Spatio-temporal variation in the wintering associations of an alpine bird.

Many animals make behavioural changes to cope with winter conditions, being gregariousness a common strategy. Several factors have been invoked to explain why gregariousness may evolve during winter, with individuals coming together and separating as they trade off the different costs and benefits of living in groups. These trade-offs may, however, change over space and time as a response to varying environmental conditions. Despite its importance, little is known about the factors triggering gregarious behaviour during winter and its change in response to variation in weather conditions is poorly documented. Here, we aimed at quantifying large-scale patterns in wintering associations over 23 years of the white-winged snowfinch Montifringilla nivalis nivalis. We found that individuals gather in larger groups at sites with harsh wintering conditions. Individuals at colder sites reunite later and separate earlier in the season than at warmer sites. However, the magnitude and phenology of wintering associations are ruled by changes in weather conditions. When the temperature increased or the levels of precipitation decreased, group size substantially decreased, and individuals stayed united in groups for a shorter time. These results shed light on factors driving gregariousness and points to shifting winter climate as an important factor influencing this behaviour.

Effects of fertilization and irrigation on vascular plant species richness, functional composition and yield in mountain grasslands.

Land-use intensification is a major threat to biodiversity in agricultural grasslands and fertilization is one of the main drivers. The effects of fertilization on biodiversity and plant functional composition (community-weighted mean traits and mean ecological indicator values) are well studied in lowland regions, but have received less attention in mountain grasslands. Moreover, in inner-alpine dry valleys, fertilizer is often applied in combination with irrigation, and irrigation effects are less well known. We experimentally tested the effects of fertilization and irrigation on vascular plant species richness and the functional composition of mountain grasslands in the Swiss Alps. After five years, fertilization increased yield but the relationship was quadratic with maximum yield reached at intermediate fertilizer levels (58 kg N ha-1year-1). The species richness of all vascular plants and forbs decreased, on average, by 6 and 5 species respectively, per 50 kg N of extra fertilizer (ha-1 year-1) applied. Fertilization also favored fast-growing plants (increased mean specific leaf area) and plants typically found in productive environments (increased mean indicator values for soil productivity and moisture). In contrast, we found no effects of irrigation on plant community composition, which suggests that irrigation does not affect vascular plant diversity to the same extent as fertilization in these mesic mountain hay meadows, at least in the mid-term. Our finding that maximum yield can be achieved at intermediate fertilizer levels is very important from an applied, agronomical and conservation point of view. It suggests that without loss of yield, farming costs and at the same time environmental pollution and negative effects on biodiversity can be reduced by applying less fertilizer. We therefore recommend maintaining non-intensive land use and keeping fertilizer inputs as low as possible to maintain the high plant diversity of mountain grasslands.

Species interactions and climate change: How the disruption of species co-occurrence will impact on an avian forest guild.

Interspecific interactions are crucial in determining species occurrence and community assembly. Understanding these interactions is thus essential for correctly predicting species' responses to climate change. We focussed on an avian forest guild of four hole-nesting species with differing sensitivities to climate that show a range of well-understood reciprocal interactions, including facilitation, competition and predation. We modelled the potential distributions of black woodpecker and boreal, tawny and Ural owl, and tested whether the spatial patterns of the more widespread species (excluding Ural owl) were shaped by interspecific interactions. We then modelled the potential future distributions of all four species, evaluating how the predicted changes will alter the overlap between the species' ranges, and hence the spatial outcomes of interactions. Forest cover/type and climate were important determinants of habitat suitability for all species. Field data analysed with N-mixture models revealed effects of interspecific interactions on current species abundance, especially in boreal owl (positive effects of black woodpecker, negative effects of tawny owl). Climate change will impact the assemblage both at species and guild levels, as the potential area of range overlap, relevant for species interactions, will change in both proportion and extent in the future. Boreal owl, the most climate-sensitive species in the guild, will retreat, and the range overlap with its main predator, tawny owl, will increase in the remaining suitable area: climate change will thus impact on boreal owl both directly and indirectly. Climate change will cause the geographical alteration or disruption of species interaction networks, with different consequences for the species belonging to the guild and a likely spatial increase of competition and/or intraguild predation. Our work shows significant interactions and important potential changes in the overlap of areas suitable for the interacting species, which reinforce the importance of including relevant biotic interactions in predictive climate change models for increasing forecast accuracy.

