Questions: (i) Do species richness and turnover across forest edges change with edge age and management intensity of adjacent lands? (ii) Does edge species composition respond to aging and landscape management and what are the environmental factors explaining this response?Location: Agricultural landscapes of the Picardy region, N France. Methods:We sampled forest edges differing in age (from a few decades to several centuries) and embedding landscape matrix (from slightly managed 'bocages' to intensively cultivated open fields). We recorded vascular plant species and a set of environment, landscape and historical variables along transects oriented perpendicularly to forest edges. We used mixed models to assess the impact of edge age and landscape type on edge species richness and turnover. We investigated the relationship between edge community composition and explanatory variables using redundancy analyses and a split-plot design.Results: Species richness decreased with both increased edge age and increased landscape management intensity, while species turnover was not influenced by any of these factors. Edge maturation was characterized by a specialization of the flora over the entire transect, which is likely a response to increased shade, litter layer thickness and soil acidity. As landscape management was more intensive, true forest species were replaced by nitrophilous and/or calciphilous non-forest species, which might be more tolerant of agrochemical and lime drift and are able to disperse through a hostile matrix. Conclusion:Although edge aging was associated with the progressive development of environmental gradients, especially light availability and litter thickness, plant communities poorly reflect these gradients under the constraint of neighbouring landscape management. On the contrary, the stronger the management intensity, the sharper the edge-interior gradient.

Aim: To propose a species distribution modelling framework and its companion "iSDM" R package for predicting the potential and realized distributions of invasive species within the invaded range.Location: Northern France. Methods:The non-equilibrium distribution of invasive species with the environment within the invaded range affects the environmental representativeness of species presenceabsence data collected from the field and introduces uncertainty in observed absences as these may either reflect unsuitable sites or be incidental. To address these issues, we here propose an environmental systematic sampling design to collect presence-absence data from the field and a probability index to sort and subsequently separate environmental absences (EAs: reflecting environmentally unsuitable sites) from dispersal-limited absences (DLAs: reflecting sites out of dispersal reach). We first conducted a comprehensive test based on a virtual species to evaluate the performance of our framework. Then, we applied it on different life stages of a non-native tree species (Prunus serotina Ehrh.) invasive in Europe. Results:Regarding the potential distribution, we found higher model performances for both the virtual species (true skill statistics (TSS) > 0.75) and P. serotina (TSS ≥ 0.68) after carefully selecting absences with a low probability to be DLAs compared with classical models that incorporate both EAs and DLAs (e.g. TSS = 0.11 for P. serotina with 80% of DLAs). On the contrary, both EAs and DLAs as well as dispersal-related covariates were needed to capture the realized distribution of both the virtual species and P. serotina. Main Conclusions:Our framework helps overcoming the conceptual and methodological limitations of the disequilibrium in species' distribution models inherent to invasive species and enables managers to robustly estimate both the realized and potential distributions of invasive species. Although more relevant for modelling the distribution of non-native species, this framework can also be applied to native species. K E Y W O R D Salien species, biological invasions, dispersal limitations, potential niche, realized niche, species distribution modelling, virtual species

Aim: In response to environmental changes and to avoid extinction, species may either track suitable environmental conditions or adapt to the modified environment. However, whether and how species adapt to environmental changes remains unclear. By focusing on the realized niche (i.e. the actual space that a species inhabits and the resources it can access as a result of limiting biotic factors present in its habitat), we here examine shifts in the realized-niche width (i.e. ecological amplitude) and position (i.e. ecological optimum) of 26 common and widespread forest understorey plants across their distributional ranges. Location: Temperate forests along a ca. 1800-km-long latitudinal gradient from northern France to central Sweden and Estonia. Methods: We derived species' realized-niche width from a -diversity metric, which increases if the focal species co-occurs with more species. Based on the concept that species' scores in a detrended correspondence analysis (DCA) represent the locations of their realized-niche positions, we developed a novel approach to run species-specific DCAs allowing the focal species to shift its realized-niche position along the studied latitudinal gradient while the realized-niche positions of other species were held constant. Results: None of the 26 species maintained both their realized-niche width and position along the latitudinal gradient. Few species (9 of 26: 35%) shifted their realized-niche width, but all shifted their realized-niche position. With increasing latitude, most species (22 of 26: 85%) shifted their realized-niche position for soil nutrients and pH towards nutrient-poorer and more acidic soils. Main conclusions: Forest understorey plants shifted their realized niche along the latitudinal gradient, suggesting local adaptation and/or plasticity. This macroecological pattern casts doubt on the idea that the realized niche is stable in space and time, which is a key assumption of species distribution models used to predict the future of biodiversity, hence raising concern about predicted extinction rates

