Polygyny in the Willow Warbler Phylloscopus trochilus in Swedish Lapland

The breeding sex ratio of Willow Warblers Phylloscopus trochilus in a subalpine birch forest was strongly female biased. Very few territorial males remained unmated and most became polygynous. Bigyny was the rule, but some males probably had three females. Male assistance at secondary nests varied and was probably dependent on temporal distance between the females' nests. There was no significant reduction in reproductive output for these females. Variation in the sex ratio of breeding birds can be explained, at least partly, by variation in the timing of male settlement.     

The presence of non-territorial males in Willow Warbler Phylloscopus trochilus populations–a removal study

A total of six (unmarked) and 24 (individually colour-ringed) male Willow Warblers Phylloscopus trochilus were removed from their territories in two study areas: subalpine birch and lowland mixed forest, respectively. The removals were made during three separate periods throughout the breeding season: (1) pair-formation and egg-laying; (2) early incubation; and (3) middle and second half of the incubation period. During Period 1 all of the males removed ( n = 10) were quickly replaced by new males, while during Period 2 eight of the 11 males removed were replaced. The new males defended basically the same territorial borders as the males they replaced. During Period 3 only one of the nine males removed was replaced. Since the replacements occurred within a few hours of the removals, it is suggested that the new settlers were non-territorial males that were already present in the respective areas before the experimental removals were made. The mean wing-lengths of the two groups of males indicated that the initial occupants of the territories were 2 years old or older in both study areas, whereas the new settlers in the subalpine birch forest (mean Willow Warbler density: 55 territories/km²) were 1-year-old birds, and two years old, or older, in the mixed forest area (mean Willow Warbler density: 133 territories/km²).     

Willow Warbler Phylloscopus trochilus habitat in woods with different structure and management in southern England

Capsule Use of Light Detection and Ranging (LiDAR) data identified suitable Willow Warbler habitat based on mean vegetation height. This habitat model provided maps of distribution and occupation of suitable habitat.

Aims To identify habitat associations in woods with different vegetation structure and management systems during a period of low Willow Warbler populations.

Methods Locations of all Willow Warblers were mapped during the breeding season in three woods of contrasting management; recent low intervention, actively coppiced woodland and high forest with clear‐fells. Height profile models of each wood were derived from airborne LiDAR. The mean vegetation height at locations with Willow Warblers and a sample from the rest of the wood were used to produce models of optimum habitat and breadth of habitat occupied in each wood. The habitat model was then used to produce maps of suitable habitat.

Results The habitat models did not differ between woods, with highest probability of Willow Warbler occurrence in mean vegetation heights of 3.7–5.3 m. Habitat of heights 6–11 m appeared less suitable, being only partly occupied. Habitat maps showed that habitat of suitable height was only occupied when it occurred as large patches; smaller patches (mostly <0.5 ha) and edges along rides and fields were not used.

Conclusion The use of LiDAR derived measures of vegetation height identified areas of suitable habitat for Willow Warblers. Willow Warblers occupied areas of low mean vegetation height either as early successional or open canopy woodland in all woods. Height‐based habitat maps can identify areas of suitable habitat within larger expanses of heterogeneous woodland and are a potentially useful tool in assessing changes in extent of what are often temporary patches of habitat.     

Habitat selection, foraging niches and horizontal spacing of Willow Warbler Phylloscopus trochilus and Chiffchaff P. collybita in an area of sympatry

Habitat selection of the Willow Warbler and the Chiffchaff was studied along a succession of grey alder forest in Central Norway (63^oN) where both species breed. Both were found in all serai stages, but there was a slight tendency for the Willow Warbler to be found more often in the voungest stages than was the Chiffchaff.
Foraging niches and horizontal spacing were studied within one particular forest. Most of the records of both Willow Warbler and Chiffchaff came from the tree layer, and the overlap between the species in their foraging niches was considerable. However, the species were found to occupy almost mutually exclusive territories.     

