45Evidence from an international survey in the Atlantic biogeographic region of Europe 46indicates that chronic nitrogen deposition is reducing plant species richness in acid 47 grasslands. Across the deposition gradient in this region (2 to 44 kg N ha -1 yr -1 ) species 48 richness showed a curvilinear response, with greatest reductions in species richness when 49 deposition increased from low levels. This has important implications for conservation 50 policies, suggesting that to protect the most sensitive grasslands resources should be 51 focussed where deposition is currently low. Soil pH is also an important driver of species 52 richness indicating that the acidifying effect of nitrogen deposition may be contributing to 53 species richness reductions. The results of this survey suggest that the impacts of nitrogen 54 deposition can be observed over a large geographical range.

Recent studies from mountainous areas of small spatial extent (<2500 km(2) ) suggest that fine-grained thermal variability over tens or hundreds of metres exceeds much of the climate warming expected for the coming decades. Such variability in temperature provides buffering to mitigate climate-change impacts. Is this local spatial buffering restricted to topographically complex terrains? To answer this, we here study fine-grained thermal variability across a 2500-km wide latitudinal gradient in Northern Europe encompassing a large array of topographic complexities. We first combined plant community data, Ellenberg temperature indicator values, locally measured temperatures (LmT) and globally interpolated temperatures (GiT) in a modelling framework to infer biologically relevant temperature conditions from plant assemblages within <1000-m(2) units (community-inferred temperatures: CiT). We then assessed: (1) CiT range (thermal variability) within 1-km(2) units; (2) the relationship between CiT range and topographically and geographically derived predictors at 1-km resolution; and (3) whether spatial turnover in CiT is greater than spatial turnover in GiT within 100-km(2) units. Ellenberg temperature indicator values in combination with plant assemblages explained 46-72% of variation in LmT and 92-96% of variation in GiT during the growing season (June, July, August). Growing-season CiT range within 1-km(2) units peaked at 60-65°N and increased with terrain roughness, averaging 1.97 °C (SD = 0.84 °C) and 2.68 °C (SD = 1.26 °C) within the flattest and roughest units respectively. Complex interactions between topography-related variables and latitude explained 35% of variation in growing-season CiT range when accounting for sampling effort and residual spatial autocorrelation. Spatial turnover in growing-season CiT within 100-km(2) units was, on average, 1.8 times greater (0.32 °C km(-1) ) than spatial turnover in growing-season GiT (0.18 °C km(-1) ). We conclude that thermal variability within 1-km(2) units strongly increases local spatial buffering of future climate warming across Northern Europe, even in the flattest terrains.

Summary1. Semi-natural habitats have been shaped by human disturbance regimes for centuries. Spatially and temporally heterogeneous land-use practices, such as cutting, burning, grazing and turf-cutting, have resulted in complex mosaic landscapes that are of high priority for conservation in Europe. Contemporary conservation subjects these systems to management regimes that are generally less diverse, in terms of disturbances and finescale temporal and spatial variability, than traditional land use, but the ecological consequences of these simplifications are unclear. 2. We investigated the interactive effects of fire and grazing on plant species composition and diversity along local environmental (moisture) gradients in coastal heathlands. A replicated series of post-fire successions ( n = 12) was initiated in three heathland habitats and the areas subjected to two grazing regimes. Floristic and environmental data were recorded in permanent plots over a 5-year period. Community data were analysed using multivariate ordination techniques (principal components analysis, partial redundancy analyses, and principal response curves) and generalized linear models. 3. Fire induced strong successional trends in the species composition of the heathlands. These trends differed among heathland habitats, and with grazing. Strong interactions between fire, habitat and grazing implied that the effect of grazing on the successional dynamics differed among habitats. Species diversity decreased in the first year after fire but increased beyond the pre-fire levels during succession. This trend was not affected by local environment or grazing, although there were main effect differences in diversity between environments and grazing regimes. 4. Synthesis and applications . Our results demonstrate that the two management practices do not have simple additive effects within the semi-natural system studied, as grazing created ecological opportunities for additional sets of species, increased variability among habitats, and added complexity to the post-fire successional dynamics. In order to preserve diversity, conservation management should thus aim to preserve the level of complexity of the traditional management regimes, both in terms of the actual disturbances (e.g. fire and grazing) as well as the spatial scales at which they are applied. Further, the considerable change in these effects along the local environmental gradient brings into question the efficiency of general management prescriptions, and indicates that local environmental variability should be taken into account in the conservation of semi-natural habitats.

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Abstract. Successional dynamics in plant community assembly may result from both deterministic and stochastic ecological processes. The relative importance of different ecological processes is expected to vary over the successional sequence, between different plant functional groups, and with the disturbance levels and land-use management regimes of the successional systems. We evaluate the relative importance of stochastic and deterministic processes in bryophyte and vascular plant community assembly after fire in grazed and ungrazed anthropogenic coastal heathlands in Northern Europe. A replicated series of post-fire successions (n = 12) were initiated under grazed and ungrazed conditions, and vegetation data were recorded in permanent plots over 13 years. We used redundancy analysis (RDA) to test for deterministic successional patterns in species composition repeated across the replicate successional series and analyses of co-occurrence to evaluate to what extent species respond synchronously along the successional gradient. Change in species co-occurrences over succession indicates stochastic successional dynamics at the species level (i.e., species equivalence), whereas constancy in co-occurrence indicates deterministic dynamics (successional niche differentiation). The RDA shows high and deterministic vascular plant community compositional change, especially early in succession. Co-occurrence analyses indicate stochastic species-level dynamics the first two years, which then give way to more deterministic replacements. Grazed and ungrazed successions are similar, but the early stage stochasticity is higher in ungrazed areas. Bryophyte communities in ungrazed successions resemble vascular plant communities. In contrast, bryophytes in grazed successions showed consistently high stochasticity and low determinism in both community composition and species co-occurrence. In conclusion, stochastic and individualistic species responses early in succession give way to more niche-driven dynamics in later successional stages. Grazing reduces predictability in both successional trends and species-level dynamics, especially in plant functional groups that are not well adapted to disturbance.