Winter territoriality and its implications for the breeding ecology of White-throated Dippers Cinclus cinclus

Capsule: Pairs of White-throated Dippers Cinclus cinclus which defended winter territories bred earlier than non-territorial individuals, but there was no difference in reproductive success.

Aims: The effect of winter territoriality on breeding ecology has rarely been studied in resident birds. We carried out a preliminary investigation of whether winter territorial behaviour and territory size affect the timing of reproduction, breeding territory size and reproductive success in a riverine bird, the White-throated Dipper.

Methods: We monitored an individually marked population of White-throated Dippers in the UK. Wintering individuals were classified as either territorial or ‘floaters’ according to their patterns of occurrence and behaviour, and their nesting attempts were closely monitored in the subsequent months. Winter and breeding territory sizes were measured by gently ‘pushing’ birds along the river and recording the point at which they turned back.

Results: All birds defending winter territories did so in pairs, but some individuals changed partners before breeding. Territorial pairs that were together throughout the study laid eggs significantly earlier than pairs containing floaters and those comprising territorial birds that changed partners. However, there were no significant differences in clutch size, nestling mass or the number of chicks fledged. There was no relationship between winter territory length and lay date or any measure of reproductive success, although sample sizes were small. Winter territories were found to be significantly shorter than breeding territories.

Conclusion: Winter territoriality may be advantageous because breeding earlier increases the likelihood that pairs will raise a second brood, but further study is needed. Territories are shorter in winter as altitudinal migrants from upland streams increase population density on rivers, but this may also reflect seasonal changes in nutritional and energetic demands.     

Structure of tree vegetation may reduce costs of territory defence in Eurasian Nuthatch Sitta europaea

Capsule In pastured old oak woodland, breeding pairs of Eurasian Nuthatch selected territories with significantly higher densities of trees and cavities.

Aims To define territorial borders that exist within a nuthatch population and to compare densities of trees and cavities between used and non-used parts of their available habitat in order to determine habitat preferences.

Methods The number and location of territories was estimated by mapping the individual positions of colour-ringed birds. We measured habitat characteristics in regular sampling grid (number of trees and number of cavities per 50?×?50?m) during three successive seasons and modelled probability of occurrence of nuthatch territories in the area.

Results Breeding pairs were observed to defend territories of median size between 1.46 and 2.93?ha. These sizes correspond well to high-quality habitat territories for this species. We found that pairs tended to occupy territories with higher densities of trees and cavities. Based on generalized linear mixed-effects model estimates, we found that in pastured oak woodland habitat with density of 60 trees per ha, the estimated probability of occurrence (with 95% confidence interval) of a nuthatch territory was about 96% (83?99). For cavity density, the probability of territory occurrence was about 74% (52?89) at the level of 60 cavities per ha.

Conclusion We suggest that higher tree and cavity densities reduce territory defence costs in the species because birds may use less energy during foraging, predation avoidance and competition for a nest hole.     

Nest site selection and patterns of nest re-use in the Hooded Crow Corvus cornix

Capsule: Hooded Crows Corvus cornix selected nesting trees based on species, height, grouping and distance from an occupied house. Nest re-use was common and pairs that re-used old nests produced more fledglings than those that built a new nest.

Aims: To determine the features of trees that influenced whether they were used by Hooded Crows as nest sites, to establish what factors influenced nest re-use between years and to explore potential costs or benefits of nest re-use.

Methods: In a large area of Orkney, Scotland, the features of trees that contained a Hooded Crow nest were compared to those of trees where nests were absent. Patterns of nest re-use between years were examined in relation to the availability of alternative sites, previous nesting success and the number of equivalent options to the tree used previously within 200?m of this site.

Results: Hooded Crows favoured spruce and pine trees as nest sites, above the most locally abundant tree species, elder and willow. Preference for trees increased with tree height, local tree density and distance from occupied houses. Over half of the crows studied re-used an old nest when one was available and crows that re-used an old nest fledged more offspring than those that built a new nest. The likelihood of a new nest being built increased as the number of potential locations to build increased. Territories where a nest survived the winter were more likely to be reoccupied the following year than those where nests fell, while territories with fewer trees around the old site were most likely to be abandoned, suggesting that those were territories of lower quality.

Conclusions: Hooded Crows displayed strong preferences for nest sites that might favour nesting success by offering concealment, shelter and protection from ground-based predators. Nest re-use was common, especially when alternative sites were scarce, and appeared to facilitate greater reproductive output.     

