The Mitogenome Relationships and Phylogeography of Barn Swallows (Hirundo rustica)

The barn swallow (Hirundo rustica) poses a number of fascinating scientific questions, including the taxonomic status of postulated subspecies. Here, we obtained and assessed the sequence variation of 411 complete mitogenomes, mainly from the European H. r. rustica, but other subspecies as well. In almost every case, we observed subspecies-specific haplogroups, which we employed together with estimated radiation times to postulate a model for the geographical and temporal worldwide spread of the species. The female barn swallow carrying the Hirundo rustica ancestral mitogenome left Africa (or its vicinity) around 280 thousand years ago (kya), and her descendants expanded first into Eurasia and then, at least 51 kya, into the Americas, from where a relatively recent (<20 kya) back migration to Asia took place. The exception to the haplogroup subspecies specificity is represented by the sedentary Levantine H. r. transitiva that extensively shares haplogroup A with the migratory European H. r. rustica and, to a lesser extent, haplogroup B with the Egyptian H. r. savignii. Our data indicate that rustica and transitiva most likely derive from a sedentary Levantine population source that split at the end of the Younger Dryas (YD) (11.7 kya). Since then, however, transitiva received genetic inputs from and admixed with both the closely related rustica and the adjacent savignii. Demographic analyses confirm this species’ strong link with climate fluctuations and human activities making it an excellent indicator for monitoring and assessing the impact of current global changes on wildlife.     

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。     

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。     

High brood patch temperature of less colourful,less pheomelanic female Barn Swallows Hirundo rustica

In birds, even a minor difference in egg temperature (1–1.5 °C) has been shown to affect the fitness of offspring by changing hatching success, incubation period and nestling quality. Female, but not male, passerines develop brood patches. Thus if there are traits, such as plumage ornamentation, that indicate optimal egg temperature, males should pair with females that exhibit those traits. However, no study has yet investigated the relationship between female brood patch temperature, which would directly affect egg temperature, and female plumage ornamentation. In this study, we examined the surface temperature of female brood patches during nocturnal incubation and examined its relationship with female plumage ornaments in Asian Barn Swallows Hirundo rustica gutturalis. After controlling for ambient air temperature, brood patch temperature was negatively associated with colour saturation of the female throat patch. No other female ornaments, such as tail‐length, white tail spots or throat patch size, predicted brood patch temperature. When oral (mouth) temperature was statistically controlled, females with less colourful throats and longer tails showed higher brood patch temperature, indicating that these females had hotter brood patches in relation to the temperature of other body parts. Furthermore, we found a negative relationship between pheomelanin pigmentation and brood patch temperature after controlling for ambient air temperature or oral temperature. To our knowledge, this is the first study to show that female ornaments can predict the absolute/relative thermal investment in brood patches. This relationship, together with other aspects of female quality, may affect male mate preference and female ornamentation.     

Segregation in the African wintering ranges of English and Swiss Swallow Hirundo rustica populations: a stable isotope study

Capsule Stable isotope analysis of Swallow feathers, grown in Africa, revealed significant differences between populations breeding in Switzerland and England.

Aims To investigate the extent to which Swallow populations breeding in Switzerland and England are separated on their African wintering grounds.

Methods Swallows were caught at breeding colonies, biometric measurements were taken and feathers, grown in Africa, were collected. Feathers were combusted in a Carlo Erba C/N/S analyser and the δ¹³C and δ¹⁵N signatures were measured using a mass spectrometer.

Results The δ¹³C signatures of Swiss birds were significantly more depleted than those of birds from England. The δ¹⁵N signatures did not differ between the two populations.

Conclusion Birds from Switzerland and England probably winter in geographically distinct parts of Africa. The Swiss birds probably feed on prey that are more reliant on C₃ vegetation, from woodlands, than the prey of English birds, which are more reliant on C₄ vegetation, from grasslands.     

Latency in response of barn swallow Hirundo rustica populations to changes in breeding habitat conditions

Inclusion of past, in addition to current ecological conditions can improve the realism of animal‐habitat models, because populations may show delayed responses to changing environments. We studied the relationships between the distribution of barn swallows (Hirundo rustica L.) in 2001 and ecological factors between 1986 and 2001. Livestock farming was the best predictor of swallow distribution. Maximum predictive power of swallow abundance was achieved by livestock farming data 7 years before the census and significantly earlier than 2001, as shown by a bootstrap analysis. Breeding philopatry, social facilitation of breeding habitat choice may delay response of barn swallow populations to rapidly changing ecological conditions in anthropogenic habitats. By bootstrapping the set of farms and applying our predictive distribution models to livestock farming data in the past and those predicted for the near future we forecast no significant change in swallow populations until 2015. This study suggests that population studies should incorporate historical ecological data.     

