Breeding phenology,reproductive traits and apparent survival of sympatric Marsh Warbler Acrocephalus palustris and Blyth’s Reed Warbler Acrocephalus dumetorum

Capsule: Sympatric Marsh Warblers Acrocephalus palustris and Blyth’s Reed Warblers Acrocephalus dumetorum differ significantly in their life history traits.

Aims: To provide a direct comparison of demographic parameters among two sympatric populations of the closely related Marsh Warbler and Blyth’s Reed Warbler.

Methods: We examined breeding phenology and reproductive traits at a 25?ha study plot. We use program MARK to estimate daily nest survival and adult apparent survival rates.

Results: On average, Marsh Warblers laid the first eggs 3 days later than Blyth’s Reed Warblers. Mean clutch size in the Marsh Warbler was significantly lower than in the Blyth’s Reed Warbler. There are no significant differences between the two species for nest daily survival, duration of incubation and nestling periods. Apparent survival of adults was slightly higher in Marsh Warblers than in Blyth’s Reed Warblers.

Conclusion: Our findings suggest that two ecologically similar sympatric species differ significantly in terms of life history traits. We assume that observed differences could be the result of adaptations to environmental factors in the central parts of the species’ ranges or due to differences in mortality on migratory pathways or wintering grounds.     

Habitat-dependent population recovery in the Dartford Warbler Sylvia undata following a severe winter episode

Capsule?After a population crash following a long period of winter freezing and snow blanketing the ground, Dartford Warblers underwent a population recovery, positive in heathlands but non-existent in nearby early-growth forest.

Aims?To estimate the impact of a severe winter episode on the population size of Dartford Warblers in two habitats, heathland and early-growth forest.

Methods?We analysed data collected on 124 point counts in three successive years, 1 year before and 2 years after the cold spell. We first estimated trends in detected numbers in the two habitats. Second, we used distance sampling and capture–recapture approaches to estimate global population sizes corrected for detection probability, verifying that the observed trends were not biased by detection probability.

Results?Warbler populations crashed after the severe winter in 2008–9, while numbers increased in spring 2010 only on heathlands, not in early-growth forests. Variation in detection probability alone could not explain this difference.

Conclusion After the severe winter episode, Dartford Warblers showed a short-term population recovery only in heathland.     

Song in two cryptic species: comparative analysis of Large-billed Reed Warblers Acrocephalus orinus and Blyth’s Reed Warblers Acrocephalus dumetorum

Capsule: Songs of Large-billed Reed Warblers Acrocephalus orinus and Blyth’s Reed Warblers Acrocephalus dumetorum differed in quantitative parameters. Blyth’s Reed Warbler used different modes of singing on breeding grounds and migration stopovers.

Aims: To compare the songs of two cryptic species of reed warblers. To compare Blyth’s Reed Warbler songs in different parts of the breeding range and on migration stopovers in central Asia. To investigate the status of Blyth’s Reed Warblers in central Asia.

Methods: We analysed song recordings of individual singing males and conducted field observations of singing behaviour at migration stopover and breeding sites. The status of Blyth’s Reed Warblers in central Asia was reviewed from the historical Russian literature and other sources.

Results: We found differences between six out of eight variables in songs of Large-billed and Blyth’s Reed Warblers on breeding sites. Blyth’s Reed Warbler songs recorded on migration stopovers showed differences in five variables from Large-billed Reed Warblers and in six variables from breeding Blyth’s Reed Warblers. On migration stopovers, Blyth’s Reed Warblers sang actively but did not stay more than 1–2 days. The evidence suggests that Blyth’s Reed Warbler does not breed in central Asia.

Conclusion: Songs of Large-billed and Blyth’s Reed Warblers could be distinguished by quantitative analysis. Blyth’s Reed Warblers used different modes of singing on breeding sites and migration stopovers. We believe vocalizations of Blyth’s Reed Warblers on migration to be plastic song.     

POLYGYNY AND BREEDING ECOLOGY OF THE CETTI'S WARBLER CETTIA CETTI

Male Cetti's Warblers bred simultaneously with up to three females. Harem size was correlated with size of male but not of his territory. Males occupied large usually exclusive ranges and devoted much time to patrolling and singing often far from their nests. Incubation was solely by the female who also provided most to all of the food for the young. Incubation and fledging periods were prolonged compared with similar sized birds but first nests were unusually successful. Polygynously mated females laid larger clutches than monogamous ones. Males with fewer females took a greater share of feeding but chick growth and survival were not impaired by polygyny, nor did foods brought to the young differ accordingly. The existence of this unusual breeding system is discussed with emphasis on the co-evolution of sexual dimorphism and non-equal sex ratios resulting from malemale competition. It is postulated that the linear dispersion of its habitat may especially have favoured operation of the Verner-Orians polygyny model in the evolutionary history of the Cetti's Warbler.     

Nest-site selection and predation in Savi's Warblers Locustella luscinioides

Capsule Savi's Warblers preferred sites dominated by high and dense Juncus and Rubus with thick litter, and avoided pure reeds; predation was higher in sites with less litter.

Aim To describe nest-site selection and identify which habitat and nest features influence predation risk.

Methods Comparisons were made between habitat characteristics measured at nest-sites and random points, and between predated and successful nests.

Results Total vegetation cover, litter thickness, habitat type, undergrowth height and undergrowth cover, which were higher or denser at nest-sites, were the most important variables influencing the distribution of Savi's Warbler nests. Nests were most often predated in sites with low litter thickness.

Conclusion Savi's Warblers are dependent on the presence of dense low vegetation and litter for nesting, which should be maintained close to or within reed-beds for their benefit.     

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 use and densities of co‐existing migrant Willow Warblers Phylloscopus trochilus and resident eremomelas Eremomela spp. in Zimbabwe

Capsule Migrant Willow Warblers occupy more woodland types and occur at higher densities than ecologically‐similar resident Afrotropical warblers.

Aims To compare population densities of Willow Warblers and eremomelas in adjacent acacia, mopane and miombo woodlands, and assess the abundance of potential invertebrate prey in each habitat type, in order to investigate whether Palearctic migrants use more open habitats and are more flexible in habitat use than their Afrotropical counterparts in the same feeding guild.

Methods Using distance sampling we carried out four replicated sets of point counts in acacia woodland and three sets of counts in miombo and mopane between December 1999 and February 2000. We noted the tree species in which we saw warblers foraging and took beating‐tray samples of potential arthropod prey present on tree foliage in each of the three habitats.

Results Willow Warbler density in acacia woodland increased from 1.80 ± 0.54 (se) birds/ha in early December to 7.15 ± 1.41 birds/ha in late January after influxes of later arrivals. Densities of Willow Warblers in miombo and mopane were much lower (1.14 ± 0.28 and 0.38 ± 0.23 birds/ha, respectively) and did not show significant changes. Burnt‐necked Eremomelas averaged 0.74 ± 0.34 birds/ha in acacia woodland, and in miombo densities of Green‐capped and Yellow‐bellied Eremomelas were 0.23 ± 0.17 and 0.34 ± 0.26 birds/ha, respectively. Densities in mopane were too low to estimate reliably. Willow Warblers and Green‐capped Eremomelas showed some apparent preferences in tree species used for foraging but differences in tree use were not obviously related to the abundance of arthropod taxa present as potential prey.

Conclusion Willow Warblers occupied more habitats at greater density than similar Afrotropical warblers. They appear to favour acacia, but their settlement patterns and the reasons for disparities between densities of immigrants and residents are unclear.     

Endemic Cyprus Warbler Sylvia melanothorax and colonizing Sardinian Warbler Sylvia melanocephala show different habitat associations

Anthropogenic habitat change and assisted colonization are promoting range expansions of some widespread species with potential consequences for endemic fauna. The recent colonization of Cyprus by breeding Sardinian Warblers Sylvia melanocephala has raised concerns that it might be displacing the closely related and endemic Cyprus Warbler Sylvia melanothorax. Habitat associations of both species were examined using models of abundance within the 95% density kernel of the Sardinian Warbler’s range and also outside this range for Cyprus Warbler. Within the Sardinian Warbler’s range, the two species were associated with subtly different scrub habitats. Outside the Sardinian Warbler’s range, the Cyprus Warbler differed again in its habitat association, but this probably resulted from marked differences in habitat extent and availability in different parts of the island rather than from competitive displacement, as none of the habitat or land‐use elements differentially associated with Cyprus Warblers was positively associated with Sardinian Warbler occurrence. This suggests that the Sardinian Warbler has exploited a different niche, rather than displacing the endemic species, and has perhaps benefitted from changing land‐use patterns, particularly recent fallows and abandoned agriculture, in contrast to the stronger association of Cyprus Warblers with semi‐natural scrub.     

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.     

Timing of breeding and nestling diet of Wood Warbler Phylloscopus sibilatrix in relation to changing food supply

Capsule Wood Warblers did not match their reproduction to the caterpillar peak

Aims To study the timing of Wood Warblers’ breeding and nestling diet in relation to caterpillar abundance under primeval conditions in the Bia?owie?a National Park, Poland.

Methods Observations of food brought by parents. Inter‐year and seasonal changes in availability of folivorous caterpillars were assessed by direct counts and caterpillar frass collection.

Results Maximum food requirements occurred two weeks after the peak of caterpillar abundance. The mismatch had no effect on nestlings’ development. Diet varied little across years and habitats, but varied strongly within a season. Following the decline of ‘green’ caterpillars, their proportion in the Wood Warbler diet strongly declined. Caterpillars were replaced by winged insects. Small nestlings received more spiders than older ones.

Conclusion Timing of Wood Warbler breeding in Bia?owie?a National Park was constrained by the females’ arrival time. Birds did not match their reproduction to the caterpillar peak. ‘Green’ caterpillars were the preferred food for nestlings; birds responded to the caterpillar decrease by feeding smaller proportions of them and switching to alternative prey, namely winged insects.     

Nest defence,enemy recognition and nest inspection behaviour of experimentally parasitized Reed Warblers Acrocephalus scirpaceus

Capsule Reed Warblers in a regularly parasitized population do not recognize Cuckoo Cuculus canorus as a special enemy and do not change their behaviour at nest immediately after being parasitized.

Aims To assess if an intruder near the nest influences the behaviour of the Cuckoo host.

Methods Host responses to Cuckoo, control Pigeon dummies and human intruder were observed. Host behaviour at 71 nests was video-recorded for 30 minutes at four experimental groups of nests: Cuckoo dummy, Cuckoo dummy + Cuckoo egg, Pigeon dummy, human intruder.

Results Reed Warblers did not respond differently to the Cuckoo and the control species. The experimental procedure had no significant effect on the behaviour of hosts during the study period. We were unable to find any differences in the time spent at the nest, clutch inspection behaviour and nest defence behaviour between morning and afternoon experimental groups. Our results do not support the hypothesis that afternoon laying by the Cuckoo is maintained by a selection pressure from the host. We observed no ejection or egg-pecking during the 30-min period after the experimental parasitism.

Conclusions Low aggression and non-specificity of host responses in our study area are in line with the fact that the Reed Warbler is an intermediate rejecter of Cuckoo eggs as expected from the spatial habitat structure hypothesis.     

Why Seychelles Warblers fail to recolonize nearby islands: unwilling or unable to fly there?

The Seychelles Warbler Acrocephalus sechellensis is a rare island endemic which, from 1920 to 1988, occurred only on Cousin Island (29 ha) in the Seychelles. Despite the saturated nature of this population and the possibility of obtaining higher reproductive success on new nearby islands, inter-island dispersal by Seychelles Warblers is extremely rare (0.10%). We test the hypothesis that Seychelles Warblers show an adaptation typical for island birds: a low-cost reduced-size flight apparatus. We compared the anatomy of the flight apparatus (wing shape, wing loading, skeletal parts and musculature) of Seychelles Warblers with that of three closely related migratory Acrocephalus species (Eurasian Reed Warbler A. scirpaceus , Australian Reed Warbler A. australis and African Reed Warbler A. baeticatus ). Seychelles Warblers do not differ from the migratory warblers in pectoral mass and skeletal attachment area relative to body mass, wing shape and wing loading. Seychelles Warblers show the morphological structures required for sustained flight, but may have the behavioural reluctance to cross what they may regard as extensive bodies of water.     

Habitat associations of Wood Warblers Phylloscopus sibilatrix breeding in Welsh oakwoods

Capsule Territory locations, density and the change in numbers over 20 years were associated with characteristics of the canopy, understorey structure and field-layer vegetation cover.

