Associations between abundances of free-roaming gamebirds and common buzzards Buteo buteo are not driven by consumption of gamebirds in the buzzard breeding season.

Releasing gamebirds in large numbers for sport shooting may directly or indirectly influence the abundance, distribution and population dynamics of native wildlife. The abundances of generalist predators have been positively associated with the abundance of gamebirds. These relationships have implications for prey populations, with the potential for indirect impacts of gamebird releases on wider biodiversity. To understand the basis of these associations, we investigated variation in territory size, prey provisioning to chicks, and breeding success of common buzzards Buteo buteo, and associations with variation in the abundances of free-roaming gamebirds, primarily pheasants Phasianus colchicus, and of rabbits Oryctolagus cuniculus and field voles Microtus agrestis, as important prey for buzzards. The relative abundance of gamebirds, but not those of rabbits or voles, was weakly but positively correlated with our index of buzzard territory size. Gamebirds were rarely brought to the nest. Rabbits and voles, and not gamebirds, were provisioned to chicks in proportion to their relative abundance. The number of buzzard chicks increased with provisioning rates of rabbits, in terms of both provisioning frequency and biomass, but not with provisioning rates for gamebirds or voles. Associations between the abundances of buzzards and gamebirds may not be a consequence of the greater availability of gamebirds as prey during the buzzard breeding season. Instead, the association may arise either from habitat or predator management leading to higher densities of alternative prey (in this instance, rabbits), or from greater availability of gamebirds as prey or carrion during the autumn and winter shooting season. The interactions between gamebird releases and associated practices of predator control and shooting itself require better understanding to more effectively intervene in any one aspect of this complex social-ecological system.

Broadening the toolset for stakeholder engagement to explore consensus over wolf management.

Facilitating coexistence between people and large carnivores is critical for large carnivore conservation in human-dominated landscapes, when their presence impacts negatively on human interests. Such situations will often require novel ways of mediating between different values, worldviews and opinions about how carnivores should be managed. We report on such a process in an agricultural area of recent wolf recovery in central Italy where unsolved social tensions over wolf presence have radicalized opinions on either side of the wolf debate, resulting in a stalemate. Where previous mitigation policies based on top-down damage compensation have failed, we tested the potential for applying a participatory approach to engage different stakeholder groups in a dialogue aimed at sharing a deep understanding of the problem and co-creating potential solutions. We based our approach on the theory of meta-consensus, using a decision support tool known as Multi Criteria Decision Analysis (MCDA). Over the course of three months, we carried out five workshops with stakeholder representatives from farming, hunting and environmental associations, and one biologist. Stakeholders shared several objectives and agreed over many management interventions, including the management of free-ranging dogs, the implementation of damage prevention measures, and a damage compensation system suitable for farmers. The process facilitated agreement over actions aimed at improving relations between stakeholders and enhancing the state of knowledge on the issues at stake. Most importantly, we recorded positive social and relationship outcomes from the workshops, and observed a willingness from participants to engage in further discussions over disputed management preferences. Overall, we found MCDA to be a useful tool for laying the groundwork for further participatory and deliberative processes on wolf management. However, challenges ahead included the involvement of a larger number of representatives of different social sectors, and a simplification of the methodology which some participants found too complicated and time consuming.

Patterns of satellite tagged hen harrier disappearances suggest widespread illegal killing on British grouse moors.

Identifying patterns of wildlife crime is a major conservation challenge. Here, we test whether deaths or disappearances of a protected species, the hen harrier, are associated with grouse moors, which are areas managed for the production of red grouse for recreational shooting. Using data from 58 satellite tracked hen harriers, we show high rates of unexpected tag failure and low first year survival compared to other harrier populations. The likelihood of harriers dying or disappearing increased as their use of grouse moors increased. Similarly, at the landscape scale, satellite fixes from the last week of life were distributed disproportionately on grouse moors in comparison to the overall use of such areas. This pattern was also apparent in protected areas in northern England. We conclude that hen harriers in Britain suffer elevated levels of mortality on grouse moors, which is most likely the result of illegal killing.

Impact of wild prey availability on livestock predation by snow leopards.

