed their body mass or flight behavior over time to compensate for the detrimental effect of carrying a tag. We tested 18 great tits, randomly assigned to a control (untagged) or one of two different types of a radio‐tag as used in previous studies in the wild (0.9 g or 1.2 g; ~5% or ~6–7% of body mass, respectively), and determined their upward escape‐flight performance 1, 7, 14, and 28 days after tagging. In between experiments, birds were housed in large free‐flight aviaries. For each escape‐flight, we used high‐speed 3D videography to determine flight paths, escape‐flight speed, wingbeat frequency, and actuator disk loading (ratio between the bird weight and aerodynamic thrust production capacity). Tagged birds flew upward with lower escape‐flight speeds, caused by an increased actuator disk loading. During the 28‐day period, all groups slightly increased their body mass and their in‐flight wingbeat frequency. In addition, during this period, all groups of birds increased their escape‐flight speed, but tagged birds did so at a lower rate than untagged birds. This suggests that birds may increase their escape‐flight performance through skill learning; however, tagged birds still remained slower than controls. Our findings suggest that tagging a songbird can have a prolonged effect on the performance of rapid flight maneuvers. Given the absence of tag effects on reproduction and survival in most songbird radio‐tagging studies, tagged birds in the wild might adjust their risk‐taking behavior to avoid performing rapid flight maneuvers.     

Tail length in birds in relation to tail shape,general flight ecology and sexual selection

Relative tail length (longtailedness) of Palearctic birds was assessed by the standardized residuals of log–log regressions of tail length on wing length and tarsus length. The mean degree of tail shortening was greater than mean degree of tail lengthening, but there was a greater frequency of extreme long-tailed than short-tailed species. Longtailedness was greater in ornamental pin, lyre, deep forked and graduated shaped tails. These shapes (except graduated, for which data were lacking) were also relatively long-tailed according to shortest-rectrix lengths, this extra length potentially contributing compensatory lift. In forked tails, tail ratio increased linearly with longtailedness to above the aerodynamic optimum, and thus the most elongated forked tails were also more deeply forked. Tail shortening was marked for rounded tails, a surprising result in view of their slightly ornamental shape. Phylogenetically independent contrasts showed significantly greater longtailedness in graduated than square-tailed species, confirming the species-wide analysis. In phylogenetically independent contrasts of longtailedness and ecological factors, short-tailed species had significantly greater flight distances than medium-tailed species, but long- and medium-tailed species did not differ in migratory distance, foraging distance, overall flight distance or importance of aerial foraging. The data suggest that ecological factors, i.e. natural selection, are more important in the evolution of short-tailedness than longtailedness in birds, and that an additional influence of sexual selection on tail length and shape is also widespread.     

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.     

Sexual selection in the barn swallow Hirundo rustica. VI. Aerodynamic adaptations

Secondary sexual characters are assumed to be costly to produce and maintain, and this will select for morphological modifications that reduce the magnitude of such costs. Here we test whether a feather ornament, the sexually exaggerated outermost tail feathers of male barn swallows Hirundo rustica, a trait currently subject to a directional female mate preference, and other aspects of the morphology used for flight have been modified to increase aerodynamic performance. This was done by making comparisons among sexes within populations, among individuals varying in tail length within populations, and among populations from different parts of Europe. Male barn swallows experienced reduced drag from their elongated tail feathers by morphological modifications of the ornamental feathers as compared to females. Morphological features of the outermost tail feathers were unrelated to tail length in both males and females within populations. Wing and tail morphology (length of central tail feathers and wings, wing span, wing area, wing loading, and aspect ratio) was modified in males compared to females. Barn swallows with long tails had morphological tail and wing modifications that reduced the cost of a large ornament, and similar modifications were seen among populations. The costs of the exaggerated secondary sexual character were therefore reduced by the presence of cost-reducing morphological modifications. The assumptions of reliable signalling theory, that signals should be costly, but more so to low than to high quality individuals, were not violated because long-tailed male barn swallows had the largest cost-reducing morphological characters.     

Testing the functional significance of tail streamers

Studies of the evolution of elaborate ornaments have concentrated on their role in increasing attractiveness to mates. The classic examples of such sexually selected structures are the elongated tails of some bird species. Elongated tails can be divided into three categories: graduated tails, pin tails and streamers. There seems to be little debate about whether graduated and pin tails are ornaments; i.e. costly signals used in mate choice. However, in the case of streamers there is considerable discussion about their function. It has been suggested that tail streamers could be (i) entirely naturally selected, (ii) entirely sexually selected, (iii) partly naturally and partly sexually selected. The prime example of a species with tail streamers is the swallow (Hirundo rustica) in which both sexes have tail streamers. In this paper we discuss the aerodynamic consequences of different types of manipulation of the streamer and/or outer tail feather. We make qualitative predictions about the aerodynamic performance of swallows with manipulated tail streamers; these predictions differ depending on whether streamers have a naturally or sexually selected function. We demonstrate that these hypotheses can only be separated if tail streamers are shortened and changes in aerodynamic performance measured during turning flight.     

