The feeding ecology of Bonelli’s eagle (<Emphasis Type="Italic">Aquila fasciata</Emphasis>) floaters in southern Spain: implications for conservation

Although many studies have investigated the feeding habits of Palaearctic raptors, few have analysed non-breeding populations during dispersal. Bonelli’s eagle (Aquila fasciata), a threatened species in Western Europe, has a relatively long and critical dispersal period. We studied feeding habits, prey selection, and the influence of prey density on floater abundance in this species during its dispersal period in southern Spain. Differences were found between the diet of floaters and that of the closest breeding populations. Diet diversity was rather low for floaters, with European wild rabbits (Oryctolagus cuniculus) being the main prey and the only prey positively selected. Moreover, the number of floater Bonelli’s eagles observed in the dispersal areas was positively associated with rabbit abundance. Other prey included red-legged partridges (Alectoris rufa) and pigeons (Columba sp.). We propose measures to match suitable prey availability for floater eagles in settlement areas and the use of such areas as human hunting fields.     

Availability of optimal‐sized prey affects global distribution patterns of the golden eagle Aquila chrysaetos

Climate and landscape change are expected to significantly affect trophic interactions, which will especially harm top predators such as the golden eagle Aquila chrysaetos. Availability of optimal prey is recognized to influence reproductive success of raptors on a regional scale. For the golden eagle, medium‐sized prey species between 0.5 and 5 kg are widely considered to be optimal prey during the breeding season, whereas smaller and larger species are deemed as energetically sub‐optimal. However, knowledge about the effects of optimal prey availability is still scarce on larger scales. To decrease this apparent knowledge gap, we combined biogeographical information on range margins with information about the foraging behaviour and reproductive success of golden eagles from 67 studies spanning the Northern Hemisphere. We hypothesized that availability of optimal prey will affect foraging behaviour and breeding success and, thus, distribution patterns of the golden eagle not only on a local but also on a continental scale. We correlated the diet breadth quantifying foraging generalism, breeding success and proportions of small (< 0.5 kg), medium (0.5–5 kg) and large‐sized (> 5 kg) prey species within the diet with the minimum distance of the examined eagles to the actual species distribution boundary. Closer to the range edge, we observed decreased proportions of medium‐sized prey species and decreasing breeding success of golden eagles. Diet breadth as well as proportions of small and large‐sized prey species increased, however, towards the range edge. Thus, availability of optimal‐sized prey species seems to be a crucial driver of foraging behaviour, breeding success and distribution of golden eagles on a continental scale. However, underlying effects of landscape characteristics and human influence on optimal prey availability has to be investigated in further large‐scale studies to fully understand the major threats facing the golden eagle and possibly other large terrestrial birds of prey.     

Diet specificity is not associated with increased reproductive performance of Golden Eagles Aquila chrysaetos in Western Scotland

Amongst raptor species, individuals with specialized diets are commonly observed to have higher reproductive output than those with general diets. A suggested cause is that foraging efficiency benefits accrue to diet specialists. This diet specificity hypothesis thus predicts that diet breadth and reproductive success should be inversely related within species. We highlight, however, that a prey availability hypothesis also makes the same prediction in some circumstances. Hence, when high diet specificity results from high encounter rates with an abundant, preferred prey, then prey availability may affect reproductive success, with diet specialization as an incidental correlate. Using three insular study areas in western Scotland, we examine diet specificity and reproductive success in Golden Eagles Aquila chrysaetos. Diet breadth and breeding productivity were not negatively related in any of our study areas, even though birds with specific diets did tend to have a higher incidence of preferred prey (grouse and lagomorphs) in the diet. Indeed, in two study areas there was evidence that diet generalists had higher breeding productivity. Our results therefore failed to support the diet specificity hypothesis but were consistent with the prey availability hypothesis. We highlight that although many other studies are superficially consistent with the diet specificity hypothesis, our study is not alone in failing to provide support and that the hypothesis does not provide a generic explanation for all relevant results. Diet specificity in predators can be at least partially a response to prey diversity, availability and distribution, and benefits associated with different prey types, so that being a generalist is not necessarily intrinsically disadvantageous. We suggest that the available evidence is more consistent with variation in prey abundance and availability as a more influential factor explaining spatial and temporal variation in breeding productivity of ‘generalist’ species such as the Golden Eagle. Under this argument, prey abundance and availability are the main drivers of variation in reproductive output. Diet specificity is a consequence of variation in prey availability, rather than a substantial cause of variation in reproductive success.     

