Predation as a landscape effect: the trading off by prey species between predation risks and protection benefits

1. Predators impose costs on their prey but may also provide benefits such as protection against other (e.g. nest) predators. The optimal breeding location in relation to the distance from a nesting raptor varies so as to minimize the sum of costs of adult and nest predation. We provide a conceptual model to account for variation in the relative predation risks and derive qualitative predictions for how different prey species should respond to the distance from goshawk Accipiter gentilis nests. 2. We test the model predictions using a comprehensive collection of data from northern Finland and central Norway. First, we carried out a series of experiments with artificial bird nests to test if goshawks may provide protection against nest predation. Second, we conducted standard bird censuses and nest-box experiments to detect how the density or territory occupancy of several prey species varies with distance from the nearest goshawk nest. 3. Nest predation rate increased with distance from goshawk nest indicating that goshawks may provide protection for birds' nests against nest predation. Abundance (or probability of presence) of the main prey species of goshawks peaked at intermediate distances from goshawk nests, reflecting the trade-off. The abundance of small songbird species decreased with distance from goshawk nests. The goshawk poses little risk to small songbirds and they may benefit from goshawk proximity in protection against nest predation. Finally, no pattern with distance in pied flycatcher territory (nest box) occupation rate or the onset of egg-laying was detected. This is expected, as flycatchers neither suffer from marked nest predation risk nor are favoured goshawk prey. 4. Our results suggest that territory location in relation to the nest of a predator is a trade-off situation where adult birds weigh the risk of themselves being predated against the benefits accrued from increased nest survival. Prey species appear able to detect and measure alternative predation risks, and respond adaptively. From the prey perspective, the landscape is a mosaic of habitat patches the quality of which varies according to structural and floristic features, but also to the spatial distribution of predators.     

Competitive interactions among raptors in boreal forests

We examined inter-specific interactions among goshawks (Accipiter gentilis), common buzzards (Buteo buteo) and honey buzzards (Pernis apivorus) in western Finland in 1983–1996. Because goshawks are among the largest birds of prey species in boreal forests they may take over the nest of smaller and less-competitive forest-dwelling raptors when searching for suitable places for breeding. Accordingly, more than half of newly established goshawk territories were found on the territories previously occupied by the common buzzard and the honey buzzard. Otherwise, territory sharing between these species was rare. Fledgling production of honey buzzards was not associated with the presence of goshawks, probably owing to the almost 2 months later onset of breeding. This probably decreases competitive interactions between these two species. An intensive interference competition, instead, seemed to be evident between common buzzards and goshawks, because the fledgling production of common buzzards was decreased by 20% as a result of failures during incubation and nestling period in the vicinity (<1 km) of occupied goshawk nests. Similarly, territory occupancy of common buzzards till the next breeding season was significantly reduced in the presence of goshawks. Relatively high proportions of occupied buzzard territories (17%) in the study area were shared by breeding goshawks on the same territory. This suggests that although their diets are dissimilar they inhabit similar habitats and might compete for the available prime nesting habitats within forest landscapes. In addition, goshawks benefit from taking over the complete nests of other raptors, imposing upon the original owners of the nest, because building a large stick nest is probably energetically costly. As a large raptor, the goshawk apparently has a competitive advantage over smaller ones, and may have an ever-increasing impact on smaller birds of prey, if there is a lack of sheltered forests inducing competition for the available nest sites.     

Northern Goshawk Habitat: an Intersection of Science,Management, and Conservation

