Effect of Sky Lark plots and additional tramlines on territory densities of the Sky Lark Alauda arvensis in an intensively managed agricultural landscape

Capsule: Sky Lark plots and additional tramlines increase Sky Lark Alauda arvensis territory densities in winter crops.

Aims: To analyse the effects on Sky Lark territory density of Sky Lark plots and additional tramlines in winter cereals and oilseed rape.

Methods: We mapped Sky Lark territories on fields with Sky Lark plots or additional tramlines as well as on adjacent control sites, from 2010 to 2013 in Saxony, Germany, where agricultural land use is intensive, dominated by winter-sown crops and takes place in large fields.

Results: In test sites with Sky Lark plots, 5.6 and 3.1 territories per 10?ha were found in the early (April/May) and late periods (June/July) respectively, compared to 3.3 and 1.4 territories per 10?ha in control sites. Sky Lark territory densities in fields with additional tramlines were 1.6 times higher in the early period (4.2 versus 2.6 territories per 10?ha) and 2.2 times higher in the late period (3.6 versus 1.6 territories per 10?ha).

Conclusion: Sky Lark plots and additional tramlines improve large fields for the Sky Lark and have the potential to increase the Sky Lark population.     

Assessing breeding success in common woodland birds using a novel method

Capsule Repeated counts of fledged broods can provide a useful estimate of breeding success for most common woodland birds.

Aims To assess the efficacy of comparing fledged-brood survey data with territory mapping using simple mark–recapture analysis techniques to provide an estimate of breeding success for common woodland birds that does not involve finding nests.

Methods Three observers undertook territory mapping surveys of adults, followed by counts of fledged broods four times a week during May–July 2007 in two 15 ha woods each, both in southern England. Using known fledging to maturity periods, these counts were used to calculate daily detection probabilities for broods of ubiquitous species. These enabled fledged brood territory occupancy probabilities (i.e. brood to territory ratios) to be estimated that take account of the possibility that broods were present but missed by surveys.

Results Of the 19 species found in all six woods, mean daily detection probability estimates for fledged broods of 17 species ranged from 0.17 to 0.50 with significant variation between woods for 12 species, but within region/observer for four species. The mean probability of detecting a brood at least once was over 75% using four visits per week and over 50% using two visits. Only for Great Spotted Woodpeckers Dendrocopos major and Garden Warblers Sylvia borin was the fledging period too short and the daily detection probability too low to provide a reasonable estimate of the territory occupancy probability.

Conclusion Daily detection probabilities for fledged broods of most common woodland birds were sufficiently high to enable useable estimates of fledged-brood territory occupancy probabilities to be made based on a survey programme involving two or three visits per week between late May and the end June. The method used may have application as a means of providing a relatively easily derived productivity index for woodland bird monitoring programmes or for research studies.     

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.     

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.     

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.     

Effect of boundary vegetation and landscape features on diversity and abundance of breeding bird communities of abandoned crop fields in southwest Poland

Capsule The number of territories of birds nesting in abandoned crop fields was most strongly positively affected by field size and the presence of a railway embankment.

Aims To assess the influence of different habitat features on the number of birds nesting in abandoned crop fields.

Methods A breeding bird survey was conducted in spring 2002 using the mapping method on 67 abandoned fields located in southwest Poland. The associations between eight habitat features of the abandoned fields and the numbers of particular species and groups of birds, as well as the whole bird community, were tested with a glm.

Results The models for eight individual species revealed the following ranking, in terms of number of species affected by each predictor: area of fields (5), railway embankments and adjacent crop fields (4), roads and power lines (2), and number of trees (1). The number of the most numerous group of birds (species nesting on the ground, 79% of all breeding pairs) was affected by four predictors: field area, railway embankments, power lines and roads. The number of all birds was affected by six predictors: field area, railway embankments, power lines, roads, ditches and trees.

Conclusion Limited effects of trees, hedgerows and ditches on the number of territories suggest that abandoned fields without woody vegetation are attractive breeding habitats for many farmland birds. One of the effective ways of protecting farmland bird species in central Europe may be the preservation of abandoned fields in their present form, especially those <5 ha in area.     