Life history,sexual dimorphism and ‘ornamental’ feathers in the mesozoic bird Confuciusornis sanctus

The life history of Confuciusornis sanctus is controversial. Recently, the species’ body size spectrum was claimed to contradict osteohistological evidence for a rapid, bird-like development. Moreover, sexual size dimorphism was rejected as an explanation for the observed bimodal size distribution since the presence of elongated rectrices, an assumed ‘male’ trait, was uncorrelated with size. However, this interpretation (i) fails to explain the size spectrum of C. sanctus which is trimodal rather than bimodal, (ii) requires implausible neonate masses and (iii) is not supported by analogy with sexual dimorphisms in modern birds, in which elongated central rectrices are mostly sex-independent. Available information on C. sanctus is readily reconciled if we assume a bird-like life history, as well as a pronounced sexual size dimorphism and sexually isomorphic extravagant feathers as frequently observed in extant species.     

Aerodynamic costs of long tails in male barn swallows Hirundo rustica and the evolution of sexual size dimorphism

Exaggerated tail feathers of birds constitute a standard exampleof evolution of extravagant characters due to sexual selection.Such secondary sexual traits are assumed to be costly to produceand maintain, and they usually are accompanied by morphologicaladaptations that tend to reduce their costs. The aerodynamiccosts for male barn swallows Hirundo rustica of having longtails were quantified using aerodynamics theory applied to morphologicaldata from seven European populations. Latitudinal differencesin tail length were positively correlated with differences inflight costs predicted by aerodynamics theory. A positive relationshipbetween aerodynamic costs of long tails and the degree of sexualsize dimorphism was found among populations. Latitudinal differencesin foraging costs may result in tail length being relativelysimilar in males and females in southern populations, whereasthe low foraging costs for males in northern populations mayallow them to cope with higher aerodynamic costs, giving riseto large sexual size dimorphism. Enlargement of wingspan inmales can alleviate but not eliminate the costs of tail exaggeration,and therefore differences in aerodynamic costs of male ornamentswere maintained among populations. Sexual size dimorphism in thebarn swallow arises as a consequence of latitudinal differencesin the advantages of sexual selection for males and the costsof long tails for males and females.     

The importance of tail length for habitat use in the Bearded Tit Panurus biarmicus: an experimental study

The original function of the tail in birds is likely to be related to flight performance or bipedal locomotion. In this study, we examined the role of the long graduated tail of male Bearded Tits Panurus biarmicus , focusing on the function of the tail as a balancing organ when moving in reeds, the habitat in which this species spends its whole life. We performed an aviary experiment using males with shortened, elongated or control (unmanipulated) tails and recorded their use of different reed heights and preferences for different sitting postures. Tail manipulation did not change the use of different reed heights between treatments. The head-up posture was the preferred sitting technique and was independent of tail length. We found that males with elongated tails showed less variation in the use of different sitting postures along the reed. In addition, the use of the 'straddle' sitting technique varied in relation to tail length, tending to be less frequent when tails were experimentally shortened. These results suggest that tail length influences how Bearded Tits move and may exploit their habitat. The role of the tail as a balancing organ when using small twigs or branches may have been underestimated.     

HOW TO COMPENSATE FOR COSTLY SEXUALLY SELECTED TAILS: THE ORIGIN OF SEXUALLY DIMORPHIC WINGS IN LONG-TAILED BIRDS

Recent work on birds suggests that certain morphological differences between the sexes may have evolved as an indirect consequence of sexual selection because they offset the cost of bearing extravagant ornaments used for fighting or mate attraction. For example, long-tailed male sunbirds and widowbirds also have longer wings than females, perhaps to compensate for the aerodynamic costs of tail elaboration. We used comparative data from 57 species to investigate whether this link between sexual dimorphism in wing and tail length is widespread among long-tailed birds. We found that within long-tailed families, variation in the extent of tail dimorphism was associated with corresponding variation in wing dimorphism. One nonfunctional explanation of this result is simply that the growth of wings and tails is controlled by a common developmental mechanism, such that long-tailed individuals inevitably grow long wings as well. However, this hypothesis cannot account for a second pattern in our data set: as predicted by aerodynamic theory, we found that, comparing across long-tailed families, sexual dimorphism in wing length varied with tail shape as well as with sex differences in tail length. Thus, wing dimorphism was generally greater in species with aerodynamically costly graduated tails than in birds with cheaper, streamer-shaped tails. This result was not caused by confounding phylogenetic effects, because it persisted when phylogeny was controlled for, using an independent comparisons method. Our findings therefore confirm that certain aspects of sexual dimorphism may sometimes have evolved through selection for traits that reduce the costs of elaborate sexually selected characters. We suggest that future work aimed at understanding sexual selection by investigating patterns of sexual dimorphism should attempt to differentiate between the direct and indirect consequences of sexual selection.     