Differential habitat selection by immature and adult Grey Eagle-buzzards Geranoaetus melanoleucus

During the 1992 breeding season, we studied the habitat selection, behaviour, aggressive interactions and diet of immature and adult Grey Eagle-buzzards Geranoaetus melanoleucus in an area of the Argentinean Patagonia. Immature eagles selected areas rich in prey, had no preference for flat or rugged areas and did not avoid areas close to active nests or those used by adult eagles. The density of the European Hare Lepus europaeus was the best predictor of the distribution of immature eagles. Adult eagles preferred rugged to flat areas but were not significantly affected by prey distribution. Immature eagles used flapping flight more frequently than did adult eagles. Wing-flapping frequency of immatures did not differ when flying over slopes or plains. In contrast, adults used a more economic flight with a lower wing-flapping frequency than that of immatures, especially when flying over slopes. Adults, in contrast to immatures, tended to select slopes when flying over predominantly flat country. Only in 22% of the occasions when adult and immatures were seen together were immatures attacked by adults, a rate of aggressive encounters similar to that observed between immatures (32%). This similarity indicates that adults do not actively exclude immatures from certain areas. The main prey of both immature and adult eagles was the introduced European Hare followed by native rodents and birds. Immatures ate significantly more carrion and fewer birds than did adults. These observations suggest that differences in flight behaviour and in the flight silhouette between adult and immature eagles may be responsible for their different habitat selection in relation to topography. While immatures apparently cue on prey density for habitat selection, adults select areas that allow more economic foraging flights     

A community-level evaluation of the impact of prey behavioural and ecological characteristics on predator diet composition

Although predation avoidance is the most commonly invoked explanation for vertebrate social evolution, there is little evidence that individuals in larger groups experience lower predation rates than those in small groups. We compare the morphological and behavioural traits of mammal prey species in the Taï forest, Ivory Coast, with the diet preferences of three of their non-human predators: leopards, chimpanzees and African crowned eagles. Individual predators show marked differences in their predation rates on prey species of different body sizes, but clear patterns with prey behaviour were apparent only when differences in prey habitat use were incorporated into the analyses. Leopard predation rates are highest for terrestrial species living in smaller groups, whereas eagle predation rates are negatively correlated with group size only among arboreal prey. When prey predation rates are summed over all three predators, terrestrial species incur higher predation rates than arboreal species and, within both categories, predation rates decline with increasing prey group size and decreasing density of groups in the habitat. These results reveal that it is necessary to consider anti-predator strategies in the context of a dynamic behavioural interaction between predators and prey.     

Taphonomic signature of golden eagles (Aquila chrysaetos) on bone prey remains

Abstract

The golden eagle (Aquila chrysaetos) is one of the most important birds of prey in the Northern Hemisphere. This raptor is used to building large nests in high cliffs to which they return for several breeding years accumulating important amounts of their prey skeletal remains. This makes the golden eagle one of the major predators able to accumulate faunal remains in archaeological sites. Despite this fact, the taphonomic signature of golden eagles has not been properly characterized. Here we present the analysis of ingested and non-ingested faunal remains predated and accumulated by this raptor in two different nesting areas from the Iberian Peninsula. Results show how the faunal taxonomic record may vary depending on the ecological zone. Leporids and terrestrial carnivores are the best represented. The observed anatomical representation, breakage and bone surface modification patterns are discussed for different taxa. The taphonomic pattern varies depending on the type of prey and the origin of skeletal materials (non-ingested vs. pellets). Finally, after comparing our results with marks left by other predators, several characteristic features are noted to recognise golden eagles as agents of animal bones accumulations in the fossil record.     