The biannual primary moult of Willow Warblers Phylloscopus trochilus in Europe and Africa

The Willow Warbler Phylloscopus trochilus is one of the few bird species that undergoes two primary moults a year, a post-nuptial moult in the breeding area and a moult in the wintering area. Primary-moult data for Willow Warblers from Finland, Sweden, Britain, the Netherlands, Belgium. Guinea-Bissau, Uganda, Kenya, Malawi, Zambia, Zimbabwe, Botswana and South Africa are analysed. The parameters of primary moult (mean starting date, standard deviation of starting date, and duration) are estimated using the techniques of Underhill & Zucchini (T.988 Ibis 1 30: 358–372) and Underhill, Zucchini & Summers (1990 Ibis 132: 118-12 3). The scheduling of moult in relation to theother main components of the annual cycle, breeding and migration, is considered. The mean durations of post-nuptial moult for P. t. trochilus and P. t. acredula are 36.5 and 38.3 days, respectively; the start and termination of moult for P. t. trochilus are about 3.5 days later for each degree of latitude northwards, and the start and termination of moult for P. t. acredula, are about 10 days later than that of the most northerly populations of P. t. trochilus studied. Females start their postnuptial moult about 10 days later than males. Southward migration commences as soon as post-nuptial moult is complete. There is an increasing constraint on the timing of breeding and post-nuptial moult events at higher latitudes, leading to overlap between them. The duration of pre-nuptial moult is longer than that of post-nuptial moult, and is completed shortly prior to northward migration.     

Using stable isotopes to link breeding population trends to winter ecology in Willow Warblers,Phylloscopus trochilus

Capsule Stable isotope composition of feathers grown in Africa indicates that Willow Warblers, Phylloscopus trochilus, breeding in areas of Britain experiencing different population trends may vary in their distribution, timing of moult and/or use of prey resources during the non-breeding season.

Aims To compare stable isotope ratios of feathers of Willow Warblers breeding in parts of Britain with differing rates of population change.

Methods Feathers were collected from three regions with differing recent population trends; northern Scotland (population increased), western Wales (declined slightly) and eastern England (declined rapidly). Collection occurred at ringing sites run by British Trust for Ornithology (BTO) volunteers as part of the Constant Effort Sites (CES) scheme between April and July in 2008 and 2009.

Results Willow Warblers from northern Scotland had significantly higher δ ¹⁵N and δ ¹³C values than those from eastern England, although the regional mean values did not differ greatly. There was no significant difference in δ ¹⁵N and δ ¹³C values between the sexes, but δ ¹⁵N values were significantly greater in individuals caught later in the breeding season.

Conclusions During the premigratory moult period in Africa, Willow Warblers from different parts of Britain may vary in their distribution, timing of moult and/or use of prey resources.     

Moult, flight muscle "hypertrophy" and premigratory lipid deposition of the juvenile Willow warbler, Phylloscopus trochilus

In England juvenile Willow warblers undergo premigratory lipid deposition in August. Prior to this there is a moult of the juvenile plumage which results in a reduction in both the glycogen content and the wet weight of the pectoralis muscles, the latter reflecting a decrease in the lipid-free dry weight. Muscle lipid levels are apparently unaffected by this moult. The decrease in the weight of the lipid-free dry fraction of the body extends also to components other than these muscles at this time. A correlation between the wet weight and the glycogen content of the pectoralis muscles suggests that cold stress may be the cause of the low glycogen levels found in the middle of the moult; part of this increased demand for thermogenesis may be due to lower body insulation brought about by feather loss during the moult. The muscle "hypertrophy" found in the premigratory period is simply a return to pre-moult weights.
The uniformity of total body weights during and after the moult indicates that at the end of this moult the water fraction of the body decreases in weight. The pectoralis muscles of juvenile Willow warblers are not fully grown on fledging; these unmoulted birds may also have higher body lipid levels than during the following moult.     

Woodland structure,rather than tree identity,determines the breeding habitat of Willow Warblers Phylloscopus trochilus in the northwest of England

Capsule Woodland structure, rather than tree species, is the most important determinant of breeding habitat selection by Willow Warblers in North West England.

Aims To examine how habitat characteristics predict the occurrence of male Willow Warbler territories.

Methods Woodland structure (trunk density, trunk diameter, canopy cover and understory cover), tree species and food abundance were compared between woodland areas within and outside of male territories at a site in the UK.