Sexual selection and tail streamers in the barn swallow

The functional significance of elongated, narrow tips of the tail feathers of certain birds, so-called tail streamers, has recently been discussed from an aerodynamic point of view, and the effects of sexual selection on such traits have been questioned. We review our long-term field studies using observational and experimental approaches to investigate natural and sexual selection in the barn swallow, Hirundo rustica, which has sexually size-dimorphic outermost tail feathers. Experimental manipulation of the length of the outermost tail feathers has demonstrated sexual selection advantages of tail elongation and disadvantages of tail shortening, with opposite effects for natural selection in terms of foraging efficiency, haematocrit and survival. These findings are contrary to the prediction of a general deterioration from both shortening and elongation, if the tail trait was determined solely by its effects on aerodynamic efficiency and flight manoeuvrability. Patterns of sexual selection in manipulated birds conform with patterns in unmanipulated birds, and selection differentials for different components of sexual selection in manipulated birds are strongly positively correlated with differentials in unmanipulated birds. Age and sex differences in tail length, and geographical patterns of sexual size dimorphism, are also consistent with sexual selection theory, but inconsistent with a purely natural selection advantage of long outermost tail feathers in male barn swallows.     

辽西早白垩世今鸟类一基干有喙新属种(英文)

根据发现于辽宁西部建昌早白垩世九佛堂组一件保存较为完好的标本,描述了原始今鸟类一新属种,李氏叉尾鸟(Schizooura lii gen.et sp.nov.)。系统发育分析表明,它较建昌鸟和古喙鸟进步,但又较热河生物群中其他已知的今鸟类原始。这一新种叉骨呈Ⅴ型,且具有一短的叉骨突,这是这一特征首次在早白垩世今鸟中发现。其胸骨加长,后缘无窗孔,且无很深的凹口。肱骨三角肌脊膨大,近乎达到肱骨长的一半。新鸟的头骨保存较好,显示其具喙的特征,前颌骨直接与额骨相关节。值得关注的是,这件标本首次展示了在中生代鸟类中,除扇形尾羽之外的第二种尾羽类型。新标本具叉形尾羽,中间分隔明显。对现生鸟类的研究表明,叉尾型的空气动力学效率比扇尾型要低,但在性选择方面具有优势。这一新的发现说明,在热河生物群生活的森林环境中,今鸟类的基干种类在尾羽性状上或许已经存在着不同的选择策略。     

The avian tail reduces body parasite drag by controlling flow separation and vortex shedding.

The aerodynamic effect of the furled avian tail on the parasite drag of a bird's body was investigated on mounted, frozen European starling Sturnus vulgaris in a wind tunnel at flight speeds between 6 and 14 m s(-1). Removal of tail rectrices and dorsal and ventral covert feathers at the base of the tail increased the total parasite drag of the body and tail by between 25 and 55%. Flow visualization and measurements of dynamic pressure in the tail boundary layer showed that in the intact bird a separation bubble forms on the ventral side of the body, and reattaches to the ventral side of the tail. This bubble is a consequence of the morphology of the body, with a rapid contraction posterior to the pelvis and hind legs. The tail and the covert feathers at its base act as a combined splitter plate and wedge to control vortex shedding and body wake development, and thereby are important to minimize drag. This hitherto unsuspected mechanism is central to understanding the morphology of the avian body, and may have had a significant influence on the evolution of avian tail morphology by pre-adapting the tail for radiation and specialization as an aerodynamic lifting structure and as an organ of communication in sexual selection.     

Experimental analyses of sexual and natural selection on short tails in a polygynous warbler

We believe that no experimental study has yet tested Darwin's idea that, as well as generating trait elaboration, intersexual selection might sometimes drive sex-biased trait reduction. Here we present the results of two experiments exploring the negative relationship between tail length and reproductive success in male golden-headed cisticolas (Cisticola exilis). In the first experiment, artificially shortening a male's tail produced a dramatic increase in his reproductive success, measured as either the number of females nesting or number of chicks Hedged on his territory. A second experiment, in which manipulated birds were flown through a maze, revealed that short tails also impose costs by reducing aerodynamic performance during slow-speed foraging flight. Because tail shortening yields reproductive benefits and viability costs, we conclude it has evolved via sexual selection. Disentangling exactly how short tails enhance male reproductive success is more difficult. Male-male competition appears partly responsible: aerodynamic theory predicts that tail reduction enhances high-speed flight and, in line with this, shortened-tail males spent more time engaged in high-speed aerial chases of rivals and defended higher-quality territories. However, shortened-tail males had higher reproductive success independent of territory quality and spent more time in aerial displays which may be directed at females. This suggests that tail shortening is also favoured via female choice based on male phenotype.     