Effect of neighbour visibility on nest attendance patterns of Barn Swallows Hirundo rustica in loose colonies

To assess the benefits of nesting at a site hidden from neighbours in a loosely colonial species, the Barn Swallow Hirundo rustica, we carried out two field experiments, obstruction removal and mirror placement, both replicating a situation in which a nest is made visible from another nest. Under manipulated conditions in both experiments, females increased the length of time they stayed at their nests during the egg‐laying and late incubation stages, while males extended their duration of stay during the egg‐laying and early nestling stages in mirror placement experiments only. The results suggest that Barn Swallows conceal their nests to reduce fitness costs imposed by neighbours nesting in view and that hiding the nest can reduce the amount of time spent guarding the nest during certain stages of the breeding period.     

Structural coloration and sexual selection in the barn swallow Hirundo rustica

Structural coloration has been hypothesized to play a role insexual selection, and we tested whether this was the case ina field study of the barn swallow Hirundo rustica. The dorsaliridescent plumage of barn swallows has a strong reflectancein the ultraviolet (UV) region, with adult males on averagereflecting 8-9% more than adult females, as revealed by a 2-yearstudy in southwestern Spain. The correlation between structural coloration (described by the reflectance in the UV part of thespectrum, UV chroma and blue chroma) and three other secondarysexual characters significantly associated with male matingsuccess (tail length, tail asymmetry, and red facial coloration)was weak and generally nonsignificant. Nor was there a significantrelationship between color parameters and body condition. Wetested for an association between structural coloration of the dorsal plumage and sexual selection in a number of independenttests. Arrival date of males was not significantly relatedto color; there was no significant relationship between colorationand probability of survival or age; mated males did not havestronger reflectance than unmated males; and the duration ofthe premating period was not significantly related to color.Reproductive success was not significantly correlated withplumage coloration in males, nor was the feeding rate of offspringby brightly colored males higher than that of males with lessbright plumage. Given that sample sizes were large, and the power of statistical tests high, we conclude that current sexualselection on the coloration of the dorsal plumage in the barnswallow is, at best, weak.     

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.     

Breeding Barn Swallows Hirundo rustica on organic and conventional arable farms in the Netherlands

Populations of farmland birds are under pressure as a result of agricultural intensification. It has been proposed that less intensive approaches to farming, such as organic farming methods, may halt these population declines. In addition, organic farmers may have a more positive attitude towards nature and the environment which can possibly also have positive effects on the populations of breeding birds. We have compared the attitude of conventional and organic farmers towards the presence of Barn Swallows Hirundo rustica and the abundance of breeding Barn Swallows on organic and conventional arable farms in the Netherlands. We found that the abundance of breeding Barn Swallows did not differ between these two types of farms an that both organic and conventional farmers had a positive attitude towards the presence of Barn Swallows on their farms. Our results show that organic farming does not attract more Barn Swallows. However, agricultural intensification could have resulted in lower breeding success and, consequently, population declines, although there may be other contributory factors as well.     

Phenotypic quality and molt in the barn swallow, Hirundo rustica

Phenotypic quality may determine the development and expressionof secondary sexual characters. We studied the relationshipbetween molt and several measures of phenotypic quality in thesexually size-dimorphic barn swallow (Hirundo rustica) in itswinter quarters in Namibia. Males were in a more advanced stageof molt than females and juveniles, and the speed of molt asdetermined from the residual of the regression of the size ofthe gap in wings caused by missing and growing feathers on wingmolt score (residual wing raggedness) was also higher in malesthan in females and juveniles. Male barn swallows with longand symmetric tail feathers had a more advanced stage of moltand molted at a higher speed than males with short and asymmetrictails. Long-tailed females had a delayed molt, and females withasymmetric tails had less advanced molt and lower rates of feathergrowth than females with symmetric tails. Molt of secondariesin juveniles also appeared to be less advanced if they had longtails. Adult barn swallows molted their tail feathers in anirregular sequence with the longest, outermost tail featherusually replaced before the second or the third outermost feathers.Good body condition was positively associated with a high moltscore for some feather tracts and a rapid wing molt in adultfemales and tail molt in juveniles. Mallophaga were only weaklynegatively associated with primary and secondary molt scorein adult females and speed of wing molt in adult males. In conclusion,phenotypic quality of adult male barn swallows as reflectedby the expression of their secondary sexual character duringthe previous molt reliably reflected stage and speed of currentmolt.     