Population estimates of the Endangered Hawaiʻi ʻĀkepa (Loxops coccineus) in different habitats on windward Mauna Loa

Endangered Hawai‘i ?ākepas (Loxops coccineus) are endemic to Hawai‘i island, where they occur in five spatially distinct populations. Data concerning the status and population trends of these unique Hawaiian honeycreepers are crucial for assessing the effectiveness of recovery and management actions. In 2016, we used point‐transect distance sampling to estimate the abundance of Hawai‘i ?ākepas in portions of Hawai‘i Volcanoes National Park (HAVO) and the Ka?ū Forest Reserve (KFR) on Mauna Loa volcano. We then compiled the survey data from four other populations to provide a global population estimate. In our HAVO and KFR study area, we mapped habitat classes to determine the population densities in each habitat. Densities were highest (1.03 birds/ha) in open‐canopy montane ?ōhi?a (Metrosideros polymorpha) woodland. In contrast, densities of the largest ?ākepa population on Mauna Kea volcano were highest in closed‐canopy ?ōhi?a and koa (Acacia koa) forest where the species is dependent on nest cavities in tall (> 15 m), large (> 50‐cm diameter at breast height) trees. We surveyed potential nesting habitat in HAVO and KFR and found only one cavity in the short‐stature montane ?ōhi?a woodland and five cavities in the tall‐stature forest. Differences in densities between the Mauna Kea and Mauna Loa populations suggest that Hawai‘i ?ākepas may exhibit different foraging and nesting behaviors in the two habitats. The estimated overall population density in the HAVO and KFR study area was 0.52 birds/ha, which equates to 3663 (95% CI 1725–6961) birds in their 11,377‐ha population range. We calculated a global population of 16,428 (95% CI 10,065–25,198) birds, which is similar to an estimate of 13,892 (95% CI 10,315–17,469) birds made in 1986. Our results suggest that populations are stable to increasing in the two largest populations, but the three other populations are smaller (range = 77–1443 birds) and trends for those populations are unknown.     

The effect of woodland area on avian community composition in a fragmented southern UK landscape and associated management recommendations

ABSTRACT

Capsule: Smaller woodlands not only support fewer species but also show different avian community composition due to loss of woodland interior and an increase in edge habitat.

Aims: To use observed community composition changes, rather than traditional total species richness-area relationships, to make area-specific management recommendations for optimizing woodland habitat for avian communities in fragmented landscapes.

Methods: 17 woodlands were selected in Oxfordshire, UK, with areas between 0.2 and 120 ha. Three dawn area searches were conducted in each woodland between 1st April and 28th May 2016 to record encounter rates for each species. The impact of internal habitat variation on woodland comparability was assessed using habitat surveys.

Results: Woodlands with area less than 3.6 ha showed a significant positive relationship between total avian species richness and woodland area. Woodlands with area over 3.6 ha were all consistent with a mean (± se) total richness of 25.4?±?0.6 species, however the number of woodland specialists continued to increase with woodland area. Woodland generalists dominated the total encounter rate across the area range, however the fractional contribution of woodland specialists showed a significant positive correlation with woodland area, while the fractional contribution of non-woodland species significantly decreased. Non-woodland species numbers peaked in mid-sized woodlands with enhanced habitat heterogeneity.

Conclusions: Community composition analysis enabled more targeted recommendations than total species richness analysis, specifically: large woodlands (over 25?ha) in southern UK should focus conservation efforts on providing the specific internal habitats required by woodland specialists; medium-sized woodlands (between approximately 4 and 25?ha) should focus on promoting internal habitat variety, which can benefit both woodland species and non-woodland species of conservation concern in the surrounding landscape; small woodlands (under 4?ha) should focus on providing nesting opportunities for non-woodland species and on improving connectivity to maximize habitat for woodland generalists and facilitate movement of woodland specialists.     

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.     

Group size and composition are correlated with population density in the group-territorial blue korhaan (Eupodotis caerulescens)

The blue korhaan (Eupodotis caerulescens) is a poorly known bird endemic to the grasslands of southern Africa. It lives in permanently held territories defended by groups of individuals. During 2001 and 2002, 61 transects were carried out along 544 km of roads in Mpumalanga province (South Africa) to collect data on territory density and group size and composition in this species. Habitat, altitude and topography variables were measured to evaluate their influence on population density. Average territory density was 1.56 ± 0.19 (SE) territories per 5 km and maximum densities attained 5.6 territories per 5 km. Within its area of occurrence, higher densities occurred along transects with larger proportion of short grassland (either burned or grazed) and flatter topography. Mean group size was 3.03 ± 0.11 birds (n = 119), with 87.4% of the groups having one to four birds. Most common group composition was one male and one female‐like (34.5%), followed by two male and one female‐like (17.2%) and two male and two female‐like (13.8%). Single individuals were always males, and the male/female‐like ratio peaked at a group size of three and decreased thereafter. Mean group size increased with territory density (r_s = 0.42, P = 0.007, n = 41). There were no differences in group size across habitat types. The occurrence of habitat saturation and cooperative breeding in this species could explain the observed results.     