Abstract: We responded to the claim by Greenwald et al. (2005) that the management recommendations for the northern goshawk in the Southwestern United States (MRNG; Reynolds et al. 1992), a food web-based conservation plan that incorporated both northern goshawk (Accipiter gentilis) and multiple prey habitats, may be inadequate to protect goshawks. Greenwald et al. (2005) based this claim on their review of 12 telemetry studies of goshawk habitat selection and 5 nontelemetry studies of the effects of vegetation structure at the home range scale on goshawk nest occupancy and reproduction that appeared after the 1992 publication of the MRNG. Greenwald et al. (2005) summarized their review as showing that 1) goshawks were habitat specialists limited to forests with mature and old-growth structures including large trees, high canopy cover, multiple canopy layering, and abundant woody debris; 2) habitats were not selected on the basis of prey abundance and, therefore, managing for prey habitats diluted goshawk habitats; and 3) selection for openings, edges, and habitat diversity was inconclusive. Our review found that when the studies' respective authors pooled their radiotagged goshawks there were weak to strong selections for old forest structures. However, the studies also documented extensive variation in use of vegetation types and structures by individual goshawks; some avoided openings, edges, young forests, and old forests, whereas others selected for these characteristics. Additionally, by virtue of their wide geographic distribution, the studies showed that the focal populations themselves occurred in a variety of forest types, some with large structural differences. We found no evidence in Greenwald's et al. (2005) review that the MRNG are inadequate to protect goshawks. Rather, the studies reviewed by Greenwald et al. (2005), as well as many studies they missed, supported the MRNG. The suggestion of inadequacy by Greenwald et al. (2005) appeared rooted in misunderstandings of goshawk habitats described in the MRNG, a discounting of the extent of variation in vegetation structural and seral stages used by goshawks, a limited understanding of the extent to which prey limits goshawks, a failure to recognize the dynamic nature of forests, and an incomplete review of the literature. We believe the MRNG are adequate because they maximize the sustainable amount of mature and old forests in goshawk home ranges and specify the kinds and intermixtures of prey habitats within home ranges. Implementation of MRNG should reduce the likelihood that the availability of vegetation structures suited to goshawk nesting and foraging, as well as abundance and availability of prey, will limit goshawk nest occupancy and reproduction.     

Density‐dependent increase in superpredation linked to food limitation in a recovering population of northern goshawks Accipiter gentilis

A better understanding of the mechanisms driving superpredation, the killing of smaller mesopredators by larger apex predators, is important because of the crucial role superpredation can play in structuring communities and because it often involves species of conservation concern. Here we document how the extent of superpredation has changed over time, and assessed the impact of such temporal variation on local mesopredator populations using 40 yr of dietary data collected from a recovering population of northern goshawks Accipiter gentilis, an archetypical avian superpredator. We then assessed which mechanisms were driving variation in superpredation, e.g. was it opportunistic, a response to food becoming limited (due to declines in preferred prey) or to reduce competition. Raptors comprised 8% of goshawk diet on average in years when goshawk abundance was high, which is higher than reported elsewhere. Additionally, there was a per capita increase in superpredation as goshawks recovered, with the proportion of goshawk diet comprising raptors increasing from 2 to 8% as the number of goshawk home‐ranges increased from ≤ 14 to ≥ 25. This increase in superpredation coincided with a population decline in the most commonly killed mesopredator, the Eurasian kestrel Falco tinnunculus, which may represent the reversal of the ‘mesopredator release’ process (i.e. mesopredator suppression) which occurred after goshawks and other large raptors declined or were extirpated. Food limitation was the most likely driver of superpredation in this system given: 1) the substantial decline of two main prey groups in goshawk diet, the increase in diet diversity and decrease in goshawk reproductive success are all consistent with the goshawk population becoming food‐limited; 2) it's unlikely to be purely opportunistic as the increase in superpredation did not reflect changes in the availability of mesopredator species; and 3) the majority of mesopredators killed by goshawks do not compete with goshawks for food or nest sites.     

Interactions between common buzzard Buteo buteo and goshawk Accipiter gentilis: trade-offs revealed by a field experiment

