Habitat selection by adult Golden Eagles Aquila chrysaetos during the breeding season and implications for wind farm establishment

Capsule: Global Positioning System (GPS)-tagged adult Golden Eagles Aquila chrysaetos breeding in forests in northern Sweden selected clear-cuts, coniferous forests with lichens and steep slopes during the breeding season but avoided wetlands and mixed forest.

Aims: To investigate the habitat selection patterns of tree-nesting Golden Eagles, and identify how potential conflicts with wind farm development could be minimized.

Methods: The study is based on GPS tracking data from 22 adult eagles. We estimated home range sizes using a biased random bridge approach and habitat selection patterns using resource selection functions following a use-availability design.

Results: Core home range size among adults was variable during the breeding season (5–30?km²). Individual movement extents were variable, but sexes did not significantly differ in their scale of movement. At the landscape scale, individuals selected for clear-cuts and coniferous forest with ground lichens, whereas wetland, water bodies and mixed forest were avoided. Steeper and south facing slopes were selected for, whereas, north facing slopes were avoided.

Conclusions: Potential conflicts between eagles and wind energy establishment can be reduced if wind farms are placed away from steep slopes, minimizing areas that are clear-cut during construction, and locating turbines within dense, young and other less favoured forest habitats.     

Comparative nest habitat characteristics of sympatric White‐tailed Haliaeetus albicilla and Golden Eagles Aquila chrysaetos in western Scotland

Capsule Golden and White‐tailed Eagles selected different habitats for nesting.

Aim To investigate differences in nesting habitat used by sympatrically breeding eagles in western Scotland, following reintroduction of White‐tailed Eagles from 1975 onwards.

Methods Nest‐site locations from national surveys in 2003–05 were entered into a geographical information system (GIS) in order to measure a set of geographic parameters for each nest site. Binary logistic regression with backwards deletion of non‐significant terms was used to derive minimum adequate models at two spatial scales of the likelihood of an eagle nest belonging to one species or the other. We compared changes in occupancy between 1992 and 2003 of Golden Eagle territories inside and outside a GIS model of potential White‐tailed Eagle habitat and according to proximity to White‐tailed Eagle nests.

Results White‐tailed Eagles nested at lower altitudes than Golden Eagles, in more wooded habitats with more open water close by, tending to nest in trees where these were present. There were 3359 km² of potential White‐tailed Eagle nesting habitat within 25 km of existing White‐tailed Eagle nests, containing 54 Golden Eagle territory centres, but we found no difference in change of occupancy for Golden Eagle territories close to White‐tailed Eagles compared with those further away.

Conclusion White‐tailed and Golden Eagles appear to partition nesting habitat in the west of Scotland by altitude. This corresponds with behaviour in western Norway and with the situation described in historical accounts of nest‐sites in western Scotland prior to extinction of White‐tailed Eagles. It is also consistent with recent studies showing little overlap in breeding season diet of Golden and White‐tailed Eagles in western Scotland, and likely partitioning of foraging habitat by altitude. We conclude that the likelihood of competitive exclusion is less than previously suggested.     

Variation in quality of Golden Eagle territories and a management strategy for wind farm projects in northern Sweden

Capsule Breeding productivity of 86 occupied Golden Eagle territories in northern Sweden varied greatly, with the upper quartile of the most productive territories being responsible for 44% of the total productivity of the studied territories. This information is used in a regional strategy for managing wind farm project proposals in areas inhabited by Golden Eagles, whereby the more productive territories are provided with a greater degree of protection than their less productive counterparts.     

Factors influencing the decline of a Bonelli's eagle Hieraaetus fasciatus population in southeastern Spain: demography, habitat or competition?

