Haematocrit correlates with tail ornament size in three populations of the Barn Swallow (Hirundo rustica)

1. Handicap models of sexual selection propose that male ornaments are indicators of male quality and that honesty is enforced by the costs imposed by the exaggerated ornamental traits. In long-distance migratory birds that feed on the wing, the aerodynamic cost of exaggerated ornamental characters should be particularly high because the size of the ornaments deviates from the natural selection optimum. During migration, birds are expected to raise their oxygen consumption in relation to the energetic demands imposed by their morphology. An increase of haematocrit is an adaptive response to enhance oxygen uptake and efficiency of transfer to the muscular tissues during spells of intense muscular activity.
2. The change of haematocrit of Barn Swallows ( Hirundo rustica ) after their arrival to the breeding sites, and the relationships between haematocrit values recorded after migration and the size of ordinary and sexually selected morphological characters in three Barn Swallow populations were analysed.
3. Males had higher haematocrit values than females. Individual haematocrit values declined after arrival to the breeding sites. Haematocrit values of males were significantly and positively correlated with the size of their ornamental tail but not correlated with other characters, thus suggesting that well-ornamented males, in order to arrive early, have to raise their haematocrit above the level of short-tailed males.
4. Males and females of similar tail length did not differ in their haematocrit, thus suggesting that sexual dimorphism in haematocrit might be functionally related to dimorphism in tail length.
5. Our results are consistent with the handicap principle because long-tailed males experience lower mortality and larger seasonal reproductive success compared with short-tailed males.     

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.     

Barn Swallow Hirundo rustica parents work harder when foraging conditions are good

In altricial birds, the great effort involved in supplying food to nestlings can create trade‐offs in the allocation of resources between the current brood and parental self‐maintenance. In poor foraging conditions, parents have to adjust their energy expenditure in relation to the increased foraging costs. However, intra‐specific variation in parental energy expenditure has rarely been evaluated in the context of these trade‐offs. Here, we quantified the daily energy expenditure (DEE) of parent Barn Swallows Hirundo rustica during the nestling period in relation to foraging conditions while controlling for differences in brood size and nestling age. DEE varied substantially with environmental conditions, increasing by 10 kJ/day per 5 °C in ambient temperature, and by 11 kJ/day per hour in day length. Parent birds did not compensate for a poor aerial insect supply on cool days, but reduced their DEE. Parents only slightly buffered a negative energy balance during chick provisioning with stored body reserves. They did not sacrifice their own energy demands to keep up a high energy flow to the brood when foraging conditions were poor. Instead they worked harder when foraging conditions allowed a surplus intake, fully compensating for their additional efforts, and made maximum use of the rich food supply, allowing the brood to accrue body reserves to compensate for low food intake on cold days. This strategy of energy management may have evolved in the context of the adaptation to the aerial foraging mode and to the ephemeral nature of aerial food resources.     

哈尔滨家燕mandschurica亚种繁殖生态特征

许多鸟种因具有较强的适应性而分布广泛,例如家燕(Hirundo rustica)。在不同纬度地区研究其繁殖生态学特征有助于了解家燕对不同环境的适应。家燕mandschurica亚种在国内仅分布于黑龙江省,且该亚种的繁殖资料少有报道,为此,于2016和2017年每年的4至10月,在哈尔滨对其繁殖生态特征开展研究,并与国内目前已发表的家燕gutturalis亚种繁殖生态特征进行对比。家燕mandschurica亚种4月底或5月初迁来哈尔滨,9月下旬或10月初南迁;窝卵数为4～6枚(n=19);卵长径(18.7±1.4)mm(15.9～22.5 mm)、卵短径(13.0±0.5)mm(12.2～14.2 mm),卵重(1.6±0.1)g(1.3～1.9 g),n=35;孵卵期为(16±2)d(14～18 d,n=19);育雏期为(17±1)d(16～18 d,n=6)。雏鸟的体长、翅长及尾羽长的生长曲线能与Logistic较好地拟合;体重、嘴峰和跗跖在5日龄左右生长最快,体长和翅长在7日龄左右增长最快,雏鸟的生长模式符合能量分配假说。与我国南方的家燕gutturalis亚种相比,在哈尔滨繁殖的mandschurica亚种的卵更小、更轻,二者孵卵期相似,但后者育雏期更短。这或许与不同地区食物丰富度及亲鸟喂食策略有关。     