Factors influencing diet variation in a generalist predator across its range distribution

Factors linked with intraspecific variation in trophic diversity are still poorly understood in generalist species like the Montagu’s harrier (Circus pygargus) but may have important implications for conservation management at a wide scale. We described geographic patterns of Montagu’s harrier diet across Eurasia, gathering diet data from 30 studies in 41 areas from 11 countries. We grouped prey as invertebrates, reptiles, small mammals, large mammals, eggs, small birds and large birds, and calculated the contribution of each prey type to the diet (as % biomass) and Shannon’s Diversity Index for each study site. We analysed qualitative estimates of prey abundance in relation to latitude and longitude, then diet composition in relation to habitat of the study area and prey abundance estimates. Diet diversity of Montagu’s harriers increased from north to south, while abundance of all prey groups other than small mammals showed the opposite trends. Agricultural areas in northern latitudes seemed to hold high densities of small mammals, but low densities of alternative prey. Overall, birds were the main prey in most of Montagu’s harrier’s distribution range, although the relative importance of each prey type in the diet was significantly explained by its local abundance and habitat, confirming the opportunistic foraging strategy of this raptor species. Consumption of mammals was an exception to this trend, being negatively associated with the abundance of alternative prey, suggesting that this prey is not preferred. Trophic diversity in this species could be influenced by land-use changes through variations in the abundance and availability of prey, which could impact its population dynamics. This may be particularly important for northern populations of Montagu’s harriers breeding in agricultural habitats, where trophic diversity is already low.     

Comparative breeding performance of Marsh Harriers Circus aeruginosus along a gradient of land‐use intensification and implications for population management

Assessing variation in breeding performance in relation to habitat characteristics may provide insights into predicting the consequences of land‐use change on species ecology and population dynamics. We compared four Marsh Harrier Circus aeruginosus populations subject to similar environmental conditions, but which differed in habitat composition, ranging from natural habitats to intensively cultivated areas. Using a 6‐year dataset, we characterized breeding habitat and diet in these four study sites, and analysed breeding performance in relation to this gradient of land‐use intensification. There was minimal variation in breeding performance between study years but consistent variation between study sites. Unexpectedly, Marsh Harriers breeding in intensively cultivated habitats had higher reproductive success than those breeding in more natural habitats, which, however, hosted higher breeding densities, so overall net population productivity (fledglings per unit area) was similar across sites. This resulted from combined effects of density‐dependence and different predation rates between study sites. The colonization of intensive farmland habitats may not necessarily impact negatively on population sustainability when breeding success and population density are traded against each other. However, our findings should not mask longer‐term conservation issues for populations breeding in these intensively managed areas, and further studies should assess potential long‐term negative effects of occupancy of human‐altered habitat.     

Wildflower areas within revitalized agricultural matrices boost small mammal populations but not breeding Barn Owls

Agro-ecosystems have recently experienced dramatic losses of biodiversity due to more intensive production methods. In order to increase species diversity, agri-environment schemes provide subsidies to farmers who devote a fraction of their land to ecological compensation areas (ECAs). Several studies have shown that invertebrate biodiversity is actually higher in ECAs than in nearby intensively cultivated farmland. It remains poorly understood, however, to what extent ECAs also favour vertebrates, such as small mammals and their predators, which would contribute to restoring functional food chains within revitalised agricultural matrices. We studied small mammal populations among eight habitat types—including wildflower areas, a specific ECA in Switzerland—and habitat selection (radiotracking) by the Barn Owl Tyto alba, one of their principal predators. Our prediction was that habitats with higher abundances of small mammals would be more visited by foraging Barn Owls during the period of chicks’ provisioning. Small mammal abundance tended to be higher in wildflower areas than in any other habitat type. Barn Owls, however, preferred to forage in cereal fields and grassland. They avoided all types of crops other than cereals, as well as wildflower areas, which suggests that they do not select their hunting habitat primarily with respect to prey density. Instead of prey abundance, prey accessibility may play a more crucial role: wildflower areas have a dense vegetation cover, which may impede access to prey for foraging owls. The exploitation of wildflower areas by the owls might be enhanced by creating open foraging corridors within or around wildflower areas. Wildflower areas managed in that way might contribute to restore functional links in food webs within agro-ecosystems.     

