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.

Isotopic Characterization of Mercury Atmosphere-Foliage and Atmosphere-Soil Exchange in a Swiss Subalpine Coniferous Forest.

To understand the role of vegetation and soil in regulating atmospheric Hg0, exchange fluxes and isotope signatures of Hg were characterized using a dynamic flux bag/chamber at the atmosphere-foliage/soil interfaces at the Davos-Seehornwald forest, Switzerland. The foliage was a net Hg0 sink and took up preferentially the light Hg isotopes, consequently resulting in large shifts (-3.27‰) in δ202Hg values. The soil served mostly as net sources of atmospheric Hg0 with higher Hg0 emission from the moss-covered soils than from bare soils. The negative shift of δ202Hg and Δ199Hg values of the efflux air relative to ambient air and the Δ199Hg/Δ201Hg ratio among ambient air, efflux air, and soil pore gas highlight that Hg0 re-emission was strongly constrained by soil pore gas evasion together with microbial reduction. The isotopic mass balance model indicates 8.4 times higher Hg0 emission caused by pore gas evasion than surface soil photoreduction. Deposition of atmospheric Hg0 to soil was noticeably 3.2 times higher than that to foliage, reflecting the high significance of the soil to influence atmospheric Hg0 isotope signatures. This study improves our understanding of Hg atmosphere-foliage/soil exchange in subalpine coniferous forests, which is indispensable in the model assessment of forest Hg biogeochemical cycling.

Soil-plant interactions modulated water availability of Swiss forests during the 2015 and 2018 droughts.

Central Europe has been experiencing unprecedented droughts during the last decades, stressing the decrease in tree water availability. However, the assessment of physiological drought stress is challenging, and feedback between soil and vegetation is often omitted because of scarce belowground data. Here we aimed to model Swiss forests' water availability during the 2015 and 2018 droughts by implementing the mechanistic soil-vegetation-atmosphere-transport (SVAT) model LWF-Brook90 taking advantage of regionalized depth-resolved soil information. We calibrated the model against soil matric potential data measured from 2014 to 2018 at 44 sites along a Swiss climatic and edaphic drought gradient. Swiss forest soils' storage capacity of plant-available water ranged from 53 mm to 341 mm, with a median of 137 ± 42 mm down to the mean potential rooting depth of 1.2 m. Topsoil was the primary water source. However, trees switched to deeper soil water sources during drought. This effect was less pronounced for coniferous trees with a shallower rooting system than for deciduous trees, which resulted in a higher reduction of actual transpiration (transpiration deficit) in coniferous trees. Across Switzerland, forest trees reduced the transpiration by 23% (compared to potential transpiration) in 2015 and 2018, maintaining annual actual transpiration comparable to other years. Together with lower evaporative fluxes, the Swiss forests did not amplify the blue water deficit. The 2018 drought, characterized by a higher and more persistent transpiration deficit than in 2015, triggered widespread early wilting across Swiss forests that was better predicted by the SVAT-derived mean soil matric potential in the rooting zone than by climatic predictors. Such feedback-driven quantification of ecosystem water fluxes in the soil-plant-atmosphere continuum will be crucial to predicting physiological drought stress under future climate extremes.

Assessing the response of forest productivity to climate extremes in Switzerland using model-data fusion.

The response of forest productivity to climate extremes strongly depends on ambient environmental and site conditions. To better understand these relationships at a regional scale, we used nearly 800 observation years from 271 permanent long-term forest monitoring plots across Switzerland, obtained between 1980 and 2017. We assimilated these data into the 3-PG forest ecosystem model using Bayesian inference, reducing the bias of model predictions from 14% to 5% for forest stem carbon stocks and from 45% to 9% for stem carbon stock changes. We then estimated the productivity of forests dominated by Picea abies and Fagus sylvatica for the period of 1960-2018, and tested for productivity shifts in response to climate along elevational gradient and in extreme years. Simulated net primary productivity (NPP) decreased with elevation (2.86 ± 0.006 Mg C ha-1  year-1  km-1 for P. abies and 0.93 ± 0.010 Mg C ha-1  year-1  km-1 for F. sylvatica). During warm-dry extremes, simulated NPP for both species increased at higher and decreased at lower elevations, with reductions in NPP of more than 25% for up to 21% of the potential species distribution range in Switzerland. Reduced plant water availability had a stronger effect on NPP than temperature during warm-dry extremes. Importantly, cold-dry extremes had negative impacts on regional forest NPP comparable to warm-dry extremes. Overall, our calibrated model suggests that the response of forest productivity to climate extremes is more complex than simple shift toward higher elevation. Such robust estimates of NPP are key for increasing our understanding of forests ecosystems carbon dynamics under climate extremes.

