Directional turnover towards larger-ranged plants over time and across habitats.

Species turnover is ubiquitous. However, it remains unknown whether certain types of species are consistently gained or lost across different habitats. Here, we analysed the trajectories of 1827 plant species over time intervals of up to 78 years at 141 sites across mountain summits, forests, and lowland grasslands in Europe. We found, albeit with relatively small effect sizes, displacements of smaller- by larger-ranged species across habitats. Communities shifted in parallel towards more nutrient-demanding species, with species from nutrient-rich habitats having larger ranges. Because these species are typically strong competitors, declines of smaller-ranged species could reflect not only abiotic drivers of global change, but also biotic pressure from increased competition. The ubiquitous component of turnover based on species range size we found here may partially reconcile findings of no net loss in local diversity with global species loss, and link community-scale turnover to macroecological processes such as biotic homogenisation.

Long-term effects of climate change on carbon storage and tree species composition in a dry deciduous forest.

Forest vegetation and soils have been suggested as potentially important sinks for carbon (C) with appropriate management and thus are implicated as effective tools in stabilizing climate even with increasing anthropogenic release of CO2 . Drought, however, which is often predicted to increase in models of future climate change, may limit net primary productio (NPP) of dry forest types, with unknown effects on soil C storage. We studied C dynamics of a deciduous temperate forest of Hungary that has been subject to significant decreases in precipitation and increases in temperature in recent decades. We resampled plots that were established in 1972 and repeated the full C inventory by analyzing more than 4 decades of data on the number of living trees, biomass of trees and shrubs, and soil C content. Our analyses show that the decline in number and biomass of oaks started around the end of the 1970s with a 71% reduction in the number of sessile oak stems by 2014. Projected growth in this forest, based on the yield table's data for Hungary, was 4.6 kg C/m2 . Although new species emerged, this new growth and small increases in oak biomass resulted in only 1.9 kg C/m2 increase over 41 years. The death of oaks increased inputs of coarse woody debris to the surface of the soil, much of which is still identifiable, and caused an increase of 15.5%, or 2.6 kg C/m2 , in the top 1 m of soil. Stability of this fresh organic matter input to surface soil is unknown, but is likely to be low based on the results of a colocated woody litter decomposition study. The effects of a warmer and drier climate on the C balance of forests in this region will be felt for decades to come as woody litter inputs decay, and forest growth remains impeded.

Unexpected westward range shifts in European forest plants link to nitrogen deposition.

Climate change is commonly assumed to induce species' range shifts toward the poles. Yet, other environmental changes may affect the geographical distribution of species in unexpected ways. Here, we quantify multidecadal shifts in the distribution of European forest plants and link these shifts to key drivers of forest biodiversity change: climate change, atmospheric deposition (nitrogen and sulfur), and forest canopy dynamics. Surprisingly, westward distribution shifts were 2.6 times more likely than northward ones. Not climate change, but nitrogen-mediated colonization events, possibly facilitated by the recovery from past acidifying deposition, best explain westward movements. Biodiversity redistribution patterns appear complex and are more likely driven by the interplay among several environmental changes than due to the exclusive effects of climate change alone.

Long-term nitrogen deposition reduces the diversity of nitrogen-fixing plants.

Biological nitrogen fixation is a fundamental part of ecosystem functioning. Anthropogenic nitrogen deposition and climate change may, however, limit the competitive advantage of nitrogen-fixing plants, leading to reduced relative diversity of nitrogen-fixing plants. Yet, assessments of changes of nitrogen-fixing plant long-term community diversity are rare. Here, we examine temporal trends in the diversity of nitrogen-fixing plants and their relationships with anthropogenic nitrogen deposition while accounting for changes in temperature and aridity. We used forest-floor vegetation resurveys of temperate forests in Europe and the United States spanning multiple decades. Nitrogen-fixer richness declined as nitrogen deposition increased over time but did not respond to changes in climate. Phylogenetic diversity also declined, as distinct lineages of N-fixers were lost between surveys, but the "winners" and "losers" among nitrogen-fixing lineages varied among study sites, suggesting that losses are context dependent. Anthropogenic nitrogen deposition reduces nitrogen-fixing plant diversity in ways that may strongly affect natural nitrogen fixation.