Habitat preferences of the Syrian Woodpecker Dendrocopos syriacus in urban environments: an ambiguous effect of pollution

Capsule The occurrence of the Syrian Woodpecker Dendrocopos syriacus in an urbanized habitat is determined by the number of trees and pollutant emissions. Air pollution may weaken trees and increase the number of insects inhabiting them.

Aims To determine the habitat preferences of the Syrian Woodpecker in the highly urbanized environment of the city of Kraków.

Methods A set of 13 habitat and environmental parameters of 50 breeding territories and 50 random points were evaluated. A logistic regression was used to determine the probability of species occurrence, and Akaike's information criterion was used for model selection.

Results The number of trees, coverage of woody vegetation, total vegetation cover and level of pollutant emissions were significantly higher in Syrian Woodpecker breeding territories than in the random points. The model best explaining the probability of species occurrence consisted of four parameters: the number of trees, coverage of the total built-up area, total vegetation cover and pollutant emissions. The parameters best explaining a high probability of species occurrence were high number of trees and high pollutant emissions.

Conclusion Air pollution may weaken trees and potentially increase the number of insects inhabiting them, an important component of the Syrian Woodpecker's diet. However, the negative impact of air pollution on birds may impact on their condition. Urban habitats could, therefore, act as an ecological trap for this species.     

Do bridge construction activities influence birds using the River Mersey,in northwest England?

Capsule: Bridge construction works over a major river did not adversely impact the overall wintering and breeding bird assemblage, although contrasting species-specific trends in breeding number were noted.

Aims: To determine whether the wintering water-bird and breeding bird assemblage using the River Mersey, in northwest England, was influenced by bridge construction works.

Methods: Common bird census and wintering bird surveys were carried out before and during construction of the Mersey Gateway bridge. The species abundance and richness of wintering water-birds, assigned to a group (‘Waterfowl’, ‘Wader’, ‘Wetland species’ and ‘Gull’) and breeding birds, assigned to a group (‘Salt marsh species’, ‘Waterfowl species’, ‘Wader species’, ‘Warbler species’ and ‘Other passerine species’), were separately compared before and during the works.

Results: We found little evidence that the bridge construction displaced wintering water-birds and instead found that for most bird groups, their assemblages were higher during the construction works, compared to before. Similarly, we found no evidence that the bridge construction reduced breeding density of any bird group. Contrasting species-specific trends were, however, noted, with some species only recorded breeding during the construction works, and other species recorded in higher breeding densities before the works.

Conclusion: The inadvertent increase in the mud-flats around the bridge structure since the works resulted in more mud-flat being exposed for a greater amount of time, and thus increased the foraging opportunities for many birds. This may at least partly explain why overall wintering bird abundance and richness increased since the works began. Although most breeding birds either increased or remained stable, in number since the works, a small number reduced. This may have been a consequence of the removal of suitable nesting habitat. This study provides a preliminary assessment of the influence of bridge construction on birds.     

Habitat associations of British breeding farmland birds

Capsule Territory distribution for ten species was most strongly positively influenced by the presence of hedges and woodland edge.

Aims To describe and rank the importance of different habitat predictors on the distribution of bird territories.

Methods We derived territory maps for ten bird species across 25 sites on English lowland farmland in 2002. We related habitat predictors to the distribution of these species using information theoretic methods.

Results Habitat predictors were ranked as follows (numbers in parentheses indicate the number of species with a strong effect): hedge presence (8), boundary height (7), woodland edge (6), tree presence in boundary (4), brassica (mainly oil seed rape) (3), within‐field vegetation height (3), boundary strip (3), boundary width (3), tilled fields (3), winter set‐aside (2), ditch (1), winter stubble fields (1).

Conclusions Non‐cropped habitats had the most consistent positive effects across all ten species, with crop types and their margins exerting smaller effects.     

Exploring the effects of truncated,pooled and sexed data in distance sampling estimation of breeding bird abundance

Capsule Field methods used by the UK's Breeding Bird Survey (BBS) provide a practical approach to estimating breeding bird abundance and this paper discusses how they might be adapted to increase accuracy.

Aims Using Salisbury Plain as a case study, examine the use of distance sampling to produce estimates of breeding bird abundance.

Methods During 2005, 157 1 km squares were surveyed on Salisbury Plain using the UK’s BBS methods, with the exception that all birds were sexed whenever possible and allocated to one of five distance bands (0–10 m/10–25 m/25–100 m/100–250 m/ >250 m). Data were modelled to investigate the effects of truncated, pooled and sexed bird data in estimating breeding populations.