Viability Is Associated with Melanin-Based Coloration in the Barn Swallow (Hirundo rustica)

Pigmentation of body surface in animals can have multiple determinants and accomplish diverse functions. Eumelanin and pheomelanin are the main animal pigments, being responsible of yellow, brownish-red and black hues, and have partly common biosynthetic pathways. Many populations of vertebrates show individual variation in melanism, putatively with large heritable component. Genes responsible for eu- or pheomelanogenesis have pleiotropic but contrasting effects on life-history traits, explaining the patterns of covariation observed between melanization and physiology (e.g. immunity and stress response), sexual behavior and other characters in diverse taxa. Yet, very few studies in the wild have investigated if eu- and pheomelanization predict major fitness traits like viability or fecundity. In this correlative study, by contrasting adult barn swallows (Hirundo rustica) matched for age, sex, breeding site, and year and date of sampling, we show that males but not females that survived until the next year had paler, relatively more eu- than pheomelanic pigmentation of ventral body feathers. Better performance of individuals that allocate relatively more to eumelanogenesis was expected based on previous evidence on covariation between eumelanic pigmentation and specific traits related to immunity and susceptibility to stress. However, together with the evidence of no covariation between viability and melanization among females, this finding raises the question of the mechanisms that maintain variation in genes for melanogenesis. We discuss the possibility that eu- and pheomelanization are under contrasting viability and sexual selection, as suggested by larger breeding and sperm competition success of darker males from other barn swallow subspecies.     

White tail spots in breeding Barn Swallows Hirundo rustica signal body condition during winter moult

The determinants and function of pigmentation of feathers and other tissues have been the focus of a large number of studies, particularly with respect to socio‐sexual communication. However, many birds exhibit depigmented white spots or bars on their feathers whose function is poorly understood. Here we assess whether white feather spots reflect phenotypic condition at the time of moult by investigating the covariation between spot size or shape and condition‐dependent feather growth rate, as gauged by width of the growth bars on the tail feathers of Barn Swallows. We found that feathers with higher growth rates had larger, less rounded white spots. In addition, variance in spot perimeter for a given spot area was larger in males than in females. This study is the first to provide evidence that features of white markings on feathers directly reflect body condition at the time of moult and can therefore reliably signal phenotypic quality in the context of socio‐sexual communication. In addition, the study highlights the potential communication function of the shape and not just the size of colour signals.     

A Trade-Off between Reproduction and Feather Growth in the Barn Swallow (Hirundo rustica)

Physiological trade-offs mediated by limiting energy, resources or time constrain the simultaneous expression of major functions and can lead to the evolution of temporal separation between demanding activities. In birds, plumage renewal is a demanding activity, which accomplishes fundamental functions, such as allowing thermal insulation, aerodynamics and socio-sexual signaling. Feather renewal is a very expensive and disabling process, and molt is often partitioned from breeding and migration. However, trade-offs between feather renewal and breeding have been only sparsely studied. In barn swallows (Hirundo rustica) breeding in Italy and undergoing molt during wintering in sub-Saharan Africa, we studied this trade-off by removing a tail feather from a large sample of individuals and analyzing growth bar width, reflecting feather growth rate, and length of the growing replacement feather in relation to the stage in the breeding cycle at removal and clutch size. Growth bar width of females and length of the growing replacement feather of both sexes were smaller when the original feather had been removed after clutch initiation. Importantly, in females both growth bar width and replacement feather length were negatively predicted by clutch size, and more strongly so for large clutches and when feather removal occurred immediately after clutch completion. Hence, we found strong, coherent evidence for a trade-off between reproduction, and laying effort in particular, and the ability to generate new feathers. These results support the hypothesis that the derived condition of molting during wintering in long-distance migrants is maintained by the costs of overlapping breeding and molt.     

Experimental manipulation of tail length in female barn swallows (Hirundo rustica) affects their future reproductive success

Previous studies have shown no significant effect of experimentaltail length manipulation in female barn swallows (Hirundo rustica)at the beginning of a breeding season on reproductive successor behavior during that breeding season. In the present study,we investigate if tail length manipulation had any effect onreproductive performance the following year, the so-called long-termeffect, in contrast to the short-term effects already studied.We found that females with experimentally elongated externaltail feathers at the beginning of a breeding season producedless offspring during the breeding season the following yearthan did females with shortened or unmanipulated tails. Theseresults suggest that tail elongation caused flight deficienciesthat deteriorated the condition of females and eventually reducedreproductive success. The finding of long-term effects but nosignificant short-term effects for female tail elongation suggeststhat female barn swallows have the ability to adjust immediateparental investment. Detrimental effects of long tails in femalesin terms of decreased reproductive success might explain whyfemale tails are not as long as those of males. Finally, femalesmated to long-tailed (sexually attractive) males decreased theirreproductive success the following year more than did femalesmated to short-tailed males, possibly owing to differentialparental effort causing a deterioration of their condition.     

