Widespread breakdown in masting in European beech due to rising summer temperatures.

Climate change effects on tree reproduction are poorly understood, even though the resilience of populations relies on sufficient regeneration to balance increasing rates of mortality. Forest-forming tree species often mast, i.e. reproduce through synchronised year-to-year variation in seed production, which improves pollination and reduces seed predation. Recent observations in European beech show, however, that current climate change can dampen interannual variation and synchrony of seed production and that this masting breakdown drastically reduces the viability of seed crops. Importantly, it is unclear under which conditions masting breakdown occurs and how widespread breakdown is in this pan-European species. Here, we analysed 50 long-term datasets of population-level seed production, sampled across the distribution of European beech, and identified increasing summer temperatures as the general driver of masting breakdown. Specifically, increases in site-specific mean maximum temperatures during June and July were observed across most of the species range, while the interannual variability of population-level seed production (CVp) decreased. The declines in CVp were greatest, where temperatures increased most rapidly. Additionally, the occurrence of crop failures and low seed years has decreased during the last four decades, signalling altered starvation effects of masting on seed predators. Notably, CVp did not vary among sites according to site mean summer temperature. Instead, masting breakdown occurs in response to warming local temperatures (i.e. increasing relative temperatures), such that the risk is not restricted to populations growing in warm average conditions. As lowered CVp can reduce viable seed production despite the overall increase in seed count, our results warn that a covert mechanism is underway that may hinder the regeneration potential of European beech under climate change, with great potential to alter forest functioning and community dynamics.

Simulating tree growth response to climate change in structurally diverse oak and beech forests.

This study aimed to simulate oak and beech forest growth under various scenarios of climate change and to evaluate how the forest response depends on site properties and particularly on stand characteristics using the individual process-based model HETEROFOR. First, this model was evaluated on a wide range of site conditions. We used data from 36 long-term forest monitoring plots to initialize, calibrate, and evaluate HETEROFOR. This evaluation showed that HETEROFOR predicts individual tree radial growth and height increment reasonably well under different growing conditions when evaluated on independent sites. In our simulations under constant CO2 concentration ([CO2]cst) for the 2071-2100 period, climate change induced a moderate net primary production (NPP) gain in continental and mountainous zones and no change in the oceanic zone. The NPP changes were negatively affected by air temperature during the vegetation period and by the annual rainfall decrease. To a lower extent, they were influenced by soil extractable water reserve and stand characteristics. These NPP changes were positively affected by longer vegetation periods and negatively by drought for beech and larger autotrophic respiration costs for oak. For both species, the NPP gain was much larger with rising CO2 concentration ([CO2]var) mainly due to the CO2 fertilisation effect. Even if the species composition and structure had a limited influence on the forest response to climate change, they explained a large part of the NPP variability (44% and 34% for [CO2]cst and [CO2]var, respectively) compared to the climate change scenario (5% and 29%) and the inter-annual climate variability (20% and 16%). This gives the forester the possibility to act on the productivity of broadleaved forests and prepare them for possible adverse effects of climate change by reinforcing their resilience.

Responses of forest ecosystems in Europe to decreasing nitrogen deposition.

Average nitrogen (N) deposition across Europe has declined since the 1990s. This resulted in decreased N inputs to forest ecosystems especially in Central and Western Europe where deposition levels are highest. While the impact of atmospheric N deposition on forests has been receiving much attention for decades, ecosystem responses to the decline in N inputs received less attention. Here, we review observational studies reporting on trends in a number of indicators: soil acidification and eutrophication, understory vegetation, tree nutrition (foliar element concentrations) as well as tree vitality and growth in response to decreasing N deposition across Europe. Ecosystem responses varied with limited decrease in soil solution nitrate concentrations and potentially also foliar N concentrations. There was no large-scale response in understory vegetation, tree growth, or vitality. Experimental studies support the observation of a more distinct reaction of soil solution and foliar element concentrations to changes in N supply compared to the three other parameters. According to the most likely scenarios, further decrease of N deposition will be limited. We hypothesize that this expected decline will not cause major responses of the parameters analysed in this study. Instead, future changes might be more strongly controlled by the development of N pools accumulated within forest soils, affected by climate change and forest management.