Results Pooling of the inner distance bands made little difference to estimates, while truncation over 100 m affected them by up to 25%, generally leading to an increase in density and the width of the confidence limits. The national BBS does not distinguish between bird sexes and therefore any density estimate produced relates to individuals. Our analysis suggests that halving the number of individuals (assuming an equal sex ratio) could lead to significant underestimation of population. This is particularly the case for species where there are differences in detectability between sexes or skewed sex ratios. In such cases, the density of males may represent a more accurate assessment of the population.

Conclusions Bird surveys incorporate distance sampling because it assesses changes in detection probability and this paper suggests how these methods might be adapted to increase accuracy. Changes include increasing both survey effort (e.g. number of visits or distance covered), the number of distance bands, and recording sexed bird data. Numbers of pairs or territories could then be estimated by either halving the density of individuals, or using the density of males, whichever is highest. These suggestions have not been tested against a known population and further work is desirable.     

Value of a broken umbrella: abandoned nest sites of the black stork (Ciconia nigra) host rich biodiversity

Protecting habitats for charismatic vertebrates can provide an ‘umbrella’ for less conspicuous organisms, especially when these are threatened by the same processes. However, such a conservation scheme is vulnerable to the extirpation of the focal species. We studied wider biodiversity values in long protected black stork (Ciconia nigra) nest sites, which were abandoned by the bird and thus legally subject to de-listing. In 20 abandoned nest sites in Estonia, we (i) mapped breeding birds within 600 m from the stork nest, and (ii) carried out time-limited surveys of lichens, polypore fungi, vascular plants and bryophytes in 2-ha plots. The breeding bird assemblages (64 species recorded) included 19 red-listed species, and showed no clear aggregation to the immediate surroundings of the stork nest. We recorded 740 plant and fungal species, of which 134 (18%) were of conservation concern (nationally protected, red-listed or extremely rare). Across the 2-ha plots, the numbers of the species of conservation concern varied more than three-fold (maximum 42 species), being affected notably by dead wood accumulation over time and presence of nemoral broad-leaved trees. The results demonstrate that many abandoned nest sites of the black stork have broader biodiversity significance, both due to the bird’s habitat requirements and the natural development during the protection. Expanding the umbrella function to sites abandoned by a focal species, but intact from anthropogenic degradation, can thus be a cost-effective conservation approach due to its low additional administrative burden. In most jurisdictions, the assessment procedure for such situations should be formalized, however.

     

Breeding of Tawny Owls Strix aluco in rural and urban habitats in southern Finland

Capsule The annual average breeding frequency, clutch size, offspring production and chick survival of Tawny Owls did not differ between rural and urban nesting territories.

Aims To determine whether the general intensity of human habitation in the territory affects breeding.

Methods Clutch size, offspring production, breeding frequency and prey abundance were determined from 210 rural and 60 urban nesting territories monitored between 1994 and 2006.

Results Fluctuations in the annual average clutch size did not differ between habitats. Clutch size and offspring production paralleled each other in rural habitats but not in urban ones. Annual average clutch size followed the regional spring abundance of small mammals in rural Tawny Owls but not in urban ones. The breeding frequency was higher after mild winters in rural environments but not in urban ones.

Conclusion Over an extended time period, rural and urban habitats were largely of equal quality. In urban environments, however, owls seem to be less affected by the pronounced regional abundance fluctuations of small mammals and weather conditions of the preceding winter that largely govern the breeding of owls elsewhere.     

Winter habitat associations of seed-eating passerines on Scottish farmland

CapsuleExtensive surveys highlighted the importance of weedy fodder brassicas, stubbles and open farmland landscapes to declining birds.

Aims To determine habitat associations of seed-eating passerines on Scottish farmland in winter, and recommend appropriate conservation measures.

Methods Transect surveys were carried out on farmland in 100 1-km squares, containing 2885 fields, across Scotland. Birds, crops, field boundary features and weeds were recorded on three visits. Bird–habitat associations were examined using logistic regression models.

Results Seed-eating passerines were highly concentrated: 100 fields (1.4% of the area surveyed) held half the total count. The highest field densities of birds were recorded in fodder brassica crops and stubbles, averaging 12 and 3.3 birds/ha respectively. In fodder crops, weedier fields held many more birds. Multiple regression analysis emphasized the importance of fodder crops, stubbles and farmyards and suggested that declining bird species preferred more open landscapes. Several bird species were associated with fields containing very weedy patches, and/or weeds of the family Chenopodiaceae. These features were commonest in fodder brassicas. Many of the crop types found to be important as bird habitats in winter have declined in area in Scotland; the availability of seed food in such habitats may also have diminished. Such changes may partly explain recent population declines.