I examined the behavioural interactions between common buzzard Buteo buteo and goshawk Accipiter gentilis and their effects on buzzard breeding success and brood defence with a two-year field experiment using dummies and playback calls. A priori I showed through an extensive nest site analysis that there is considerable nesting habitat overlap between the two species and hence potential for interspecific competition for prime nesting habitat. Buzzards had a significantly lower breeding success when presented with a goshawk dummy compared to control broods but there was no effect of buzzard dummies on reproductive success. Buzzards failing with their breeding attempt tended to select another nest site while successful buzzards more frequently used the same nest again. Buzzard pairs were less often attacked by common crows Corvus corone while exposed to goshawk dummies compared to buzzard dummies. The decision to desert a nest seems to be a trade-off between predation risk on the one hand and protection against crows on the other. Goshawks proved far more aggressive against an intraspecific dummy than buzzards. Buzzards adjusted their level of brood defence against both intra- and interspecific dummies according to the age of offspring but not offspring number, with an increasing brood defence level with increasing offspring age. Thus the behaviour of buzzards towards goshawks is a result of a complex system of trade-offs between predation risk, competition for prime nesting habitat and protection from crows on which brood value acts as a temporal modifier.     

Large-scale forest composition influences northern goshawk nesting in Wisconsin

The northern goshawk (Accipiter gentilis atricapillus) is a woodland raptor that uses a variety of forest types for nesting across its breeding range, but strongly depends on older forests with large trees and open understories. Goshawks may select nesting locations by maximizing the convergence of nesting and foraging habitats. Insights into goshawk responses to heterogeneous landscapes can be gained by examining the location of active nest sites through time and at multiple spatial scales. We examined the landscape-scale forest conditions that influenced the probability of active goshawk nests in the United States Forest Service, Chequamegon-Nicolet National Forest (CNNF) in northern Wisconsin. We used goshawk nest survey and monitoring data from 1997 to 2006 to determine the probability of an active nest site over time in relation to forest composition and road density at 3 scales (200-m, 500-m, and 1,000-m radii). Goshawk nests were located primarily in upland hardwood (64%), conifer (23%), and older aspen–birch (≥26 yrs old; 11%) habitat cover types. We used Bayesian temporal autoregressive models of nest locations across multiple spatial scales to analyze these data. The probability of active goshawk nest occurrence increased with increasing conifer cover (1,000 m) and decreased with increasing cover of older aspen–birch and density of primary roads (500 m). In addition, lesser proportions of older aspen–birch at intermediate scales around goshawk nests had a stronger effect on the probability of a nest being active than conifer and primary roads. Thus, the ratio of conifer cover (within 1,000 m) to older aspen–birch cover (within 500 m) in landscapes surrounding nest sites was the key driver in predicting the probability of an active nest site. This finding can be used by forest managers to help sustain the active status of a goshawk nesting area through time (i.e., annually), and foster goshawk nesting activity in areas where active nesting is not currently occurring. Published 2013. This article is a U.S. Government work and is in the public domain in the USA.     

Delayed numerical response of goshawks to population fluctuations of forest grouse

Delayed density-dependent mortality induced by delayed numerical response of predators can drive prey populations to fluctuate in high-amplitude cycles. We studied numerical response of goshawks Accipiter gentilis to varying densities of their main prey (forest grouse) in western Finland during 1979–1996. Occupancy rate of goshawk territories tracked grouse numbers with a two year lag. Occupancy rate of goshawk territories and pooled number of adult and young goshawks correlated negatively with a 1–2 year lag to the chick production of grouse. Goshawk to grouse ratio was negatively related to grouse density. This suggests that goshawk predation on grouse is inversely dependent on grouse density. We conclude that in northern Europe with few alternative preys, goshawk predation might contribute to the generation of multiannual cycles of forest grouse. This should be tested experimentally.     

Effect of Japanese lesser sparrowhawks Accipiter gularis on the nest site selection of azure-winged magpies Cyanopica cyana through their nest defending behavior

The frequency of azure-winged magpies Cyanopica cyana that nest in association with breeding Japanese lesser sparrowhawks Accipiter gularis was compared between the 1990s and 2000s. During 1990–1994 azure-winged magpies nested within 50 m of their nests in all of the hawk nest sites studied. During 2005–2006, however, the magpies nested in only two of nine hawk nest sites. During the 1990s Japanese lesser sparrowhawks chased jungle crows away when they intruded within 50 m of their nest. During the 2000s, on the other hand, the hawks attempted to expel few crows at a distance of more than 10 m from their nest. As a result of it, the predation rate of simulated magpie nests located in the hawk nest sites was significantly higher during the 2000s. This suggests that the magpies have the ability to assess the defending behavior of the hawks and judge whether they can rely on their defense. Most azure-winged magpies may stop nesting in association with breeding Japanese lesser sparrowhawks because of the reduced defending range of their nest sites.     