Author Correction: Environment and host as large-scale controls of ectomycorrhizal fungi.

Change history: In the HTML version of this Article, author 'Filipa Cox' had no affiliation in the author list, although she was correctly associated with affiliation 3 in the PDF. In addition, the blue circles for 'oak' were missing from Extended Data Fig. 1. These errors have been corrected online.

Environment and host as large-scale controls of ectomycorrhizal fungi.

Explaining the large-scale diversity of soil organisms that drive biogeochemical processes-and their responses to environmental change-is critical. However, identifying consistent drivers of belowground diversity and abundance for some soil organisms at large spatial scales remains problematic. Here we investigate a major guild, the ectomycorrhizal fungi, across European forests at a spatial scale and resolution that is-to our knowledge-unprecedented, to explore key biotic and abiotic predictors of ectomycorrhizal diversity and to identify dominant responses and thresholds for change across complex environmental gradients. We show the effect of 38 host, environment, climate and geographical variables on ectomycorrhizal diversity, and define thresholds of community change for key variables. We quantify host specificity and reveal plasticity in functional traits involved in soil foraging across gradients. We conclude that environmental and host factors explain most of the variation in ectomycorrhizal diversity, that the environmental thresholds used as major ecosystem assessment tools need adjustment and that the importance of belowground specificity and plasticity has previously been underappreciated.

The response of soil solution chemistry in European forests to decreasing acid deposition.

Acid deposition arising from sulphur (S) and nitrogen (N) emissions from fossil fuel combustion and agriculture has contributed to the acidification of terrestrial ecosystems in many regions globally. However, in Europe and North America, S deposition has greatly decreased in recent decades due to emissions controls. In this study, we assessed the response of soil solution chemistry in mineral horizons of European forests to these changes. Trends in pH, acid neutralizing capacity (ANC), major ions, total aluminium (Altot ) and dissolved organic carbon were determined for the period 1995-2012. Plots with at least 10 years of observations from the ICP Forests monitoring network were used. Trends were assessed for the upper mineral soil (10-20 cm, 104 plots) and subsoil (40-80 cm, 162 plots). There was a large decrease in the concentration of sulphate (SO42-) in soil solution; over a 10-year period (2000-2010), SO42- decreased by 52% at 10-20 cm and 40% at 40-80 cm. Nitrate was unchanged at 10-20 cm but decreased at 40-80 cm. The decrease in acid anions was accompanied by a large and significant decrease in the concentration of the nutrient base cations: calcium, magnesium and potassium (Bc = Ca2+  + Mg2+  + K+ ) and Altot over the entire dataset. The response of soil solution acidity was nonuniform. At 10-20 cm, ANC increased in acid-sensitive soils (base saturation ≤10%) indicating a recovery, but ANC decreased in soils with base saturation >10%. At 40-80 cm, ANC remained unchanged in acid-sensitive soils (base saturation ≤20%, pHCaCl2 ≤ 4.5) and decreased in better-buffered soils (base saturation >20%, pHCaCl2 > 4.5). In addition, the molar ratio of Bc to Altot either did not change or decreased. The results suggest a long-time lag between emission abatement and changes in soil solution acidity and underline the importance of long-term monitoring in evaluating ecosystem response to decreases in deposition.

Increasing trends of dissolved organic nitrogen (DON) in temperate forests under recovery from acidification in Flanders, Belgium.

