Densities and population estimates of breeding Skylarks Alauda arvensis in Britain in 1997

We present the results of a national survey of breeding Skylarks Alauda arvensis in Britain in 1997 carried out by the British Trust for Ornithology (BTO). Numbers of Skylarks and land-use types were recorded by volunteers in 608 1-km squares, selected using random stratification based on the Institute of Terrestrial Ecology's (ITE) Landscape Classification to avoid over- or under-sampling particular habitat types. The results suggest a maximum national population of around 1000 000 pairs. This agrees extremely well with the national population estimate of around 1046 000 pairs derived from the Breeding Bird Survey (BBS) of the BTO/Joint Nature Conservation Committee/Royal Society for the Protection of Birds, and provides further evidence for the robustness of the BBS as a national monitoring scheme for common and widespread species. The figure is approximately half that of the most recent published estimate of 2000 000 in 1988–91. However, the scale of this discrepancy is likely to reflect a bias in Atlas field methodology and site selection since there has been a real decline of approximately 9% since 1990. Arable squares supported the highest densities of Skylarks; 4.6–6.0 pairs per km², and 46–49% of the British breeding population was associated with arable areas. Marginal upland and upland areas supported lower densities but still accounted for approximately 34% of the estimated national breeding population. Differences in density at broad habitat scales were reflected in geographical differences across Britain, with southern and eastern arable regions supporting much higher densities than western and northern ones. At a finer scale, Skylarks occurred at highest densities on, and showed highest habitat preferences for, set-aside and various types of ungrazed grassland. Winter cereal, improved grassland and set-aside held the highest proportion of the Skylark population on farmland in England and Wales; grazed pasture, winter cereals and spring cereals held the highest proportion in Scotland.     

Conservation tillage encourages early breeding by Skylarks Alauda arvensis

Capsule Increased crop residue on the soil surface, left as a result of conservation tillage, encouraged earlier nesting in winter wheat fields.     

Factors affecting the territory distribution of Skylarks Alauda arvensis breeding on lowland farmland

The distribution of breeding Skylarks on lowland farmland was examined on a monthly basis on 13 farms in southern England in 1996. General linear models identified crop type and field area, shape and boundary characteristics as significant independent predictors of Skylark territory numbers in most or all months. Models predicted territory numbers best during the height of the breeding season and less well during territory establishment in March and abandonment in July. From March to May, crop height had no significant effect on Skylark territory distribution, but in June and July it had a highly significant quadratic effect, models suggesting an optimal vegetation height of around 0.55 m in these months. Set-aside held high territory densities, permanent pasture low densities. Spring cereals held higher territory densities than winter cereals. This could best be explained by differences between the two cereal types in crop structure. There was evidence of a positive correlation between crop height diversity and territory density at the farm scale. The results are discussed with reference to the species' recent severe population decline.     

Density and productivity of breeding Skylarks Alauda arvensis in relation to crop type on agricultural lands in western France

Capsule Small field size and the maintenance of set-aside and lucerne are important to ensure high breeding pair densities and productivity.

Aims To investigate the effects of crop types and their attributes on density and productivity of breeding Skylark.

Methods At each of four selected study sites in western France, territory density, vegetation height, vegetation cover and field size was estimated by field and attempts were made to find nests. Crop types included winter and spring cereals, oilseed rape, sunflower, maize, grass, lucerne, set-aside, and bare ground.

Results About 80% of Skylark territories included more than one crop type. Birds preferred small fields and territory density decreased with increasing field size. Density was highest in crops with low vegetation height and cover. Set-aside, lucerne and grass supported highest territory density. Fledging productivity was highest in set-aside and lucerne, and was zero on bare ground. Skylark density decreased throughout the breeding season (-26% in 1999 and -29% in 2000), suggesting an instability in territory distribution or activity in intensive farmland.

Conclusions Farming systems that decrease field size and increase set-aside and lucerne instead of oilseed rape, maize and sunflower will benefit Skylark and other declining farmland species.     

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.     