We investigated three possible causes of territory desertion among Bonelli's eagles Hieraaetus fasciatus in Murcia (southeastern Spain): low demographic parameters, low habitat quality and competition with Golden eagles Aquila chrysaetos. From 1983 to 1997, we surveyed a Bonelli's eagle population. Abandoned and occupied territories were compared to find differences in demographic parameters (flight rate, productivity and mortality) or habitat characteristics. Mortality was significantly higher in abandoned territories. Abandoned territories also had larger areas of forest and extensive agriculture, while occupied territories had more shrublands. Competition with Golden eagles was not a determinant of territorial abandonement but interacted with human persecution of the species. Management implications are discussed.     

Complex effects of habitat loss on Golden Eagles Aquila chrysaetos

The development of commercial forests presents potential threats to large raptors that rely on prey caught in open country. We examined the effect of afforestation of breeding habitat used by a population of Golden Eagles Aquila chrysaetos in Scotland where, over the last 50 years, extensive stands of exotic conifers have been planted. Using data for 31 years on territory occupancy and breeding success, together with spatiotemporally dynamic mapping of forest cover and predicted areas of territory‐use in a Geographical Information System, we examined relationships between forest cover and Eagle ecology at landscape and individual territory scales. Several territories were abandoned during the earliest phases of forest planting, but relatively few were apparently lost to later plantings. Territories with poorer breeding productivity appeared to be more vulnerable to abandonment than territories with better breeding productivity. At the landscape scale, temporal differences in breeding productivity were negatively related to the extent of forest cover, although productivity of individual territories showed no clear relationship with forest cover. Several territories with less than a 5% increase in forest cover experienced reduced productivity; however, territories least constrained by neighbouring pairs of Eagles showed an increase in productivity. Territories experiencing the greatest increases in forest cover showed a greater use of spatially separated nest‐sites by occupying pairs. Hence, pairs that were less constrained by neighbours appeared to compensate for loss of open habitat by shifting their territory‐use, whereas pairs that were more constrained could not compensate for open habitat loss and suffered reduced productivity (and, probably in some cases, abandoned the territory). We suggest that simple guidelines based on the extent and locations of habitat loss are inadequate when predicting effects on large territorial raptors such as Golden Eagles. Consideration should also be given to the ‘quality’ of a territory or occupying pair, as well as the extent to which territory‐use is constrained by neighbouring pairs or other ‘unsuitable habitat’ which may have been affected by previous episodes of open habitat loss.     

Spatial distribution of undulating flight displays of territorial Golden Eagles Aquila chrysaetos in Lewis,Scotland

ABSTRACT

Capsule: The undulating display flight of Golden Eagles Aquila chrysaetos has a territorial function on current consensus: we found that displays could occur at any location within a home range of territorial birds and were not more frequent at the territorial limits (as documented by a previous study) or around the nest site.

Aims: To test the null hypothesis that display activity does not spatially differ within the home range of territorial Golden Eagles.

Methods: We used 1488?h of observations of Golden Eagles between April 2003 and October 2004 (excluding November through January inclusive) on the Isle of Lewis, northwest Scotland involving primarily four territories. Territorial boundaries and nest sites were determined by the behaviour of the focal territorial birds. All flight activity was cast into three spatial categories (near-boundary, near-nest and elsewhere) and the relative occurrence of displays was referenced by overall flight activity recorded in these spatial categories.

Results: We failed to reject the null hypothesis. Prior findings of greater display rate at territory limits were not confirmed. Territorial Golden Eagles apparently displayed wherever they happened to be in their home range. There was no difference in display frequency between successful and failed breeding pairs and, within pairs, between successful and failed breeding seasons. Monthly changes in display activity did not differ from monthly changes in the presence of ‘intruding’ sub-adults. A peak in non-territorial bird activity in August and September was not reflected by a peak in display, however, we could not distinguish recently fledged ‘non-threatening’ birds in this period.

Conclusions: Undulating flight displays of Golden Eagles focused neither on territory boundaries nor the nest. Displays may be spontaneous or occur in response to the detection of potential trespassers or neighbours, however, we could not rule out any role of pair-bonding.     