Aim: Macroclimate is a major determinant of large-scale diversity patterns. However, the influence of smaller-scale factors on local diversity across large spatial extents is not well documented. Here, we quantify the relative importance of local (patch-scale), landscape-scale and macroclimatic drivers of herbaceous species diversity in small forest patches in agricultural landscapes across Europe. Location: Deciduous forest patches in eight regions along a macroclimatic gradient from southern France to central Sweden and Estonia. Methods: The diversity of forest specialists and generalists at three levels (whole forest patch, sampling plots within patches and between scales) was related to patch-scale (forest area, age, abiotic and biotic heterogeneity), landscape-scale (amount of forest, grasslands and hedgerows around the patch, patch isolation) and macroclimatic variables (temperature and precipitation) using generalized linear mixed models and variation partitioning for each group of variables. Results: The total amount of explained variation in diversity ranged from 8% for plot-scale diversity of generalists to 54% for patch-scale diversity of forest specialists. Patch-scale variables always explained more than 60% of the explained variation in diversity, mainly due to the positive effect of within-patch heterogeneity on patch-scale and between-scale diversities and to the positive effect of patch age on plot-scale diversity of forest specialists. Landscape-scale variables mainly contributed to the amount of explained variation in plot-scale diversity, being more important for forest specialists (21%) than for generalists (18%). Macroclimatic variables contributed a maximum of 11% to the plot-scale diversity of generalists. Main conclusions: Macroclimate poorly predicts local diversity across Europe, and herbaceous diversity is mainly explained by habitat features, less so by landscape structure. We show the importance of conserving old forest patches as refugia for typical forest species, and of enhancing the landscape context around the patches by reducing the degree of disturbance caused by agriculture

Summary 1.Although habitat fragmentation is recognized as a major threat to biodiversity, few studies have examined the relative importance of local, landscape and historical factors in controlling local species assemblages, and how these factors interact, in patchy ecosystems. We quantified the direct and indirect effects of patch size, patch heterogeneity, agricultural intensity and patch age on plant species richness and composition of forest patches embedded in agricultural landscapes. 2. In six 5 · 5 km-sampling landscape windows, we surveyed each forest patch for vascular plant species and collected three sets of independent variables, describing patch size and heterogeneity, landscape composition and history. The six windows were arranged along a gradient of agriculture intensity in rural landscapes of the Picardy region (N France). 3. We used non-metric multidimensional scaling (NMS) to detect major environmental gradients underlying variation in species composition among patches. We then constructed structural equation models (SEM) to quantify the direct and indirect effects of the three sets of variables on local plant diversity, which was successively incorporated as patch scores along the first three NMS axes, woody species richness, forest herb species richness, and non-forest herb species richness. 4. A major influence of the landscape matrix on local species composition was revealed by NMS and subsequent SEM, mainly through non-forest herb species, which explained most of the between patch floristic dissimilarity. Species richness increased with patch heterogeneity, whereas patch area never had a direct effect. Forest herbs were more responsive to patch age and connectivity than other species, whereas non-forest and woody species were more influenced by agriculture intensity in the surrounding matrix. 5. Synthesis. We used one of the largest data sets ever collected in temperate fragmented forests to build, for the first time, a structural model incorporating all suspected drivers of local plant communities. We showed that the number and identity of local species coexisting in successional fragments of a forest metacommunity at a given time is controlled by a unique combination of interacting local, landscape and historical factors. Preserving the largest, oldest fragments and favouring species movements in the surrounding matrix is the best way to conserve forest specialists in changing rural landscapes.