Population declines and reproductive performance of Skylarks Alauda arvensis in different regions and habitats of the United Kingdom

Declines in the number of breeding Skylarks Alauda aruensis and changes in their reproductive performance were analysed using data from two long-running surveys co-ordinated by the British Trust for Ornithology: the Common Birds Census and the Nest Record Scheme. In the UK, the number of breeding Skylarks declined by approximately 55% between 1975 and 1994. This decline was steepest in agricultural habitats and in regions associated with intensive agriculture. In contrast, Skylark reproductive performance per nest, in terms of clutch size, brood size and post-hatching survival rate of nests, showed a general improvement over time. This improvement was greatest in intensively farmed agricultural habitats. Therefore changes in reproductive performance per nesting attempt were probably not responsible for the decline in numbers. It is inferred that possible causes of the decline of the Skylark are: reductions in the number of breeding attempts per pair per season, reductions in the proportion of birds attempting to breed, and increased mortality outside the breeding season.     

Extra-pair paternity in the Skylark Alauda arvensis

We present the first quantitative data on the genetic breeding system of a lark (Alaudidae), the Skylark Alauda arvensis . Using a set of eight microsatellite loci isolated in a variety of passerine species, we genotyped 171 offspring from 52 broods of Skylark and detected 35 extra-pair offspring (20%), in 14 different broods (27%). All offspring matched their putative mother, so there was no evidence of intraspecific brood parasitism. Previous non-genetic studies had suggested that the species was predominantly socially monogamous, with only rare occurrences of social polygyny and polyandry, although some behaviours, such as mate guarding, did suggest the possibility of extra-pair copulations. The relatively high level of extra-pair paternity in this species is likely to affect the variation in male reproductive success because extra-pair paternity was non-randomly distributed amongst males, with those with shorter wings more likely to be cuckolded.     

Sustainable Arable Farming For an Improved Environment (SAFFIE): managing winter wheat sward structure for Skylarks Alauda arvensis

Research has shown a close correlation between the decline of the UK Skylark Alauda arvensis population and the replacement of spring-sown cereals with winter-sown varieties, in which advanced sward development prevents successful multiple nesting attempts and reduces access for foraging. Widescale reversal of sowing times is unlikely for commercial reasons, so research has recently focused on ways of manipulating the sward structure of winter wheat to prolong access to nest-sites and food. An RSPB pilot study investigated leaving small 'undrilled patches' in otherwise conventionally managed winter wheat crops. This option was later incorporated into a fully replicated experimental design, as part of the Sustainable Arable Farming For an Improved Environment (SAFFIE) project. This large consortium-led project aims to test solutions for improving biodiversity within winter-cereal-dominated rotations. The experiment described here ran over 2002–3, with three field-scale 'treatments' on 15 sites in the first year. The treatments compare (1) conventional winter wheat, (2) winter wheat sown in double-normal width (25 cm) wide-spaced rows (WSR) and (3) winter wheat with two 4-m by 4-m undrilled patches per hectare (UP). Results from the 2002 breeding season showed that undrilled patch treatments supported more breeding Skylarks for longer, most likely by aiding accessibility of food. WSR rows were little used by Skylarks and did not improve the abundance of favoured seed and invertebrate food items over conventional crops. Nesting performance and foraging patterns are discussed with reference to invertebrate food abundance and its accessibility, as determined by sward structure.     

The song of Skylarks Alauda arvensis indicates the deterioration of an acoustic environment resulting from wind farm start‐up

The rapid development of wind energy may have negative effects on bird populations, including collisions with turbines, displacement due to disturbance or habitat loss, indirect effects of reduced breeding success and barrier effects. This challenging conservation issue has attracted a great deal of interest, but the noise generated by turbines has been largely overlooked. Here, we studied acoustic behaviour of Skylarks Alauda arvensis in relation to wind farm start‐up to assess whether a change in song parameters can indicate a deterioration in the acoustic environment. We recorded territorial males displaying close to operating and non‐operating turbines and at a control site without turbines. In the following breeding season, we undertook replications at the same sites, except that the non‐operating turbines were now in operation. We found that Skylarks displaying at the wind farm were affected by wind turbine noise. Males singing close to operating wind turbines sang higher‐frequency songs than males from a control site and those that displayed near non‐operating turbines. In addition, an upward frequency shift in songs was observed when non‐operating turbines started to operate in the consecutive season. We therefore conclude that the frequency shift observed did not result from turbine presence, but from the noise they started to generate. This shows that a change in song parameters may reliably and within a relatively short time indicate a significant deterioration of the acoustic environment as a consequence of wind farm start‐up. This may help conservation biologists to identify species and populations that are particularly susceptible to wind farm noise.     