Spatial consistency in susceptibility of prey species to predation by two Accipiter hawks

Predators impose strong selection on their prey, regulate prey populations and engage in coevolutionary interactions with their prey. The intensity of selection and the strength of coevolutionary interactions will depend on how stringent predators are in their choice of prey. We estimated susceptibility of different species of birds to predation by two common raptors, the northern goshawk Accipiter gentilis and the Eurasian sparrowhawk A. nisus, in an agricultural landscape in Denmark and boreal forests in Finland. We estimated susceptibility to predation as the deviation of the log₁₀‐transformed observed frequency of prey of different species from the log₁₀‐transformed expectation based on population density during the breeding season. We found a high degree of consistency in susceptibility to predation by the goshawk in two areas in Finland. More importantly, there was significant consistency in susceptibility to predation between Denmark and Finland, albeit the degree of consistency in the goshawk was higher than in the sparrowhawk. There was considerable overlap in susceptibility to predation between goshawk and sparrowhawk in Denmark, but not in Finland, implying differences in intensity of interspecific competition as reflected by a much higher extent of goshawk predation on sparrowhawks in Denmark than in Finland. Our findings suggest that hawks impose similar selection pressures on their prey populations, and that the degree of consistency has implications for intensity of interspecific killing.     

Habitat Effects on the Breeding Performance of Three Forest-Dwelling Hawks

Habitat loss causes population declines, but the mechanisms are rarely known. In the European Boreal Zone, loss of old forest due to intensive forestry is suspected to cause declines in forest-dwelling raptors by reducing their breeding performance. We studied the boreal breeding habitat and habitat-associated breeding performance of the northern goshawk (Accipiter gentilis), common buzzard (Buteo buteo) and European honey buzzard (Pernis apivorus). We combined long-term Finnish bird-of-prey data with multi-source national forest inventory data at various distances (100–4000 m) around the hawk nests. We found that breeding success of the goshawk was best explained by the habitat within a 2000-m radius around the nests; breeding was more successful with increasing proportions of old spruce forest and water, and decreasing proportions of young thinning forest. None of the habitat variables affected significantly the breeding success of the common buzzard or the honey buzzard, or the brood size of any of the species. The amount of old spruce forest decreased both around goshawk and common buzzard nests and throughout southern Finland in 1992–2010. In contrast, the area of young forest increased in southern Finland but not around hawk nests. We emphasize the importance of studying habitats at several spatial and temporal scales to determine the relevant species-specific scale and to detect environmental changes. Further effort is needed to reconcile the socioeconomic and ecological functions of forests and habitat requirements of old forest specialists.     

Spatial relationships and mechanisms of coexistence between dominant and subordinate top predators

Most forest ecosystems contain a diverse community of top‐level predators. How these predator species interact, and how their interactions influence their spatial distribution is still poorly understood. Here we studied interactions among top predators in a guild of diurnal forest raptors in order to test the hypothesis that predation among competing predators (intraguild predation) significantly affects the spatial distribution of predator species, causing subordinate species to nest farther away from the dominant ones. The study analyzed a guild in southwestern Europe comprising three raptor species. For 8 years we studied the spatial distribution of used nests, breeding phenology, intraguild predation, territory occupancy, and nest‐builder species and subsequent nest‐user species. The subordinate species (sparrowhawk Accipiter nisus) nested farther away from the dominant species (goshawk A. gentilis), which preyed on sparrowhawks but not on buzzards Buteo buteo, and closer to buzzards, with which sparrowhawks do not share many common prey. This presumably reflects an effort to seek protection from goshawks. This potential positive effect of buzzards on sparrowhawks may be reciprocal, because buzzards benefit from old sparrowhawk nests, which buzzards used as a base for their nests, and from used sparrowhawk nests, from which buzzards stole prey. Buzzards occasionally occupied old goshawk nests. These results support our initial hypothesis that interspecific interactions within the raptor guild influence the spatial distribution of predator species in forest ecosystems, with intraguild predation as a key driver. We discuss several mechanisms that may promote the coexistence of subordinate and dominant predators and the spatial assembly of this raptor guild: spatial refuges, different breeding phenology, spatial avoidance, low territory occupancy between neighboring nesting territories, nest concealment and protection, and diet segregation.     