Results Territories contained higher trunk numbers, had a narrow range of trunk diameters, and intermediate canopy cover. Food abundance did not differ with occupancy. Willow and alder were the most common trees within territories, in contrast to birch which has been found in previous studies. The habitat structure matches young woodlands, where birches often grow. However, at the study site the birches were large and mature, and therefore unsuitable. Moreover, woodland structure variables were better predictors of occupancy than any particular tree genera.

Conclusion The results indicate that vegetation structure, but not tree species or food availability, influence breeding habitat selection by Willow Warblers. The preferred structure is similar to coppice woodlands; therefore, the Willow Warbler decline may be linked to the loss of this traditional management across south England.     

Extra-pair paternity and song characteristics in the willow warbler Phylloscopus trochilus

Complexity in bird song is often argued to be advantageous in processes of sexual selection, and numerous studies show that characteristics of song are associated with increased success in territory defence or mate attraction. Less evidence exists on the relationship between bird song characteristics and patterns of extra-pair paternity. We tested whether males that suffered from extra-pair paternity differed in song characteristics from males with no extra pair paternity in the willow warbler Phylloscopus trochilus . In the Scottish population that we studied, we found that 23.5% of young were not related to the social father, and that 47% of nests contained at least one extra pair young. Older males were less likely to suffer paternity loss. While song repertoire size was not related to loss of paternity, males with short songs suffered higher paternity loss than males with long songs. Although arrival date is a good correlate of social mate choice in this population, it was not related to extra-pair paternity. These results suggest that females use song length, or a trait correlated with this song characteristic, as a cue for choosing extra-pair partners in this species, or alternatively that variance in the success of mate guarding or female coercion is related to this song variable.     

甘肃莲花山暗绿柳莺的孵卵行为

2001年5～7月,在甘肃省莲花山自然保护区对暗绿柳莺Phylloscopus trochiloides的孵卵行为进行了初步研究.结果 表明,孵化期内雌鸟日活动期长度平均为(848.5±14.8) min (n=17),每天离巢(16.0±3.0)次(n=15),每次离巢时间为(12.3±5.0) min (n=251),每次在巢时间为(43.6±21.9) min (n=236),在巢率为(78.8±2.4)%.雌鸟在巢时卵温平均为(31.3±3.5)℃ (n=10646),离巢时卵温平均为(26.6±4.8)℃ (n=2876);夜晚的平均卵温为(30.6±3.5)℃ (n=9239).孵卵温度在孵卵期有逐渐上升的趋势.     

湖南壶瓶山发现峨眉柳莺

正2018年5月5日,在湖南石门县壶瓶山国家级自然保护区中部的神水溪先后观察到6只峨眉柳莺(Phylloscopus emeiensis),拍摄到照片(图1),并录制了鸣唱声(图2)。同年5和6月,又先后在壶瓶山自然保护区内的平洞、望月湖、剩头、渫水、油榨湾和壶瓶山主峰共观察到41只峨眉柳莺(表1)。经查阅相关资料(郑作新等2010,邓学建等2013,郑光美2017),确认该鸟种为湖南鸟类分布新记录种。观察发现,     

Estimation of the Genetic Diversity of Willow Warbler Populations of the Subspecies <Emphasis Type="Italic">Phylloscopus trochilus acredula</Emphasis> (L.) in Different Parts of Its Nesting Area in the European Part of Russia

The genetic diversity (GD) of the Phylloscopus trochilus acredula subspecies populations was studied in different parts of the breeding area in the European part of Russia using microsatellite analysis. It was detected that the level of GD sufficient for the preservation of population genetic structure of the species with optimal ratio of intra- and interpopulation components of the gene diversity and minimal deviations from panmixia is maintained in the studied populations. The maximal values of GD indices were registered in an ecologically suboptimum zone on the periphery of the breeding range.     

