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The Mediterranean is among the regions predicted to be most affected by climate change due to rising temperatures and increasing frequency as well as intensity of extreme weather events, such as drought. These shifts in climatic conditions might lead to changes in species community composition by enabling the increase of drought-tolerant species at the expense of less tolerant ones. This hypothesis was tested in the current study using chlorophyll fluorescence data from a 21-year precipitation exclusion experiment in a Mediterranean forest for two co-dominant Quercus ilex L. and Phillyrea latifolia L. species with contrasting low and high levels of drought tolerance, respectively. The maximum potential quantum efficiency of photosystem II (PSII) (Fv/Fm), photochemical efficiency of PSII (yield) and non-photochemical quenching (NPQ) varied seasonally. The Fv/Fm and NPQ levels were positively correlated with air temperature and standardized precipitation-evapotranspiration index (SPEI), whereas yield, which was greater under drought treatment, was negatively associated with vapor pressure deficit and SPEI. The Fv/Fm values showed similar increase in the two species over the 21-year study period regardless of treatment and in parallel to progressive warming. By contrast, yield values were higher in Q. ilex than in P. latifolia, while NPQ values were higher in P. latifolia than in Q. ilex. Notably, high yield values were also observed in the drought-treated plots. Throughout the study, plants in the drought-treated plots exhibited decreased basal area, leaf biomass and aerial cover due to high stem mortality. In addition, a continuous increase in temperature was detected in summer and autumn, which might explain the observed increase in Fv/Fm values over the study period. Higher yield and lower NPQ detected in Q. ilex could be attributed to less competition for resources in the drought-treated plots and acclimation of Q. ilex plants over the study period. Our results indicate that reduction in stem density could improve forest resilience to climate change-induced drought conditions.

Global net land carbon uptake or net biome production (NBP) has increased during recent decades1. Whether its temporal variability and autocorrelation have changed during this period, however, remains elusive, even though an increase in both could indicate an increased potential for a destabilized carbon sink2,3. Here, we investigate the trends and controls of net terrestrial carbon uptake and its temporal variability and autocorrelation from 1981 to 2018 using two atmospheric-inversion models, the amplitude of the seasonal cycle of atmospheric CO2 concentration derived from nine monitoring stations distributed across the Pacific Ocean and dynamic global vegetation models. We find that annual NBP and its interdecadal variability increased globally whereas temporal autocorrelation decreased. We observe a separation of regions characterized by increasingly variable NBP, associated with warm regions and increasingly variable temperatures, lower and weaker positive trends in NBP and regions where NBP became stronger and less variable. Plant species richness presented a concave-down parabolic spatial relationship with NBP and its variability at the global scale whereas nitrogen deposition generally increased NBP. Increasing temperature and its increasing variability appear as the most important drivers of declining and increasingly variable NBP. Our results show increasing variability of NBP regionally that can be mostly attributed to climate change and that may point to destabilization of the coupled carbon–climate system.

Consistent information on the current elemental composition of vegetation at global scale and the variables that determine it is lacking. To fill this gap, we gathered a total of 30 912 georeferenced records on woody plants foliar concentrations of nitrogen (N), phosphorus (P) and potassium (K) from published databases, and produced global maps of foliar N, P and K concentrations for woody plants using neural networks at a resolution of 1 km(2). We used data for climate, atmospheric deposition, soil and morphoclimatic groups to train the neural networks. Foliar N, P and K do not follow clear global latitudinal patterns but are consistent with the hypothesis of soil substrate age. We additionally built generalized linear mixed models to investigate the evolutionary history effect together with the effects of environmental effects. In this comparison, evolutionary history effects explained most of the variability in all cases (mostly > 60%). These results emphasize the determinant role of evolutionary history in foliar elemental composition, which should be incorporated in upcoming dynamic global vegetation models.