Habitat amount mediates the effect of fragmentation on a pollinator's reproductive performance, but not on its foraging behaviour.

Agricultural intensification, with its associated habitat loss and fragmentation, is among the most important drivers of the ongoing pollination crisis. In this quasi-experimental study, conducted in intensively managed vineyards in southwestern Switzerland, we tested the separate and interdependent effects of habitat amount and fragmentation on the foraging activity and reproductive performance of bumblebee Bombus t. terrestris colonies. Based on a factorial design, we selected a series of spatially replicated study sites across a dual gradient of habitat amount (area of ground-vegetated vineyards) and fragmentation (density of ground-vegetated vineyard fields) in a landscape predominantly consisting of vineyards with bare grounds. The foraging activity of individual bumblebees was measured using the radio frequency identification (RFID) technology, and we assessed final colony size to measure reproductive performance. We found an interactive effect of habitat amount and fragmentation on colony size. More specifically, the degree of fragmentation had a negative effect on bumblebee colony size when the amount of habitat was low, while it had a weak positive effect on colony size in landscapes with high amounts of habitat. At the level of individual vineyard fields, ground vegetation cover exerted a positive effect on bumblebee colony size. Fragmentation, but not habitat amount, significantly influenced foraging activity, with more foraging trips in sites with lower degrees of fragmentation. Our results emphasise the importance of studying the separate and interdependent effects of habitat amount and fragmentation to understand their influence on pollinators, providing guidance for optimising the spatial configuration of agricultural landscapes from a biodiversity viewpoint.

Larval and phenological traits predict insect community response to mowing regime manipulations.

For the restoration of biodiversity in agricultural grasslands, it is essential to understand how management acts as an ecological filter on the resident species. Mowing constitutes such a filter: only species that possess functional traits enabling them to withstand its consequences can persist in the community. We investigated how the timing of mowing modulates this filtering effect for insects. We predicted that two traits drive species responses. Species with larval development within the meadow vegetation will suffer more from mowing than species whose larvae develop in or on the ground, or outside the meadows, while species with a later phenology should benefit from later mowing. We conducted a five-year experiment, replicated at 12 sites across the Swiss lowlands, applying three different mowing regimes to low-intensity hay meadows: (1) first cut of the year not earlier than 15 June (control regime); (2) the first cut delayed until 15 July; and (3) leaving an uncut grass refuge on 10-20% of the meadow area (after earliest first cut on 15 June). Before the first cut in years 4 or 5, we sampled larvae of Lepidoptera and sawflies, and adults of moths, parasitoid wasps, wild bees, hoverflies, ground beetles, and rove beetles. Overall, before the first cut of the year, abundances of species with vegetation-dwelling larvae were higher in meadows with delayed mowing or an uncut grass refuge, with some taxon-specific variation. In contrast, species whose larval development is independent of the meadow vegetation showed no differences in abundance between mowing regimes. Species richness did not differ among regimes. For species with vegetation-dwelling larvae, a fourth-corner analysis showed an association between early phenology and the control regime. No associations were found for the other functional groups. Our results show that slight modifications of mowing regimes, easily implementable in agri-environmental policy schemes, can boost invertebrate abundance, potentially benefitting insectivorous vertebrates.

Sex-Specific Heterogeneity in Fixed Morphological Traits Influences Individual Fitness in a Monogamous Bird Population.

Theoretical work has emphasized the important role of individual traits on population dynamics, but empirical models are often based on average or stage-dependent demographic rates. In this study on a monogamous bird, the Eurasian hoopoe (Upupa epops), we show how the interactions between male and female fixed and dynamic heterogeneity influence demographic rates and population dynamics. We built an integral projection model including individual sex, age, condition (reflecting dynamic heterogeneity), and fixed morphology (reflecting fixed heterogeneity). Fixed morphology was derived from a principal component analysis of six morphological traits. Our results revealed that reproductive success and survival were linked to fixed heterogeneity, whereas dynamic heterogeneity influenced mainly the timing of reproduction. Fixed heterogeneity had major consequences for the population growth rate, but interestingly, its effect on population dynamics differed between the sexes. Female fixed morphology was directly linked to annual reproductive success, whereas male fixed morphology also influenced annual survival, being twice higher in large than in small males. Even in a monogamous bird with shared parental care, large males can reach 10% higher fitness than females. Including the dynamics of male and female individual traits in population models refines our understanding of the individual mechanisms that influence demographic rates and population dynamics and can help in identifying differences in sex-specific strategies.

