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Summary This study investigated the factors underlying the variability of needle and soil elemental composition and stoichiometry and their relationships with growth in Pinus sylvestris forests throughout the species' distribution in Europe by analysing data from 2245 forest stands. Needle N concentrations and N:P ratios were positively correlated with total atmospheric N deposition, whereas needle P concentrations were negatively correlated. These relationships were especially pronounced at sites where high levels of N deposition coincided with both higher mean annual temperature and higher mean annual precipitation. Trends towards foliar P deficiency were thus more marked when high N deposition coincided with climatic conditions favourable to plant production. Atmospheric N deposition was positively correlated with soil solution NO3− , SO42− , K+, P and Ca2+ concentrations, the soil solution NO3−:P ratio, total soil N and the total soil N:Olsen P ratio and was negatively correlated with soil Olsen P concentration. Despite these nutrient imbalances, during the period studied (1990–2006), N deposition was positively related with Pinus sylvestris absolute basal diameter (BD) growth, although only accounting for the 10% of the total variance. However, neither N deposition nor needle N concentration was related with relative annual BD growth. In contrast, needle P concentration was positively related with both absolute and relative annual BD growth. These results thus indicate a tendency of European P. sylvestris forests to store N in trees and soil in response to N deposition and unveil a trend towards increased nutrient losses in run‐off as a consequence of higher soil solution N concentrations. Overall, the data show increasing ecosystem nutrient imbalances with increasingly limiting roles of P and other nutrients such as K in European P. sylvestris forests, especially in the centre of their distribution where higher levels of N deposition are observed. Thus, although the data show that N deposition has had an overall positive effect on P. sylvestris growth, the effect of continuous N deposition, associated with decreasing P and K and increasing N:P in leaves and in soil, may in the future become detrimental for the growth and competitive ability of P. sylvestris trees. Lay Summary

1. Atmospheric nitrogen (N) deposition and climate warming are two major components of global change that drive species richness and composition in plant communities. However, their combined effects have been insufficiently investigated across large spatial and temporal scales particularly in high-elevation, nutrient-limited ecosystems. 2. We examine whether and how N deposition and climate warming have altered the plant richness and the composition of subalpine semi-natural, extensively grazed grasslands of the Pyrenees, using two complementary approaches: (i) analysis of 553 relevés to explore vegetation changes across large ecological gradients including temperature and N deposition (spatial approach) and (ii) a re-sampling of a subset of 40 sites among the 553 sites to assess temporal changes over the past decades (temporal approach). 3. Both approaches showed that the vascular plant species richness increased when temperature and cumulative N deposition increase, shifting the species composition towards more thermophilic and eutrophic communities. 4. Synthesis. We hypothesize that the release from abiotic constraints (milder temperature and higher nitrogen availability) due to global changes and long-standing extensive grazing counteracting the negative effects of nitrogen deposition have been responsible for the diversity and compositional changes of plant communities over the last decades in the Pyrenees. Thus, in contrast with other grasslands, high-elevation grazed grasslands may increase in species diversity with nitrogen deposition under climate warming.

Aims: Given the significance of the herbaceous understorey layer for temperate forest ecosystem biodiversity and functioning, it is important to have a thorough understanding of its dynamics in response to environmental change. However, site-specific factors such as soil type may cause differential temporal herb layer changes within a single study area under comparable external pressure. Surprisingly, relatively few studies have examined herb layer changes on different site types within the same forest complex. The aim of this study is to determine the temporal differences in herb vegetation and to compare these for two differing site types in the same forest complex. Location: Ancient broad-leaved woodlands traditionally managed as coppice with standards in South Belgium, an area with secondary deposits and a mild temperate climate. Methods: In 2008 we re-inventoried 43 vegetation recordings from 1953 to 1954 in forests on two site types (acid and neutral forests). First, we described differences in soil pH, litter and overstorey characteristics between the sites. Next, multivariate analysis, Ellenberg indicator value and CSR signature were used to determine differences in herb layer diversity and composition between both the two survey times and the two site types. Finally, we analysed temporal differences in individual species abundances and frequencies. Results: The two site types clearly differed with respect to soil, litter and over-storey characteristics. Temporal changes between the surveys were mostly highly significant on the neutral sites and mostly not significant on the acid sites. On both site types, the species pool size declined. On the neutral site, the mean plot species richness also declined. Based on the plant-derived characteristics (Ellenberg indicator values) of the plots, light availability decreased and nitrogen availability significantly increased on the neutral sites. Conclusion: Input of eutrophying deposits and management changes are seen as key drivers of herb layer changes in the study area over the last half-century. However, our study suggests that site characteristics such as parent material and soil pH should also be considered. Together with the key drivers, incorporating differences in site characteristics may improve our understanding of temporal shifts in herb layer vegetation in response to environmental change.