Observer variation in estimates of Meadow Pipit Anthus pratensis and Skylark Alauda arvensis abundance on moorland

Capsule Counts least susceptible to observer effects were those within 25 m of a transect and the total number along a transect, with observer effects greater for distance sampling.     

Habitat selection,reproduction and predation of wintering lemmings in the Arctic

Snow cover has dramatic effects on the structure and functioning of Arctic ecosystems in winter. In the tundra, the subnivean space is the primary habitat of wintering small mammals and may be critical for their survival and reproduction. We have investigated the effects of snow cover and habitat features on the distributions of collared lemming (Dicrostonyx groenlandicus) and brown lemming (Lemmus trimucronatus) winter nests, as well as on their probabilities of reproduction and predation by stoats (Mustela erminea) and arctic foxes (Vulpes lagopus). We sampled 193 lemming winter nests and measured habitat features at all of these nests and at random sites at two spatial scales. We also monitored overwinter ground temperature at a subsample of nest and random sites. Our results demonstrate that nests were primarily located in areas with high micro-topography heterogeneity, steep slopes, deep snow cover providing thermal protection (reduced daily temperature fluctuations) and a high abundance of mosses. The probability of reproduction increased in collared lemming nests at low elevation and in brown lemming nests with high availability of some graminoid species. The probability of predation by stoats was density dependent and was higher in nests used by collared lemmings. Snow cover did not affect the probability of predation of lemming nests by stoats, but deep snow cover limited predation attempts by arctic foxes. We conclude that snow cover plays a key role in the spatial structure of wintering lemming populations and potentially in their population dynamics in the Arctic.     

Sex-ratio of Skylark Alauda arvensis broods in relation to timing of breeding

Capsule Earlier broods tend to be more male biased than later broods     

Habitat selection of wintering passerines in salt marshes of the German Wadden Sea

The salt marshes of the Wadden Sea are important wintering areas for some species of granivorous passerines, which have declined considerably since the 1960s. We investigated the habitat choice of all wintering passerines in eight study areas in German salt marshes with special consideration of human impact on these habitats. Granivorous species that almost exclusively winter in salt marshes, Shorelark (Eremophila alpestris), Snow Bunting (Plectrophenax nivalis) and Twite (Carduelis flavirostris) were concentrated in the lower salt marsh vegetation and in the driftlines, while all other species preferred the high upper salt marsh communities, although Rock Pipits (Anthus petrosus littoralis) fed in muddy areas along ditches. Shorelarks switched habitat in conditions where seeds were scarce to feed instead on arthropods in upper salt marshes. Intensively sheep-grazed upper salt marshes resemble lower salt marshes in their vegetation and were therefore mainly visited by Shorelarks, Snow Buntings and Twites. In winter, the driftline is preferred by the two former species, while in autumn and spring more birds foraged in the salt marshes. Twites prefer to feed mainly on seeds of Salicornia. Areas with S. europaea are visited mainly in late autumn and early winter, while areas with S. stricta are used throughout the winter because of a steady supply of seeds. Several years after embankment, polders are hardly used any more by the lower salt marsh species as the habitat changes into freshwater marshes. Large embankment projects since the early 1960s have included salt marshes and intertidal flats, and the resultant loss of habitat is responsible for the decline of lower salt marsh species. For other passerine species the effects of reclamation are unknown. The effects of intensified grazing on the wintering populations of Shorelark, Snow Bunting and Twite are still unresolved. Although grazing supports lower salt marsh vegetation, the seed production per plant is much lower there and some important seed producers hardly occur. Since grazing was reduced and embankment projects have been stopped, the salt marsh areas (especially lower salt marshes) have increased and so have the wintering populations of Shorelark, Snow Bunting and Twite. For the other species, the consequences of habitat changes are unknown, although it is suggested that reduced grazing will support them. Reducing the human impact on salt marshes will, in the long run, probably lead to a natural salt marsh with much variety in elevation and in its corresponding vegetation and bird communities. Meanwhile, management by grazing might be required in parts of the salt marshes.     