Evaluating the influence of diet-related variables on breeding performance and home range behaviour of a top predator

Diet composition is linked to reproductive performance directly or indirectly by other life-history traits, including home range behaviour. The relationships between prey abundance, diet and individual fitness have often been explored. However, these relationships are complex and difficult to disentangle, especially in vertebrate top predators. Here, we present the results of a long-term study using multi-model inference procedures to elucidate the influence of diet-related variables on breeding parameters and home range behaviour of a top predator, the eagle owl Bubo bubo. Superpredation, diet diversity, rat biomass and rabbit mean weight were the most important variables when analysing reproductive parameters, suggesting that less diverse diets with greater rabbit biomass percentage may benefit reproductive performance, whereas rat biomass percentage is apparently associated with greater variation of breeding success. Earlier laying dates seem to be associated with the consumption, on average, of smaller rabbits. On the other hand, edge density was the most relevant factor determining the variation in home range behaviour, with individual characteristics, such as age and sex, also being important. Although the relative importance of the diet-related variables was generally low, mean weight of alternative prey, diet diversity and rabbit biomass also helped to explain home range parameters. In an optimal foraging context, centred on the abundance of the main prey species, our results suggest that when rabbits are less available eagle owls may increase home range size in order to obtain alternative prey, increasing at the same time their dietary diversity, which may also require higher movement speed.     

Avian top predator and the landscape of fear: responses of mammalian mesopredators to risk imposed by the golden eagle

Top predators may induce extensive cascading effects on lower trophic levels, for example, through intraguild predation (IGP). The impacts of both mammalian and avian top predators on species of the same class have been extensively studied, but the effects of the latter upon mammalian mesopredators are not yet as well known. We examined the impact of the predation risk imposed by a large avian predator, the golden eagle (Aquila chrysaetos, L.), on its potential mammalian mesopredator prey, the red fox (Vulpes vulpes, L.), and the pine marten (Martes martes, L.). The study combined 23 years of countrywide data from nesting records of eagles and wildlife track counts of mesopredators in Finland, northern Europe. The predation risk of the golden eagle was modeled as a function of territory density, density of fledglings produced, and distance to nearest active eagle territory, with the expectation that a high predation risk would reduce the abundances of smaller sized pine martens in particular. Red foxes appeared not to suffer from eagle predation, being in fact most numerous close to eagle nests and in areas with more eagle territories. This is likely due to similar prey preferences of the two predators and the larger size of foxes enabling them to escape eagle predation risk. Somewhat contrary to our prediction, the abundance of pine martens increased from low to intermediate territory density and at close proximity to eagle nests, possibly because of similar habitat preferences of martens and eagles. We found a slightly decreasing trend of marten abundance at high territory density, which could indicate that the response in marten populations is dependent on eagle density. However, more research is needed to better establish whether mesopredators are intimidated or predated by golden eagles, and whether such effects could in turn cascade to lower trophic levels, benefitting herbivorous species.     