Significant gaps remain in understanding the response of plant reproduction to environmental change. This is partly because measuring reproduction in long-lived plants requires direct observation over many years and such datasets have rarely been made publicly available. Here we introduce MASTREE+, a data set that collates reproductive time-series data from across the globe and makes these data freely available to the community. MASTREE+ includes 73,828 georeferenced observations of annual reproduction (e.g. seed and fruit counts) in perennial plant populations worldwide. These observations consist of 5971 population-level time-series from 974 species in 66 countries. The mean and median time-series length is 12.4 and 10 years respectively, and the data set includes 1122 series that extend over at least two decades (≥20 years of observations). For a subset of well-studied species, MASTREE+ includes extensive replication of time-series across geographical and climatic gradients. Here we describe the open-access data set, available as a.csv file, and we introduce an associated web-based app for data exploration. MASTREE+ will provide the basis for improved understanding of the response of long-lived plant reproduction to environmental change. Additionally, MASTREE+ will enable investigation of the ecology and evolution of reproductive strategies in perennial plants, and the role of plant reproduction as a driver of ecosystem dynamics. EEA Santa Cruz Fil: Hacket-Pain, Andrew. University of Liverpool. School of Environmental Sciences. Department of Geography and Planning; Reino Unido Fil: Foest, Jessie J. University of Liverpool. School of Environmental Sciences. Department of Geography and Planning; Reino Unido Fil: Pearse, Ian S. U.S. Fort Collins Science Center. Geological Survey; Estados Unidos Fil: LaMontagne, Jalene M. DePaul University. Department of Biological Sciences; Estados Unidos Fil: Koenig, Walter D. University of California Berkeley. Hastings Reservation; Estados Unidos Fil: Vacchiano, Giorgio. University of Milan. Milan Department of Agricultural and Environmental Sciences; Italia. Fil: Bogdziewicz, Michał. Adam Mickiewicz University. Faculty of Biology. Institute of Environmental Biology; Polonia Fil: Bogdziewicz, Michał. University Grenoble Alpes. INRAE, LESSEM; Francia Fil: Caignard, Thomas. Université Bordeaux. INRAE, BIOGECO; Francia Fil: Celebias, Paulina. Adam Mickiewicz University. Faculty of Biology. Institute of Environmental Biology; Polonia Fil: Peri, Pablo Luis. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Santa Cruz; Argentina. Fil: Peri, Pablo Luis. Universidad Nacional de la Patagonia Austral; Argentina. Fil: Peri, Pablo Luis. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Fil: Ascoli, Davide. University of Torino. Department of Agricultural. Forest and Food Sciences (DISAFA); Italia Fil: Gallo, Leonardo Ariel. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Bariloche; Argentina Fil: Gallo, Leonardo Ariel. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Investigaciones Forestales y Agropecuarias Bariloche; Argentina Fil: Marchelli, Paula. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Bariloche; Argentina Fil: Marchelli, Paula. Consejo Nacional de Investigaciones Cientificas y Tecnicas. Instituto de Investigaciones Forestales y Agropecuarias Bariloche; Argentina

Applying industrial waste amendments, such as steel slag and biochar, to soils is an increasingly common practice to improve soil fertility, crop yield, and soil carbon (C) pool storage and stability. However, the effects of separate and combined applications of slag and biochar on total and active soil organic C (SOC) pools and the associated relationships between soil microorganisms and C cycling are unclear. Therefore, this study examined the effects of steel slag, rice straw biochar, and slag+biochar amendments to subtropical early and late rice paddy soils on total and active SOC concentrations and microbial abundance and diversity. The results showed that slag+biochar increased SOC content in early and late rice, with increases in dissolved organic C of 28.7 and 52.6 % in the early and late rice jointing stages, respectively. Soil organic C was positively correlated with soil C: N ratios in the two crop seasons. Applications of slag and biochar alone increased the diversity of soil bacteria in early rice but decreased in late rice. Meanwhile, the combined application of slag and biochar decreased diversity of soil bacterial community diversity and increased the relative abundance of some beneficial bacterial taxa, such as Pseudomonas, Bacillus, Flavisolibacter, and Ferruginibacte which are related to SOC cycling and sequestration. The results highlight that the combined slag and biochar amendments can improve soil specific properties, such as pH and SOC stocks, which elicit alterations in soil bacterial composition, especially of some keystone genera, altogether resulting in enhanced soil C sequestration and C pool stability.

The leaf economics spectrum1,2 and the global spectrum of plant forms and functions3 revealed fundamental axes of variation in plant traits, which represent different ecological strategies that are shaped by the evolutionary development of plant species2. