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.     

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.     

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.     

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.     

Sexual dimorphism in wing and tail length as shown by the Swallow Hirundo rustica

In both sexes of Hirundo rustica two metric traits are significantly different. One of these traits, tail length, shows a positive correlation between mates and a negative correlation with the date of first egg laying. On the other hand, wing lengths are not significantly related in any of the corresponding tests. Similar relations were recorded both for all analysed birds and for at least two-year-old individuals. It is suggested that sexual selection, as proposed by Darwin (1871) for monogamous birds, is responsible for sexual dimorphism in the tail length of the Swallow. It seems that intersexual difference in wing length could be a result of such mechanisms as stabilizing the body sizes of males and females at different energetically optimum levels by natural selection.     

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.     

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.     

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.     

Effects of an Immune Challenge on Multiple Components of Song Display in Barn Swallows Hirundo rustica: Implications for Sexual Selection

Bird song is believed to honestly reveal male quality including the ability of singers to face parasitism. In a natural population of barn swallows Hirundo rustica, we experimentally imposed a cost on song production by an immune challenge. We therefore vaccinated a group of reproducing males with an antigen (Newcastle disease virus), and injected phosphate‐buffered saline to a control group. Immune challenge significantly reduced one song feature, rattle duration. This decrease was related to male quality, as measured by tail length, because males with short tails reduced the duration of their rattle significantly more than males with long tails. In addition, another song feature, strophe duration, decreased in the control group, while it remained constant in the challenged group. Duration of the rattle has previously been found to be positively related to testosterone level, and it may hence reflect male competitive ability. Thus, male barn swallows may not have the potential to produce long rattles when their immune system is challenged. By maintaining their strophe duration after an immune challenge, males may compensate for the decrease in rattle duration. Our results suggest that different song features may convey different types of information, and that barn swallow song, in particular their rattle, may reliably reveal information about activation of the immune system.     

Parental care trade‐offs in the inter‐brood phase in Barn Swallows Hirundo rustica

In seasonal environments with limited time and energy resources, double‐brooded birds face trade‐offs in the timing of their two reproductive attempts and in the effort allocated to the first and the second broods. In the Barn Swallow Hirundo rustica a long care period for the first brood enhances the survival of first‐brood chicks, but also delays the start of the second brood, which in turn reduces the survival prospects of second‐brood chicks. Probably as a response to this trade‐off, double‐brooded Barn Swallows reduce the period of post‐fledging care for first‐brood fledglings. By radiotracking whole families, we investigated the determinants of this behaviour and its consequences for the survival of the first‐brood fledglings. The end of the females’ investment in post‐fledging care of the first brood was related to the beginning of egg synthesis for the second clutch. With the start of egg synthesis, females significantly reduced provisioning rates to the first‐brood fledglings to less than one‐fifth of the previous rates, while the proportion of time they spent foraging remained high. Assuming that the females’ foraging success was constant, we conclude that their energy income was allocated to egg production rather than fledgling provision. Males did not compensate for the females’ reduced feeding rates. Thus the start of egg production for the second clutch had a marked effect on the quantity of food received by first‐brood fledglings. In parallel with the changes in parental behaviour and provisioning rates, we observed a marked drop in the daily survival rate of first‐brood chicks. These results support the hypothesis that females face a strong trade‐off in the allocation of energy to subsequent broods. Energy allocation to a second clutch involves a cost in terms of reduced provisioning, and as a result the survival of first‐brood chicks is compromised. This is probably outweighed by the improved success of an early second brood.     

Parental Care and Sexual Selection in the Bluethroat, Luscinia s. svecica: A Field-experimental Test of the Differential Allocation Hypothesis

One of the processes by which secondary sexual characters in monogamous species with biparental care may evolve is described by the differential allocation hypothesis. It predicts that mates adjust their parental effort to the attractiveness of their partner. The more attractive sex is expected to withhold parental effort while the less attractive sex is expected to compensate for the partner's decreased effort thus preventing a reduction in breeding success. Asymmetry in attractiveness then imposes costs on the less attractive sex which are thought to be exceeded by the indirect (i.e. genetic) benefits which result from sharing offspring with a highly attractive partner. This idea was tested with bluethroats, Luscinia s. svecica , a bird species in which the male is characterized by a complex multiple-coloured plumage ornament on the throat and chest. We manipulated male attractiveness by attaching conspicuous coloured leg bands which matched the blue and chestnut components of the male ornament. Control males were banded with green and yellow bands. The results from another field experiment on mate-guarding behaviour demonstrate the effectiveness of this treatment. As measures of parental effort we determined: 1. clutch size; 2. feeding rates by video recording feeding activity of both parents; and 3. brood mass up to day 9. The results do not support the differential allocation hypothesis. The experimental and control groups differed in none of the above variables. On the basis of these results we discuss the ecological factors which may influence the differential allocation process in monogamous birds.     