The effects of temperature, altitude and latitude on the arrival and departure dates of the swallow Hirundo rustica in the Slovak Republic

The (barn) swallow Hirundo rustica is a traditional harbinger of spring in many countries of the Northern Hemisphere. This paper uses information on the arrival and departure dates of the swallow in the Slovak Republic for the 30 years 1961–1985 and 1996–2000. Records were taken at 19 locations throughout the Republic representing an altitude range from 105 m to 760 m. Monthly temperature data were constructed from six meteorological stations. With the use of regression techniques, trends towards later arrival, earlier departure and the effects of latitude, altitude and temperature are all apparent. Received: 25 October 2000 / Revised: 23 April 2001 / Accepted: 23 April 2001     

Geographical and seasonal variation in the intensity of sexual selection in the barn swallow Hirundo rustica: a meta‐analysis

Sexual selection arises from competition among individuals for access to mates, resulting in the evolution of conspicuous sexually selected traits, especially when inter‐sexual competition is mediated by mate choice. Different sexual selection regimes may occur among populations/subspecies within the same species. This is particularly the case when mate choice is based on multiple sexually selected traits. However, empirical evidence supporting this hypothesis at the among‐populations level is scarce. We conducted a meta‐analysis of the intensity of sexual selection on the largest database to date for a single species, the barn swallow (Hirundo rustica), relying on quantitative estimates of sexual selection. The intensity of sexual selection was expressed as the strength (effect size) of the relationships between six plumage ornaments (tail length, tail asymmetry, size of white spots on tail, ventral plumage colour, throat plumage colour and throat patch size) and several fitness proxies related to reproduction, parental care, offspring quality, arrival date from spring migration, and survival. The data were gathered for four geographically separated subspecies (H. r. rustica, H. r. erythrogaster, H. r. gutturalis, H. r. transitiva). The overall mean effect size (Z_r = 0.214; 95% confidence interval = 0.175–0.254; N = 329) was of intermediate magnitude, with intensity of sexual selection being stronger in males than in females. Effect sizes varied during the breeding cycle, being larger before egg deposition, when competition for access to mates reaches its maximum (i.e. in the promiscuous part of the breeding cycle), and decreasing thereafter. In addition, effect sizes from experiments were not significantly larger than those from correlative studies. Finally, sexual selection on different sexually dimorphic traits varied among subspecies. This last result suggests that morphological divergence among populations has partly arisen from divergent sexual selection, which may eventually lead to speciation.     

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.     

Viability and expression of sexual ornaments in the barn swallow Hirundo rustica: a meta‐analysis

Sexual selection results in the evolution of exaggerated secondary sexual characters that can entail a viability cost. However, in species where sexual ornaments honestly reflect individual quality, the viability cost of secondary sexual characters may be overwhelmed by variation in individual quality, leading to expect that individuals with the largest secondary sexual characters show higher, rather than lower viability. Here, we used meta‐analyses to test whether such expected positive relationship between sexual ornamentation and viability exists in the barn swallow Hirundo rustica, which is one of the most studied model species of sexual selection under field conditions. We found a mean positive effect size of viability in relation to the expression of secondary sexual characters of 0.181 (CI: 0.084–0.278), indicating that in this species the more ornamented individuals are more viable, and therefore of high quality. Analyses of moderator variables showed similar effects in males and females, the H. r. rustica subspecies rather than others and tail length rather than other secondary sexual characters. Future research emphasis on other subspecies than the European one and secondary sexual characters than tail length may help identify the sources of heterogeneity in effect sizes.     

三江源国家公园草地地上生物量时空动态及其气候影响

三江源区位于青藏高原腹地,作为长江、黄河、澜沧江三大河流的发源地,是我国重要的生态安全屏障。基于三江源区域草地AGB的野外调查数据,本研究采用多种机器学习算法集成分析的方式构建模型,实现了高精度的三江源国家公园草地AGB时空估算。基于AGB时空模拟结果,分析了近19年(2000—2018年)三江源国家公园区域草地AGB的时空动态变化。研究结果显示：(1)通过多种机器学习结合贝叶斯平均模型,草地AGB模拟值与实测值的r为0.88,RMSE为71.60g/m²,表明多模型集成分析的方式对草地AGB估算获得了较好的模拟效果。(2)三江源国家公园区域草地AGB的空间分布具有明显的空间异质性,呈从东南向西北递减的趋势。(3)2000—2018年长江源国家公园、黄河源国家公园和澜沧江国家公园区域草地AGB多年平均值分别为82.96 g/m²、117.54g/m²和168.39 g/m²。(4)近19年间,在黄河和长江源园区受到温度上升的影响草地AGB呈现出非显著性上升趋势;澜沧江区域,由于2015和2016年的...