Diet,nesting density,and breeding success of rough-legged buzzards (Buteo lagopus) on the Nenetsky Ridge,Arctic Russia

The rough-legged buzzard, a circumpolar avian predator, is usually defined as rodent specialist in the tundra but as a generalist in the boreal zone, leaving open the question of where the shift in feeding strategy occurs. Here, we investigated the diet and breeding biology of buzzards as well as abundance of their possible prey during 5 years in the low-Arctic shrub tundra on the Nenetsky Ridge, Russia. We employed three complementary methods to assess the diet of this Arctic predator—pellet dissection, identification of prey remains on nests, and stable isotope analysis—to overcome their respective limitations. We documented fluctuations in abundances of the likely prey, namely rodents, ptarmigans, and hares. Nesting density of buzzards changed substantially over the years, but did not track the abundance cycle of the rodents. The number of buzzard fledglings was relatively low (1.08 ± 0.3) and did not change according to the density of rodents. In the year when rodents were at their lowest abundance, diet analyses of nestlings documented a shift from rodents to alternative prey, with a decrease in the proportion of tundra voles and an increase in proportion of hares, ptarmigans, and ducks. Here, we argue that buzzards may adopt different feeding strategies along the gradient from generalists to specialists. While the rough-legged buzzard is usually considered a small rodent specialist, our study shows that it can shift to alternative prey where or when rodents are scarce and when alternative prey are sufficiently abundant to provide subsistence for breeding.     

Impacts of birds of prey on gamebirds in the UK: a review

The influence of predators on the distribution, density and dynamics of their prey species has long been of interest to ecologists and wildlife managers. Where the prey population is also utilized by humans, conflicts may arise through competition for a limited resource. Because gamebird shooting in the UK provides employment, recreation and income, the impact of birds of prey on gamebird populations has been the subject of intense debate for many years. Various approaches have been used to assess the impacts that raptors have on gamebird populations. Here we review the applicability and limitations of the methods used and assess the scientific evidence for population-level and economic impacts of raptors on gamebird populations in the UK. Raptors may, in some situations, take large numbers of gamebirds and may be an important proximate cause of mortality, although few studies have addressed the impacts of raptors on either breeding or pre-shooting densities. Two exceptions are studies of Hen Harrier Circus cyaneus , Peregrine Falcon Falco peregrinus predation on Red Grouse Lagopus lagopus scoticus on moorland in Scotland and Sparrowhawk Accipiter nisus predation on Grey Partridge Perdix perdix on farmland in England. Both these studies suggested that raptors could have population-level impacts when their gamebird prey was already at low density. Studies on predation of captively bred gamebirds suggest that numbers taken by raptors at release pens vary considerably and in a few cases raptors have been documented killing relatively large numbers. On the whole, however, it appears that raptors account for a relatively small proportion of mortality among released birds and the impact on subsequent shooting bags is unknown. We summarize important gaps in current knowledge and recommend specific areas for future research.     

Birds of prey as limiting factors of gamebird populations in Europe: a review

Whether predators can limit their prey has been a topic of scientific debate for decades. Traditionally it was believed that predators take only wounded, sick, old or otherwise low-quality individuals, and thus have little impact on prey populations. However, there is increasing evidence that, at least under certain circumstances, vertebrate predators may indeed limit prey numbers. This potential role of predators as limiting factors of prey populations has created conflicts between predators and human hunters, because the hunters may see predators as competitors for the same resources. A particularly acute conflict has emerged over the past few decades between gamebird hunters and birds of prey in Europe. As a part of a European-wide research project, we reviewed literature on the relationships between birds of prey and gamebirds. We start by analysing available data on the diets of 52 European raptor and owl species. There are some 32 species, mostly specialist predators feeding on small mammals, small passerine birds or insects, which never or very rarely include game animals (e.g. hares, rabbits, gamebirds) in their diet. A second group (20 species) consists of medium-sized and large raptors which prey on game, but for which the proportion in the diet varies temporally and spatially. Only three raptor species can have rather large proportions of gamebirds in their diet, and another seven species may utilise gamebirds locally to a great extent. We point out that the percentage of a given prey species in the diet of an avian predator does not necessarily reflect the impact of that predator on densities of prey populations. Next, we summarise available data on the numerical responses of avian predators to changing gamebird numbers. In half of these studies, no numerical response was found, while in the remainder a response was detected such that either raptor density or breeding success increased with density of gamebirds. Data on the functional responses of raptors were scarce. Most studies of the interaction between raptors and gamebird populations give some estimate of the predation rate (per cent of prey population taken by predator), but less often do they evaluate the subsequent reduction in the pre-harvest population or the potential limiting effect on breeding numbers. The few existing studies indicate that, under certain conditions, raptor predation may limit gamebird populations and reduce gamebird harvests. However, the number and extent of such studies are too modest to draw firm conclusions. Furthermore, their geographical bias to northern Europe, where predator-prey communities are typically simpler than in the south, precludes extrapolation to more diverse southern European ecosystems. There is an urgent need to develop further studies, particularly in southern Europe, to determine the functional and numerical responses of raptors to gamebird populations in species and environments other than those already evaluated in existing studies. Furthermore, additional field experiments are needed in which raptor and possibly also mammalian predator numbers are manipulated on a sufficiently large spatial and temporal scale. Other aspects that have been little studied are the role of predation by the non-breeding part of the raptor population, or floaters, on the breeding success and survival of gamebirds, as well as the effect of intra-guild predation. Finally there is a need for further research on practical methods to reduce raptor predation on gamebirds and thus reduce conflict between raptor conservation and gamebird management.     