Response to Comment on "Forest microclimate dynamics drive plant responses to warming".

Bertrand et al question our interpretation about warming effects on the thermophilization in forest plant communities and propose an alternative way to analyze climatic debt. We show that microclimate warming is a better predictor than macroclimate warming for studying forest plant community responses to warming. Their additional analyses do not affect or change our interpretations and conclusions.

Response to Comment on "Forest microclimate dynamics drive plant responses to warming".

Schall and Heinrichs question our interpretation that the climatic debt in understory plant communities is locally modulated by canopy buffering. However, our results clearly show that the discrepancy between microclimate warming rates and thermophilization rates is highest in forests where canopy cover was reduced, which suggests that the need for communities to respond to warming is highest in those forests.

Forest microclimate dynamics drive plant responses to warming.

Climate warming is causing a shift in biological communities in favor of warm-affinity species (i.e., thermophilization). Species responses often lag behind climate warming, but the reasons for such lags remain largely unknown. Here, we analyzed multidecadal understory microclimate dynamics in European forests and show that thermophilization and the climatic lag in forest plant communities are primarily controlled by microclimate. Increasing tree canopy cover reduces warming rates inside forests, but loss of canopy cover leads to increased local heat that exacerbates the disequilibrium between community responses and climate change. Reciprocal effects between plants and microclimates are key to understanding the response of forest biodiversity and functioning to climate and land-use changes.

Replacements of small- by large-ranged species scale up to diversity loss in Europe's temperate forest biome.

Biodiversity time series reveal global losses and accelerated redistributions of species, but no net loss in local species richness. To better understand how these patterns are linked, we quantify how individual species trajectories scale up to diversity changes using data from 68 vegetation resurvey studies of seminatural forests in Europe. Herb-layer species with small geographic ranges are being replaced by more widely distributed species, and our results suggest that this is due less to species abundances than to species nitrogen niches. Nitrogen deposition accelerates the extinctions of small-ranged, nitrogen-efficient plants and colonization by broadly distributed, nitrogen-demanding plants (including non-natives). Despite no net change in species richness at the spatial scale of a study site, the losses of small-ranged species reduce biome-scale (gamma) diversity. These results provide one mechanism to explain the directional replacement of small-ranged species within sites and thus explain patterns of biodiversity change across spatial scales.

Influence of human impacts on trace metal accumulation in soils of two Hungarian cities.

Despite the advanced activity of urban soil research, comparison of available trace metal contents in the soil of settlements has not yet been well investigated. First aim of research was the comparison of human impacts on urban soils in two Hungarian cities with different structure and development. To detect the sources of contamination, Szombathely and Sopron cities were separated into urban, suburban and peri-urban areas. Altogether 192 topsoil samples were collected at 0-20 cm in order to measure the physicochemical properties of the soil. Instead of total element contents, plant-available trace metal concentrations - Cd, Co, Cu, Ni, Pb, Zn - were determined using ICP-OES method. Due to different geological conditions, urban soils of the investigated settlements had similar properties in the case of pH, CaCO3 and texture. In the case of plant available trace metals, limit excesses were found in urban and peri-urban areas mostly, but high values were recorded in the suburb. The amount of Cdavailable, Coavailable and Niavailable were negligible. Downtown area was the most contaminated in both cities, but extremes appeared in suburban areas. In Szombathely, Cuavailable values were <40 mg Cu/kg, but extremely high Cuavailable results were found in the peri-urban viticulture areas of Sopron (>90 mg Cu/kg). The samples taken alongside busy roads near the city centre were Pbavailable-contaminated due to traffic. The Pbavailable concentration decreased slightly towards the suburban areas of the cities. The Znavailable level exceeded 40 mg Zn/kg in soils taken from the watercourses of Szombathely. The highest trace metal pollution levels were recorded in soils of public parks of Sopron. Urban soils of Szombathely were more homogenized than urban soils of Sopron. Based on statistical analyses and comparison of the relations of urban, suburban, and peri-urban areas of Sopron to Szombathely, Szombathely's urban soils show more homogeneity.