Hoopoe males experience intra-seasonal while females experience inter-seasonal reproductive costs.

Reproductive and survival costs due to reproductive investment are a central element for the evolution of life histories. Both intra- (reduction of reproductive performance of second brood due to investment in first brood) and inter-seasonal costs (reduction of reproductive performance or annual survival due to reproductive investment in preceding year) may appear in multiple breeding species. Knowledge about how trade-offs within and between seasons shape individual trajectories and influence fitness are crucial in life-history evolution, yet intra- and inter-seasonal reproductive costs are rarely analysed simultaneously. We investigated sex-specific differences in intra- and inter-seasonal reproductive and survival costs in response to previous reproductive effort in a monogamous, double-brooding bird, the hoopoe (Upupa epops), accounting for heterogeneity in individual and annual quality. Intra-seasonal reproductive costs were detected in males and inter-seasonal reproductive and survival costs were detected in females. In males, the probability of being a successful double breeder was negatively correlated with the number of hatchlings produced in the first brood. In females, the number of fledglings raised in the first brood was negatively correlated with the reproductive effort in the preceding season. Female annual survival was also negatively influenced by the number of broods produced in the previous reproductive season. Most of these reproductive costs were detected only in years with low productivity, suggesting that costs become evident when environmental conditions are harsh. Our results illustrate how different investment in current vs. future reproduction and survival shape different life-history strategies in males and females of a monogamous bird species.

Impacts of nitrogen addition on plant biodiversity in mountain grasslands depend on dose, application duration and climate: a systematic review.

Although the influence of nitrogen (N) addition on grassland plant communities has been widely studied, it is still unclear whether observed patterns and underlying mechanisms are constant across biomes. In this systematic review, we use meta-analysis and metaregression to investigate the influence of N addition (here referring mostly to fertilization) upon the biodiversity of temperate mountain grasslands (including montane, subalpine and alpine zones). Forty-two studies met our criteria of inclusion, resulting in 134 measures of effect size. The main general responses of mountain grasslands to N addition were increases in phytomass and reductions in plant species richness, as observed in lowland grasslands. More specifically, the analysis reveals that negative effects on species richness were exacerbated by dose (ha(-1) year(-1) ) and duration of N application (years) in an additive manner. Thus, sustained application of low to moderate levels of N over time had effects similar to short-term application of high N doses. The climatic context also played an important role: the overall effects of N addition on plant species richness and diversity (Shannon index) were less pronounced in mountain grasslands experiencing cool rather than warm summers. Furthermore, the relative negative effect of N addition on species richness was more pronounced in managed communities and was strongly negatively related to N-induced increases in phytomass, that is the greater the phytomass response to N addition, the greater the decline in richness. Altogether, this review not only establishes that plant biodiversity of mountain grasslands is negatively affected by N addition, but also demonstrates that several local management and abiotic factors interact with N addition to drive plant community changes. This synthesis yields essential information for a more sustainable management of mountain grasslands, emphasizing the importance of preserving and restoring grasslands with both low agricultural N application and limited exposure to N atmospheric deposition.

Rodent-avoidance, topography and forest structure shape territory selection of a forest bird.

BACKGROUND: Understanding the factors underlying habitat selection is important in ecological and evolutionary contexts, and crucial for developing targeted conservation action in threatened species. However, the key factors associated to habitat selection often remain poorly known. We evaluated hypotheses related to abiotic and biotic factors thought to affect territory selection of the wood warbler Phylloscopus sibilatrix, a passerine living in an unpredictable environment owing to irregular rodent outbreaks and showing long-term declines particularly in Western Europe. RESULTS: Comparing breeding territories to unoccupied areas located close-by revealed that territory occupancy in north-western Switzerland was positively related to slope steepness (topographic hypothesis supported) as well as to numbers of tussocks and trees, respectively, while it showed a unimodal relationship to cover of herb layer (forest structure hypothesis supported). Furthermore, a strong negative correlation between breeding territory occupancy and rodent numbers was found, suggesting that wood warblers avoid areas with high rodent densities (rodent-avoidance hypothesis supported). Comparing breeding territories to abandoned territories showed that breeding territories were located on steeper slopes (topography hypothesis supported), at larger distance from the forest edge (anthropogenic disturbance hypothesis supported) and harboured more trees (forest structure hypothesis supported) than abandoned territories. CONCLUSIONS: Aside from structural and topographic features of the habitat, wood warblers are affected by rodent numbers when settling, making habitat selection unpredictable from year to year. Forestry practices promoting relatively high tree densities, few bushes and an intermediate low-growing ground vegetation cover would enhance habitat quality for this declining passerine. In contrast, forestry practices aiming at increasing light in forests (selective thinning, group-felling) or keeping forest stands permanently covered with shrubs, bushes and trees of various sizes (continuous cover forestry) do not benefit the wood warbler.