Diet variation of common buzzards in Finland supports the alternative prey hypothesis

The regional synchrony of short-term population fluctuations of small rodents and small game has usually been explained by varying impacts of generalist predators subsisting on both voles and small game (the "alternative prey hypothesis" APH). APH says that densities of predators increase as a response to increasing vole densities and then these predators shift their diet from the main prey to the alternative prey when the main prey decline and vice versa. We studied the diet composition of breeding common buzzards Buteo buteo during 1985-92 in western Finland. Microtus voles were the main prey and water voles, shrews, forest grouse, hares and small birds the most important alternative prey. Our data from the between-year variation in the diet composition of buzzards fulfilled the main predictions of APH. The yearly proportion of main prey (Microtus voles) in the diet was higher in years of high than low vole abundance. The proportion of grouse in the diet of buzzards was negatively related to the abundance of Microtus voles in the field and was nearly independent of grouse abundance in the field. In addition, buzzards mainly took grouse chicks and young hares which is consistent with the prediction of APH. Therefore, we conclude that buzzards are able to shift their diet in the way predicted by the APH and that buzzards, together with other generalist predators, may reduce the breeding success of small game in the decline phase of the vole cycle, and thus substantially contribute to the existence of short-term population cycles of small game.     

Functional responses of the rough-legged buzzard in a multi-prey system

The functional response is a key element of predator–prey interactions. Basic functional response theory explains foraging behavior of individual predators, but many empirical studies of free-ranging predators have estimated functional responses by using population-averaged data. We used a novel approach to investigate functional responses of an avian predator (the rough legged-buzzard Buteo lagopus Pontoppidan, 1763) to intra-annual spatial variation in rodent density in subarctic Sweden, using breeding pairs as the sampling unit. The rough-legged buzzards responded functionally to Norwegian lemmings (Lemmus lemmus L. 1758), grey-sided voles (Myodes rufocanus Sundevall, 1846) and field voles (Microtus agrestis L. 1761), but different rodent prey were not utilised according to relative abundance. The functional response to Norwegian lemmings was a steep type II curve and a more shallow type III response to grey-sided voles. The different shapes of these two functional responses were likely due to combined effects of differences between lemmings and grey-sided voles in habitat utilisation, anti-predator behaviour and size-dependent vulnerability to predation. Diet composition changed less than changes in relative prey abundance, indicating negative switching, with high disproportional use of especially lemmings at low relative densities. Our results suggest that lemmings and voles should be treated separately in future empirical and theoretical studies in order to better understand the role of predation in this study system.     

Variation in the diet of red foxes on Scottish moorland in relation to prey abundance

The diet of red fox Vulpes vulpes was investigated through analysis of 340 scats collected during 1992-1996 from moorland in south-west Scotland. Rodents, game-birds, lagomorphs, carrion and insectivores were the most frequently occurring food types. Fox diet and prey abundance were compared between heather dominant and grass dominant habitat types within the study area. Rodents were the most frequently occurring food type in each habitat and occurred in 63% of seats overall. Rodents occurred more often in scats from grass dominant sites while gamebirds and lagomorphs were more frequent in scats from heather dominant sites. The occurrence of rodents in winter fox scats increased with rodent abundance. In contrast, the occurrence of gamebirds in winter fox scats was unrelated to gamebird abundance but negatively related to rodent abundance. It appeared that foxes switched to gamebirds in years, or habitats, where rodents were uncommon.     