Aims To identify habitat characteristics associated with territory locations and density of Wood Warblers Phylloscopus sibilatrix in Welsh oakwoods in 2009–2011, and the change in abundance between 1982–1984 and 2003–2004.

Methods In 2009–2011, habitat characteristics were compared between 106 territories and 226 unoccupied points in 19 woods. Mean wood-scale habitat values were related to density in 27 woodland blocks. The change in Wood Warbler numbers between 1982–1984 and 2003–2004 was related to initial habitat quality in the 1980s and the change in habitat characteristics between the two time periods.

Results The location of territories in 2009–2011 was positively associated with canopy height, and with intermediate values of slope steepness, field-layer vegetation cover, canopy cover, the proportion of Oak Quercus in the tree community and subcanopy cover. Density was positively associated with slope, subcanopy cover at 0.5–2 m height and a landscape dominated by coniferous plantation and moorland; and with intermediate values of the proportion of oaks in the tree community. Wood Warblers declined by 24.4% in the two Welsh regions between 1982–1984 and 2003–2004, and trends were positively associated with the initial cover of Bramble and, in Gwynedd only, canopy cover.

Conclusions Wood Warblers were associated with a number of structural habitat variables, which could be related to the past management of the study woods. Management should be targeted at restoring habitat quality for Wood Warblers through the introduction of a moderate grazing regime.     

Cross‐decades stability of an avian hybrid zone

Hybrid zones are particularly valuable for understanding the evolution of partial reproductive isolation between differentiated populations. An increasing number of hybrid zones have been inferred to move over time, but in most such cases zone movement has not been tested with long‐term genomic data. The hybrid zone between Townsend's Warblers (Setophaga townsendi) and Hermit Warblers (S. occidentalis) in the Washington Cascades was previously inferred to be moving from northern S. townsendi southwards towards S. occidentalis, based on plumage and behavioural patterns as well as a 2000‐km genetic wake of hermit mitochondrial DNA (mtDNA) in coastal Townsend's Warblers. We directly tested whether hybrid zone position has changed over 2–3 decades by tracking plumage, mtDNA and nuclear genomic variation across the hybrid zone over two sampling periods (1987–94 and 2015–16). Surprisingly, there was no significant movement in genomic or plumage cline centres between the two time periods. Plumage cline widths were narrower than expected by neutral diffusion, consistent with a ‘tension zone’ model, in which selection against hybrids is balanced by movement of parental forms into the zone. Our results indicate that this hybrid zone is either stable in its location or moving at a rate that is not detectable over 2–3 decades. Despite considerable gene flow, the stable clines in multiple phenotypic and genotypic characters over decades suggest evolutionary stability of this young pair of sister species, allowing divergence to continue. We propose a novel biogeographic scenario to explain these patterns: rather than the hybrid zone having moved thousands of kilometres to its current position, inland Townsend's met coastal Hermit Warbler populations along a broad front of the British Columbia and Alaska coast and hybridization led to replacement of the Hermit Warbler plumage with Townsend's Warbler plumage patterns along this coastline. Hence, hybrid zones along British Columbia and Alaska moved only a short distance from the inland to the coast, whereas the Hermit Warbler phenotype appears stable in Washington and further south. This case provides an example of the complex biogeographic processes that have led to the distribution of current phenotypes within and among closely related species.     

Survival of British Sedge Warblers Acrocephalus schoenobaenus in relation to west African rainfall

Fluctuations in the population levels and annual adult survival rates of British Sedge Warblers Acrocephalus schoenobaenus since the late 1960s are strongly correlated with indices of wet season rainfall in the west African winter quarters. Population changes are unrelated to estimates of breeding productivity in the previous year. Habitat availability in the winter quarters has probably been the main factor limiting the size of the Sedge Warbler population in Britain during the period of study.     

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.     