Interactions between habitat heterogeneity and food affect reproductive output in a top predator

1. Habitat heterogeneity has important repercussions for species abundance, demography and life-history patterns. While habitat effects have been more thoroughly studied in top-down situations (e.g. in association with predation), their role in bottom-up situations (e.g. in association with food abundance) has been less explored and the underlying mechanism(s) behind the ecological patterns have not commonly been identified. 2. With material from 1993 to 2003, we test the hypothesis that the reproduction of Finnish northern goshawks Accipiter gentilis (L.) is bottom-up limited by habitat composition, especially in situations where the density of their main prey (grouse) is low. Special emphasis was placed on identifying the mechanism(s) behind potential habitat effects. 3. While laying date and large-scale variation in the main prey density (but not habitat composition) were related to the number of eggs goshawks laid, small-scale differences in alternative prey density between different territories later influenced how many young were fledged via the mechanism of habitat-dependent partial-brood loss. As a result of this mechanism, a difference in nestling condition also arose between goshawk territories with differing habitat compositions. 4. As the relative proportions of different landscape elements in a given landscape is a function of large-scale differences in geomorphology and land use, this means that the reproductive performance of goshawks as averaged over larger scales can be understood correctly only in respect to the fact that habitat gradients differ across landscapes. 5. In addition to being one of the first papers identifying the mechanism of partial brood loss as being primarily responsible for the habitat-specific differences in the production of young, this study further illustrates the need to identify small-scale mechanisms to correctly understand the large-scale patterns of reproductive performance in territorial species. The repercussions of the observed habitat effect for local population development are discussed.     

Analysis of nest occupancy and nest reproduction in two sympatric raptors: common buzzard Buteo buteo and goshawk Accipiter gentilis

Nest site selection can have important fitness consequences in birds. I analysed the habitat characteristics of 392 nests of two sympatric raptor species (common buzzard Buteo buteo and goshawk Accipiter gentilis ) in Germany and their relation to nest occupation rate and nest reproductive success. For common buzzard, multivariate models explained only small proportions of the variance in nest occupation rate and nest reproductive success (13–19%). Important variables related to nest occupation rate were human disturbances, intra- and interspecific neighbour density, the amount of forested area and nest tree crown cover. Variables related to nest site reproductive success also included human disturbance, intra- and interspecific neighbour density and nest tree crown cover as well as nest distance to the nearest forest edge. In contrast, models for the goshawk explained a much higher proportion of the variation in nest occupation rate and nest reproductive success (41–43%). Important variables related to nest occupation rate were the remoteness of the nest site and direct human disturbance. Variables related to nest site reproductive success were remoteness of the nest site and good hunting habitat. Goshawks seem to be more sensitive to human disturbance than buzzards. A multiple discriminant analysis showed that nest site characteristics substantially overlapped between the species and there is a good evidence that competition for optimal nest sites occurs. Thus, buzzards might be constrained by the dominant goshawk in their nest site selection.     

Resource partitioning by three species of newts during their aquatic phase

This paper describes the multidimensional niche of syntopic smooth newts Triturus vulgaris. alpine newts Triturus alpestris. and Italian crested newts Triturus carnifex. which constitute a complex community of eight components (the three species and their larvae, plus juvenile and neotenic alpine newts) All the potentially limiting resources (seasonal time, diel time, habitat and food) are analyzed as single niche dimensions, and then the real multidimensional niche is calculated Resource states are objectively defined, using an algorithm that groups the significant resource categories The larvae were segregated by prey, season, habitat, and water depth, in order of decreasing importance, their overlap was very low when their complete multidimensional niche was considered, and therefore they have few opportunities for present–day competition The adults were largely segregated by season and water depth, and to a lesser degree by habitat and prey, their multidimensional niches overlapped to a certain degree, thus leaving conditions for ongoing competition Considenng all the species and the life stages together, the order of importance for segregation was season, prey, habitat, water depth and diel activity, with a low overlap in the complete niche Differences m prey size were probably due to morphological constraints, differences in diel activity had very little effect on segregation except m separating larvae from the adults, and were likely promoted by need to avoid mutual predation and cannibalism It remains to be tested whether seasonal, habitat, and water depth differences are due to competitive pressures or to ecophysiological constraints     