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.     

Does variation of sex ratio enhance reproductive success of offspring in tawny owls (Strix aluco)

Tawny owls, Strix aluco, laid female-biased clutches on territories with more abundant prey (field voles) in June, the month that chicks fledge. This appeared to enhance the subsequent reproductive success of fledglings, as in 1995 there was a significant correlation between the number of chicks fledged by adult females and the June vole abundance in the territory on which they were reared as chicks. This relationship did not hold for males. Since tawny owls lay eggs in March, these results indicate that owls are able to predict the June vole numbers on their territory, and respond by producing more of the sex most likely to gain a long-term benefit when resources are good.     

Flexibility in a changing arctic food web: Can rough‐legged buzzards cope with changing small rodent communities?

Indirect effects of climate change are often mediated by trophic interactions and consequences for individual species depend on how they are tied into the local food web. Here we show how the response of demographic rates of an arctic bird of prey to fluctuations in small rodent abundance changed when small rodent community composition and dynamics changed, possibly under the effect of climate warming. We observed the breeding biology of rough‐legged buzzards (Buteo lagopus) at the Erkuta Tundra Monitoring Site in southern Yamal, low arctic Russia, for 19 years (1999–2017). At the same time, data on small rodent abundance were collected and information on buzzard diet was obtained from pellet dissection. The small rodent community experienced a shift from high‐amplitude cycles to dampened fluctuations paralleled with a change in species composition toward less lemmings and more voles. Buzzards clearly preferred lemmings as prey. Breeding density of buzzards was positively related to small rodent abundance, but the shift in small rodent community lead to lower numbers relative to small rodent abundance. At the same time, after the change in small rodent community, the average number of fledglings was higher relative to small rodent abundance than earlier. These results suggest that the buzzard population adapted to a certain degree to the changes in the major resource, although at the same time density declined. The documented flexibility in the short‐term response of demographic rates to changes in structure and dynamics of key food web components make it difficult to predict how complex food webs will be transformed in a warmer Arctic. The degree of plasticity of functional responses is indeed likely to vary between species and between regions, depending also on the local food web context.     

Numerical responses of an avian predator to prey fluctuations in a temperate latitude: breeders vs. entire population

Distinct numerical responses of predators to fluctuations in the abundance of their prey are often observed in northern regions but occur more rarely in temperate latitudes. This statement is, however, mostly based on observations of breeding populations, while in some predators, for example in raptors, numerous non-breeding floaters can occur. I estimated the breeding density and reproductive performance (nest survey) as well as the density of entire population (transects with distance sampling) of the common buzzard Buteo buteo in western Poland (52°N) in the years 2005–2014 to test the hypotheses that in temperate latitudes: (1) the breeding population of these birds does not show any numerical response, understood as annual changes in their abundance; (2) its reproductive success, however, changes with the abundance of main prey, the common vole Microtus arvalis; and (3) hence the entire buzzard population (including potential immature floaters) present in a given area during the nesting period responds numerically with some time delay. The reproductive success of buzzards was positively correlated with their prey abundance. Contrary to my predictions, however, the breeding population of buzzards showed a slight numerical response with a 3-year lag and the entire population tracked vole fluctuations without any time delay. The immediate numerical response of the entire buzzard population was probably caused by large-scale movements of floaters. Such rapid numerical responses of some predators may contribute to the relative stability of prey populations in temperate latitudes compared to northern regions.     