Habitat associations and breeding success of the Chaffinch Fringilla coelebs

Capsule Population trends for Chaffinch on farmland are unlikely to be explained by their preference for non-crop habitats alone.

Aims To investigate the importance of non-cropped habitats for Chaffinch territory distribution, breeding success and foraging habitat selection in Scottish farmland.

Methods Territory distribution, nesting success and foraging behaviour of adults feeding chicks at the nest were recorded and related to habitat composition on two Scottish lowland farms.

Results Higher Chaffinch territory densities were associated with the presence of hedgerows, trees and grass leys, whereas lower densities were found adjacent to winter Barley. Nests were predominantly located in conifer trees and hedgerows. Trees were selected preferentially for foraging. Poor breeding success was detected in nests associated with cereal crops and wider field margins. Farmland in Scotland supported much lower territory densities than in England.

Conclusion Chaffinches rely predominantly on non-cropped habitats for territory establishment, nesting and foraging habitats. Differences in habitat composition between England and Scotland may explain differences in territory densities. However, the Chaffinch's preference for non-crop habitats is unlikely to explain its population trends alone. Immigration from other habitats and/or a decrease in inter-specific competition for resources may also have contributed to the increase in the national farmland Chaffinch population.     

Postfledging habitat selection of juvenile middle spotted woodpeckers: a multi-scale approach

Despite its relevance for the persistence of populations, the ecological mechanisms underlying habitat use decisions of juvenile birds are poorly understood. We examined postfledging habitat selection of radio-tracked juvenile middle spotted woodpeckers Dendrocopos medius at multiple hierarchically-nested spatial scales in NW Spain. At the landscape and home range scales, old oak forest was the most used and selected habitat, young oak forests and pine plantations were avoided, and riverside forests were used as available. At a lower scale, birds selected larger diameter trees for foraging. Home ranges had higher densities of large deciduous trees (mainly oaks Quercus spp., but also poplars Populus spp. and willows Salix spp. >22  cm and >33  cm DBH) selected for foraging by juveniles than non-used areas. These results suggest that foraging conditions may drive, at least partly, habitat use decisions by juvenile birds. We also discuss the potential influence of intraspecific competition, the search for a future breeding territory in the early postfledging period and predation avoidance on habitat use decisions by juvenile birds. Contrary to previous studies on migrant forest birds, postfledging juvenile woodpeckers selected the same habitat as for the breeding adults (i.e. old oak forest), indicating that migrant and resident specialist avian species may require different conservation actions. Conservation strategies of woodpecker populations should consider the protection of old oak forests with high densities of large trees to provide suitable habitat to breeding adults and postfledging juveniles. The habitat improvement for this indicator and umbrella species would also favour other organisms that depend on characteristics of old-growth oak forests.     

Structure and utilisation of the early oakwoods

Summary

The paper explores whether any of the familar range of current oakwood structures in the west of Scotland can be used as realistic analogues for the prehistoric oakwoods. Even for the early historic oak woodlands we have no detailed knowledge of previous structures and composition.

The structure of today's oak woodlands, and to some extent even the legacy of species within that habitat, are a type of biocultural heritage. The oak woodlands that we see today are the product of a long interaction: between man's use of the woodlands and the ongoing natural processes within the oakwood ecosystem.

Grazing animals, especially domestic cattle and red deer, have played a key part in determining oak woodland structure in historic times, and perhaps have a future role too. Are there lessons to be drawn from knowledge of past management and regeneration of oak which are relevant to the present day, when most managers are finding it difficult to regenerate oak woodlands on any scale?

Understanding past utilisation and structure of oak woodlands is valuable mainly as a guide to future management. Indeed one might ask which models and structures are relevant for managing an oak woodland resource now regarded mainly as a wildlife habitat?     

Effects of variation in breeding habitat on Ring Ouzel Turdus torquatus productivity and chick condition

Capsule Territory habitat influenced clutch size and within-brood variability of chick condition in Ring Ouzel Turdus torquatus.