We evaluated trends (2005-2013) and patterns of dissolved organic nitrogen (DON) and its ratio with dissolved organic carbon (DOC), DOC:DON in atmospheric deposition and soil solution of five Level II plots of the International Co-operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests (ICP Forests) in Flanders, Northern Belgium. The primary aim was to confirm positive postulated trends in DON levels and DOC:DON under on-going recovery from acidification. The DON concentrations (0.95-1.41 mg L(-1)) and fluxes (5.6-8.3 kg ha(-1)y(-1)) in throughfall were about twice as high compared to precipitation in the open field (0.40-0.48 mg L(-1), 3.0-3.9 kg ha(-1)y(-1)). Annual soil profile leaching losses of DON varied between 1.2 and 3.7 kg ha(-1)y(-1). The highest soil DON concentrations and fluxes were observed beneath the O horizon (1.84-2.36 mg L(-1), 10.1-12.3 kg ha(-1)y(-1)). Soil solution concentrations and fluxes of DON showed significant increasing trends. Temporarily soil solution DOC:DON rose following an exceptionally long spring drought in 2007, suggesting an effect of drying and rewetting on DOM composition. Further research is needed to test the dependence of DON and DOC:DON on factors such as latitude, forest cover, length of the growing season, hydrology and topography. Nonetheless, even with considerable variation in soil type, level of base saturation, and soil texture in the five included ICP Forests Level II plots, all data revealed a proportionally larger positive response of DON flux than DOC to recovery from acidification.

Tree mineral nutrition is deteriorating in Europe.

The response of forest ecosystems to increased atmospheric CO2 is constrained by nutrient availability. It is thus crucial to account for nutrient limitation when studying the forest response to climate change. The objectives of this study were to describe the nutritional status of the main European tree species, to identify growth-limiting nutrients and to assess changes in tree nutrition during the past two decades. We analysed the foliar nutrition data collected during 1992-2009 on the intensive forest monitoring plots of the ICP Forests programme. Of the 22 significant temporal trends that were observed in foliar nutrient concentrations, 20 were decreasing and two were increasing. Some of these trends were alarming, among which the foliar P concentration in F. sylvatica, Q. Petraea and P. sylvestris that significantly deteriorated during 1992-2009. In Q. Petraea and P. sylvestris, the decrease in foliar P concentration was more pronounced on plots with low foliar P status, meaning that trees with latent P deficiency could become deficient in the near future. Increased tree productivity, possibly resulting from high N deposition and from the global increase in atmospheric CO2, has led to higher nutrient demand by trees. As the soil nutrient supply was not always sufficient to meet the demands of faster growing trees, this could partly explain the deterioration of tree mineral nutrition. The results suggest that when evaluating forest carbon storage capacity and when planning to reduce CO2 emissions by increasing use of wood biomass for bioenergy, it is crucial that nutrient limitations for forest growth are considered.

Environmental drivers of ectomycorrhizal communities in Europe's temperate oak forests.

Ectomycorrhizal fungi are major ecological players in temperate forests, but they are rarely used in measures of forest condition because large-scale, high-resolution, standardized and replicated belowground data are scarce. We carried out an analysis of ectomycorrhizas at 22 intensively monitored long-term oak plots, across nine European countries, covering complex natural and anthropogenic environmental gradients. We found that at large scales, mycorrhizal richness and evenness declined with decreasing soil pH and root density, and with increasing atmospheric nitrogen deposition. Shifts in mycorrhizas with different functional traits were detected; mycorrhizas with structures specialized for long-distance transport related differently to most environmental variables than those without. The dominant oak-specialist Lactarius quietus, with limited soil exploration abilities, responds positively to increasing nitrogen inputs and decreasing pH. In contrast, Tricholoma, Cortinarius and Piloderma species, with medium-distance soil exploration abilities, show a consistently negative response. We also determined nitrogen critical loads for moderate (9.5-13.5 kg N/ha/year) and drastic (17 kg N/ha/year) changes in belowground mycorrhizal root communities in temperate oak forests. Overall, we generated the first baseline data for ectomycorrhizal fungi in the oak forests sampled, identified nitrogen pollution as one of their major drivers at large scales and revealed fungi that individually and/or in combination with others can be used as belowground indicators of environmental characteristics.