Floater interference reflects territory quality in the Spanish Imperial Eagle Aquila adalberti: a test of a density‐dependent mechanism

We report on an 11‐year study of floater interference in a population of Spanish Imperial Eagles Aquila adalberti. We analysed changes over the years in the productivity of 15 territories to test predictions of two hypotheses of density‐dependent productivity in relation to the presence of floaters (birds without territories). According to the ‘interference' hypothesis, the frequency of intrusion by floaters increases with density, resulting in a decrease in productivity. Thus, in a high‐density population a negative relationship between floater intrusions and productivity of the territory is expected. In contrast, under the ‘habitat heterogeneity' hypothesis, as density increases a higher proportion of individuals is forced to occupy lower quality habitats. Support of this hypothesis requires that floaters detect differences in quality among territories and preferentially visit the better quality territories. Consequently, a positive relationship between floater intrusions and productivity is expected. Results showed that floaters tended to visit their natal area at the beginning of the breeding season. Among floater Eagles, males made significantly more intrusions per day than did females, but females stayed in the natal population for longer each year than males. Floater intrusions and productivity were highly positively correlated, supporting the ‘habitat heterogeneity' hypothesis; individuals were apparently able to assess the quality of a territory and, at the frequencies observed, their interference with the breeding pair had no obvious negative effect on productivity.     

Nesting habitat overlap between the Common Buzzard Buteo buteo and the Lesser Spotted Eagle Clanga pomarina for conservation planning in Natura 2000 sites

Capsule: The nesting habitat of the Common Buzzard Buteo buteo and Lesser Spotted Eagle Clanga pomarina extensively overlap, indicating that they exploit similar resources.

Aim: We aimed to determine the overlap in the nest platforms, nest trees and nest stands used by these raptors, find any evidence for the avoidance of the larger Lesser Spotted Eagle by the smaller Common Buzzard, and provide conservation implications for habitat protection of the former species in habitats that overlap extensively.

Methods: Nest sites were mapped during 2012–2014 in the Bir?ai Forest Spatial Protection Area, northern Lithuania. Fifty-three nest sites occupied by Common Buzzards and 26 by Lesser Spotted Eagles were compared.

Results: The nest platforms of both raptors were similarly placed in the tree canopies. Most Lesser Spotted Eagle nests were built in spruce, while the Common Buzzard usually nested in birch. The nest stands of the eagles were on wetter soil and located closer to the forest edge than those of the buzzards, otherwise, the nest stands did not differ significantly. There was no evidence for spatial avoidance of the larger raptor by the Common Buzzard.

Conclusions: The different components of the nesting habitats extensively overlapped, and the distribution of the interspecific pairs lacked spatial avoidance. We suggest that the nest sites of both raptors were a largely shared resource, especially if located close to the forest edges. We propose, as a rule of thumb, applying protection by way of buffer zones around buzzard nest sites if they are located close to eagle nest sites and the forest edge.     

Wintering Golden Eagles on the coastal plain of South Carolina

Golden Eagles (Aquila chrysaetos) are rare winter residents in eastern North America, with most found along the Appalachian Mountains and few reported on the coastal plain of the Carolinas. We used remote cameras baited with wild pig (Sus scrofa) and white‐tailed deer (Odocoileus virginianus) carcasses to detect, age, and individually identify Golden Eagles on the U.S. Department of Energy's Savannah River Site on the coastal plain of South Carolina. We identified eight individual Golden Eagles during the winters of 2013–2014 and 2014–2015, with one detected during both winters. We detected eagles for 19 and 66 calendar days during the winters of 2013–2014 and 2014–2015, respectively, with two adult eagles detected for 30 and 31 calendar days in 2014–2015. Eagles typically scavenged on carcasses for a few days, left, and then returned when cameras were baited with another carcass, suggesting they had remained in the area. These observations suggest that large tracts of forests on the coastal plain may be important wintering areas for some Golden Eagles and, further, that other areas in the coastal plain of the southeastern United States may also harbor wintering eagles. Identification of wintering areas of Golden Eagles in the east will be an important step in the conservation of this protected species, and camera traps baited with carcasses can be an effective tool for such work.     

