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 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.

Article (refereed) -postprintTipping, E.; Benham, S.; Boyle, J.F.; Crow, P.; Davies, J.; Fischer, U.; Guyatt, H.; Helliwell, R.; Jackson-Blake, L.; Lawlor, A.J.; Monteith, D.T.; Rowe, E.C.; Toberman, H. 2014. Atmospheric deposition of phosphorus to land and freshwater. Environmental Science: Processes and Impacts, 16 (7). 1608-1617. 10.1039/c3em00641g Contact CEH NORA team at noraceh@ceh.ac.ukThe NERC and CEH trademarks and logos ('the Trademarks') are registered trademarks of NERC in the UK and other countries, and may not be used without the prior written consent of the Trademark owner. Oceania, and South-Central America. The deposition rates are log-normally distributed, 30 and for the whole data set the geometric mean deposition rates are 0.027, 0.019 and 31 0.14 g m -2 a -1 for TP, FTP and PO 4 -P respectively. At smaller scales there is little 32 systematic spatial variation, except for high deposition rates at some sites in Germany, 33 likely due to local agricultural sources. In cases for which PO 4 -P was determined as well 34 as one of the other forms of P, strong parallels between logarithmic values were found. 35Based on the directly-measured deposition rates to land, and published estimates of P 36 deposition to the oceans, we estimate a total annual transfer of P to and from the 37 atmosphere of 3.7 Tg. However, much of the phosphorus in larger particles (principally 38 primary biological aerosol particles) is probably redeposited near to its origin, so that 39 long-range transport, important for tropical forests, large areas of peatland and the 40 oceans, mainly involves fine dust from deserts and soils, as described by the simulations 41of Mahowald et al. (Global Biogeochemical Cycles 22, GB4026, 2008). We suggest that 42 local release to the atmosphere and subsequent deposition bring about a pseudo-43 diffusive redistribution of P in the landscape, with P-poor ecosystems, for example 44 ombrotrophic peatlands and oligotrophic lakes, gaining at the expense of P-rich ones. 45Simple calculations suggest that atmospheric transport could bring about significant local 46 redistribution of P among terrestrial ecosystems.Although most atmospherically 47 transported P is natural in origin, local transfers from fertilised farmland to P-poor 48 ecosystems may be significant, and this requires further research. 49 50

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.

Local biodiversity trends over time are likely to be decoupled from global trends, as local processes may compensate or counteract global change. We analyze 161 long-term biological time series (15-91 years) collected across Europe, using a comprehensive dataset comprising 6,200 marine, freshwater and terrestrial taxa. We test whether (i) local long-term biodiversity trends are consistent among biogeoregions, realms and taxonomic groups, and (ii) changes in biodiversity correlate with regional climate and local conditions. Our results reveal that local trends of abundance, richness and diversity differ among biogeoregions, realms and taxonomic groups, demonstrating that biodiversity changes at local scale are often complex and cannot be easily generalized. However, we find increases in richness and abundance with increasing temperature and naturalness as well as a clear spatial pattern in changes in community composition (i.e. temporal taxonomic turnover) in most biogeoregions of Northern and Eastern Europe.

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Litterfall is a major, yet poorly studied, process within forest ecosystems globally. It is important for carbon dynamics, edaphic communities, and maintaining site fertility. Reliable information on the carbon and nutrient input from litterfall, provided by litter traps, is relevant to a wide audience including policy makers and soil scientists. We used litterfall observations of 320 plots from the pan‐European forest monitoring network of the “International Co‐operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests” to quantify litterfall fluxes. Eight litterfall models were evaluated (four using climate information and four using biomass abundance). We scaled up our results to the total European forest area and quantified the contribution of litterfall to the forest carbon cycle using net primary production aggregated by bioregions (north, central, and south) and by forest types (conifers and broadleaves). The 1,604 analyzed annual litterfall observations indicated an average carbon input of 224 g C · m−2 · year−1 (annual nutrient inputs 4.49 g N, 0.32 g P, and 1.05 g K · m−2), representing a substantial percentage of net primary production from 36% in north Europe to 32% in central Europe. The annual turnover of carbon and nutrient in broadleaf canopies was larger than for conifers. The evaluated models provide large‐scale litterfall predictions with a bias less than 10%. Each year litterfall in European forests transfers 351 Tg C, 8.2 Tg N, 0.6 Tg P, and 1.9 Tg K to the forest floor. The performance of litterfall models may be improved by including foliage biomass and proxies for forest management.

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