Predator abundance in relation to small game management in southern Portugal: conservation implications

The interaction between hunting interests and legally protected predators is often a contentious conservation problem, requiring detailed understanding of predator responses to game management. This issue was addressed in southern Portugal in a treatment-control natural experiment, whereby the abundances of small game, corvids, birds of prey and carnivores were compared in 12 game estates (>500 ha) and 12 matching areas with similar sizes and land uses but no game management. European rabbits (Oryctolagus cuniculus), Iberian hares (Lepus granatensis) and, less so, red-legged partridges (Alectoris rufa) were far more numerous in game estates than elsewhere. Among legally controlled species, there were less Eurasian jays (Garrulus glandarius) but more red foxes (Vulpes vulpes) in game estates, though the latter were primary targets of predator culling. Fox abundance within game estates varied inversely with an index of management intensity (density of small game feeding sites) and increased along with hare abundance. As for protected species, only common kestrels (Falco tinnunculus) and genets (Genetta genetta) were fewer in game estates. The abundance of raptors within game estates varied inversely with gamekeeper density, whereas that of common buzzards (Buteo buteo) increased along with rabbit abundance. Overall, there was little evidence that game management reduced local predator abundances, except in the most intensively managed estates. Game estates provided concentrations of prey that was scarce elsewhere, which may have favoured increased abundances of some predators. Further investigations are needed to find out whether high prey densities may attract predators to game estates with increased mortality risk, which may thus become population sinks for protected species.     

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.     

The effects of released pheasants on invertebrate populations in and around woodland release sites

The release of gamebirds for recreational shooting exerts a series of effects on the ecosystems into which they are placed. Pheasants (Phasianus colchicus) are omnivorous and eat invertebrates, especially when young or, if females, when breeding. Consequently, the release of large numbers of pheasants into woodland release pens may affect local invertebrate populations. Previous studies have reported mixed evidence. We conducted pitfall trapping at 13 sites (49 pens) in central England over 2 years (totaling 65 pen measures), comprising three surveys annually, immediately prior to releases in mid‐summer, 4 weeks later when most birds were still in the pens, and another 5 weeks later when most birds had dispersed. We compared traps inside and 25 m outside pens in the same wood. We considered release densities and whether the birds had prior experience of eating invertebrates. While accounting for overall seasonal declines in invertebrates trapped, we detected changes for total invertebrate biomass and total counts indicative of released pheasants causing local decreases inside pens, either directly by predation or indirectly by modifying vegetation. There were also relative decreases outside pens when the pheasants start to disperse, suggesting that the released pheasants may be affecting invertebrates in those nearby areas or that their earlier effects inside the pen, modifying vegetation or increasing invertebrate activity, increased the relative susceptibility of invertebrates there to trapping. However, these effects were not seen for specific invertebrate groups believed to be especially susceptible to pheasants. For slugs and detritivores, we detected small population increases inside pens. Across the study period, decreases for most measures were greater outside pens than inside them. We conclude that any effects pheasants have on invertebrate communities at release sites in woodlands are complex and that long‐term and taxon‐specific studies should be considered to understand the local net ecological effects of gamebird releases on invertebrates.     

Coping with fast climate change in northern ecosystems: mechanisms underlying the population‐level response of a specialist avian predator

Northern ecosystems are facing unprecedented climate modifications, which pose a major threat for arctic species, especially the specialist predator guild. However, the mechanisms underlying responses of predators to climate change remain poorly understood. Climate can influence fitness parameters of predators either through reduced reproduction or survival following adverse weather conditions, or via changes in the population dynamics of their main prey. Here, we combined three overlapping long‐term datasets on the breeding density and parameters of a rodent‐specialist predator, the rough‐legged buzzard Buteo lagopus, its main prey population dynamics and climate variables, collected in subarctic areas of Finland and Norway, to assess the impact of changing climate on the predator reproductive response. Rough‐legged buzzards responded to ongoing climate change by advancing their laying date (0.1 d yr^?1 over the 21 yr of the study period), as a consequence of earlier snowmelt. However, we documented for the same period a decrease in breeding success, which principally resulted from an indirect effect of changes in the dynamics of their main prey, i.e. grey‐sided voles Microtus oeconomus, and not from the expected negative effect of unfavorable weather conditions during the brood‐rearing period on nestling survival. Additionally, we showed the striking impact of autumn and winter weather conditions on vole population growth rates in subarctic ecosystems, with a strong positive correlation between mean snow depth in autumn and winter and both winter and summer population growth rates. Our results highlighted that, in northern ecosystems, ongoing climate change has the potential to impact specialist predator species through two mechanistic linkages, which may in the long‐run, threaten the viability of their populations, and lead to potential severe cascading trophic effects at the ecosystem level.     