Sequential movement into coastal habitats and high spatial overlap of predator and prey suggest high predation pressure in protected areas

In theory, predators should attempt to match the distribution of their prey, and prey to avoid areas of high predation risk. However, there is a scarcity of empirical knowledge on predator and prey spatial use when both are moving freely in their natural environment. In the current study, we use information collated on a predators’ diet, its population structure, as well as predator and prey relative abundance, and track the movements of predator and prey simultaneously to compare habitat use and evaluate predation pressure. The study was conducted in elasmobranch protected areas of coastal Tasmania, Australia. The species considered were the broadnose sevengill shark Notorynchus cepedianus, the apex predator in the area, and five chondrichthyan prey species. Notorynchus cepedianus and its prey show similar seasonality in the use of these coastal areas: more abundant in warmer months and absent in winter. Predator and prey also showed high spatial overlap and similar habitat use patterns. These similar movement patterns of predator and prey combined with the additional ecological information (diet, population structure of predator, relative abundance of predator and prey) suggests that N. cepedianus move into coastal areas to exploit seasonally abundant prey. Also, while in protected areas, chondrichthyans are subjected to high predation pressure. Overall, results illustrate the value of simultaneously recording and integrating multiple types of information to explore predator–prey relationships and predation pressure.     

Large-scale spatio-temporal shifts in the diet of a predator mediated by an emerging infectious disease of its main prey

Aim To explore the influence of an emerging infectious disease (EID) affecting a prey species on the spatial patterns and temporal shifts in the diet of a predator over a large geographical scale. We reviewed studies on the diet of Bonelli’s eagles (Hieraaetus fasciatus) in order to determine the repercussions of the reduction in the density of its main prey, the rabbit (Oryctolagus cuniculus), caused by outbreaks of rabbit haemorrhagic disease (RHD) since 1988. Location Western continental Europe. Methods We compiled published and unpublished information on the diet of breeding Bonelli’s eagles from Portugal, Spain and France for a 39‐year study period (1968–2006). Nonparametric tests were used in order to analyse temporal shifts in diet composition and trophic diversity (H′) between the periods of ‘high’ (before outbreak of RHD) and ‘low’ rabbit density (after outbreak of RHD). A combination of hierarchical agglomerative clustering and non‐metric multidimensional scaling (NMDS) analyses were used to test for the existence of geographical patterns in the diet of Bonelli’s eagles in each period. Results The diet of the Bonelli’s eagle consisted of rabbit (28.5%), pigeons (24.0%), partridges (15.3%), ‘other birds’ (11.6%), ‘other mammals’ (7.1%), corvids (7.0%), and herptiles (6.4%). However, RHD had large consequences for its feeding ecology: the consumption of rabbits decreased by one‐third after the outbreak of RHD. Conversely, trophic diversity (H′) increased after outbreak of RHD. At the same time, the analyses showed clear geographical patterns in the diet of the Bonelli’s eagle before, but not after, RHD outbreak. Main conclusions Geographical patterns in the diet of the Bonelli’s eagle in western Europe seem to be driven mainly by spatio‐temporal variation in the abundance of rabbits and, to a lesser extent, by the local (territorial) environmental features conditioning the presence and density of alternative prey species. We show that an EID can disrupt predator–prey relationships at large spatial and temporal scales through a severe decline in the population of the main prey species. Hence we argue that strict guidelines should be drawn up to prevent human‐aided dissemination of ‘pathogen pollution’, which can threaten wildlife not only at the population and species level but also at the community and ecosystem scale.     

Diet composition and foraging success in generalist predators: Are specialist individuals better foragers?

Factors affecting individual diet specialization in generalist populations and the relationship between diet and foraging success remain poorly studied, particularly in terrestrial wide-ranging predators. We studied whether individual variations in diet in Montagu's harrier males (determined through a combination of direct foraging observations and pellet analysis) were associated with patterns of foraging habitat selection and foraging success of 12 radiotracked males during the breeding period. We found important differences in diet composition and breadth between individuals. Diet diversity was negatively related to hunting success: the most efficient individuals in terms of hunting success had the most specialized diet. This study also suggests an important role of individual foraging habitat selection in explaining individual diet, as the proportion of different prey types in the diet was associated with habitat composition within the home range, with higher proportion of those habitats that held higher abundances of their more frequent prey. This study thus provides evidence of individual diet specialization having a knock-on effect on foraging efficiency in a wide-ranging raptor and highlights the role of individual behaviour as a driving force of intra-population niche variation.     