An increasing proportion of the world's poor is rearing livestock today, and the global livestock population is growing. Livestock predation by large carnivores and their retaliatory killing is becoming an economic and conservation concern. A common recommendation for carnivore conservation and for reducing predation on livestock is to increase wild prey populations based on the assumption that the carnivores will consume this alternative food. Livestock predation, however, could either reduce or intensify with increases in wild prey depending on prey choice and trends in carnivore abundance. We show that the extent of livestock predation by the endangered snow leopard Panthera uncia intensifies with increases in the density of wild ungulate prey, and subsequently stabilizes. We found that snow leopard density, estimated at seven sites, was a positive linear function of the density of wild ungulates-the preferred prey-and showed no discernible relationship with livestock density. We also found that modelled livestock predation increased with livestock density. Our results suggest that snow leopard conservation would benefit from an increase in wild ungulates, but that would intensify the problem of livestock predation for pastoralists. The potential benefits of increased wild prey abundance in reducing livestock predation can be overwhelmed by a resultant increase in snow leopard populations. Snow leopard conservation efforts aimed at facilitating increases in wild prey must be accompanied by greater assistance for better livestock protection and offsetting the economic damage caused by carnivores.

Parasitized mates increase infection risk for partners.

Individuals can gain fitness benefits and costs through their mates. However, studies on sexual selection have tended to focus on genetic benefits. A potentially widespread cost of pairing with a parasitized mate is that doing so will increase an individual's parasite abundance. Such a cost has been overlooked in systems in which parasites are indirectly transmitted. We manipulated the abundance of the nematode parasite Trichostrongylus tenuis, an indirectly transmitted parasite, within pairs of wild red grouse Lagopus lagopus scoticus in spring. Parasite levels were correlated within pairs before the experiment. We removed parasites from males, females, or both members of the pair and evaluated individual parasite uptake over the subsequent breeding period. At the end of the breeding season, an individual's parasite abundance was greater when its mate had not been initially purged of parasites. This cost appeared to be greater for males. We discuss the implications of our results in relation to the costs that parasites may have on sexual selection processes.

The ornament-condition relationship varies with parasite abundance at population level in a female bird.

Environmental heterogeneity is expected to create variation in the ornament-condition relationship. This topic has been studied in males with less attention being given to females. Here, we explore inter-population variation in the relationship between the size of a male-like trait, supra-orbital combs, and body mass in female red grouse Lagopus lagopus scoticus. We used the abundance of the nematode Trichostrongylus tenuis, a parasite with strong negative effects on this species, as a proxy of environmental conditions. We studied six populations over 5 years and showed that the comb size-body mass relationship varied with mean parasite abundance, with stronger ornament-condition relationships in populations with higher parasite infection levels. Our study supports the idea that environmental conditions, and in particular parasite infection levels, may affect the reliability of female ornaments as condition indicators.

Low intensity, mixed livestock grazing improves the breeding abundance of a common insectivorous passerine.

Livestock grazing is a major driver of ecosystem change and has been associated with significant declines in various bird species in Britain and worldwide. However, there is little experimental evidence to show how grazing affects bird populations. We manipulated livestock densities in a replicated field experiment and found that mixed sheep and cattle grazing, at low intensity, improved the breeding abundance of a common upland passerine, the meadow pipit Anthus pratensis, after two years. Plots stocked with sheep alone (at high or low density) or not stocked at all held fewer pipit territories. Despite a year-on-year decline in pairs of meadow pipits in intensively grazed plots, we found no effect of sheep number on breeding abundance. Our results support the hypothesis that mixed species of herbivores generate greater heterogeneity in vegetation structure, which modifies prey availability, resulting in a greater abundance of birds. The results of our study should inform the management of grassland areas and enhance the abundance of some bird species, particularly in areas that have seen significant shifts from mixed livestock grazing to grazing dominated by single species of animals.

Testing the role of parasites in driving the cyclic population dynamics of a gamebird.

The role of parasites in regulating populations has been the subject of debate. We tested whether parasites caused population cycles in red grouse by manipulating parasite intensities in four, paired 1 km(2) study areas during cyclic population declines over 4 years. Parasite reductions led to (1) larger grouse broods, (2) higher population densities in both autumn and spring, (3) reduced autumn population declines in one of two regions, and (4) reduced spring declines, but only in the first year. We infer that a single trophic interaction between a parasite and its host does not explain cyclic dynamics in spring breeding density in this species, although it contributed to the start of a cyclic decline. Another process was operating to drive the populations down. Together with our other results these findings emphasize that both trophic and intrinsic processes may act within populations to cause unstable dynamics.

Interactions between intrinsic and extrinsic mechanisms in a cyclic species: testosterone increases parasite infection in red grouse.