Estimating occupancy to monitor northern goshawk in the central Rocky Mountains

Designing monitoring programs to evaluate trends in low-density wildlife species at regional scales is challenging given difficulties detecting uncommon organisms distributed in potential habitats over large spatial extents. The northern goshawk (Accipiter gentilis) has been petitioned for listing under the Endangered Species Act and the review of the petition indicated a need for information on population trend. To evaluate trends in goshawk populations, the U.S. Forest Service developed the Northern Goshawk Bioregional Monitoring Design to estimate goshawk occupancy over broad spatial extents. We adapted and implemented this design to approximately 30,600 km² of 88,128 km² of National Forest System lands in the Forest Service Rocky Mountain Region, including portions of Colorado, Wyoming, and South Dakota. We developed a stratified random design to monitor goshawk occupancy in sampling units, defined by primary and secondary habitat quality as well as accessibility. To define habitat quality, we examined a time series for 58 previously located nesting territories. Using logistic regression, we found that the dominant conifer species and status of aspen in postfledging zones best characterized high-quality goshawk nesting habitat. We applied model results to stratify 4,445 sampling units based on habitat quality and further stratified sampling units based on accessibility into easy and difficult access categories. We conducted field sampling during the goshawk breeding season in the summer of 2006 to estimate detection probabilities and occupancy rates. Within our sampling frame, we sampled 51 sampling units and estimated goshawk occupancy of 0.329 (95% CI: 0.213–0.445). Occupancy within primary strata (high quality) sampling units was 0.811 (SE = 0.113), whereas occupancy in secondary strata (lower quality) sampling units was 0.124 (SE = 0.067). Future implementation of this monitoring program can achieve 0.8 power to detect 30–40% declines in with 140 sampling units. Our implementation of a stratified sampling design to monitor occupancy of goshawks at a region-wide scale reduced the number of sampling units in each administrative unit and focused our efforts on those areas most likely to have goshawks. © 2011 The Wildlife Society.     

Buteo Nesting Ecology: Evaluating Nesting of Swainson’s Hawks in the Northern Great Plains

Swainson’s hawks (Buteo swainsoni) are long-distance migratory raptors that nest primarily in isolated trees located in areas of high grassland density. In recent years, anthropogenic conversion of grassland habitat has raised concerns about the status of the breeding population in the northern Great Plains. In 2013, we initiated a study to investigate the influence of extrinsic factors influencing Swainson’s hawk nesting ecology in north-central South Dakota and south-central North Dakota. Using ground and aerial surveys, we located and monitored nesting Swainson’s hawk pairs: 73 in 2013 and 120 in 2014. We documented 98 successful breeding attempts that fledged 163 chicks; 1.52 and 1.72 fledglings per successful nest in 2013 and 2014, respectively. We used Program MARK to evaluate the influence of land cover on nest survival. The top model, S _{Dist2Farm+%Hay}, indicated that nest survival (fledging at least one chick) decreased as nests were located farther from farm sites and as the percent of hay cover increased within 1200-m of the nest site (34.4%; 95% CI = 27.6%–42.3%). We used logistic regression analysis to evaluate the influence of landscape variables on nest-site selection; Swainson’s hawks selected for nest sites located closer to roads. We suggest that tree belts associated with farm sites, whether occupied or not, provide critical breeding sites for Swainson’s hawks. Additionally, poor breeding success may be related to the late migratory behavior of this species which requires them to occupy marginal habitat due to other raptors occupying the most suitable habitat prior to Swainson’s hawks arriving to the breeding grounds.     

