Birds bias offspring sex ratio in response to livestock grazing.

Livestock grazing, which has a large influence on habitat structure, is associated with the widespread decline of various bird species across the world, yet there are few experimental studies that investigate how grazing pressure influences avian reproduction. We manipulated grazing pressure using a replicated field experiment, and found that the offspring sex ratio of a common upland passerine, the meadow pipit Anthus pratensis, varied significantly between grazing treatments. The proportion of sons was lowest in the ungrazed and intensively grazed treatments and highest in treatments grazed at low intensity (by sheep, or a mixture of sheep and cattle). This response was not related to maternal body condition. These results demonstrate the sensitivity of avian reproductive biology to variation in local conditions, and support growing evidence that too much grazing, or the complete removal of livestock from upland areas, is detrimental for common breeding birds.

The functional response of a generalist predator.

BACKGROUND: Predators can have profound impacts on the dynamics of their prey that depend on how predator consumption is affected by prey density (the predator's functional response). Consumption by a generalist predator is expected to depend on the densities of all its major prey species (its multispecies functional response, or MSFR), but most studies of generalists have focussed on their functional response to only one prey species. METHODOLOGY AND PRINCIPAL FINDINGS: Using Bayesian methods, we fit an MSFR to field data from an avian predator (the hen harrier Circus cyaneus) feeding on three different prey species. We use a simple graphical approach to show that ignoring the effects of alternative prey can give a misleading impression of the predator's effect on the prey of interest. For example, in our system, a "predator pit" for one prey species only occurs when the availability of other prey species is low. CONCLUSIONS AND SIGNIFICANCE: The Bayesian approach is effective in fitting the MSFR model to field data. It allows flexibility in modelling over-dispersion, incorporates additional biological information into the parameter priors, and generates estimates of uncertainty in the model's predictions. These features of robustness and data efficiency make our approach ideal for the study of long-lived predators, for which data may be sparse and management/conservation priorities pressing.

Effect of local cultural context on the success of community-based conservation interventions.

Conservation interventions require evaluation to understand what factors predict success or failure. To date, there has been little systematic investigation of the effect of social and cultural context on conservation success, although a large body of literature argues it is important. We investigated whether local cultural context, particularly local institutions and the efforts of interventions to engage with this culture significantly influence conservation outcomes. We also tested the effects of community participation, conservation education, benefit provision, and market integration. We systematically reviewed the literature on community-based conservation and identified 68 interventions suitable for inclusion. We used a protocol to extract and code information and evaluated a range of measures of outcome success (attitudinal, behavioral, ecological, and economic). We also examined the association of each predictor with each outcome measure and the structure of predictor covariance. Local institutional context influenced intervention outcomes, and interventions that engaged with local institutions were more likely to succeed. Nevertheless, there was limited support for the role of community participation, conservation education, benefit provision, and market integration on intervention success. We recommend that conservation interventions seek to understand the societies they work with and tailor their activities accordingly. Systematic reviews are a valuable approach for assessing conservation evidence, although sensitive to the continuing lack of high-quality reporting on conservation interventions.

Can parasites drive population cycles in mountain hares?

Understanding the drivers of population fluctuations is a central goal of ecology. Although well-established theory suggests that parasites can drive cyclic population fluctuations in their hosts, field evidence is lacking. Theory predicts that a parasite that loosely aggregates in the host population and has stronger impact on host fecundity than survival should induce cycling. The helminth Trichostrongylus retortaeformis in the UK's only native lagomorph, the mountain hare, has exactly these properties, and the hares exhibit strong population fluctuations. Here we use a host-parasite model parametrized using the available empirical data to test this superficial concordance between theory and observation. In fact, through an innovative combination of sensitivity and stability analyses, we show that hare population cycles do not seem to be driven by the parasite. Potential limitations in our parametrization and model formulation, together with the possible secondary roles for parasites in determining hare demography, are discussed. Improving our knowledge of leveret biology and the quantification of harvesting emerge as future research priorities. With the growing concern over the present management of mountain hares for disease control in Scotland, understanding their population drivers is an important prerequisite for the effective management of this species.

Sensitivity to assumptions in models of generalist predation on a cyclic prey.