莲花山云南柳莺的孵卵行为

20 0 2年 6～ 7月 ,在甘肃省莲花山自然保护区对云南柳莺的孵卵行为进行了研究。结果表明 ,雌鸟孵卵的日活动期为 (80 0 5± 42 8)min (n =1 5 ) ,每天离巢 (3 2 7± 3 9)次 (n =1 5 ) ,每次离巢时间 (6 6±1 8)min (n =5 99) ,每次坐巢时间 (1 8 4± 9 2 )min (n =5 83 ) ,坐巢率为 (73 1± 1 9) %。雌鸟每次坐巢时间和离巢时间的长度均与气温显著相关。日活动期雌鸟在巢的平均卵温为 3 2 3℃ ,夜晚的平均卵温为3 2 7℃。整个孵卵期卵温在发育临界值 2 8℃以上的时间比例为 92 7%。在孵卵后期卵温有逐渐上升的趋势。     

Functional infertility among territorial males in two passerine species, the willow warbler Phylloscopus trochilus and the bluethroat Luscinia svecica

Information about male infertility in free-living birds is scarce, but anecdotal and circumstantial evidence suggests that it does occur regularly at a low frequency. In this paper we document three cases of azoospermia in two passerine species, the willow warbler Phylloscopus trochilus and the bluethroat Luscinia svecica at their breeding grounds in South Norway. In willow warblers, two males out of a sample of 50 territory holders had no sperm in their seminal glomera, the storage site of sperm ready for ejaculation. The two males also had very small testes. One out of 48 bluethroat males also had no sperm in the seminal glomera. This male had an extreme asymmetry of the testes, with the right testis being about twice as large as the left. He also failed to fertilize any eggs in his own nest, as well as in neighbouring nests, as revealed by microsatellite genotyping. Thus, the proportion of males without sperm seems to be at a magnitude of a few (2–4) percent in both species. These are among the first estimates of the frequency of azoospermia in wild birds. Our results indicate a significant risk for sexually monogamous females of laying unfertilized eggs, which could favour the evolution of extra-pair copulation as a fertility insurance strategy in females.     

Male parental care, female reproductive success, and extrapair paternity

Birds differ considerably in the degree of male parental care,and it has been suggested that interspecific variation in extrapairpaternity is determined by the relative importance of benefitsto females from male parental care and good genes from extrapairsires. I estimated the relationship between extrapair paternityand the importance of male parental care for female reproductivesuccess mainly based on male removal studies, using a comparativeapproach. The reduction in female reproductive success causedby the absence of a male mate was positively correlated withthe male contribution to feeding offspring. The frequency ofextrapair paternity was negatively related to the reductionin female reproductive success caused by the absence of a mate.This was also the case when potentially confounding variablessuch as developmental mode of offspring and sexual dichromatismwere considered. A high frequency of extrapair paternity occursparticularly in bird species in which males play a minor rolein offspring provisioning and in which attractive males providerelatively little parental care. Bird species with frequentextrapair paternity thus appear to be those in which direct fitness benefits from male care are small, females can readilycompensate for the absence of male care, and indirect fitnessbenefits from extrapair sires are important.     

Characterization of a divergent chromosome region in the willow warbler Phylloscopus trochilus using avian genomic resources

Genome scans have made it possible to find outlier markers thought to have been influenced by divergent selection in almost any wild population. However, the lack of genomic information in nonmodel species often makes it difficult to associate these markers with certain genes or chromosome regions. Furthermore, the extent of linkage disequilibrium (LD) in the genome will determine the density of markers required to identify the genes under selection. In this study, we investigated a chromosome region in the willow warbler Phylloscopus trochilus surrounding a single marker previously identified in a genome scan. We first located the marker in the assembled genome of another species, the zebra finch Taeniopygia guttata, and amplified surrounding sequences in Fennoscandian willow warblers. Within an investigated chromosome region of 7.3 Mb as mapped to the zebra finch genome, we observed elevated genetic differentiation between a southern and a northern population across a 2.5-Mb interval comprising numerous coding genes. Within the southern and northern populations, higher values of LD were mostly found between SNPs within the same locus, but extended across distantly situated loci when the analyses were restricted to sampling sites showing intermediate allele frequencies of southern and northern alleles. Our study shows that cross-species genome information is a useful resource to obtain candidate sequences adjacent to outlier markers in nonmodel species.