BackgroundThe tick Ixodes ricinus has considerable impact on the health of humans and other terrestrial animals because it transmits several tick-borne pathogens (TBPs) such as B. burgdorferi (sensu lato), which causes Lyme borreliosis (LB). Small forest patches of agricultural landscapes provide many ecosystem services and also the disservice of LB risk. Biotic interactions and environmental filtering shape tick host communities distinctively between specific regions of Europe, which makes evaluating the dilution effect hypothesis and its influence across various scales challenging. Latitude, macroclimate, landscape and habitat properties drive both hosts and ticks and are comparable metrics across Europe. Therefore, we instead assess these environmental drivers as indicators and determine their respective roles for the prevalence of B. burgdorferi in I. ricinus.MethodsWe sampled I. ricinus and measured environmental properties of macroclimate, landscape and habitat quality of forest patches in agricultural landscapes along a European macroclimatic gradient. We used linear mixed models to determine significant drivers and their relative importance for nymphal and adult B. burgdorferi prevalence. We suggest a new prevalence index, which is pool-size independent.ResultsDuring summer months, our prevalence index varied between 0 and 0.4 per forest patch, indicating a low to moderate disservice. Habitat properties exerted a fourfold larger influence on B. burgdorferi prevalence than macroclimate and landscape properties combined. Increasingly available ecotone habitat of focal forest patches diluted and edge density at landscape scale amplified B. burgdorferi prevalence. Indicators of habitat attractiveness for tick hosts (food resources and shelter) were the most important predictors within habitat patches. More diverse and abundant macro- and microhabitat had a diluting effect, as it presumably diversifies the niches for tick-hosts and decreases the probability of contact between ticks and their hosts and hence the transmission likelihood.ConclusionsDiluting effects of more diverse habitat patches would pose another reason to maintain or restore high biodiversity in forest patches of rural landscapes. We suggest classifying habitat patches by their regulating services as dilution and amplification habitat, which predominantly either decrease or increase B. burgdorferi prevalence at local and landscape scale and hence LB risk. Particular emphasis on promoting LB-diluting properties should be put on the management of those habitats that are frequently used by humans. In the light of these findings, climate change may be of little concern for LB risk at local scales, but this should be evaluated further.Electronic supplementary materialThe online version of this article (10.1186/s13071-017-2590-x) contains supplementary material, which is available to authorized users.

1. Global forest loss and fragmentation have strongly increased the frequency of forest patches smaller than a few hectares. Little is known about the biodiversity and ecosystem service supply potential of such small woodlands in comparison to larger forests. As it is widely recognized that high biodiversity levels increase ecosystem functionality and the delivery of multiple ecosystem services, small, isolated woodlands are expected to have a lower potential for ecosystem service delivery than large forests hosting more species.2. We collected data on the diversity of six taxonomic groups covering invertebrates, plants and fungi, and on the supply potential of five ecosystem services and one disservice within 224 woodlands distributed across temperate Europe. We related their ability to simultaneously provide multiple ecosystem services (multiservice delivery potential) at different performance levels to biodiversity of all studied taxonomic groups (multidiversity), forest patch size and age, as well as habitat availability and connectivity within the landscape, while accounting for macroclimate, soil properties and forest structure.3. Unexpectedly, despite their lower multidiversity, smaller woodlands had the potential to deliver multiple services at higher performance levels per area than | 5Journal of Applied Ecology VALDÉS et AL.

BackgroundThe castor bean tick (Ixodes ricinus) transmits infectious diseases such as Lyme borreliosis, which constitutes an important ecosystem disservice. Despite many local studies, a comprehensive understanding of the key drivers of tick abundance at the continental scale is still lacking. We analyze a large set of environmental factors as potential drivers of I. ricinus abundance. Our multi-scale study was carried out in deciduous forest fragments dispersed within two contrasting rural landscapes of eight regions, along a macroclimatic gradient stretching from southern France to central Sweden and Estonia. We surveyed the abundance of I. ricinus, plant community composition, forest structure and soil properties and compiled data on landscape structure, macroclimate and habitat properties. We used linear mixed models to analyze patterns and derived the relative importance of the significant drivers.ResultsMany drivers had, on their own, either a moderate or small explanatory value for the abundance of I. ricinus, but combined they explained a substantial part of variation. This emphasizes the complex ecology of I. ricinus and the relevance of environmental factors for tick abundance. Macroclimate only explained a small fraction of variation, while properties of macro- and microhabitat, which buffer macroclimate, had a considerable impact on tick abundance. The amount of forest and the composition of the surrounding rural landscape were additionally important drivers of tick abundance. Functional (dispersules) and structural (density of tree and shrub layers) properties of the habitat patch played an important role. Various diversity metrics had only a small relative importance. Ontogenetic tick stages showed pronounced differences in their response. The abundance of nymphs and adults is explained by the preceding stage with a positive relationship, indicating a cumulative effect of drivers.ConclusionsOur findings suggest that the ecosystem disservices of tick-borne diseases, via the abundance of ticks, strongly depends on habitat properties and thus on how humans manage ecosystems from the scale of the microhabitat to the landscape. This study stresses the need to further evaluate the interaction between climate change and ecosystem management on I. ricinus abundance.Electronic supplementary materialThe online version of this article (doi:10.1186/s12898-017-0141-0) contains supplementary material, which is available to authorized users.