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.     

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。     

Vane emargination of outer tail feathers improves flight manoeuvrability in streamerless hirundines, Hirundinidae

Recent studies have suggested that the proximal part of the swallow (Hirundo rustica) tail streamer appears to aid turning flight, as expected if streamers evolved initially purely through natural selection for enhanced manoeuvrability. However, the evolution of slender aerodynamically advantageous streamers is also predicted by an alternative hypothesis, which suggests that such a trait could develop primarily to ameliorate the aerodynamic cost of a long size-dimorphic tail. To distinguish between these hypotheses, we have investigated for the effect on manoeuvrability of trimming the tips of the outer tail feathers into short streamers, without lengthening these feathers, in two streamerless hirundine species--the house martin (Delichon urbica) and the sand martin (Riparia riparia). This allowed us to examine the aerodynamic costs and benefits of streamers at an early evolutionary stage that predates elongation of the outermost tail feathers through female choice. We showed that such initial streamers enhance manoeuvrability in streamerless hirundines, confirming the findings of recent studies. However, in contrast to these studies, we showed that improved manoeuvrability resulting from streamers could arise before the outermost tail feathers have become elongated (e.g. owing to female choice). The occurrence of such an aerodynamic advantage depends on the ancestral shape of a forked tail. This provides support for the hypothesis that streamers, like those in the barn swallow, might evolve initially purely through natural selection for enhanced manoeuvrability.     

Size scaling and stiffness of avian primary feathers: implications for the flight of Mesozoic birds

The primary feathers of birds are subject to cyclical forces in flight causing their shafts (rachises) to bend. The amount the feathers deflect during flight is dependent upon the flexural stiffness of the rachises. By quantifying scaling relationships between body mass and feather linear dimensions in a large data set of living birds, we show that both feather length and feather diameter scale much closer to predictions for geometric similarity than they do to elastic similarity. Scaling allometry also indicates that the primary feathers of larger birds are relatively shorter and their rachises relatively narrower, compared to those of smaller birds. Two-point bending tests indicated that larger birds have more flexible feathers than smaller species. Discriminant functional analyses (DFA) showed that body mass, primary feather length and rachis diameter can be used to differentiate between different magnitudes of feather bending stiffness, with primary feather length explaining 63% of variance in rachis stiffness. Adding fossil measurement data to our DFA showed that Archaeopteryx and Confuciusornis do not overlap with extant birds. This strongly suggests that the bending stiffness of their primary feathers was different to extant birds and provides further evidence for distinctive flight styles and likely limited flight ability in Archaeopteryx and Confuciusornis.     

A comparative study of the chromosomes of birds

Karyotype analysis and morphometric measurement of the chromosomes of eleven species of Indian birds are described. The unequivocal identification of W chromosome in the females of five species of the present investigation further strengthens the generalisation that, at least in Carinatae, the sex chromosome constitutions are of ZZ and ZW types in males and females respectively. — The chromosomes of different species of birds so far worked out in each order have been compared using quantitative methods and tentative conclusions have been drawn regarding chromosomal affinities between species of different taxonomic categories.     

Sex-limited expression of ornamental feathers in birds

Extravagant secondary sexual characters show sexual size dimorphismin some species but are completely sex limited in others. Sexualornamentation has been hypothesized to benefit mainly malesthrough sexual selection, but the costs of secondary sexualcharacters initially would be experienced by both sexes. Theevolution of sexual size dimorphism of ornaments and, eventually, the complete sex-limited expression of these characters, willdepend on the effects of sexual and natural selection on thetwo sexes. A phylogenetic analysis controlling for similaritiesdue to common ancestry of 60 independent evolutionary originsof feather ornamentation in birds was used to investigate ecologicalfactors correlated with sexual size dimorphism and sex-limited expression of secondary sexual characters. When the size ofan ornament is large relative to body size, the trait willbe particularly costly for females, resulting in selectionfor increased sexual size dimorphism of the ornament. Indeed,sexual size dimorphism of ornaments was positively related to the relative size of male ornaments but was unrelated torelative size of female ornaments. Species with polygynousand lekking mating systems with little or no male parentalcare (in particular nest building and incubation) demonstratedsex-limited expression of ornaments as compared to monogamous species. Species with no food provisioning of offspring by themale showed a trend for increased sexual size dimorphism ofornaments. Therefore, large natural selection costs duringreproduction imposed by the expression of secondary sexualcharacters are related to the evolution of sexual size dimorphismof ornaments and eventually their complete loss from females.     