Conclusions Agri-environment measures for seed-eating passerines should support the widespread provision of small areas of seed-rich wintering habitats, such as weedy fodder brassicas and stubbles, especially in open (0–10% woodland and scrub) landscapes in areas with mild winters.     

Abundance of breeding farmland birds on arable and set-aside fields at forest edges

The aim with this study was to investigate whether abundance of farmland birds on fields at forest edges were associated with (I) type of field (young set-aside vs arable fields), (n) the length and structure of the field-forest edge zone, and/or (m) with residual habitats such as habitat islands, ditches, roads etc Twenty-eight farmland bird species (all nesting and/or foraging on open fields) were censused during the breeding season on 48 plots (open fields with adjoining forest edges) in the central parts of Sweden, covering a total area of 595 ha Skylark Alauda arvensis , linnet Carduelis cannabina , whitethroat Sylvia communis and whinchat Saxicola rubetra were found in significantly higher numbers in set-aside-plots than cereal ones However, the most important factor explaining variation in the abundance of most species was the structure of the field-forest ecotone, with the length of shrubby southern deciduous forest edges being the most important factor in 7 of the species Mixed forest edges seemed to be of some importance for the abundance of 3 species, while associations between abundance and length of the other deciduous and coniferous field-forest ecotones only were significant for one species each Skylarks, white wagtails Motaalla alba and whinchats were positively associated to ditches and yellowhammers Emberiza citrinella and linnets were significantly associated to habitat islands The observed preferences for set-asides and shrubby field forest edges are suggested to be results of reduced predation risk and increased food abundance     

Habitat use by three Acrocephalus warblers in an intensively used farmland area: the influence of breeding patch and its surroundings

Habitat use by the sedge warbler Acrocephalus schoenobaenus, reed warbler A. scirpaceus and marsh warbler A. palustris was studied during 1998–2000 in an intensively cultivated agricultural landscape in western Poland. The birds occupied mid-field marsh patches (0.05–9.84 ha) and drainage ditches. Using logistic regression models, habitat preferences for particular patch type, characteristics of breeding patch vegetation and surrounding crop type were investigated. All habitat measurements were done within 50-m radius circles around breeding territory centers. The most important factors that influenced species distributions were the proportions of particular habitats within patch area and the type of surrounding crops. Sedge and reed warblers preferred areas with a high proportion of reeds and meadows. Marsh warblers avoided emergents other than reeds, and favored herbaceous vegetation and bushes. Open water areas positively affected reed warbler distribution. Crop type had a significant influence on within-year territory distribution and changes in between-year occupation pattern. In general, birds preferred to settle near fields of oil seed rape, while root crops and spring cereals were avoided. Breeding patch type and structure of reedbeds had relatively little influence on the distribution of breeding territories. The results obtained show that the surrounding landscape may significantly influence habitat use of species breeding in habitat islands in farmland. Depending on their structure and food resources, crops might have different value to the birds.     

Identifying important bird habitats in a sub-arctic area undergoing rapid land-use change

Capsule Habitats in lowland South Iceland sustain bird populations of international importance, with highest densities in wet habitats.

Aims In areas important for biodiversity there is an urgent need to assess large-scale variation in the biodiversity value of habitats to inform management. We carried out a large-scale survey to assess the conservation value of sub-arctic, Icelandic bird habitats.

Methods Bird counts were carried out on 200 transects in the five most common vegetated habitat types in South Iceland. Based on these counts, breeding bird density and diversity were compared between habitats and total population sizes of common species in these habitats were calculated.

Results Overall, eight species (seven waders and Meadow Pipit) composed over 95% of all birds counted. The combined density of those species exceeded 275 birds/km² in all habitats. The two wettest habitat types had the highest density of birds.

Conclusion Wet habitats in lowland South Iceland held particularly high densities of breeding birds, notably waders, which constitute populations of international importance. Wet habitat types are generally of higher value for more species, than dryer ones.     

Winter use of wild bird cover crops by passerines on farmland in northeast England

Capsule Declining farmland species were more abundant in these crops which can be matched to the birds' requirements.

Aims To assess the use of seed-bearing crop by a range of bird species in winter.

Methods The study was carried out over three winters at a site in County Durham (England). Eleven bird species were represented, five of which are nationally targeted for conservation action.