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.

Allometric equations for integrating remote sensing imagery into forest monitoring programmes.

Remote sensing is revolutionizing the way we study forests, and recent technological advances mean we are now able - for the first time - to identify and measure the crown dimensions of individual trees from airborne imagery. Yet to make full use of these data for quantifying forest carbon stocks and dynamics, a new generation of allometric tools which have tree height and crown size at their centre are needed. Here, we compile a global database of 108753 trees for which stem diameter, height and crown diameter have all been measured, including 2395 trees harvested to measure aboveground biomass. Using this database, we develop general allometric models for estimating both the diameter and aboveground biomass of trees from attributes which can be remotely sensed - specifically height and crown diameter. We show that tree height and crown diameter jointly quantify the aboveground biomass of individual trees and find that a single equation predicts stem diameter from these two variables across the world's forests. These new allometric models provide an intuitive way of integrating remote sensing imagery into large-scale forest monitoring programmes and will be of key importance for parameterizing the next generation of dynamic vegetation models.

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.

Temperature and leaf form drive contrasting sensitivity to nitrogen deposition across European forests.

Raised emissions of biologically reactive nitrogen (N) have intensified N deposition, enhancing tree productivity globally. Nonetheless, the drivers of forest sensitivity to N deposition remain unknown. We used stem growth data from 62,000 trees across Europe combined with N deposition data to track the effects of air temperature and precipitation on tree growth's sensitivity to N deposition and how it varied depending on leaf form over the past 30 years. Overall, N deposition enhanced conifer growth (until 30 kg N ha-1 yr-1) while decreasing growth for broadleaved angiosperms. Lower temperatures led to higher growth sensitivity to N deposition in conifers potentially exacerbated by N limitation. In contrast, higher temperatures stimulated growth sensitivity to N deposition for broadleaves. Higher precipitation equally increased N deposition sensitivity in all leaf forms. We conclude that air temperature and leaf form are decisive in disentangling the effect of N deposition in European forests, which provides crucial information to better predict the contribution of N deposition to land carbon sink enhancement.

Fungal community composition predicts forest carbon storage at a continental scale.

Forest soils harbor hyper-diverse microbial communities which fundamentally regulate carbon and nutrient cycling across the globe. Directly testing hypotheses on how microbiome diversity is linked to forest carbon storage has been difficult, due to a lack of paired data on microbiome diversity and in situ observations of forest carbon accumulation and storage. Here, we investigated the relationship between soil microbiomes and forest carbon across 238 forest inventory plots spanning 15 European countries. We show that the composition and diversity of fungal, but not bacterial, species is tightly coupled to both forest biotic conditions and a seven-fold variation in tree growth rates and biomass carbon stocks when controlling for the effects of dominant tree type, climate, and other environmental factors. This linkage is particularly strong for symbiotic endophytic and ectomycorrhizal fungi known to directly facilitate tree growth. Since tree growth rates in this system are closely and positively correlated with belowground soil carbon stocks, we conclude that fungal composition is a strong predictor of overall forest carbon storage across the European continent.

Forest tree growth is linked to mycorrhizal fungal composition and function across Europe.

Most trees form symbioses with ectomycorrhizal fungi (EMF) which influence access to growth-limiting soil resources. Mesocosm experiments repeatedly show that EMF species differentially affect plant development, yet whether these effects ripple up to influence the growth of entire forests remains unknown. Here we tested the effects of EMF composition and functional genes relative to variation in well-known drivers of tree growth by combining paired molecular EMF surveys with high-resolution forest inventory data across 15 European countries. We show that EMF composition was linked to a three-fold difference in tree growth rate even when controlling for the primary abiotic drivers of tree growth. Fast tree growth was associated with EMF communities harboring high inorganic but low organic nitrogen acquisition gene proportions and EMF which form contact versus medium-distance fringe exploration types. These findings suggest that EMF composition is a strong bio-indicator of underlying drivers of tree growth and/or that variation of forest EMF communities causes differences in tree growth. While it may be too early to assign causality or directionality, our study is one of the first to link fine-scale variation within a key component of the forest microbiome to ecosystem functioning at a continental scale.