The ground flora (vascular plants < 2 m high, excluding trees and shrubs) was recorded in 1974 and 1991 from 163 permanent 10 m x 10 m plots arranged on a systematic grid across Wytham Woods, near Oxford (UK). The Woods cover about 320 ha, are predominantly deciduous, but of varying ages and management types. The total number of species found (173, 167 respectively), the mean richness per plot (16.7, 17.2) and the breakdown of the species list between different species types (ancient woodland indicators, other woodland species, non-woodland species) showed no significant differences between 1974 and 1991, but mean ground cover declined from 80% to 64%. Ancient woodland indicators as a group showed less change between the years than species associated with the open glades and grassland patches in the wood. Some species increased in frequency across the woodland including Arum maculatum, Brachypodium sylvaticum, Deschampsia cespitosa and Poa trivialis; while others such as Ajuga reptans, Angelica sylvestris and Circaea lutetiana declined. Mean cover of Rubus fruticosus per plot declined from 35% to 6% and of Mercurialis perennis from 32 to 24%. More species were lost from plantations than from semi-natural stands. The results are based on only two times, so inferences on possible causes must be drawn with caution. However the results are consistent with the effects of (1) the changing conditions associated with stand growth, particularly in the plantations, and (2) an increase in browsing/ grazing by deer. No evidence was found for an effect of changing soil nitrogen levels on the vegetation. Changes in the ground flora as well as the woody layers, and in managed as well as unmanaged stands should be monitored, if nature conservation objectives are to be met.

The regeneration of Japanese black pine (Pinus thunbergii) seedlings is inhibited in a black locust (Robinia pseudoacacia)-dominated area. We examined the presence of pathogenic fungi in Japanese black pine seedlings in the area in order to determine the effect of pathogenic fungi on the inhibition of regeneration. When Japanese black pine seedlings were planted in the soil obtained from a black locust-dominated area, all of the seedlings died under low-intensity light conditions, whereas 84% of the seedlings survived in the soil obtained from a Japanese black pine-dominated area under the same light conditions. One fungus was isolated from 48.7% of the dead pine seedlings and was identified as Cylindrocladium pacificum Kang, Crous & Schoch, based on the morphological characteristics, growth, and DNA analysis. This fungus was also isolated from 50% of the dead pine seedlings in 2005 and 66.7% of the seedlings in 2006--both were planted in a black locust-dominated area. The virulence of this fungus increased under high-nitrogen and/or low-intensity light conditions. These results reveal the possibility that the soil eutrophication and shading by the black locust are conducive to a severe damping-off disease and threaten the survival and regeneration of Japanese black pine seedlings.

Pine forests are declining because of pine wilt disease and Robinia pseudoacacia, a nitrogen-fixing species, is dominating coastal forests along the Sea of Japan. We examined the effects of R. pseudoacacia on the regeneration of a native pine species, Pinus thunbergii. Two 200 m² plots were set up at the border of a P. thunbergii and R. pseudoacacia-dominated area in a coastal forest. We conducted monthly censuses of emergence, distribution and survival of pine seedlings in the plots from May 2003 to December 2004. Light intensity and soil properties were also measured to analyze the relationships between the survival of pine seedlings and environmental conditions using the Mantel test and the structural equation model. Pinus thunbergii seedlings emerged in spring-early summer and in late autumn. Survival of pine seedlings in the R. pseudoacacia-dominated subplots was less than half that in the pine-dominated subplots. Survival of pine seedlings emerging in May 2003 was significantly reduced by the lower light intensity and higher soil nitrogen in R. pseudoacacia-dominated subplots. The tendency was the same for seedlings emerging from April to May 2004. We concluded that R. pseudoacacia reduced the intensity of light during the growing season and increased the nitrogen content of soil, which resulted in inhibition of the natural regeneration of P. thunbergii.

We studied the influence of a car park on soil and vegetation within Richmond Park, UK, before and after imposition of fenced boundaries restricted public access. Soil and vegetation samples were taken before (once) and after (twice) access restrictions were enforced. The over-riding trend in all the data was for soil adjacent to the car park to be less acidic and more fertile than pristine local soils, accompanied by a more eutrophic grassland community (Lolium perenne/Trifolium repens, compared with Agrostis/Festuca grassland away from the car park). The chemical influence of the car park extended at least 50 m from its boundary. A common acidophilic collembole Folsomia quadrioculata was replaced by Cryptopygus thermophilus adjacent to the car park. There was little evidence from the vegetation data that car park closure benefitted the ecosystem, but chemical data showed signs of progressive recovery in the 2 years following restrictions. Possible explanations for the car park's influence on the local landscape are suggested to include calcareous chippings and canine faecal deposits.