Results Bird abundance was significantly higher where wild bird cover crops were present. Kale Brassica napus and Quinoa Chenopodium quinoa were the most used crop species, although cereals and linseed were also used.

Conclusion Simple crop mixtures can be designed to meet the needs of farmland bird communities. Our results suggest that seed production may be limited within government agri-environment schemes by restrictions on fertilizer use. The agronomy of seed-bearing crops for birds requires further investigation.     

Habitat and diet variability of two coexisting tit species in central European forests

Capsule: Blue Tits and Great Tits occupied different habitats within forests in Central Europe but their nestlings shared a similar diet.

Aims: To quantify the differences in offspring diet and territory habitat between Great Tits Parus major and Eurasian Blue Tits Cyanistes caeruleus in two European forests, and to test whether the ecological niches of the two species overlap.

Methods: Research was conducted on Great Tits and Eurasian Blue Tits, breeding in nest boxes in two forests near Kraków, Poland, during years: 2009, 2011 and 2012. Nine days after hatching, food items were collected from offspring using neck-collars. Habitat parameters surrounding each nest box were quantified.

Results: Great Tit territories were in old Oak-Hornbeam forest, whereas Blue Tits often nested in mixed forest. There were no significant differences between the two bird species in the variation in their caterpillar diets for which both species were highly variable. Great Tits collected more caterpillars of Noctuidae per nest than did Blue Tits in 2009 and 2011 in Niepo?omice Forest; Blue Tits collected more Tortricidae in 2011 and more spiders every year. In Krzyszkowice Forest in 2012, tits fed their nestlings in different periods and did not differ in the proportion of caterpillars. Habitat affected diet differently in each species.

Conclusion: Although Great Tits and Blue Tits occupied different territories in each forest and year of research, the diets of both species’ nestlings contained similar species of invertebrates. The overlaps of the birds’ environmental needs are specific at a local scale.     

The breeding status of Peregrine Falcons Falco peregrinus in the UK and Isle of Man in 2002

Capsule Numbers of breeding Peregrines are now higher than ever recorded and the recovery of the species within the UK has continued since 1991.

Aims To estimate the current Peregrine breeding population and its changes over time by recording breeding evidence and nesting range occupancy at known and potential Peregrine eyries.

Methods Attempts were made to check all potentially suitable Peregrine nesting locations for signs of Peregrine occupation. Causes of nest failure were documented where possible. Estimates of occupancy of nesting ranges that could not be visited were based on the proportion of ranges occupied from visited nesting ranges in the same region.

Results In 2002, 1530 Peregrine nesting ranges were estimated to be occupied, comprising 1456 pairs or single birds confirmed as in occupation, and 74 estimated extras (5% of total occupied nesting ranges) that were not visited. Including non‐breeding singletons holding nesting ranges, the UK and Isle of Man figure of ‘Ratcliffe pairs’ was 1437. This showed a 12% increase from 1991 and a 64% increase from the 1930s. However, there were geographical differences in change, with rapid expansion in the south of England tempered by declines in northern Scotland.

Conclusions The primary reasons for the population increase is primarily because of range expansion driven by a plentiful food supply and perhaps an increased tolerance of human disturbance. In areas where the species has declined, various factors (separately or in combination) including reduced food supply and persecution are likely to be involved.     

Identifying factors that influence bird richness and abundance on farms

Capsule: Farmers can influence species richness and abundance of typical farmland birds positively, even on rather small farms (20–50?ha) within intensively farmed areas.

Aims: To assess the impact of farm settings, farm characteristics and heterogeneity of habitats on bird species richness and abundance, and to indicate which actions and measures farmers can take to promote farmland birds at a farm level.

Methods: Farmland bird species richness and abundance were modelled as a function of farm settings, farm characteristics and semi-natural habitats on 133 farms. The data were analysed at the farm scale, as this is the ‘operating range’ of a farmer, but also at the territory scale, which represents the range birds (mainly passerines) use during the breeding season. Additionally, effects of the farm variables on species abundance/occurrence were investigated for nine widespread species.

Results: Farmland bird species abundance (but not richness) was elevated on organic compared to non-organic farms. Farmland bird species richness and abundance increased with decreasing mean field size. Crop diversity had positive effects on five species at the territory scale. Several semi-natural habitats, especially hedgerows, were associated with higher bird species richness and abundance at both farm and territory scales. Settlement revealed rather negative effects at the farm scale, but several positive relations at the territory scale.