Breeding habitat selection in the Swallow Hirundo rustica

This paper investigates the factors which influence nest site selection in Swallows. In years with peak numbers, more pairs settle on favoured farms rather than establish new 'colonies' at sub-optimal sites.     

家燕和金腰燕的卵胚胎心率比较

胚胎心率是衡量胚胎新陈代谢速率的重要指标。鸟类的胚胎心率随新陈代谢的增加而呈上升趋势。对早成性鸟类的种间比较发现,胚胎心率平均值随卵重量的增大而减小,卵体积小的种类具有相对较高的胚胎心率。国内有关野生鸟类胚胎心率的研究较少。2014年5~8月,在黑龙江扎龙国家级自然保护区,利用红外胚胎心率测量仪对两种近缘鸟类家燕(Hirundo rustica,n=14)和金腰燕(Cecropis daurica,n=14)的卵胚胎心率及其变化进行了测量与比较。两种燕均在孵卵的第2天开始出现胚胎心率,并随胚龄增加心率呈上升趋势,但在第8天及第11~14天家燕的胚胎心率显著低于金腰燕(第8天:z=﹣2.602,P=0.009;第11天:z=﹣2.497,P=0.013;第12天:z=﹣2.354,P=0.019;第13天:z=3.424,P=0.001;第14天:z=﹣3.380,P=0.001)。家燕卵胚胎日均增长心率(19.0±3.1)次/min,金腰燕卵胚胎日均增长心率(16.1±3.4)次/min,二者差异不显著(z=﹣1.792,P=0.073)。两种燕的胚胎心率与卵容量和卵重均不存在显著相关性[家燕:卵容量(1.73±0.09)cm3,r=0.192,P=0.511;卵重(1.74±0.09)g,r=0.128,P=0.663。金腰燕:卵容量(1.74±0.08)cm3,r=0.040,P=0.891;卵重(1.51±0.09)g,r=0.054,P=0.855]。这可能表明,卵大小和卵重量对家燕与金腰燕的胚胎心率均影响不明显。     

Early-Life Telomere Dynamics Differ between the Sexes and Predict Growth in the Barn Swallow (Hirundo rustica)

Telomeres are conserved DNA-protein structures at the termini of eukaryotic chromosomes which contribute to maintenance of genome integrity, and their shortening leads to cell senescence, with negative consequences for organismal functions. Because telomere erosion is influenced by extrinsic and endogenous factors, telomere dynamics may provide a mechanistic basis for evolutionary and physiological trade-offs. Yet, knowledge of fundamental aspects of telomere biology under natural selection regimes, including sex- and context-dependent variation in early-life, and the covariation between telomere dynamics and growth, is scant. In this study of barn swallows (Hirundo rustica) we investigated the sex-dependent telomere erosion during nestling period, and the covariation between relative telomere length and body and plumage growth. Finally, we tested whether any covariation between growth traits and relative telomere length depends on the social environment, as influenced by sibling sex ratio. Relative telomere length declined on average over the period of nestling maximal growth rate (between 7 and 16 days of age) and differently covaried with initial relative telomere length in either sex. The frequency distribution of changes in relative telomere length was bimodal, with most nestlings decreasing and some increasing relative telomere length, but none of the offspring traits predicted the a posteriori identified group to which individual nestlings belonged. Tail and wing length increased with relative telomere length, but more steeply in males than females, and this relationship held both at the within- and among-broods levels. Moreover, the increase in plumage phenotypic values was steeper when the sex ratio of an individual’s siblings was female-biased. Our study provides evidence for telomere shortening during early life according to subtly different dynamics in either sex. Furthermore, it shows that the positive covariation between growth and relative telomere length depends on sex as well as social environment, in terms of sibling sex ratio.     

Contrasting sensitivity of nestling and fledgling Barn Swallow Hirundo rustica body mass to local weather conditions

Local weather can influence the growth and development of young birds either indirectly, by modifying prey availability, or directly, by affecting energetic trade-offs. Such effects can have lasting implications for life history traits, but the nature of these effets may vary with the developmental stage of the birds, and over timescales from days to weeks. We examined the interactive effects of temperature, rainfall and wind speed on the mass of nestling and fledgling Barn Swallows Hirundo rustica both on the day of capture and averaging weather across the time since hatching. At the daily timescale, nestling mass was negatively correlated with temperature, but the strength of this association depended on the level of rainfall and wind speed; nestlings were typically heavier on dry or windy days, and the negative effect of temperature was strongest under calm or wet conditions. At the early lifetime timescale (i.e. from hatching to pre-fledging), nestling mass was negatively correlated with temperature at low wind speed. Fledgling body mass was less sensitive to weather; the only weather effect evident was a negative correlation with temperature at the daily scale under high rainfall that became slightly positive under low rainfall. These changes are consistent with weather effects on the availability and distribution of insects within the landscape (e.g. causing high concentrations of flying insects) and with the effects of weather variation on nest microclimate. These results together demonstrate the impacts of weather on chick growth, over immediate (daily) and longer term (nestling/fledgling lifetime) timescales. This shows that sensitivity to local weather conditions varies across the early lifetime of young birds (nestling–fledgling stages) and illustrates the mechanisms by which larger scale (climate) variations influence the body condition of individuals.     

Sexual Dimorphism in Melanin Pigmentation,Feather Coloration and Its Heritability in the Barn Swallow (Hirundo rustica)

Melanin is the main pigment in animal coloration and considerable variation in the concentrations of the two melanin forms (pheo- and eumlanin) in pigmented tissues exists among populations and individuals. Melanin-based coloration is receiving increasing attention particularly in socio-sexual communication contexts because the melanocortin system has been hypothesized to provide a mechanistic basis for covariation between coloration and fitness traits. However, with few notable exceptions, little detailed information is available on inter-individual and inter-population variation in melanin pigmentation and on its environmental, genetic and ontogenetic components. Here, we investigate melanin-based coloration in an Italian population of a passerine bird, the barn swallow (Hirundo rustica rustica), its sex- and age-related variation, and heritability. The concentrations of eu- and pheomelanin in the throat (brown) and belly (white-to-brownish) feathers differed between sexes but not according to age. The relative concentration of either melanin (Pheo:Eu) differed between sexes in throat but not in belly feathers, and the concentrations in males compared to females were larger in belly than in throat feathers. There were weak correlations between the concentrations of melanins within as well as among plumage regions. Coloration of belly feathers was predicted by the concentration of both melanins whereas coloration of throat feathers was only predicted by pheomelanin in females. In addition, Pheo:Eu predicted coloration of throat feathers in females and that of belly feathers in males. Finally, we found high heritability of color of throat feathers. Melanization was found to differ from that recorded in Hirundo rustica rustica from Scotland or from H. r. erythrogaster from North America. Hence, present results show that pigmentation strategies vary in a complex manner according to sex and plumage region, and also among geographical populations, potentially reflecting adaptation to different natural and sexual selection regimes, and that some coloration components seem to be highly heritable.     

Sexual Dimorphism and Population Differences in Structural Properties of Barn Swallow (Hirundo rustica) Wing and Tail Feathers

Sexual selection and aerodynamic forces affecting structural properties of the flight feathers of birds are poorly understood. Here, we compared the structural features of the innermost primary wing feather (P1) and the sexually dimorphic outermost (Ta6) and monomorphic second outermost (Ta5) tail feathers of barn swallows (Hirundo rustica) from a Romanian population to investigate how sexual selection and resistance to aerodynamic forces affect structural differences among these feathers. Furthermore, we compared structural properties of Ta6 of barn swallows from six European populations. Finally, we determined the relationship between feather growth bars width (GBW) and the structural properties of tail feathers. The structure of P1 indicates strong resistance against aerodynamic forces, while the narrow rachis, low vane density and low bending stiffness of tail feathers suggest reduced resistance against airflow. The highly elongated Ta6 is characterized by structural modifications such as large rachis width and increased barbule density in relation to the less elongated Ta5, which can be explained by increased length and/or high aerodynamic forces acting at the leading tail edge. However, these changes in Ta6 structure do not allow for full compensation of elongation, as reflected by the reduced bending stiffness of Ta6. Ta6 elongation in males resulted in feathers with reduced resistance, as shown by the low barb density and reduced bending stiffness compared to females. The inconsistency in sexual dimorphism and in change in quality traits of Ta6 among six European populations shows that multiple factors may contribute to shaping population differences. In general, the difference in quality traits between tail feathers cannot be explained by the GBW of feathers. Our results show that the material and structural properties of wing and tail feathers of barn swallows change as a result of aerodynamic forces and sexual selection, although the result of these changes can be contrasting.     