Soil bacterial communities are pivotal in regulating terrestrial biogeochemical cycles and ecosystem functions. The increase in global nitrogen (N) deposition has impacted various aspects of terrestrial ecosystems, but we still have a rudimentary understanding of whether there is a threshold for N input level beyond which soil bacterial communities will experience critical transitions. Using high-throughput sequencing of the 16S rRNA gene, we examined soil bacterial responses to a long-term (13 yr), multi-level, N addition experiment in a temperate steppe of northern China. We found that plant diversity decreased in a linear fashion with increasing N addition. However, bacterial diversity responded nonlinearly to N addition, such that it was unaffected by N input below 16 g N·m−2·yr−1, but decreased substantially when N input exceeded 32 g N·m−2·yr−1. A meta-analysis across four N addition experiments in the same study region further confirmed this nonlinear response of bacterial diversity to N inputs. Substantial changes in soil bacterial community structure also occurred between N input levels of 16 to 32 g N·m−2·yr−1. Further analysis revealed that the loss of soil bacterial diversity was primarily attributed to the reduction in soil pH, whereas changes in soil bacterial community were driven by the combination of increased N availability, reduced soil pH, and changes in plant community structure. In addition, we found that N addition shifted bacterial communities toward more putatively copiotrophic taxa. Overall, our study identified a threshold of N input level for bacterial diversity and community composition. The nonlinear response of bacterial diversity to N input observed in our study indicates that although bacterial communities are resistant to low levels of N input, further increase in N input could trigger a critical transition, shifting bacterial communities to a low-diversity state.

Background and aims Sufficient soil phosphorus (P) is important for achieving optimal crop production, but excessive soil P levels may create a risk of P losses and associated eutrophication of surface waters. The aim of this study was to determine critical soil P levels for achieving optimal crop yields and minimal P losses in common soil types and dominant cropping systems in China. Methods Four long-term experiment sites were selected in China. The critical level of soil Olsen-P for crop yield was determined using the linear-plateau model. The relationships between the soil total P, Olsen-P and CaCl₂-P were evaluated using two-segment linear model to determine the soil P fertility rate and leaching change-point. Results The critical levels of soil Olsen-P for optimal crop yield ranged from 10.9 mg kg⁻¹ to 21.4 mg kg⁻¹, above which crop yield response less to the increasing of soil Olsen-P. The P leaching change-points of Olsen-P ranged from 39.9 mg kg⁻¹ to 90.2 mg kg⁻¹, above which soil CaCl₂-P greatly increasing with increasing soil Olsen-P. Similar change-point was found between soil total P and Olsen-P. Overall, the change-point ranged from 4.6 mg kg⁻¹ to 71.8 mg kg⁻¹ among all the four sites. These change-points were highly affected by crop specie, soil type, pH and soil organic matter content. Conclusions The three response curves could be used to access the soil Olsen-P status for crop yield, soil P fertility rate and soil P leaching risk for a sustainable soil P management in field.

Increasing diffuse nitrate loading of surface waters and groundwater has emerged as a major problem in many agricultural areas of the world, resulting in contamination of drinking water resources in aquifers as well as eutrophication of freshwaters and coastal marine ecosystems. Although empirical correlations between application rates of N fertilizers to agricultural soils and nitrate contamination of adjacent hydrological systems have been demonstrated, the transit times of fertilizer N in the pedosphere-hydrosphere system are poorly understood. We investigated the fate of isotopically labeled nitrogen fertilizers in a three-decade-long in situ tracer experiment that quantified not only fertilizer N uptake by plants and retention in soils, but also determined to which extent and over which time periods fertilizer N stored in soil organic matter is rereleased for either uptake in crops or export into the hydrosphere.We found that 61-65% of the applied fertilizers N were taken up by plants,whereas 12-15% of the labeled fertilizer Nwere still residing in the soil organic matter more than a quarter century after tracer application. Between 8-12% of the applied fertilizer had leaked toward the hydrosphere during the 30-y observation period. We predict that additional exports of 15N-labeled nitrate from the tracer application in 1982 toward the hydrosphere will continue for at least another five decades. Therefore, attempts to reduce agricultural nitrate contamination of aquatic systems must consider the long-term legacy of past applications of synthetic fertilizers in agricultural systems and the nitrogen retention capacity of agricultural soils.