Aims To assess the relationship of Ring Ouzel productivity and chick condition with territory habitat, in order to improve understanding of the importance of breeding habitat for population change.

Methods Productivity and chick condition were recorded for Ring Ouzel pairs in a small area of upland Scotland and these were related to vegetation and physical characteristics for all breeding territories using General Linear Models.

Results We found that clutch size and variability of chick condition varied with Ring Ouzel breeding habitat.

Clutch size was related to variation in territory habitat, broadly increasing with territory grass, sedge and rush cover. The best-supported models for within-brood variability of chick condition included fledgling number and territory habitat, with chick condition broadly becoming more variable with territory bracken cover. Relationships between breeding habitat and brood size, fledgling number or mean chick condition were not supported by the data, although statistical power was low.

Conclusion Breeding habitat may be a driver of demographic rates of Ring Ouzel, with the effects of poor habitat being passed on indirectly through chick condition. The population-level importance of these effects is unclear, but this study highlights a possible link between recent observed habitat changes and first-year survival, the demographic parameter contributing most to variation in population growth rate in UK Ring Ouzels.     

Cooperative nest‐defence behaviour and territory quality in a resident and socially monogamous passerine

Nest defence is a fundamental aspect of parental care in secondary cavity‐nesting birds, and predation or competition for nesting sites can involve different defensive behaviours. Because habitat quality determines breeding success, we were interested in whether breeding pairs of the Eurasian nuthatch, Sitta europaea, established in more favourable environment also manifest higher probability of cooperative behaviour during their nest‐site defence. To explore this relationship, we quantified behavioural displays of both parents and analysed activity budget ethogram data from simulated territorial intrusions performed in the chick‐feeding phase with one conspecific and two different heterospecific stimuli (dummies of nuthatch, starling and woodpecker). We found that paired individuals shared their roles during nest‐site defence to a considerable extent. Males had a significantly higher number of attacks on intruders than females, and females performed more threat displays and controls of the brood than males. Multinomial analysis of the cooperative behaviour suggested that pairs in a high‐quality territory had higher probability of reciprocal substitution of different roles towards a balance between attacks, threat displays and nest controls. Contrary to this, pairs in a low‐quality territory had less likely pairwise combinations of simultaneous behavioural states that are associated with effective nest‐site defence. The difference in response probability according to territory quality was, however, highly variable in view of the stimulus that was used in simulated territorial intrusion. Because individual roles and the complex behavioural repertoire of pairs altered in response to territory quality and potential nest‐site competitor or brood predator, our results suggest that the cooperative nest‐defence behaviour could be linked to the breeding success of this year‐round territorial species living in a heterogeneous forest habitat.     

Spatial Distribution of Oak Mistletoe as It Relates to Habits of Oak Woodland Frugivores

This study addresses the underlying spatial distribution of oak mistletoe, Phoradendron villosum, a hemi-parasitic plant that provides a continuous supply of berries for frugivorous birds overwintering the oak savanna habitat of California''s outer coast range. As the winter community of birds consuming oak mistletoe varies from group-living territorial species to birds that roam in flocks, we asked if mistletoe volume was spatially autocorrelated at the scale of persistent territories or whether the patterns predicted by long-term territory use by western bluebirds are overcome by seed dispersal by more mobile bird species. The abundance of mistletoe was mapped on trees within a 700 ha study site in Carmel Valley, California. Spatial autocorrelation of mistletoe volume was analyzed using the variogram method and spatial distribution of oak mistletoe trees was analyzed using Ripley''s K and O-ring statistics. On a separate set of 45 trees, mistletoe volume was highly correlated with the volume of female, fruit-bearing plants, indicating that overall mistletoe volume is a good predictor of fruit availability. Variogram analysis showed that mistletoe volume was spatially autocorrelated up to approximately 250 m, a distance consistent with persistent territoriality of western bluebirds and philopatry of sons, which often breed next door to their parents and are more likely to remain home when their parents have abundant mistletoe. Using Ripley''s K and O-ring analyses, we showed that mistletoe trees were aggregated for distances up to 558 m, but for distances between 558 to 724 m the O-ring analysis deviated from Ripley''s K in showing repulsion rather than aggregation. While trees with mistletoe were aggregated at larger distances, mistletoe was spatially correlated at a smaller distance, consistent with what is expected based on persistent group territoriality of western bluebirds in winter and the extreme philopatry of their sons.     