Site selection by swans wintering in Britain and Ireland; the importance of habitat and geographic location

Monthly surveys of Bewick's Swans Cygnus columbianus bewickii , Whooper Swans Cygnus cygnus and Mute Swans Cygnus olor in Britain and Ireland were made during the 1990–1991 winter to determine factors affecting the swans' selection of feeding sites. Geographic location and habitat both influenced site selection. Whooper Swans occurred in greatest numbers at sites in Scotland, northeastern England and Northern Ireland, whereas Bewick's Swans had a more southerly distribution, reflecting differences in the migratory routes used by these two species. The resident Mute Swans were more widespread, with large flocks occurring in southeastern England and in parts of Scotland. Whooper and Mute Swans were found mainly on permanent inland waters (68% and 61%, respectively), but the majority of Bewick's Swans (60%) were on arable land. The percentage of Bewick's Swan flocks found on permanent inland waters (42%) was higher than that found on arable fields (23%), indicating that the large number recorded on arable land was a result of the birds congregating at a comparatively small number of sites. Overall, less than 15% of Whooper Swans and 3% of Mute Swans were on arable crops during the winter, but the largest flocks were associated with arable land for all three species. Thus, although the occurrence of large flocks at particular arable sites may give an impression that swans feed mainly on farmland, the swans are in fact more widely dispersed. Regional variation in the percentage of juveniles present was recorded for all three species. Changes during the winter in the distribution of juveniles, and of the swans as a whole, are considered in relation to food supply and to migratory routes for the Bewick's and Whooper Swans.     

Occurrence of Eurasian Skylark Alauda arvensis territories in relation to urban area and heterogeneous farmland

Within heterogeneous arable habitats close to urbanized Vienna, Austria, Eurasian Skylarks Alauda arvensis occurred at slightly lower densities than in other mixed farmland areas and at a much lower density than would be found in natural habitats. Territory density was negatively correlated with the occurrence of roads, tracks and high-intensity agricultural cultivation, possibly due to the resulting increased disturbance by humans.     

Regional trends in coastal wintering waders in Britain

Regional trends in the populations of waders wintering largely on estuaries were analysed using data collected by the Wetland Bird Survey between the winters of 1969/70 and 1995/96 to assess whether trends seen at the national level were repeated across regions of Britain. Not all were. Over this period, the numbers of seven of the 12 species studied increased in south and southeast England, but remained static or declined in southwest England and south Wales. Possible explanations for this are proposed. Increases in the number of Curlews Numenius arquata in the north and northeast of Scotland occurred after 1981/82, when the species stopped being a legal quarry species. A rapid decline in Oystercatcher Haematopus ostralegus and Knot Calidris canutus numbers in east England after 1988/89 may be linked to a change in the Wash shell-fisheries. Warmer winters may be making it less essential for birds to winter in the milder west of Britain. In two pairs of regions, northeast and southeast Scotland, and in northwest England/north Wales and northeast England, waders showed similar long-term population changes. The long-term population trends of Oystercatchers in the different regions correlate to those of Ringed Plovers Charadrius hiaticula and Knot, while those of Ringed Plovers are very similar to those of Bar-tailed Godwits Limosa lapponica. In some regions of Britain, the population trends of some species varied a lot between sites, a factor that is an important consideration when determining the size and geographical position of regions for future regional indexing. The present separation of regions based on water catchments and geography has given a useful insight into population changes at a finer spatial scale but it could be improved further with added biological input.