Field studies of mechanisms involved in population regulation have tended to focus on the roles of either intrinsic or extrinsic factors, but these are rarely mutually exclusive and their interactions can be crucial in determining dynamics. Experiments on red grouse Lagopus lagopus scoticus have shown that population instability can be caused both by the effects of a parasitic nematode, Trichostrongylus tenuis, on host production or by changes in testosterone influencing aggressive behaviour and recruitment. We experimentally tested for an interaction between testosterone and T. tenuis in free-living male grouse. A total of 123 grouse were caught in autumn, treated with an anthelmintic to remove parasites, and then given either testosterone or empty, control, implants. After one month grouse were re-infected with a standard dose of parasites. We show that males with increased testosterone levels had greater parasite intensities than controls after one year. We discuss possible physiological and behavioural mechanisms linking testosterone and increased parasite intensity, and the implications for our understanding of complex, unstable population dynamics.

Separating behavioral and physiological mechanisms in testosterone-mediated trade-offs.

Testosterone often mediates trade-offs between reproduction and other life-history traits, which are usually investigated using testosterone implants. However, this approach does not distinguish between the physiological and behavioral effects of testosterone. We studied a wild game bird, the red grouse Lagopus lagopus scoticus, and took a new approach to investigate mechanisms linking elevated testosterone to increased parasite intensity. We caught males in autumn, removed their parasites, implanted them with the antiandrogen flutamide in combination with an aromatase inhibitor (FA males) or with empty implants (control males), and challenged them with parasites. The FA treatment increased testosterone concentration and physiological stress, but without enhancing testosterone-dependent behaviors, because testosterone receptors were blocked. FA males ended up with more parasites than the control males the following autumn, an effect similar to that of a testosterone treatment reported elsewhere. However, and unlike the testosterone treatment, the FA treatment did not affect home range, pairing, or breeding success. The results supported a physiological mechanism (increased susceptibility) linking elevated testosterone and increased parasite intensity. The FA treatment provided a new way of investigating testosterone-mediated trade-offs whereby testosterone concentration was increased while the effects on behavior were blocked, resulting in physiological costs without phenotypic benefits.

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.

Interactions between population processes in a cyclic species: parasites reduce autumn territorial behaviour of male red grouse.

The causes of population cycles fascinate and perplex ecologist. Most work have focused on single processes, whether extrinsic or intrinsic, more rarely on how different processes might interact to cause or mould the unstable population dynamics. In red grouse (Lagopus lagopus scoticus), two causal mechanisms have been supported: territorial behaviour (changes in autumn aggressiveness) and parasites (parasite induced reduction in fecundity). Here, we report on how these two regulatory processes might interact, by testing whether the parasite suspected to cause the grouse cycles, the nematode Trichostrongylus tenuis, reduces male autumn territorial behaviour. We either treated males with an anthelmintic, to remove parasites (dosed or D-males), or challenged them with infective T. tenuis larvae, to increase parasite intensity (challenged or C-males). We first show that dosing was effective in removing T. tenuis parasites, while parasite intensities increased in challenged birds during the autumn. Because old males initially had more parasites than young males, the treatments generated greater differences in parasite intensity in old than in young males. We also show that various aspects of territorial behaviour (increase in testosterone-dependent comb size in autumn, territorial call rate, likelihood of winning territorial interactions and over-winter survival) were significantly higher in dosed than in challenged males, but in old birds only. Our data thus supported the hypothesis that parasites reduce male aggressiveness during the autumn territorial contests, and could thereby influence recruitment. Our results also highlight that the territorial behaviour of young males, which have fewer parasites, is not as limited by parasites as that of old, previously territorial males. We discuss the implications of these findings for our understanding of the processes regulating red grouse populations and causing their complex, unstable population dynamics.

Livestock grazing affects the egg size of an insectivorous passerine.

Livestock grazing is a major driver of ecosystem change, and has been associated with significant declines in various bird species worldwide. In Britain, there is particular concern that severe grazing pressure is deleteriously affecting vegetation and birds in upland regions. However, the mechanism by which grazing affects birds is unclear. Here, we report for the first time, to our knowledge, that sheep grazing pressure affects the egg size of a common upland passerine: the meadow pipit Anthus pratensis. We manipulated sheep stocking densities in a replicated field experiment, and found that plots with the highest stocking density contained nests with the smallest eggs, and that plots with low stocking density contained nests with the largest eggs. However, eggs laid in ungrazed plots were also small, suggesting that either too many sheep or their removal from upland areas might have a detrimental effect on pipit egg size. We found no significant effect on fledging success but the reduced post-fledging survival of young from smaller eggs, as seen in other studies, could partly explain declines in upland birds.