A decade of monitoring at Swiss Long-Term Forest Ecosystem Research (LWF) sites: can we observe trends in atmospheric acid deposition and in soil solution acidity?

Trends in atmospheric acid deposition and in soil solution acidity from 1995 or later until 2007 were investigated at several forest sites throughout Switzerland to assess the effects of air pollution abatements on deposition and the response of the soil solution chemistry. Deposition of the major elements was estimated from throughfall and bulk deposition measurements at nine sites of the Swiss Long-Term Forest Ecosystem Research network (LWF) since 1995 or later. Soil solution was measured at seven plots at four soil depths since 1998 or later. Trends in the molar ratio of base cations to aluminum (BC/Al) in soil solutions and in concentrations and fluxes of inorganic N (NO(3)-N + NH(4)-N), sulfate (SO(4)-S), and base cations (BC) were used to detect changes in soil solution chemistry. Acid deposition significantly decreased at three out of the nine study sites due to a decrease in total N deposition. Total SO(4)-S deposition decreased at the nine sites, but due to the relatively low amount of SO(4)-S load compared to N deposition, it did not contribute to decrease acid deposition significantly. No trend in total BC deposition was detected. In the soil solution, no trend in concentrations and fluxes of BC, SO(4)-S, and inorganic N were found at most soil depths at five out of the seven sites. This suggests that the soil solution reacted very little to the changes in atmospheric deposition. A stronger reduction in base cations compared to aluminum was detected at two sites, which might indicate that acidification of the soil solution was proceeding faster at these sites.

From the comfort zone to crown dieback: Sequence of physiological stress thresholds in mature European beech trees across progressive drought.

Drought responses of mature trees are still poorly understood making it difficult to predict species distributions under a warmer climate. Using mature European beech (Fagus sylvatica L.), a widespread and economically important tree species in Europe, we aimed at developing an empirical stress-level scheme to describe its physiological response to drought. We analysed effects of decreasing soil and leaf water potential on soil water uptake, stem radius, native embolism, early defoliation and crown dieback with comprehensive measurements from overall nine hydrologically distinct beech stands across Switzerland, including records from the exceptional 2018 drought and the 2019/2020 post-drought period. Based on the observed responses to decreasing water potential we derived the following five stress levels: I (predawn leaf water potential >-0.4 MPa): no detectable hydraulic limitations; II (-0.4 to -1.3): persistent stem shrinkage begins and growth ceases; III (-1.3 to -2.1): onset of native embolism and defoliation; IV (-2.1 to -2.8): onset of crown dieback; V (<-2.8): transpiration ceases and crown dieback is >20%. Our scheme provides, for the first time, quantitative thresholds regarding the physiological downregulation of mature European beech trees under drought and therefore synthesises relevant and fundamental information for process-based species distribution models. Moreover, our study revealed that European beech is drought vulnerable, because it still transpires considerably at high levels of embolism and because defoliation occurs rather as a result of embolism than preventing embolism. During the 2018 drought, an exposure to the stress levels III-V of only one month was long enough to trigger substantial crown dieback in beech trees on shallow soils. On deep soils with a high water holding capacity, in contrast, water reserves in deep soil layers prevented drought stress in beech trees. This emphasises the importance to include local data on soil water availability when predicting the future distribution of European beech.

Extreme summer heat and drought lead to early fruit abortion in European beech.

Years with high fruit production, known as mast years, are the usual reproduction strategy of European beech. Harsh weather conditions such as frost during flowering can lead to pollination failure in spring. It has been assumed that mast is controlled by flowering, and that after successful pollination, high amounts of fruits and seeds would be produced. However, the extremely hot and dry European summer of 2018 showed that despite successful pollination, beechnuts did not develop or were only abundant in a few forest stands. An in-depth analysis of three forest sites of European beech from the Swiss Long-Term Forest Ecosystem Research Programme over the last 15-19 years revealed for the first time that extreme summer heat and drought can act as an "environmental veto", leading to early fruit abortion. Within the forest stands in years with fruit abortion, summer mean temperatures were 1.5 °C higher and precipitation sums were 45% lower than the long-term average. Extreme summer heat and drought, together with frost during flowering, are therefore disrupting events of the assumed biennial fruiting cycle in European beech.