The importance of key marginal habitat features for birds in farmland: an assessment of habitat preferences of Red-backed Shrikes Lanius collurio in the Italian Alps

Capsule Key marginal habitat features maintain their importance even when they occur at very low density.

Aims To assess the importance for breeding birds of key habitat elements, such as isolated shrubs, hedgerows and untilled vegetation patches.

Methods We investigated the habitat preferences of Red-backed Shrikes Lanius collurio in an agricultural environment where key habitat elements have been dramatically reduced. Based upon data from territorial maps, a fine-scale model of habitat preference was developed and the results of three different analytical approaches (binary logistic regression, multimodel inference and hierarchical partitioning) were compared.

Results The occurrence of Red-backed Shrikes was positively influenced by the extent of non-grazed/mown grassland, isolated bushes and hedgerows, and negatively influenced by woodland cover. The model highlighted the importance of bushes/hedgerows despite their very scarce occurrence in the study areas (overall, average cover only 7.94%). Breeding densities were rather low, but mean densities were slightly higher in pasture zones in which bushes/hedgerows availability was higher than elsewhere.

Conclusion These results confirm the crucial importance of these key marginal elements even in depauperated farmland landscapes. Maintaining and increasing their availability is a key management option for the encouragement of breeding Red-backed Shrikes, and potentially for other species which are also declining in these farmland habitats.     

Territorial behavior of Western Sandpipers on their nonbreeding grounds: effect of sex and foraging interference

ABSTRACT Nonbreeding shorebirds often alternate social structure between anonymous flocks and territorial behavior in response to different environmental factors. To evaluate specific drivers for one species, we studied the spacing behavior of wintering Western Sandpipers (Calidris mauri) at Bahía Santa María, northwestern Mexico, using behavioral observations. The density, population structure, and territorial behavior of Western Sandpipers differed among three habitat types. Cattail marshes supported 110 birds per ha, 49% males, and no territorial birds. Mangrove flats supported 288 birds per ha, 58% males, and 5% territorial birds. Brackish flats supported 365 birds per ha, 76% males, and 7% territorial birds. Territories consisted of rectangular strips (5–9 m long, N= 77). Territory length was not related to either bird density or number of territorial birds by plot, but was positively related to nearest bird distance. Aggression rate was inversely related to territory length, suggesting that territory length is set by the costs of defense. Foraging rate was independent of territory length, and prey densities in territories did not differ from those in areas used by nonterritorial birds. Males were more likely to be territorial and had a higher aggression rate than females, suggesting that males, which forage more on surface prey, were more affected by foraging interference. Our results suggest that the territorial behavior of Western Sandpipers in our study was an opportunistic strategy to reduce foraging interference. The variation in spacing behavior we documented provides evidence that interference competition affects the social structure of Western Sandpipers during the nonbreeding season.     

Home range and territory of the Sardinian Warbler Sylvia melanocephala in Mediterranean shrubland

Capsule Singing territories were well separated.

Aims To examine the spatial distribution of Sardinian Warbler males during the breeding period in Mediterranean shrubland and, specifically, their territories, home ranges and spatial overlaps.

Methods We studied a 12-ha plot of Mediterranean shrubland in the 1997 to 1999 breeding seasons. Sardinian Warblers were captured using mist-nets, colour-ringed and their territories mapped. In 1999, seven breeding males were radiotracked in order to map home ranges.

Results The Sardinian Warbler had an average of 7.3 breeding pairs/10 ha in the study plot. The mean territory size was 8779 m² and the mean home range was 22 321 m². A positive relationship was found between the area of the home range and singing territory. Home ranges of males born in 1998 were about half the size of those of the oldest males. Paired males who moved with a female had smaller home ranges than those that were either unpaired or whose mate was incubating. The degree of home range overlap was high with some overlap between neighbouring territories. The estimate of home range area increased by 10% when the information generated by a mapping method was added and the estimated territory area increased by 31% when data generated by radiotracking were added. Transmitters remained attached to birds for an average of 9.63 ± 3.46 days (mean ± se).

Conclusions Singing territories were segregated to a considerable degree. In contrast, the wide overlaps among home ranges was best explained by the presence of food resources that the males exploit at the same time and also by the search for extra-pair copulation in nearby territories. We consider radiotracking in this species to be feasible and valid, with no evidence of negative effects on activity levels, weight or mortality.