Partitioning the effects of livestock farming on the diet of an aerial insectivorous passerine,the Barn Swallow Hirundo rustica

Capsule The presence of farm animals increases the amount of larger insect prey (mainly large Diptera and Coleoptera) and decreases the proportion of smaller prey (Hymenoptera).

Aims To determine the relationships between presence of farm animals and diversity of diet of nestling Barn Swallows from 52 colonies located at sites with and without livestock.

Methods Diet was determined on the basis of faecal analysis. We used two approaches to quantify livestock based on their spatial distribution – a breeding site-focused approach and a village-focused approach.

Results Our results show a relationship between the presence of livestock and the size, number and composition of insect prey detected in the diet of nestling Barn Swallows. The results from the two approaches we used in our study were broadly consistent, although effects varied according to the number and types of animal being raised. The presence of domestic animals increases the amount of larger prey (mainly larger Diptera and Coleoptera) and decreases the proportion of smaller prey (Hymenoptera species). The effect of individual livestock species, mainly pigs and poultry alone, was considerably higher than the effects of pooling livestock species from Principal Component Analysis.

Conclusion Our results imply that Barn Swallows were foraging beyond the immediate area of the farm buildings in which they nest, suggest considerable adaptability in the species' diet and further emphasize the ecological role that livestock play in providing a source of prey of a required size rather specific identity. Furthermore, a statistically significant positive effect of average mass of prey (which is a direct consequence of the presence of cattle at a breeding site) on colony size suggests that livestock farming provides a more profitable prey community, which may ultimately increase local populations of Barn Swallow.     

大杜鹃对家燕的巢寄生

了解杜鹃(Cuculus spp.)对不同宿主鸟类的巢寄生,是研究杜鹃与其宿主之间协同进化的重要基础资料。大杜鹃(Cuculus canorus)和家燕(Hirundo rustica)分布遍及全国,且为同域分布,但两者之间的寄生现象尚未有过系统调查。2012年和2014年4~8月,对繁殖于吉林市昌邑区桦皮厂镇(34°58′44.18″N,126°13′26.83″E,海拔184 m)和海南岛的家燕种群进行调查,结果表明,吉林市昌邑区桦皮厂镇家燕种群的寄生率为2.4%(1/42),而在海南岛所调查的1 719个家燕巢未发现杜鹃寄生现象。同时在网络上搜集家燕巢寄生的报道案例,共记录到13巢家燕被大杜鹃寄生繁殖,均发生在北方的家燕种群。     

Sexual selection in the monogamous barn swallow (Hirundo rustica). II. Mechanisms of sexual selection

Mechanisms of sexual selection in the monogamous, sexually dimorphic barn swallow (Hirundo rustica) were studied during a seven-year period. First, the sex ratio of reproducing adults was male-biased, and mated males had significantly longer tail ornaments than unmated males. Secondly, some of the unmated individuals later committed infanticide and became mated with the mother of the killed brood. Fathers of killed broods had significantly shorter tails than other males, and there was a tendency for infanticidal males to have longer tail ornaments than other unmated males. Thirdly, long-tailed male barn swallows were more successful in acquiring extra-pair copulations than other males, and females involved in extra-pair copulations, as compared to females not involved in such copulations, had mates with shorter tail ornaments. Fourthly, male barn swallows having long tails as compared to short-tailed males acquired mates in better body condition. Females mated to long-tailed males reproduced earlier, laid more eggs and were more likely to have two clutches than were females mated to short-tailed males. Finally, females mated to long-tailed males put more effort into reproduction than did other females, as evidenced by their relatively larger contribution to feeding of offspring. Thus, at least five different components of sexual selection affected male reproductive success. Selection arising from differential success during extra-pair copulations, differential reproductive success and differential male reproductive effort thus accounted for most of the selection on tail ornaments in male barn swallows.