Disturbance of wildlife by outdoor winter recreation: allostatic stress response and altered activity-energy budgets.

Anthropogenic disturbance of wildlife is of growing conservation concern, but we lack comprehensive approaches of its multiple negative effects. We investigated several effects of disturbance by winter outdoor sports on free-ranging alpine Black Grouse by simultaneously measuring their physiological and behavioral responses. We experimentally flushed radio-tagged Black Grouse from their snow burrows, once a day, during several successive days, and quantified their stress hormone levels (corticosterone metabolites in feces [FCM] collected. from individual snow burrows). We also measured feeding time allocation (activity budgets reconstructed from radio-emitted signals) in response to anthropogenic disturbance. Finally, we estimated the related extra energy expenditure that may be incurred: based on activity budgets, energy expenditure was modeled from measures of metabolism obtained from captive birds subjected to different ambient temperatures. The pattern of FCM excretion indicated the existence of a funneling effect as predicted by the allostatic theory of stress: initial stress hormone concentrations showed a wide inter-individual variation, which decreased during experimental flushing. Individuals with low initial pre-flushing FCM values augmented their concentration, while individuals with high initial FCM values lowered it. Experimental disturbance resulted in an extension of feeding duration during the following evening foraging bout, confirming the prediction that Black Grouse must compensate for the extra energy expenditure elicited by human disturbance. Birds with low initial baseline FCM concentrations were those that spent more time foraging. These FCM excretion and foraging patterns suggest that birds with high initial FCM concentrations might have been experiencing a situation of allostatic overload. The energetic model provides quantitative estimates of extra energy expenditure. A longer exposure to ambient temperatures outside the shelter of snow burrows, following disturbance, could increase the daily energy expenditure by > 10%, depending principally on ambient temperature and duration of exposure. This study confirms the predictions of allostatic theory and, to the best of our knowledge, constitutes the first demonstration of a funneling effect. It further establishes that winter recreation activities incur costly allostatic behavioral and energetic adjustments, which call for the creation of winter refuge areas together with the implementation of visitor-steering measures for sensitive wildlife.

Landscape structure affects temporal dynamics in the bumble bee virome: Landscape heterogeneity supports colony resilience.

Virus spillovers from managed honey bees, Apis mellifera, are thought to contribute to the decline of wild pollinators, including bumble bees. However, data on the impact of such viruses on wild pollinators remain scarce, and the influence of landscape structure on virus dynamics is poorly understood. In this study, we deployed bumble bee colonies in an agricultural landscape and studied changes in the bumble bee virome during field placement under varying habitat composition and configuration using a multiscale analytical framework. We estimated prevalence of viruses and viral loads (i.e. number of viral genomic equivalent copies) in bumble bees before and after placing them in the field using next generation sequencing and quantitative PCR. The results show that viral loads and number of different viruses present increased during placement in the field and that the virus composition of the colonies shifted from an initial dominance of honey bee associated viruses to a higher number (in both viral loads and number of viruses present) of bumble bee associated viruses. Especially DWV-B, typical for honey bees, drastically decreased after the time in the field. Viral loads prior to placing colonies in the field showed no effect on colony development, suggesting low impacts of these viruses in field settings. Notably, we further demonstrate that increased habitat diversity results in a lower number of different viruses present in Bombus colonies, while colonies in areas with well-connected farmland patches decreased in their total viral load after field placement. Our results emphasize the importance of landscape heterogeneity and connectivity for wild pollinator health and that these influences predominate at fine spatial scales.

The shaping of genetic variation in edge-of-range populations under past and future climate change.