Movements and landscape use of Eastern Imperial Eagles Aquila heliaca in Central Asia

Capsule: We describe ecological factors associated with movements of a globally declining raptor species, the Eastern Imperial Eagle Aquila heliaca.

Aims: To describe the movements, habitat associations and resource selection of Eastern Imperial Eagles marked in Central Asia.

Methods: We used global positioning system (GPS) data sent via satellite telemetry devices deployed on Eastern Imperial Eagles captured in Kazakhstan to calculate distances travelled and to associate habitat and weather variables with eagle locations collected throughout the annual cycle. We also used resource selection models to evaluate habitat use of tracked birds during autumn migration. Separately, we used wing-tagging recovery data to broaden our understanding of wintering locations of eagles.

Results: Eagles tagged in Kazakhstan wintered in most countries on the Arabian Peninsula, as well as Iran and India. The adult eagle we tracked travelled more efficiently than did the four pre-adults. During autumn migration, telemetered eagles used a mixture of vegetation types, but during winter and summer, they primarily used bare and sparsely vegetated areas. Finally, telemetered birds used orographic updrafts to subsidize their autumn migration flight, but they relied on thermal updrafts during spring migration.

Conclusion: Our study is the first to use GPS telemetry to describe year-round movements and habitat associations of Eastern Imperial Eagles in Central Asia. Our findings provide insight into the ecology of this vulnerable raptor species that can contribute to conservation efforts on its behalf.     

Ranging distance of resident Golden Eagles Aquila chrysaetos in western Scotland according to season and breeding status

Capsule Home-range of resident pairs of Golden Eagle was usually smaller during a successful breeding season than during winter and during an unsuccessful breeding season.

Aims To examine how Golden Eagles use space around their nests with respect to season and breeding status, and to compare home-range-use between a high and a low density region.

Methods Nine adults in six mainland Argyll ranges were radiotracked between 1991 and 1996. On the island of Mull visual observations of range-use were obtained for five ranges between 1994 and 1998.

Results Overall, Mull ranges were smaller than the Argyll ranges, reflecting the much higher range density on Mull. In both regions there were significant differences between ranging distances with season and breeding status. In general, ranging distances were smallest during breeding seasons when young were fledged.

Conclusions Studies of range-use in Golden Eagles must be conducted across a 12-month period, as a minimum.     

Spatial and temporal patterns in age structure of Golden Eagles wintering in eastern North America

The behavior of wildlife varies seasonally, and that variation can have substantial demographic consequences. This is especially true for long-distance migrants where the use of landscapes varies by season and, sometimes, age cohort. We tested the hypothesis that distributional patterns of Golden Eagles (Aquila chrysaetos) wintering in eastern North America are age-structured (i.e., birds of similar ages winter together) through the analysis of 370,307 images collected by motion-sensitive trail cameras set over bait during the winters of 2012–2013 and 2013–2014. At nine sites with sufficient data for analysis, we documented 145 eagle visits in 2012–2013 and 146 in 2013–2014. We found significant between-year variation in age structure of wintering eastern Golden Eagles, driven largely by annual differences in the proportion of first-winter birds. However, although many other species show spatial structure in wintering behavior, our analysis revealed no latitudinal organization among age cohorts of wintering eastern Golden Eagles. The lack of age-related latitudinal segregation in wintering behavior does not exclude the possibility that these eagles have sex-based or other types of dominance hierarchies that could result in spatial or temporal segregation. Alternatively, other mechanisms such as food availability or habitat structure may determine the distribution and abundance of Golden Eagles in winter.     