Conclusion: Birds, especially passerines, are restricted to a small area during the breeding season, and so even small farms can contribute to their protection by growing diverse crops, reducing field size and managing a diversity of semi-natural, uncropped habitats. These measures should ideally be accessible within the relatively small scale of a bird territory.     

Foraging Blue Tits Parus caeruleus may trade-off calorific value of food items and distance from cover

Capsule Flowers of an invasive plant species are more visited by native birds than flowers of ornithophilous endemic plants.

Aims To describe the bird guild and its behaviour visiting the century plant Agave americana in an insular environment and to determine which factors are affecting visitation rates.

Methods We noted number and species of birds visiting inflorescences on Tenerife, Canary Islands. We used multimodel inference of generalized linear models to analyse the factors affecting the number of visits and the visitor species richness.

Results Eighty-one per cent of inflorescences were visited by eight native bird species. All species fed on nectar and only the Atlantic Canary fed also on pollen. Foraging behaviour varied among species. Visitation rate increased with density and diversity of birds and flower characteristics and decreased through the day. The number of species visiting the inflorescences increased with diversity and density of birds in the surroundings and decreased through the day.

Conclusion The native bird community uses the invasive century plant as a feeding resource at a higher rate than it uses endemic ornithophilous plants. This could have negative effects for the pollination of endemic plants, but positive effects for birds.     

Habitat associations of Eurasian Skylarks Alauda arvensis breeding on Irish farmland and implications for agri-environment planning

Capsule Skylarks breeding in Ireland prefer extensive grassland habitats and almost completely avoid tillage habitats.

Aims To describe the distribution and habitat use of breeding Skylarks in Ireland, particularly in lowland agricultural habitats, and to use this information to inform conservation measures for this species.

Methods Countryside Bird Survey (CBS) and Farmland Bird Project (FBP) data were examined to determine large-scale (national) distribution and habitat selection, in addition to smaller-scale (farm- and field-level) habitat use. The CBS is a national breeding bird monitoring scheme involving 397 1-km squares. The FBP collected detailed bird and habitat data from 122 farms.

Results CBS and FBP data both showed significant regional differences in breeding Skylark densities, with the highest relative abundances in the northwest and west. Dry grassland/grass moor habitats supported the highest densities of breeding Skylarks in the CBS, which were significantly higher than in improved grassland or tillage. At the farm-level, Skylark numbers were positively related to wetland habitats but negatively associated with trees in field boundaries, dense ground vegetation and overall density of farm boundaries. At the field-scale, larger fields and unimproved grasslands were preferred.

Conclusion Agri-environment measures tailored to region-specific requirements and to the relatively local habitat preferences of target species are required if population declines of species of conservation concern, including Skylarks, are to be reversed.     

Composition and diversity of bird communities in Swedish farmland–forest mosaic landscapes

CapsuleThe amount of forest (at local and landscape scales) and occurrence of residual habitats at the local scale are shown to be the major factors influencing bird community composition in farmland–forest landscapes in central Sweden.

Aims To investigate the importance of local habitat and landscape structure for breeding birds in farmland–forest landscapes in central Sweden.

Methods Breeding birds were censused at 292 points. A detailed habitat mapping was made within 300 m of the points. Within a 300–600 m radius only two major habitats (forests and arable fields) were identified.

Results Cluster analyses of bird communities identified three site types that also differed in habitat composition: (i) partially forested sites in forested landscapes; (ii) heterogeneous sites with residual habitats in mosaic landscapes; and (iii) field-dominated farmland sites in open landscapes. A total of 19 of 25 farmland bird species (restricted to farmland or using both farmland and forest) had the highest abundance in farmland sites with mosaics of forest and farmland, while only six farmland species had the highest abundance in field-dominated sites. The bird community changed from being dominated by farmland species to being dominated by forest species (common in forest landscapes without farmland) at small proportions (10–20%) of forest at the local scale. A major difference in habitat composition between heterogeneous and field-dominated sites was the occurrence of different residual habitats (e.g. shrubby areas and seminatural grasslands). These habitats seemed to influence bird community composition more than land-use, despite covering <10% of the area. Seminatural grasslands were important for bird community composition and species-richness, but grazing seemed to be less important. Among different land-use types, cereal crops were the least preferred fields. Set-asides with tall vegetation and short rotation coppices were positively associated with species-richness of farmland birds.

Conclusion In general, the composition of the landscape was important for bird community composition, although amount and distribution of forests, occurrence of residual habitats and land-use of fields at the local scale had the strongest influence on bird community composition. The possible implications of these patterns for managing farmland–forest landscapes are discussed.