Plasticity of Noddy Parents and Offspring to Sea-Surface Temperature Anomalies

Behavioral and/or developmental plasticity is crucial for resisting the impacts of environmental stressors. We investigated the plasticity of adult foraging behavior and chick development in an offshore foraging seabird, the black noddy (Anous minutus), during two breeding seasons. The first season had anomalously high sea-surface temperatures and ‘low’ prey availability, while the second was a season of below average sea-surface temperatures and ‘normal’ food availability. During the second season, supplementary feeding of chicks was used to manipulate offspring nutritional status in order to mimic conditions of high prey availability. When sea-surface temperatures were hotter than average, provisioning rates were significantly and negatively impacted at the day-to-day scale. Adults fed chicks during this low-food season smaller meals but at the same rate as chicks in the unfed treatment the following season. Supplementary feeding of chicks during the second season also resulted in delivery of smaller meals by adults, but did not influence feeding rate. Chick begging and parental responses to cessation of food supplementation suggested smaller meals fed to artificially supplemented chicks resulted from a decrease in chick demands associated with satiation, rather than adult behavioral responses to chick condition. During periods of low prey abundance, chicks maintained structural growth while sacrificing body condition and were unable to take advantage of periods of high prey abundance by increasing growth rates. These results suggest that this species expresses limited plasticity in provisioning behavior and offspring development. Consequently, responses to future changes in sea-surface temperature and other environmental variation may be limited.     

Kestrel-Prey Dynamic in a Mediterranean Region: The Effect of Generalist Predation and Climatic Factors

Background

Most hypotheses on population limitation of small mammals and their predators come from studies carried out in northern latitudes, mainly in boreal ecosystems. In such regions, many predators specialize on voles and predator-prey systems are simpler compared to southern ecosystems where predator communities are made up mostly of generalists and predator-prey systems are more complex. Determining food limitation in generalist predators is difficult due to their capacity to switch to alternative prey when the basic prey becomes scarce.

Methodology

We monitored the population density of a generalist raptor, the Eurasian kestrel Falco tinnunculus over 15 years in a mountainous Mediterranean area. In addition, we have recorded over 11 years the inter-annual variation in the abundance of two main prey species of kestrels, the common vole Microtus arvalis and the eyed lizard Lacerta lepida and a third species scarcely represented in kestrel diet, the great white-toothed shrew Crocidura russula. We estimated the per capita growth rate (PCGR) to analyse population dynamics of kestrel and predator species.

Principal Findings

Multimodel inference determined that the PCGR of kestrels was better explained by a model containing the population density of only one prey species (the common vole) than a model using a combination of the densities of the three prey species. The PCGR of voles was explained by kestrel abundance in combination with annual rainfall and mean annual temperature. In the case of shrews, growth rate was also affected by kestrel abundance and temperature. Finally, we did not find any correlation between kestrel and lizard abundances.

Significance

Our study showed for the first time vertebrate predator-prey relationships at southern latitudes and determined that only one prey species has the capacity to modulate population dynamics of generalist predators and reveals the importance of climatic factors in the dynamics of micromammal species and lizards in the Mediterranean region.     

Role of prey and intraspecific density dependence on the population growth of an avian top predator

Exploring predator–prey systems in diverse ecosystems increases our knowledge about ecological processes. Predator population growth may be positive when conspecific density is low but predators also need areas with prey availability, associated with competition, which increases the risk of suffering losses but stabilises populations. We studied relationships between European rabbits Oryctolagus cuniculus (prey) and adult eagle owls Bubo bubo (predators) in south-western Europe. We assessed models explaining the predator population growth and stability. We estimated the abundance of rabbits and adult eagle owls during three years in eight localities of central-southern Spain. We explored models including rabbit and adult eagle owl abundance, accounting for yearly variations and including the locality as a random variable. We found that population growth of adult eagle owls was positive in situations with low conspecific abundance and tended to be negative but approaching equilibrium in situations of higher conspecific abundance. Population growth was also positively related to previous summer rabbit density when taking into account eagle owl conspecific abundance, possibly indicating that rabbits may support recruitment. Furthermore, abundance stability of adult eagle owls was positively related to previous winter–spring rabbit density, which could suggest predator population stabilisation through quick territory occupation in high-quality areas. These results exemplify the trade-off between prey availability and abundance of adult predators related to population growth and abundance stability in the eagle owl–rabbit system in south-western Europe. Despite rabbits have greatly declined during the last decades and eagle owls locally specialise on them, eagle owls currently have a favourable conservation status. As eagle owls are the only nocturnal raptor with such dependence on rabbits, this could point out that predators may overcome prey decreases in areas with favourable climate and prey in the absence of superior competitors with similar foraging mode.     