Movements and Resource Selection of Fledgling Goshawks in Montane Forests of Southeastern British Columbia

ABSTRACT The northern goshawk (Accipiter gentilis) has been the subject of considerable interest because of the impact of logging on this species' nesting habitat. However, few studies have examined movements of fledgling birds around the nest prior to independence, and even fewer have described resource requirements of young birds during their postfledging period. Over 3 years, we followed 31 radiotagged goshawk fledglings from 15 nests in southeastern British Columbia, Canada. Of these birds, 26 survived to disperse. Between fledging and dispersal 95% of fledgling relocations (n = 1, 148) were within 450 m of the nest. Fledglings primarily remained within 298 m of the nest during the first 21 days postfledging and within 525 m of the nest between 21 days postfledging and dispersal. Fledglings' movements were highly directional, with individual and sibling movements away from any particular nest tending out in one direction. Postfledging areas averaged 36.7 ha in size (median = 23.1, inter-quartile range = 20.8–39.7 ha). Fledglings strongly avoided forest <40 years old and weakly selected young forests (40–80 yr), mature forests (>80 yr), and stands with >40% canopy cover during the first 21 days and after. We suggest forest managers wishing to conserve goshawk postfledging areas in the interior montane forests of British Columbia maintain forests >40 years old with high crown closure covering an area ≥21 ha and preferably >40 ha. This area should contain all identified occupied and alternative nest trees in a nest area. At least half this area should be forest >80 years old and contain existing nests and potential for future nest trees.     

Comparison of wing morphology in three birds of prey: correlations with differences in flight behavior

Flight is the overriding characteristic of birds that has influenced most of their morphological, physiological, and behavioral features. Flight adaptations are essential for survival in the wide variety of environments that birds occupy. Therefore, locomotor structure, including skeletal and muscular characteristics, is adapted to reflect the flight style necessitated by different ecological niches. Red-tailed hawks (Buteo jamaicensis) soar to locate their prey, Cooper's hawks (Accipiter cooperii) actively chase down avian prey, and ospreys (Pandion haliaetus) soar and hover to locate fish. In this study, wing ratios, proportions of skeletal elements, and relative sizes of selected flight muscles were compared among these species. Oxidative and glycolytic enzyme activities of several muscles were also analyzed via assays for citrate synthase (CS) and for lactate dehydrogenase (LDH). It was found that structural characteristics of these three raptors differ in ways consistent with prevailing aerodynamic models. The similarity of enzymatic activities among different muscles of the three species shows low physiological differentiation and suggests that wing architecture may play a greater role in determining flight styles for these birds.     

Fear factor: prey habitat selection and its consequences in a predation risk landscape

Predation risk influences prey use of space. However, little is known about how predation risk influences breeding habitat selection and the fitness consequences of these decisions. The nest sites of central-place foraging predators may spatially anchor predation risk in the landscape. We explored how the spatial dispersion of avian predator nests influenced prey territory location and fitness related measures. We placed 249 nest boxes for migrant pied flycatchers Ficedula hypoleuca , at distances between 10 and 630 m, around seven different sparrowhawk nests Accipiter nisus . After closely monitoring flycatcher nests we found that flycatcher arrival dates, nest box occupation rates and clutch size showed a unimodal relationship with distance from sparrowhawk nests. This relationship suggested an optimal territory location at intermediate distances between 330 and 430 m from sparrowhawk nests. Furthermore, pied flycatcher nestling quantity and quality increased linearly with distance from sparrowhawk nests. These fitness related measures were between 4 and 26% larger in flycatcher nestlings raised far from, relative to those raised nearby, sparrowhawk nests. Our results suggest that breeding sparrowhawk affected both flycatcher habitat selection and reproductive success. We propose that nesting predators create predictable spatial variation in predation risk for both adult prey and possibly their nests, to which prey individuals are able to adaptively respond. Recognising predictable spatial variation in perceived predation risk may be fundamental for a proper understanding of predator-prey interactions and indeed prey species interactions.