Ecological theory predicts that generalist predators should damp or suppress long-term periodic fluctuations (cycles) in their prey populations and depress their average densities. However, the magnitude of these impacts is likely to vary depending on the availability of alternative prey species and the nature of ecological mechanisms driving the prey cycles. These multispecies effects can be modeled explicitly if parameterized functions relating prey consumption to prey abundance, and realistic population dynamical models for the prey, are available. These requirements are met by the interaction between the Hen Harrier (Circus cyaneus) and three of its prey species in the United Kingdom, the Meadow Pipit (Anthus pratensis), the field vole (Microtus agrestis), and the Red Grouse (Lagopus lagopus scoticus). We used this system to investigate how the availability of alternative prey and the way in which prey dynamics are modeled might affect the behavior of simple trophic networks. We generated cycles in one of the prey species (Red Grouse) in three different ways: through (1) the interaction between grouse density and macroparasites, (2) the interaction between grouse density and male grouse aggressiveness, and (3) a generic, delayed density-dependent mechanism. Our results confirm that generalist predation can damp or suppress grouse cycles, but only when the densities of alternative prey are low. They also demonstrate that diametrically opposite indirect effects between pairs of prey species can occur together in simple systems. In this case, pipits and grouse are apparent competitors, whereas voles and grouse are apparent facilitators. Finally, we found that the quantitative impacts of the predator on prey density differed among the three models of prey dynamics, and these differences were robust to uncertainty in parameter estimation and environmental stochasticity.

Unstable dynamics and population limitation in mountain hares.

The regular large-scale population fluctuations that characterize many species of northern vertebrates have fascinated ecologists since the time of Charles Elton. There is still, however, no clear consensus on what drives these fluctuations. Throughout their circumpolar distribution, mountain hares Lepus timidus show regular and at times dramatic changes in density. There are distinct differences in the nature, amplitude and periodicity of these fluctuations between regions and the reasons for these population fluctuations and the geographic differences remain largely unknown. In this review we synthesize knowledge on the factors that limit or regulate mountain hare populations across their range in an attempt to identify the drivers of unstable dynamics. Current knowledge of mountain hare population dynamics indicates that trophic interactions--either predator-prey or host-parasite--appear to be the major factor limiting populations and these interactions may contribute to the observed unstable dynamics. There is correlative and experimental evidence that some mountain hare populations in Fennoscandia are limited by predation and that predation may link hare and grouse cycles to microtine cycles. Predation is unlikely to be important in mountain hare populations in Scotland as most hares occur on sporting estates where predators are controlled, but this hypothesis remains to be experimentally tested. There is, however, emerging experimental evidence that some Scottish mountain hare populations are limited by parasites and that host-parasite interactions contribute to unstable dynamics. By contrast, there is little evidence from Fennoscandia that parasitism is of any importance to mountain hare population dynamics, although disease may cause periodic declines. Although severe weather and food limitation may interact to cause periodic high winter mortality there is little evidence that food availability limits mountain hare populations. There is a paucity of information concerning the factors limiting or regulating mountain hare populations in the Alps of Central Europe or in the tundra and taiga belts of Russia. Future research on mountain hare population dynamics should focus on the interactions between predation, parasitism and nutrition with stochastic factors such as climate and anthropogenic management including harvesting.

Group size effects on foraging and vigilance in migratory Tibetan antelope.

Large group sizes have been hypothesized to decrease predation risk and increase food competition. We investigated group size effects on vigilance and foraging behaviour during the migratory period in female Tibetan antelope Pantholops hodgsoni, in the Kekexili Nature Reserve of Qinghai Province, China. During June to August, adult female antelope and yearling females gather in large migratory groups and cross the Qinghai-Tibet highway to calving grounds within the Nature Reserve and return to Qumalai county after calving. Large groups of antelope aggregate in the migratory corridor where they compete for limited food resources and attract the attention of mammalian and avian predators and scavengers. We restricted our sampling to groups of less than 30 antelopes and thus limit our inference accordingly. Focal-animal sampling was used to record the behaviour of the free-ranging antelope except for those with lambs. Tibetan antelope spent more time foraging in larger groups but frequency of foraging bouts was not affected by group size. Conversely, the time spent vigilant and frequency of vigilance bouts decreased with increased group size. We suggest that these results are best explained by competition for food and risk of predation.

Long-term ecosystem dynamics in the Serengeti: lessons for conservation.

Data from long-term ecological studies further understanding of ecosystem dynamics and can guide evidence-based management. In a quasi-natural experiment we examined long-term monitoring data on different components of the Serengeti-Mara Ecosystem to trace the effects of disturbances and thus to elucidate cause-and-effect connections between them. The long-term data illustrated the role of food limitation in population regulation in mammals, particularly in migratory wildebeest and nonmigratory buffalo. Predation limited populations of smaller resident ungulates and small carnivores. Abiotic events, such as droughts and floods, created disturbances that affected survivorship of ungulates and birds. Such disturbances showed feedbacks between biotic and abiotic realms. Interactions between elephants and their food allowed savanna and grassland communities to co-occur. With increased woodland vegetation, predators' capture of prey increased. Anthropogenic disturbances had direct (hunting) and indirect (transfer of disease to wildlife) effects. Slow and rapid changes and multiple ecosystem states became apparent only over several decades and involved events at different spatial scales. Conservation efforts should accommodate both infrequent and unpredictable events and long-term trends. Management should plan on the time scale of those events and should not aim to maintain the status quo. Systems can be self-regulating through food availability and predator-prey interactions; thus, culling may not be required. Ecosystems can occur in multiple states; thus, there may be no a priori need to maintain one natural state. Finally, conservation efforts outside protected areas must distinguish between natural change and direct human-induced change. Protected areas can act as ecological baselines in which human-induced change is kept to a minimum.