Do nomadic avian predators synchronize population fluctuations of small mammals? a field experiment

Three-to-five-year population oscillations of northern small rodents are usually synchronous over hundreds of square kilometers. This regional synchrony could be due to similarity in climatic factors, or due to nomadic predators reducing the patches of high prey density close to the average density of a larger area. We estimated avian predator and small rodent densities in 4–5 predator reduction and 4–5 control areas (c. 3 km² each) during 1989–1992 in western Finland. We studied whether nomadic avian predators concentrate at high prey density areas, and whether this decreases spatial variation in prey density. The yearly mean number of avian predator breeding territories was 0.2–1.0 in reduction areas and 3.0–8.2 in control areas. Hunting birds of prey concentrated in high prey density areas after their breeding season (August), but not necessarily during the breeding season (April to June), when they were constrained to hunt in vicinity of the nest. The experimental reduction of breeding avian predators increased variation in prey density among areas but not within areas. The difference in variation between raptor reduction and control areas was largest in the late breeding season of birds of prey, and decreased rapidly after the breeding season. These results appeared to support the hypothesis that the geographic synchrony of population cycles in small mammals may be driven by nomadic predators concentrating in high prey density areas. Predation and climatic factors apparently are complementary, rather than exclusive, factors in contributing to the synchrony.     

Apex predators and the facilitation of resource partitioning among mesopredators

Apex predators may influence carnivore communities through the suppression of competitively dominant mesopredators, however they also provide carrion subsidies that could influence foraging and competition among sympatric mesopredators when small prey is scarce. We assessed coyote Canis latrans and red fox Vulpes vulpes winter diet overlap and composition from scats collected in two study areas with 3‐fold difference in grey wolf Canis lupus density due to a wolf control program. We hypothesized that differences in diet composition would be driven by the use of carrion, and tested whether 1) apex predators facilitate resource overlap, or 2) apex predators facilitate resource partitioning. We estimated the available biomass of snowshoe hares and voles based on pellet density and vole capture rates in each study area. We used molecular analysis to confirm species identification of predator scats, and used microscopic evaluation of prey remains to analyze diet composition of 471 coyote and fox scats. Ungulate carrion, voles and snowshoe hares comprised 73% of coyote and fox diet, and differences in use of carrion and microtines accounted for nearly 60% of the dissimilarity in diet among these canids. Carrion was the top‐ranked item in the coyote diet in both study areas, whereas carrion use by red foxes declined 3‐fold in the study area with higher wolf and small prey abundance. Diet overlap tended to be lower and diet diversity tended to be higher where wolves were more abundant, though these trends were not statistically significant. Taken together, our findings indicate that carrion provisions could facilitate resource partitioning in mesocarnivore communities by alleviating exploitation competition for small mammals.     

Prey preferences and recent changes in diet of a breeding population of the Northern Goshawk Accipiter gentilis in Southwestern Europe

Capsule: Northern Goshawk Accipiter gentilis diet has changed significantly since the 1980s, probably due to changes in populations of preferred prey species.

Aims and methods: To assess changes to the breeding season diet of the Northern Goshawk in southwest Europe over three decades. We examined prey remains at and around nests and assessed avian prey availability using point count and line transect surveys.

Results: During 2008–11, Goshawks mainly ate birds, with Feral Pigeons Columba livia f. domestica being the most important prey species. Goshawks preferred prey of 100?400?g and forest prey species to non-forest species. Goshawk diet has changed significantly over recent decades: 22% of current prey items belong to species that were not part of the diet in the 1980s. We suggest that these dietary changes reflect changes in the abundance of prey species of the preferred size caused by changes in land use leading to an increase in forest cover, new prey species colonization and changes in the abundance and management of domestic prey.

Conclusion: This study emphasizes that major transformations occurring in agroforestry systems are affecting the main preferred prey of important forest predators, which may have consequences for conservation of both the predators and their prey.