Responses of dispersing GPS-tagged Golden Eagles (Aquila chrysaetos) to multiple wind farms across Scotland

Wind farms may have two broad potential adverse effects on birds via antagonistic processes: displacement from the vicinity of turbines (avoidance), or death through collision with rotating turbine blades. Large raptors are often shown or presumed to be vulnerable to collision and are demographically sensitive to additional mortality, as exemplified by several studies of the Golden Eagle Aquila chrysaetos. Previous findings from Scottish Eagles, however, have suggested avoidance as the primary response. Our study used data from 59 GPS-tagged Golden Eagles with 28 284 records during natal dispersal before and after turbine operation < 1 km of 569 turbines at 80 wind farms across Scotland. We tested three hypotheses using measurements of tag records’ distance from the hub of turbine locations: (1) avoidance should be evident; (2) older birds should show less avoidance (i.e. habituate to turbines); and (3) rotor diameter should have no influence (smaller diameters are correlated with a turbine’s age, in examining possible habituation). Four generalized linear mixed models (GLMMs) were constructed with intrinsic habitat preference of a turbine location using Golden Eagle Topography (GET) model, turbine operation status (before/after), bird age and rotor diameter as fixed factors. The best GLMM was subsequently verified by k-fold cross-validation and involved only GET habitat preference and presence of an operational turbine. Eagles were eight times less likely to be within a rotor diameter’s distance of a hub location after turbine operation, and modelled displacement distance was 70 m. Our first hypothesis expecting avoidance was supported. Eagles were closer to turbine locations in preferred habitat but at greater distances after turbine operation. Results on bird age (no influence to 5+ years) rejected hypothesis 2, implying no habituation. Support for hypothesis 3 (no influence of rotor diameter) also tentatively inferred no habituation, but data indicated birds went slightly closer to longer rotor blades although not to the turbine tower. We proffer that understanding why avoidance or collision in large raptors may occur can be conceptually envisaged via variation in fear of humans as the ‘super predator’ with turbines as cues to this life-threatening agent.     

Habitat preferences and habitat restoration options for small‐bodied and juvenile fish species in the northern Murray–Darling Basin

Degradation of instream habitats in the northern Murray–Darling Basin has occurred through numerous stressors, including siltation, clearing of bankside vegetation, intrusion of livestock and impacts of pest species. A better understanding of habitat preferences of native fish species could help guide future instream habitat restoration actions. The habitat choices of seven native fish species, juvenile Murray Cod (Maccullochella peelii), juvenile Golden Perch (Macquaria ambigua ambigua), juvenile Silver Perch (Bidyanus bidyanus), adult Murray–Darling Rainbowfish (Melanotaenia fluviatilis), adult Olive Perchlet (Ambassis agassizii), adult Un‐specked Hardyhead (Craterocephalus stercusmuscarum fulvus) and adult carp gudgeons (Hypseleotris spp.) were tested in preference troughs to help inform potential habitat restoration actions in the Condamine catchment. Each species was given a choice between pair combinations of open sandy habitat, submerged macrophytes, emergent plants and rocky rubble. Habitat preferences varied between species. Murray Cod, Golden Perch, carp gudgeons and Olive Perchlets preferred structure over open sandy habitat, whilst juvenile Silver Perch, Un‐specked Hardyhead and Murray–Darling Rainbowfish did not avoid open sandy habitats. Juvenile Murray Cod preferred rocky rubble habitat over all other habitat choices. Use of complex rock piles to provide nursery habitat for Murray Cod populations is a potential restoration option. Introduction of rock could also benefit Golden Perch and carp gudgeons. Use of emergent plants, submerged macrophytes and rocky rubble for habitat restoration all appear to have merit for one or more species of small‐bodied fishes or juvenile stages of larger sized fishes. Rocky rubble or floating attached macrophytes could be viable restoration options in areas too turbid to establish submerged macrophytes. These habitat interventions would complement existing actions such as re‐snagging and provision of fish passage to assist with sustainable management of native fish populations.     

Status of Golden Eagle Aquila chrysaetos in Britain in 2015

Capsule: A complete survey of Golden Eagles Aquila chrysaetos in Britain in 2015 found that the population had increased by 15% since 2003 to 508 territorial pairs.