Dynamic modelling of the long term behaviour of cadmium, lead and mercury in Swiss forest soils using CHUM-AM.

The applicability of the dynamic soil model CHUM-AM was tested to simulate concentrations of Cd, Pb and Hg in five Swiss forest soils. Soil cores of up to 50 cm depth were sampled and separated into two defined soil layers. Soil leachates were collected below the litter by zero-tension lysimeters and at 15 and 50 cm soil depths by tension lysimeters over two years. The concentrations of Cd, Pb and Hg in the solid phase and soil solution were measured by ICP-MS (Cd, Pb) or CV-AFS (Hg). Measured metal concentrations were compared with modelled concentrations using CHUM-AM. Additionally we ran the model with three different deposition scenarios (current deposition; maximum acceptable deposition according to the Swiss ordinance on Air Pollution Control; critical loads according to CLRTAP) to predict metal concentrations in the soils for the next 1000 years. Assuming current loads concentrations of Cd and Pb showed varying trends (increasing/decreasing) between the soils. Soils rich in organic carbon or with a high pH value showed increasing trends in Cd and Pb concentrations whereas the concentrations in the other soils decreased. In contrast Hg concentrations are predicted to further increase in all soils. Critical limits for Pb and Hg will partly be exceeded by current loads or by the critical loads proposed by the CLRTAP but the critical limits for Cd will rarely be reached within the next 1000 years. In contrast, maximal acceptable deposition will partly lead to concentrations above the critical limits for Pb in soils within the next 400 years, whereas the acceptable deposition of Cd will not lead to concentrations above the proposed critical limits. In conclusion the CHUM-AM model is able to accurately simulate heavy metal (Cd, Pb and Hg) concentrations in Swiss forest soils of various soil properties.

Atmospheric deposition and ozone levels in Swiss forests: are critical values exceeded?

Air pollution affects forest health through atmospheric deposition of acidic and nitrogen compounds and elevated levels of tropospheric ozone (O3). In 1985, a monitoring network was established across Europe and various research efforts have since been undertaken to define critical values. We measured atmospheric deposition of acidity and nitrogen as well as ambient levels of O3 on 12, 13, and 14 plots, respectively, in the framework of the Swiss Long-Term Forest Ecosystem Research (LWF) in the period from 1995 to 2002. We estimated the critical loads of acidity and of nitrogen, using the steady state mass balance approach, and calculated the critical O3 levels using the AOT40 approach. The deposition of acidity exceeded the critical loads on 2 plots and almost reached them on 4 plots. The median of the measured molar ratio of base nutrient cations to total dissolved aluminium (Bc/Al) in the soil solution was higher than the critical value of 1 for all depths, and also at the plots with an exceedance of the critical load of acidity. For nitrogen, critical loads were exceeded on 8 plots and deposition likely represents a long-term ecological risk on 3 to 10 plots. For O3, exceedance of critical levels was recorded on 12 plots, and led to the development of typical O3-induced visible injury on trees and shrubs, but not for all plots due to (1) the site specific composition of O3 sensitive and tolerant plant species, and (2) the influence of microclimatic site conditions on the stomatal behaviour, i.e., O3 uptake.

Atmospheric deposition on Swiss Long-Term Forest Ecosystem Research (LWF) plots.

Atmospheric deposition of the major elements was estimated from throughfall and bulk deposition measurements on 13 plots of the Swiss Long-Term Forest Ecosystem Research (LWF) between 1995 and 2001. Independent estimates of the wet and dry deposition of nitrogen (N) and sulfur (S) on these same plots were gained from combined simplified models. The highest deposition fluxes were measured at Novaggio (Southern Switzerland), exposed to heavy air pollution originating from the Po Plain, with throughfall fluxes averaging 29 kg ha(-1) a(-1) for N and 15 kg ha(-1) a(-1) for S. Low deposition fluxes were measured on the plots above 1800 m, with throughfall fluxes lower than 4.5 kg ha(-1) a(-1) for N and lower than 3 kg ha(-1) a(-1) for S. The wet deposition of N and S derived from bulk deposition was close to the modeled wet deposition, but the dry deposition derived from throughfall was significantly lower than the modeled dry deposition for both compounds. However, both the throughfall method and the model yielded total deposition estimates of N which exceeded the critical loads calculated on the basis of long-term mass balance considerations. These estimates were within or above the range of empirical critical loads except above 1800 m.