Incubation period and immune function: a comparative field study among coexisting birds

Developmental periods are integral components of life history strategies that can have important fitness consequences and vary enormously among organisms. However, the selection pressures and mechanisms causing variation in length of developmental periods are poorly understood. Particularly puzzling are prolonged developmental periods, because their selective advantage is unclear. Here we tested the hypotheses that immune function is stronger in species that are attacked at a higher rate by parasites and that prolonged embryonic development allows the development of this stronger immune system. Through a comparative field study among 12 coexisting passerine bird species, we show that species with higher blood parasite prevalence mounted stronger cellular immune responses than species with lower prevalence. These results provide support for the hypothesis that species facing greater selection pressure from parasites invest more in immune function. However, species with longer incubation periods mounted weaker cellular immune responses than species with shorter periods. Therefore, cellular immune responses do not support the hypothesis that longer development time enhances immunocompentence. Future studies should assess other components of the immune system and test alternative causes of variation in incubation periods among bird species.     

Ancient origins and multiple appearances of carotenoid-pigmented feathers in birds

The broad palette of feather colours displayed by birds serves diverse biological functions, including communication and camouflage. Fossil feathers provide evidence that some avian colours, like black and brown melanins, have existed for at least 160 million years (Myr), but no traces of bright carotenoid pigments in ancient feathers have been reported. Insight into the evolutionary history of plumage carotenoids may instead be gained from living species. We visually surveyed modern birds for carotenoid-consistent plumage colours (present in 2956 of 9993 species). We then used high-performance liquid chromatography and Raman spectroscopy to chemically assess the family-level distribution of plumage carotenoids, confirming their presence in 95 of 236 extant bird families (only 36 family-level occurrences had been confirmed previously). Using our data for all modern birds, we modelled the evolutionary history of carotenoid-consistent plumage colours on recent supertrees. Results support multiple independent origins of carotenoid plumage pigmentation in 13 orders, including six orders without previous reports of plumage carotenoids. Based on time calibrations from the supertree, the number of avian families displaying plumage carotenoids increased throughout the Cenozoic, and most plumage carotenoid originations occurred after the Miocene Epoch (23 Myr). The earliest origination of plumage carotenoids was reconstructed within Passeriformes, during the Palaeocene Epoch (66–56 Myr), and not at the base of crown-lineage birds.     

A comparative analysis of vigilance in birds

Individual variation in vigilance is known to vary with factors such as group size but the ecological determinants of vigilance among species have not been examined thus far in a systematic fashion. Earlier analyses suggested that vigilance should be lower in larger species and in species living in larger groups. These analyses were based on a small number of species and failed to take into account phylogenetic relationships among species. Here, I examined ecological determinants of vigilance in a large sample of bird species using a phylogenetic framework. I focused on vigilance in foraging groups of birds in the non-breeding season. Among species, vigilance by solitary foragers was not influenced by body mass. However, among species, asymptotic vigilance, the plateau reached by vigilance in larger groups, decreased with increasing group size in vegetarian clades but not in carnivorous clades. Asymptotic vigilance also increased with increasing body mass in vegetarian clades but not in carnivorous clades. Increasing group size may allow species to reduce vigilance in response to decreased predation risk. Increasing body mass may allow species to increase vigilance because more non-foraging time is available in larger species. Diet may modulate the effect of body mass and group size through factors such as within-group vigilance or foraging techniques.     

Comparing aerodynamic efficiency in birds and bats suggests better flight performance in birds

Flight is one of the energetically most costly activities in the animal kingdom, suggesting that natural selection should work to optimize flight performance. The similar size and flight speed of birds and bats may therefore suggest convergent aerodynamic performance; alternatively, flight performance could be restricted by phylogenetic constraints. We test which of these scenarios fit to two measures of aerodynamic flight efficiency in two passerine bird species and two New World leaf-nosed bat species. Using time-resolved particle image velocimetry measurements of the wake of the animals flying in a wind tunnel, we derived the span efficiency, a metric for the efficiency of generating lift, and the lift-to-drag ratio, a metric for mechanical energetic flight efficiency. We show that the birds significantly outperform the bats in both metrics, which we ascribe to variation in aerodynamic function of body and wing upstroke: Bird bodies generated relatively more lift than bat bodies, resulting in a more uniform spanwise lift distribution and higher span efficiency. A likely explanation would be that the bat ears and nose leaf, associated with echolocation, disturb the flow over the body. During the upstroke, the birds retract their wings to make them aerodynamically inactive, while the membranous bat wings generate thrust and negative lift. Despite the differences in performance, the wake morphology of both birds and bats resemble the optimal wake for their respective lift-to-drag ratio regimes. This suggests that evolution has optimized performance relative to the respective conditions of birds and bats, but that maximum performance is possibly limited by phylogenetic constraints. Although ecological differences between birds and bats are subjected to many conspiring variables, the different aerodynamic flight efficiency for the bird and bat species studied here may help explain why birds typically fly faster, migrate more frequently and migrate longer distances than bats.     