With rates of climate change exceeding the rate at which many species are able to shift their range or adapt, it is important to understand how future changes are likely to affect biodiversity at all levels of organisation. Understanding past responses and extent of niche conservatism in climatic tolerance can help predict future consequences. We use an integrated approach to determine the genetic consequences of past and future climate changes on a bat species, Plecotus austriacus. Glacial refugia predicted by palaeo-modelling match those identified from analyses of extant genetic diversity and model-based inference of demographic history. Former refugial populations currently contain disproportionately high genetic diversity, but niche conservatism, shifts in suitable areas and barriers to migration mean that these hotspots of genetic diversity are under threat from future climate change. Evidence of population decline despite recent northward migration highlights the need to conserve leading-edge populations for spearheading future range shifts.

Modulation of the adrenocortical response to acute stress with respect to brood value, reproductive success and survival in the Eurasian hoopoe.

Reproducing parents face the difficult challenge of trading-off investment in current reproduction against presumed future survival and reproduction. Glucocorticoids are supposed to mediate this trade-off because the adrenocortical response to stress disrupts normal reproductive behaviour in favour of self-maintenance and own survival. According to the brood-value hypothesis, individuals with a low survival probability until the next reproductive season have to invest in current reproduction, a process driven by a down-regulation of their adrenocortical response. If the adrenocortical response to stress effectively mediates the trade-off between current reproduction versus future survival and reproduction, we expect a negative relationship with reproductive success and a positive correlation of the adrenocortical stress response with survival. We studied the relationship between corticosterone secretion in parents and their current brood value, reproductive success and survival in a short-lived multi-brooded bird, the Eurasian hoopoe Upupa epops. The adrenocortical response to acute handling stress was correlated with the brood value within the individual (first and second broods of the year) and between individuals. Birds breeding late in the season mounted a lower total corticosterone response to acute stress than birds breeding earlier, while females showed lower levels than males. We observed a negative relationship between the adrenocortical stress response and rearing success or fledging success in females, as predicted by the brood-value hypothesis. However, we could not evidence a clear link between the adrenocortical stress response and survival. Future research testing the brood-value hypothesis and trade-offs between current reproduction and future survival should also measure free corticosterone and carefully differentiate between cross-sectional (i.e. between-individual) and individual-based experimental studies.

Conservation through connectivity: can isotopic gradients in Africa reveal winter quarters of a migratory bird?

Conservation of migratory wildlife requires knowledge of migratory connectivity between breeding and non-breeding locations. Stable isotopes in combination with geographical isotopic patterns (isoscapes) can provide inferences about migratory connectivity. This study examines whether such an approach can be used to infer wintering areas in sub-Saharan Africa, where we lack such knowledge for many species, but where this method has not been used widely. We measured δ (2)H, δ (13)C and δ (15)N in winter-grown feathers of a breeding Swiss and Spanish population of European hoopoe Upupa epops--a typical Palaearctic-Afrotropical migrant. δ (2)H values predicted that ~70 % of the hoopoes spent the non-breeding season in the western portion of their potential winter range. This was corroborated by a shallow east-west gradient in feather-δ (2)H values of museum specimens from known African origin across the potential winter range and by the recovery of Swiss hoopoes marked with geolocators. Hoopoes categorized as from eastern versus western regions of the wintering range were further delineated spatially using feather δ (13)C and δ (15)N. δ (15)N showed no trend, whereas adults were more enriched in (13)C in the western portion of the range, with eastern adults being in addition more depleted in (13)C than eastern juveniles. This suggests that eastern juveniles may have occupied more xeric habitats than sympatric adults. We demonstrated that stable isotopes, especially δ (2)H, could only very roughly delineate the winter distribution of a trans-Saharan Palaearctic migrant restricted primarily to the Sahelian and savanna belt south of the Sahara. Further refinements of precipitation isoscapes for Africa as well the development of isoscapes for δ (13)C and δ (15)N may improve assignment of this and other migrants.

Deep phylogeographic divergence and cytonuclear discordance in the grasshopper Oedaleus decorus.