Aims: The survey aimed to investigate the population size, distribution and breeding success of Golden Eagles in Britain, and to compare results with similar surveys since the early 1980s.

Methods: Every home range was visited on a minimum of three occasions between January and August 2015. First, to look for eagles or signs of their presence (January–March), then to look for evidence of breeding or further checks for occupation (April–June) and finally to record productivity of nesting pairs (July–August).

Results: The figure of 508 territorial pairs represents a 15% increase in the population from 442 pairs in 2003. The proportion of home ranges occupied was 70%. The largest increases in the proportion of occupied home ranges were in south-central Highlands (71%), northern moors and flows (38%) and northwest Highlands (29%), with modest increases of up to 10% in the other regions. Productivity was lower in 2015 than in 2003, and there was significant variation in breeding success between regions.

Conclusion: The British Golden Eagle population has increased since 2003, although the species is absent from England and Wales. The population now meets the abundance target identified to define favourable conservation status in Scotland, and while home range occupancy has increased there is regional variation, with some regions falling below the target levels. A combination of increased annual monitoring and tagging of eagles, as well as the introduction of new legislation, may serve as effective deterrents against persecution of eagles thus facilitating this population increase. However, concerns remain over low levels of home range occupancy particularly in the east Highlands, but also the proportion of sub-adult pairs holding territory in that region and in the south-central Highlands. Persecution associated with grouse moor management has been highlighted as a particular population constraint in both of these areas.     

Home range size of willow tits: a response to winter habitat loss

We examined the behavioral response to habitat loss and fragmentation of willow tits (Parus montanus) in winter in a mosaic forest landscape in northern Finland. We studied habitat preference, flock size and home range size of 16 flocks, half of which had their territories in forests fragmented by forestry and half in continuous forest. We predicted that birds would respond to habitat loss by enlarging their home range and/or diminishing group size. In addition, to compensate for fragmentation effects, willow tits might be expected to include more optimal habitat into their territories. Flocks included on average 3.9 birds and occupied territories of 12.6 ha. Willow tits avoided open areas (clear cuts and young sapling stands) and preferred mature forests and older sapling stands or pine bogs equally. Birds responded to habitat loss by enlarging their home ranges but not by reducing the group size. Large territories included a smaller proportion of mature forests, but the proportion of sapling and pine bog habitat did not change. Birds on territories that included a large proportion of open habitat localized their activity on several distinct habitat patches that were distributed over a wide area. We conclude that willow tits adjust territory use to compensate for the inclusion of unsuitable habitat within home ranges, and older sapling areas and pine bogs serve as surrogates for mature forests. However, birds did not enlarge the proportion of forest habitat in their territories with increasing habitat loss. Thus, our data do not suggest a strong effect of fragmentation, but imply that forestry practices reduce suitable wintering habitat and carrying capacity in the area. Thus winter habitat loss may explain the observed population decline of willow tits in Finland during recent decades.     

Golden Eagle Aquila chrysaetos density and productivity in relation to land abandonment and forest expansion in the Alps

Land abandonment and afforestation are causing considerable forest expansion in the Alpine chain, with consequent loss of Alpine pastures, the main foraging habitat for Golden Eagles Aquila chrysaetos. A population of 46 pairs of Golden Eagles was systematically surveyed between 1984 and 1989 in the central-eastern Italian Alps. We measured the amount of woodland within the potential hunting range of 36 pairs. Nearest-neighbour distance was correlated positively with the extent of woodland within the potential hunting range. Productivity was not significantly affected by the amount of woodland within the potential for-aging range. At current rates of forest expansion, such data suggest a 5–9% density decline in the next 20 years. However, increasing food supply and decreasing human persecution could be masking current effects of habitat loss.     

Associations between distance to forest and spatial and temporal variation in abundance of key peatland breeding bird species

Capsule Negative forest edge effects were detected for Willow Ptarmigan (Red Grouse) Lagopus lagopus and Dunlin Calidris alpina.