Extreme genetic similarity does not predict non‐breeding distribution of two closely related warblers

Detailed knowledge of migratory connectivity can facilitate effective conservation of Neotropical migrants by helping biologists understand where and when populations may be most limited. We studied the migratory behavior and non‐breeding distribution of two closely related species of conservation concern, the Golden‐winged Warbler (Vermivora chrysoptera) and Blue‐winged Warbler (Vermivora cyanoptera). Although both species have undergone dynamic range shifts and population changes attributed to habitat loss and social interactions promoting competition and hybridization, full life‐cycle conservation planning has been limited by a lack of information about their non‐breeding ecology. Because recent work has demonstrated that the two species are nearly identical genetically, we predicted that individuals from a single breeding population would have similar migratory timing and overwintering locations. In 2015, we placed light‐level geolocators on 25 males of both species and hybrids in an area of breeding sympatry at the Fort Drum Military Installation in Jefferson and Lewis counties, New York. Despite extreme genetic similarity, non‐breeding locations and duration of migration differed among genotypes. Golden‐winged Warblers (N = 2) overwintered > 1900 km southeast of the nearest Blue‐winged Warbler (N = 3) and spent nearly twice as many days in migration; hybrids (N = 2) had intermediate wintering distributions and migratory timing. Spring migration departure dates were staggered based on distance from the breeding area, and all birds arrived at the breeding site within 8 days of each other. Our results show that Golden‐winged Warblers and Blue‐winged Warblers in our study area retain species‐specific non‐breeding locations despite extreme genetic similarity, and suggest that non‐breeding locations and migratory timing vary along a genetic gradient. If the migratory period is limiting for these species, our results also suggest that Golden‐winged Warblers in our study population may be more vulnerable to population decline than Blue‐winged Warblers because they spend almost twice as many days migrating.     

Recovery of the Peregrine Falcon Falco peregrinus in Cumbria,UK, 1966–99

Capsule Since pesticide controls, it has expanded from a pre-pesticide baseline of 41 pairs to over 100 pairs.

Aims To describe the population's pattern of recovery and identify constraints on its annual productivity.

Methods Occupancy and productivity data were collected from all known and new ranges in Cumbria, UK. Statistical methods were employed to describe temporal and spatial patterns. Possible effects of weather were investigated using meteorological data.

Results The population recovered from eight pairs, with birds preferentially recolonizing 41 traditional ranges. The highest occupancy levels were 100 ranges in 1997 and 1998. Many new ranges were in peripheral and atypical locations, although some resulted from the splitting of traditional ones. The central core now has the highest recorded density (> 9 pairs per 100 km²). As occupied ranges increased, the mean nearest neighbour distance declined to an asymptote (approximately 3.5 km). Simultaneously, maximum values were observed for the percentages of known and occupied ranges in which eggs were laid. The mean number fledged by occupied ranges peaked in the early 1990s. New ranges seem to be more susceptible to the effects of May rainfall but overall there was little evidence that new and traditional ranges differ in productivity. Persecution has continued, with over 10% of examined ranges experiencing intentional disturbance in the worst years.

Conclusion The recovery of the Cumbrian population seems to be related to two processes: (1) greater productivity and survival forcing birds to occupy more peripheral and atypical sites; (2) a very high density in the central core possibly due to increased food supply resulting from the channelling of prey through its valley systems.     

Winter habitat use does not influence spring arrival dates or the reproductive success of Yellow Warblers breeding in the arctic

Winter habitat use can influence the breeding success of migratory songbirds in temperate regions due to its impact on bird condition and breeding phenology. How such carry-over effects vary with latitude is unknown. To address this question, we examined how winter habitat use, inferred from δ¹³C and δ¹⁵N signatures in winter-grown feathers, influenced the breeding phenology and productivity of Yellow Warblers (Setophaga petechia) at the extreme north of their range in the Canadian arctic (68°N) and compared this population with midlatitude Yellow Warbler (51°N) and American Redstart (Setophaga ruticilla; 44°N) populations reported in previous studies. In the arctic, we examined male arrival dates, female clutch initiation dates and the relationship between these timing variables and the number and quality of offspring produced within the season. In contrast to warblers breeding at midlatitudes, we find no support for an impact of winter habitat use on breeding phenology or productivity. Male arrival dates and female clutch initiation dates in both young and older individuals were not correlated with isotopic signatures acquired on the wintering grounds. Males with enriched δ¹⁵N signatures paired more rapidly after arrival, indicating a possible relationship between winter habitat use and condition. This relationship did not enhance annual productivity for these individuals, however, as the negative relationship between breeding phenology and reproductive success in our arctic population was significantly weaker than among Yellow Warblers breeding further south. This reduction or absence of timing effects on productivity in the north effectively removes one pathway through which carry-over effects can act.