The oribates (Oribatei) in the feathers of birds

The authors confirm the fact of disperiosn of oribate mites by birds, that was earlier noticed in acarological references. More than 400 individuals of 53 virds species belonging to different ecological groups have been examined. About 50 species alive oribates were collected in bird feathers: Liochthonius sellnicki, Brachychthonius sp., Hypochthonius rufulus, Nothrus palustris, Malaconothrus egregius, Camisia segnis, Camisia sp., Hermanniella granulata, Trhypochthonius tectorum, Oribatula tibialis, Zigoribatula exilis, Scheloribates laevigatus, Sch. latipes, Suctobelba trigona, Suctobelbella sp., Fosseremaeus laciniatus, Tectocepheus velatus, T. knullei, Nanhermannia coronata, Achpteria coleoptrata, Parachipteria punctata, Damaeus riparius, Eremaeus oblongus, Diapterobates notatus, D. humeralis, Carabodes areolatus, C. marginatus, Sphaerozetes tricuspidatus, Ceratozetes parvulus, C. cisalpinus, Mycobates sp., Punctoribates punctum, Trichoribates trimaculatus, Galumna sp., Oribella paoli, Chamobates laciniatus, Neoribates roubali, Neoliodes farinosus, Oppiella nova, O. unicarinata, O. fallax, Oppia ornata, Steganacarus striculum, Steganacarus applicatus, Tropacarus carinatus, Protoribates capucinus, Scutovertex minutus, Autognetha willmanni, A. longilamellata, Belba sp., Metabelba pulverulenta, Gustavia microcephala, Fuscozetes fuscipes [symbol: see text] Pergalumna nervosa. Carabodes marginatus were met most often, on 8 species of birds, Tectocepheus velatus, on 12 species, Oppiella unicarinata--at 7 species. Some oribate species constantly occur in bird feathers.     

Air-permeable hole-pattern and nose-droop control improve aerodynamic performance of primary feathers

Primary feathers of soaring land birds have evolved into highly specialized flight feathers characterized by morphological improvements affecting aerodynamic performance. The foremost feathers in the cascade have to bear high lift-loading with a strong bending during soaring flight. A challenge to the study of feather aerodynamics is to understand how the observed low drag and high lift values in the Reynolds (Re) regime from 1.0 to 2.0E4 can be achieved. Computed micro-tomography images show that the feather responds to high lift-loading with an increasing nose-droop and profile-camber. Wind-tunnel tests conducted with the foremost primary feather of a White Stork (Ciconia ciconia) at Re = 1.8E4 indicated a surprisingly high maximum lift coefficient of 1.5 and a glide ratio of nearly 10. We present evidence that this is due to morphologic characteristics formed by the cristae dorsales as well as air-permeable arrays along the rhachis. Measurements of lift and drag forces with open and closed pores confirmed the efficiency of this mechanism. Porous structures facilitate a blow out, comparable to technical blow-hole turbulators for sailplanes and low speed turbine-blades. From our findings, we conclude that the mechanism has evolved in order to affect the boundary layer and to reduce aerodynamic drag of the feather.     

Frequency of fault bars in feathers of birds and susceptibility to predation

Fault bars are transparent bands in the feathers of birds produced under stressful and adverse conditions. The frequency of feathers with fault bars is highly heterogeneous among species. We predicted that prey had a higher frequency of fault bars than individuals from the general population, and that a high susceptibility to predation would be associated with a low frequency of fault bars among species of birds because such species would suffer particularly high costs of producing fault bars. The frequency of fault bars in prey was almost three-fold higher than in the general population, based on a database on the frequency of fault bars and susceptibility to predation by the goshawk Accipiter gentilis L., implying intense natural selection against fault bars. A high susceptibility to predation by the sparrowhawk Accipiter nisus L. and the goshawk, relative to what would be expected from their abundance, was associated with a low frequency of fault bars across species, with long distance migration also being negatively associated with frequency of fault bars. Feathers with fault bars were more likely to break than feathers without fault bars, thereby potentially affecting the flight ability of individuals. These findings are consistent with the hypothesis that susceptibility to factors that cause production of fault bars can be modified by natural selection, as illustrated by the impact of predation on the frequency of fault bars.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 334–345.     

Amplification success of multilocus genotypes from feathers found in the field compared with feathers obtained from shot birds

Effective DNA extraction methods from bird feathers have facilitated non‐invasive sampling, leading to the suggestion that feathers are a great source for genetic studies. However, few studies have assessed whether all feathers can be used or provide equal numbers of useful templates. In this study, feathers collected in various ways from Red Grouse Lagopus lagopus were examined to establish the quality of DNA extracted. Individual samples were classified into two categories according to whether they were collected from shot birds or found in the field. DNA was extracted from all samples and genotyped at 19 microsatellite loci. PCR products were analysed on a MegaBACE 1000. A total of 93% of the ‘shot’ category produced a genotype that was considered successful (i.e. 15 of 18 loci) and 23% of the ‘collected’ category produced successful genotypes under the same criteria. There was a significant difference between shot and collected samples in genotyping success and the observed number of missing loci. Recommendations and best practices are discussed along with the utility of bird feathers as a source of DNA for population and conservation biology.     

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.