The grasshopper Oedaleus decorus is a thermophilic insect with a large, mostly south-Palaearctic distribution range, stretching from the Mediterranean regions in Europe to Central-Asia and China. In this study, we analyzed the extent of phylogenetic divergence and the recent evolutionary history of the species based on 274 specimens from 26 localities across the distribution range in Europe. Phylogenetic relationships were determined using sequences of two mitochondrial loci (ctr, ND2) with neighbour-joining and Bayesian methods. Additionally, genetic differentiation was analyzed based on mitochondrial DNA and 11 microsatellite markers using F-statistics, model-free multivariate and model-based Bayesian clustering approaches. Phylogenetic analyses detected consistently two highly divergent, allopatrically distributed lineages within O. decorus. The divergence among these Western and Eastern lineages meeting in the region of the Alps was similar to the divergence of each lineage to the sister species O. asiaticus. Genetic differentiation for ctr was extremely high between Western and Eastern grasshopper populations (F(ct)=0.95). Microsatellite markers detected much lower but nevertheless very significant genetic structure among population samples. The nuclear data also demonstrated a case of cytonuclear discordance because the affiliation with mitochondrial lineages was incongruent in Northern Italy. Taken together these results provide evidence of an ancient separation within Oedaleus and either historical introgression of mtDNA among lineages and/or ongoing sex-specific gene flow in this grasshopper. Our study stresses the importance of multilocus approaches for unravelling the history and status of taxa of uncertain evolutionary divergence.

The demographic drivers of local population dynamics in two rare migratory birds.

The exchange of individuals among populations can have strong effects on the dynamics and persistence of a given population. Yet, estimation of immigration rates remains one of the greatest challenges for animal demographers. Little empirical knowledge exists about the effects of immigration on population dynamics. New integrated population models fitted using Bayesian methods enable simultaneous estimation of fecundity, survival and immigration, as well as the growth rate of a population of interest. We applied this novel analytical framework to the demography of two populations of long-distance migratory birds, hoopoe Upupa epops and wryneck Jynx torquilla, in a study area in south-western Switzerland. During 2002-2010, the hoopoe population increased annually by 11%, while the wryneck population remained fairly stable. Apparent juvenile and adult survival probability was nearly identical in both species, but fecundity and immigration were slightly higher in the hoopoe. Hoopoe population growth rate was strongly correlated with juvenile survival, fecundity and immigration, while that of wrynecks strongly correlated only with immigration. This indicates that demographic components impacting the arrival of new individuals into the populations were more important for their dynamics than demographic components affecting the loss of individuals. The finding that immigration plays a crucial role in the population growth rates of these two rare species emphasizes the need for a broad rather than local perspective for population studies, and the development of wide-scale conservation actions.

A predictive flight-altitude model for avoiding future conflicts between an emblematic raptor and wind energy development in the Swiss Alps.

Deployment of wind energy is proposed as a mechanism to reduce greenhouse gas emissions. Yet, wind energy and large birds, notably soaring raptors, both depend on suitable wind conditions. Conflicts in airspace use may thus arise due to the risks of collisions of birds with the blades of wind turbines. Using locations of GPS-tagged bearded vultures, a rare scavenging raptor reintroduced into the Alps, we built a spatially explicit model to predict potential areas of conflict with future wind turbine deployments in the Swiss Alps. We modelled the probability of bearded vultures flying within or below the rotor-swept zone of wind turbines as a function of wind and environmental conditions, including food supply. Seventy-four per cent of the GPS positions were collected below 200 m above ground level, i.e. where collisions could occur if wind turbines were present. Flight activity at potential risk of collision is concentrated on south-exposed mountainsides, especially in areas where ibex carcasses have a high occurrence probability, with critical areas covering vast expanses throughout the Swiss Alps. Our model provides a spatially explicit decision tool that will guide authorities and energy companies for planning the deployment of wind farms in a proactive manner to reduce risk to emblematic Alpine wildlife.

Mid- and long-term responses of land snail communities to the intensification of mountain hay meadows management.

BACKGROUND: Species-rich semi-natural grasslands are impacted by the severe land-use changes that are affecting mountain regions, compromising their high biodiversity value. In particular, sprinkler irrigation and increased fertilisation stimulate vegetation growth, modifying and homogenising habitat conditions for ground-dwelling invertebrates. Among them, land snails have been largely understudied despite their commonness and vulnerability to small-scale habitat alteration. This study investigated the mid- and long-term responses of land snail communities to management intensification of montane and subalpine hay meadows. Mid-term effects were studied using a randomised block design experiment, mimicking an intensification gradient with different levels of irrigation and fertilisation applied during 5 years. Long-term effects were examined relying on an observational approach that consisted in comparing snail communities in meadows managed intensively for > 20 years with those from the 5-year experimental module. RESULTS: We show that management intensification initially boosts snail densities, but erodes species richness by - 35% in intensively-managed meadows in the long term. Contrary to our expectations, drought-tolerant (xerophilous) snails benefitted from grassland intensification, whereas mesophilous species accounted for most species losses due to intensification in the long run, indicating that the latter may be especially sensitive to the hostile microclimate conditions abruptly prevailing in a meadow after mowing. Soil pH was also a principal determinant of land snail occurrence, with almost no specimen recorded in acidic meadows (pH < 5.5), while plant diversity favoured overall snail abundance. CONCLUSIONS: Despite the fact that xerophilous snails appear tolerant to management intensification, we found that several drought-sensitive species are lost in the long term. We conclude that the preservation of species-rich land snail communities in mountain hay meadows requires the conservation and restoration of low-input grasslands on basic soils for preventing further species losses of gastropod fauna.