The breeding biology of the Pacific Swallow Hirundo tahitica in Malaysia

The breeding biology of the Pacific Swallow was studied in Malaysia. Time-budgets and preliminary energy-budgets were calculated for different stages of the breeding cycle. Breeding was seasonal, the first eggs were laid in the first week of March from 1978 to 1982. The nest was a mud cup built under an overhang, often on man-made structures over water. The mean clutch size was 2.98 eggs. So more than two successful clutches were ever raised, although up to four sets of eggs can be laid if clutches are lost. The incubation period was 16.2 days and the nestling period 19–21 days. Nestling weights showed the typical hirundine recession before fledging. The mean brood size was 2.32 and there was no evidence of undernourishment in larger broods. Nesting success was lower than for temperate hirundines but high for a tropical passerine. Incubation duties were carried out by the female alone but both parents fed the brood. The male never delivered more than 44% of the total feeds. Daily energy expenditure varied from 3.44–4.9×BMR depending upon the stage of the breeding cycle.     

Facultative prioritization of wing growth in the Welcome Swallow Hirundo neoxena

Bird species adapted to variable environments tend to have slow lean tissue growth rates and high fat deposition rates, allowing survival during food shortages. This emphasis on fat deposition may be a fixed physiological trait. Alternatively, tissue allocation may be adjusted facultatively according to the proximate food supply. We consider two models of facultative adjustment that could account for the emphasis on fat deposition: (1) the fat-priority model, in which no lean growth occurs when food is scarce, and (2) the lean-priority model, in which a minimal level of lean growth always occurs but nutrients are otherwise allocated to fat deposition. We tested these two models using Welcome Swallows Hirundo neoxena , a species we show to have a variable food supply that is influenced by weather. We reduced food supply to chicks experimentally, by enlarging broods or excluding parents from chicks, and tested for reduction in wing growth (an indicator of lean growth) and mass growth (an indicator of fat deposition). Mass growth was retarded by both manipulations, but not wing growth, corroborating the lean-priority model. This growth strategy may function not to cope with violent variation in food supply, but to maintain development and symmetry of wings and feathers in the face of moderate variation in food supply. Our results contrast with those of a similar experiment on the Black Noddy Anous minutus , a species with more severe variation in food supply.     

Water repellency and feather structure of the Blue Swallow Hirundo atrocaerulea

Rijke, A.M., Jesser, W.A., Evans, S.W & Bouwman, H. 2000. Water repellency and feather structure of the Blue Swallow Hirundo atrocaerulea. Ostrich 71 (1 & 2): 143–145.

The Blue Swallow is an endangered species in southern Africa and is probably the most endangered passerine. It is restricted to escarpments with grasslands above 1 000 m where mists are frequent. It appears to forage on the wing even in thick mist raising the question of feather wettability in relation to its adaptation. Extensive physical and behaviourial adaptations are known to occur in a wide variety of birds to deal with the problem of shedding water continuously. To study the water repellency and resistance to water penetration of Blue Swallow feathers, we have examined the microscopic structure of head, back, throat, breast and abdominal feathers as well as remiges and tail feathers by transmission light microscopy. The width (2R) and separation (2D) of rami and barbules have been measured and were used to calculate the parameter (R + D)/R that serves as an indicator ofwater shedding potential. For the remiges and tail feathers the values of the (R+D)/R range from 5 to 10 which is comparable to values for other terrestrial birds. However, for body feathers the range is from 10 (head) and 35 (abdomen)-higher than previously observed for any other bird including Swifts, Apodidae. Blue Swallow feathers are thus the most effective feather yet discovered at repelling water drops. The water repellency is highest in those feathers that are relatively shielded From the direct impact of small water drops (throat, breast, abdomen, back). By contrast, the flight feathers must possess a relatively large resistance to water penetration to avoid becoming waterlogged and this is coupled to low (R+D)/R values. Values for the barbules lay between 2 and 6—the same as found for other bird families—supporting an earlier conclusion that they have little direct effect in repelling water.