Numerical and functional responses of Common Buzzards Buteo buteo to prey abundance on a Scottish grouse moor

Predators will often respond to reductions in preferred prey by switching to alternative prey resources. However, this may not apply to all alternative prey groups in patchy landscapes. We investigated the demographic and aggregative numerical and functional responses of Common Buzzards Buteo buteo in relation to variations in prey abundance on a moor managed for Red Grouse Lagopus lagopus scotica in south‐west Scotland over three consecutive breeding and non‐breeding seasons. We predicted that predation of Red Grouse by Buzzards would increase when abundance of their preferred Field Vole Microtus agrestis prey declined. As vole abundance fluctuated, Buzzards responded functionally by eating voles in relation to their abundance, but they did not respond demographically in terms of either breeding success or density. During a vole crash year, Buzzards selected a wider range of prey typical of enclosed farmland habitats found on the moorland edge but fewer Grouse from the heather moorland. During a vole peak year, prey remains suggested a linear relationship between Grouse density and the number of Grouse eaten (a Type 1 functional response), which was not evident in either intermediate or vole crash years. Buzzard foraging intensity varied between years as vole abundance fluctuated, and foraging intensity declined with increasing heather cover. Our findings did not support the prediction that predation of Red Grouse would increase when vole abundance was low. Instead, they suggest that Buzzards predated Grouse incidentally while hunting for voles, which may increase when vole abundances are high through promoting foraging in heather moorland habitats where Grouse are more numerous. Our results suggest that declines in their main prey may not result in increased predation of all alternative prey groups when predators inhabit patchy landscapes. We suggest that when investigating predator diet and impacts on prey, knowledge of all resources and habitats that are available to predators is important.     

Variation in predator species abundance can cause variable selection pressure on warning signaling prey

Predation pressure is expected to drive visual warning signals to evolve toward conspicuousness. However, coloration of defended species varies tremendously and can at certain instances be considered as more camouflaged rather than conspicuous. Recent theoretical studies suggest that the variation in signal conspicuousness can be caused by variation (within or between species) in predators' willingness to attack defended prey or by the broadness of the predators' signal generalization. If some of the predator species are capable of coping with the secondary defenses of their prey, selection can favor reduced prey signal conspicuousness via reduced detectability or recognition. In this study, we combine data collected during three large-scale field experiments to assess whether variation in avian predator species (red kite, black kite, common buzzard, short-toed eagle, and booted eagle) affects the predation pressure on warningly and non-warningly colored artificial snakes. Predation pressure varied among locations and interestingly, if common buzzards were abundant, there were disadvantages to snakes possessing warning signaling. Our results indicate that predator community can have important consequences on the evolution of warning signals. Predators that ignore the warning signal and defense can be the key for the maintenance of variation in warning signal architecture and maintenance of inconspicuous signaling.     

Vole fluctuations,red fox responses,predation on fawns,and roe deer dynamics in a temperate latitude

The alternative prey hypothesis predicts that predators respond both functionally and numerically (with a time lag) to fluctuations in the main prey abundance, which affects the survival of alternative prey. This pattern was found in northern Europe in the community formed by voles (Microtidae), red foxes (Vulpes vulpes) and roe deer (Capreolus capreolus). We studied the same predator—prey community in a temperate latitude where, according to the predation hypothesis, only the functional response of predators to changes in main prey availability should occur. In the years 1997–2007, in western Poland, we estimated the index of common vole (Microtus arvalis) abundance (burrow counts), the density of foxes (spotlight counts), the young production in foxes (young/adult ratio), the index of fox predation on fawns (prey remains near dens) as well as the reproduction index (fawn/female ratio) and density of roe deer (total counts). The vole abundance fluctuated considerably, the young production in foxes did not correlate with the main prey availability, but the density of foxes showed direct numerical response. The index of fox predation on fawns decreased with the vole abundance and negatively affected the fawn/female ratio in roe deer. Thus, the relationships between voles and foxes were not fully consistent with the predation hypothesis. The direct numerical response of foxes should tend to stabilize this predator—prey community. It is suggested, however, that responses showed by vole-eating predators in temperate latitudes may sometimes affect their alternative prey, including animals with unfavourable conservation status.