Aims To investigate the effects of distance to forest on the abundance and changes in abundance of four key peatland breeding bird species, and to measure changes in predatory bird numbers, in the peatlands of northern Scotland.

Methods Bird surveys were carried out in 2000 at 34 plots, covering 197 km² of peatland, and 80 forestry point‐count sites, first surveyed in 1988. Habitat data were also collected in 2000. We used multi‐model inference to investigate the associations between forest distance and other habitat variables, and the abundance, and changes in abundance, of four bird species of economic or conservation importance: Red Grouse, European Golden Plover Pluvialis apricaria, Dunlin and Common Greenshank Tringa nebularia.

Results There was strong evidence that distance to forest was negatively associated with Dunlin abundance and changes in Red Grouse abundance, but only weak evidence for negative associations with Golden Plover abundance and changes in Dunlin abundance. There was no evidence of a forest distance effect on Greenshank. Among predatory birds, there were no significant increases either on peatland plots or in new forestry plantations.

Conclusions This study provides evidence that, for a given habitat quality, Dunlin densities are lower, and Red Grouse declines more likely, near to forest edges, but weak evidence only that Dunlin declines are more likely, and Golden Plover abundance lower, near to forests. These results suggest that for at least two key peatland breeding birds, forest removal is likely to benefit birds breeding on adjacent unafforested peatland.     

Habitat availability in forests and clearcuts for saproxylic beetles associated with aspen

To maintain biodiversity in managed, boreal forest in Scandinavia, aspen trees (Populus tremula) are often retained at clearcutting. In this study, the habitat availability for beetles associated with aspen coarse woody debris (CWD) was predicted for forests and clearcuts with a model of CWD dynamics. Habitat requirements of eight beetle species (Agathidium bicolor, Cerylon ferrugineum, Cyphaea curtula, Endomychus coccineus, Homalota plana, Mycetophagus fulvicollis, Ptilinus fuscus and Xylotrechus rusticus) were obtained from their occurrence patterns in relation to characteristics of CWD objects in forest and on clearcuts in a study landscape in central Sweden. Three species were more frequent in forest and three at clearcuts. Five species increased with increasing girth of the CWD. Three were more frequent on standing CWD, and two on lying CWD. From the same study area, we also obtained field data on the recruitment of CWD (i.e., tree mortality) and amounts of different types of CWD. Annual tree mortality of aspen was higher for recent clearcuts (6.3%) compared with older clearcuts (1.1%). For all species, the habitat availability was higher on clearcuts, because enhanced tree mortality increased the amount of recently dead CWD. As a conclusion, green-tree retention of aspen is a conservation effort that is beneficial for species associated with aspen CWD.     

Sex-bias and timing of natal dispersal in cooperatively breeding Puff-throated Bulbuls Alophoixus pallidus

While natal dispersal can have a significant impact on population dynamics, it is typically difficult to quantify. We investigated timing of natal dispersal of the cooperatively breeding Puff-throated Bulbul Alophoixus pallidus in a tropical evergreen forest by modelling the probability of staying in or dispersing from their natal territory whilst taking into account the effects of sex, group size, and the presence of helper(s). Birds did not disperse until the beginning of and during the breeding season following the hatching year. Dispersal was strongly female-biased both in frequency and distance: most females (95%) dispersed away from their natal territories, and of those relocated, traversed 2–7 territories. In contrast, 50% of males remained in the natal territory as helpers in their second year, while relocated dispersing males crossed 1–2 territories. Natal dispersal was not influenced by either group size or the presence of helpers. Males that fledged earlier in the breeding season exhibited higher rates of philopatry than the males that fledged later, but no correlation between fledging date and philopatry was observed in females. The probability of staying in the natal territory during the second year was 0.58 ± 0.14 SE and 0.05 ± 0.04 for males and females, respectively. These findings may add to our understanding of how natal dispersal can reflect social patterns and kin structure in cooperative breeding species from a little-studied tropical forest region.