Flight speeds of swifts (Apus apus): seasonal differences smaller than expected

We have studied the nocturnal flight behaviour of the common swift (Apus apus L.), by the use of a tracking radar. Birds were tracked from Lund University in southern Sweden during spring migration, summer roosting flights and autumn migration. Flight speeds were compared with predictions from flight mechanical and optimal migration theories. During spring, flight speeds were predicted to be higher than during both summer and autumn due to time restriction. In such cases, birds fly at a flight speed that maximizes the overall speed of migration. For summer roosting flights, speeds were predicted to be lower than during both spring and autumn since the predicted flight speed is the minimum power speed that involves the lowest energy consumption per unit time. During autumn, we expected flight speeds to be higher than during summer but lower than during spring since the expected flight speed is the maximum range speed, which involves the lowest energy consumption per unit distance. Flight speeds during spring were indeed higher than during both summer and autumn, which indicates time-selected spring migration. Speeds during autumn migration were very similar to those recorded during summer roosting flights. The general result shows that swifts change their flight speed between different flight behaviours to a smaller extent than expected. Furthermore, the difference between flight speeds during migration and roosting among swifts was found to be less pronounced than previously recorded.     

Energy expenditures for flight,aerodynamic quality,and colonization of forest habitats by birds

The power that the birds can use for flight (available power) and the power required for flight according to physical laws (requisite power) grow with an increase in body mass, the exponents of the corresponding functions being different. Small birds can follow different strategies, either improving the aerodynamic quality of the body (thereby saving the excess available power) or sacrifice aerodynamic quality in favor of morphological adaptation to factors other than the demands of flight proper, which provides the possibility of utilizing a wider range of ecological niches. A hypothesis is proposed that the high metabolic rate of passerine birds, compared to representatives of other bird orders, is an adaptation to maneuverable (i.e., relatively low-speed) flight necessary for successful colonization of forest habitats. The speed that birds of such size can develop according to the scaling theory is too high for nesting and foraging in tree crowns, and its reduction is possible in two ways: by increasing air drag or by changing the style of flight (by analogy with airplane vs. helicopter). The first way is feasible, but a high air drag due to morphological modifications (e.g., in the size of the tail or characteristics of the wing) interferes with the possibility of long-distance migration flight, as energy expenditures for it will exceed the energy potential of the bird. This is why migratory nonpasserine birds, which have used this strategy, are practically absent in forests of the temperate zone. Therefore, more promising is the second way involving transition to a new flight style and, in a certain sense, to a new morphophysiological organization. Passerines have achieved this by changing their flight style so that the wing actively generates forces (lift and thrust) only in downstroke. Such a flight requires more energy, and, to provide it in sufficient amounts, passerine birds have increased their basal metabolic rate (BMR). Thus, both their flight energy expenditures and BMR are higher than in nonpasserines. Remarkably, among approximately 8660 extant bird species known today, more than half (about 5100 species) belong to the order Passeriformes. Such a ratio, unknown in any other vertebrate class, is evidence that passerines have gained a considerable biological advantage over all other birds due to their increased BMR.     

FROM FROND TO FAN: ARCHAEOPTERYX AND THE EVOLUTION OF SHORT-TAILED BIRDS

Modern birds have extremely short tail skeletons relative to Archaeopteryx and nonavialian theropod dinosaurs. Long- and short-tailed birds also differ in the conformation of main tail feathers making up the flight surface: frond shaped in Archaeopteryx and fan shaped in extant fliers. Mechanisms of tail fanning were evaluated by electromyography in freely flying pigeons and turkeys and by electrical stimulation of caudal muscles in anesthetized birds. Results from these experiments reveal that the pygostyle, rectrices, rectricial bulbs, and bulbi rectricium musculature form a specialized fanning mechanism. Contrary to previous models, our data support the interpretation that the bulbi rectricium independently controls tail fanning; other muscles are neither capable of nor necessary for significant rectricial abduction. This bulb mechanism permits rapid changes in tail span, thereby allowing the exploitation of a wide range of lift forces. Isolation of the bulbs on the pygostyle effectively decouples tail fanning from fan movement, which is governed by the remaining caudal muscles. The tail of Archaeopteryx, however, differs from this arrangement in several important respects. Archaeopteryx probably had a limited range of lift forces and tight coupling between vertebral and rectricial movement. This would have made the tail of this primitive flier better suited to stabilization than maneuverability. The capacity to significantly alter lift and manipulate the flight surface without distortion may have been two factors favoring tail shortening and pygostyle development during avian evolution.     

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.