Accounting for predator species identity reveals variable relationships between nest predation rate and habitat in a temperate forest songbird.

Nest predation is the primary cause of nest failure in most ground-nesting bird species. Investigations of relationships between nest predation rate and habitat usually pool different predator species. However, such relationships likely depend on the specific predator involved, partly because habitat requirements vary among predator species. Pooling may therefore impair our ability to identify conservation-relevant relationships between nest predation rate and habitat. We investigated predator-specific nest predation rates in the forest-dependent, ground-nesting wood warbler Phylloscopus sibilatrix in relation to forest area and forest edge complexity at two spatial scales and to the composition of the adjacent habitat matrix. We used camera traps at 559 nests to identify nest predators in five study regions across Europe. When analyzing predation data pooled across predator species, nest predation rate was positively related to forest area at the local scale (1000 m around nest), and higher where proportion of grassland in the adjacent habitat matrix was high but arable land low. Analyses by each predator species revealed variable relationships between nest predation rates and habitat. At the local scale, nest predation by most predators was higher where forest area was large. At the landscape scale (10,000 m around nest), nest predation by buzzards Buteo buteo was high where forest area was small. Predation by pine martens Martes martes was high where edge complexity at the landscape scale was high. Predation by badgers Meles meles was high where the matrix had much grassland but little arable land. Our results suggest that relationships between nest predation rates and habitat can depend on the predator species involved and may differ from analyses disregarding predator identity. Predator-specific nest predation rates, and their relationships to habitat at different spatial scales, should be considered when assessing the impact of habitat change on avian nesting success.

Spatially explicit modeling of conflict zones between wildlife and snow sports: prioritizing areas for winter refuges.

Outdoor winter recreation exerts an increasing pressure upon mountain ecosystems, with unpredictable, free-ranging activities (e.g., ski mountaineering, snowboarding, and snowshoeing) representing a major source of stress for wildlife. Mitigating anthropogenic disturbance requires the spatially explicit prediction of the interference between the activities of humans and wildlife. We applied spatial modeling to localize conflict zones between wintering Black Grouse (Tetrao tetrix), a declining species of Alpine timberline ecosystems, and two free-ranging winter sports (off-piste skiing [including snow-boarding] and snowshoeing). Track data (snow-sports and birds' traces) obtained from aerial photographs taken over a 585-km transect running along the timberline, implemented within a maximum entropy model, were used to predict the occurrence of snow sports and Black Grouse as a function of landscape characteristics. By modeling Black Grouse presence in the theoretical absence of free-ranging activities and ski infrastructure, we first estimated the amount of habitat reduction caused by these two factors. The models were then extrapolated to the altitudinal range occupied by Black Grouse, while the spatial extent and intensity of potential conflict were assessed by calculating the probability of human-wildlife co-occurrence. The two snow-sports showed different distribution patterns. Skiers' occurrence was mainly determined by ski-lift presence and a smooth terrain, while snowshoers' occurrence was linked to hiking or skiing routes and moderate slopes. Wintering Black Grouse avoided ski lifts and areas frequented by free-ranging snow sports. According to the models, Black Grouse have faced a substantial reduction of suitable wintering habitat along the timberline transect: 12% due to ski infrastructure and another 16% when adding free-ranging activities. Extrapolating the models over the whole study area results in an overall habitat loss due to ski infrastructure of 10%, while there was a > 10% probability of human-wildlife encounters on 67% of the remaining area of suitable wintering habitat. Only 23% of the wintering habitat was thus free of anthropogenic disturbance. By identifying zones of potential conflict, while rating its relative intensity, our model provides a powerful tool to delineate and prioritize areas where wildlife winter refuges and visitor steering measures should be implemented.