Serengeti wildebeest migratory patterns modeled from rainfall and new vegetation growth.

We used evolutionary programming to model innate migratory pathways of wildebeest in the Serengeti Mara Ecosystem, Tanzania and Kenya. Wildebeest annually move from the southern short-grass plains of the Serengeti to the northern woodlands of the Mara. We used satellite images to create 12 average monthly and 180 10-day surfaces from 1998 to 2003 of percentage rainfall and new vegetation. The surfaces were combined in five additive and three multiplicative models, with the weightings on rainfall and new vegetation from 0% to 100%. Modeled wildebeest were first assigned random migration pathways. In simulated generations, animals best able to access rainfall and vegetation were retained, and they produced offspring with similar migratory pathways. Modeling proceeded until the best pathway was stable. In a learning phase, modeling continued with the ten-day images in the objective function. The additive model, influenced 25% by rainfall and 75% by vegetation growth, yielded the best agreement, with a multi-resolution comparison to observed densities yielding 76.8% of blocks in agreement (kappa = 0.32). Agreement was best for dry season and early wet season (kappa = 0.22-0.57), and poorest for the late wet season (0.04). The model suggests that new forage growth is a dominant correlate of wildebeest migration.

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.

Prevalence, intensity and aggregation of intestinal parasites in mountain hares and their potential impact on population dynamics.

Mountain hare Lepus timidus populations show unstable dynamics and since hares carry a significant helminth infection and host-parasite interactions are known to be destabilising, they have been proposed as a possible causal mechanism for the observed instability. We assessed the prevalence, intensity of infection and aggregation of the helminth parasites Graphidium strigosum and Trichostrongylus retortaeformis recovered from 589 mountain hares culled from 30 Scottish sporting estates in 1999 and 2000. Graphidium strigosum showed low prevalence and intensity of infection and was highly aggregated. In contrast, T. retortaeformis showed high prevalence and intensity of infection and a low degree of aggregation. Differences in body condition of the hares were best explained by a model including sex and month of collection and interaction terms for sex-month and intensity of infection of T. retortaeformis-month. The low degree of aggregation of T. retortaeformis and the significant negative effect of intensity of infection on body condition are in accordance with the hypothesis that the host-parasite interaction is the causative destabilising mechanism for mountain hare dynamics.

Parasite-mediated reduction in fecundity of mountain hares.

Mountain hare populations in Scotland exhibit regular 10 year fluctuations in abundance. Simple models of host-parasite population dynamics suggest that parasite-mediated reductions in host fecundity can cause a transition from stable to cyclic host population dynamics. We tested the hypothesis that parasites reduce hare fecundity by experimentally reducing parasite burdens and recording female survival, body condition and fecundity. We captured 41 adult female hares in October 2002; 22 were treated with Ivermectin to remove parasites and 19 were left untreated as controls. The treated and untreated hares were culled in May 2003 together with a second control group of nine unhandled hares. Treatment with Ivermectin significantly reduced the abundance of Trichostrongylus retortaeformis and increased-the fecundity of the hares, but had no measurable effect on body condition or over-winter survival. These results are consistent with the hypothesis that parasites may be a contributory cause of cycles in populations of mountain hares.

Analysing noisy time-series: describing regional variation in the cyclic dynamics of red grouse.

We develop a method for describing the periodicity of noisy 'quasi-cyclic' time-series based on integrals of their power spectra corresponding to different frequency intervals that we use to classify time-series as 'strongly cyclic', 'weakly cyclic' or 'non-cyclic'. We apply this analysis to over 300 time-series of shooting records of red grouse from 289 moors located in 20 regions of the UK. Time-series from 63 of these populations were not distinguishable from white noise, but significant evidence of cyclic behaviour in the 2-15 year range was detected in time-series from 183 other populations. Time-series from the remaining 43 populations, though distinguishable from white noise, did not exhibit consistently recognizable cyclic behaviour in the same period range. Cyclic populations exhibit an average periodicity of 8.3 years, but only 20% of these populations cycle with a period of four to six years. Geographically, grouse populations are remarkable more for their dynamic heterogeneity than for any observable regularity. The relationship between the location of populations and their dynamical behaviour is weak. The prevalence of cyclic time-series within a region did not significantly differ from the overall average value. Moor region explained 22% of the variation in periodicity, differing from the overall mean in three regions. Average periodicity increases significantly from 6.8 to 8.9 years from the most southerly to most northerly populations. However, latitude explains only 5.3% of the variation in periodicity of the cycles.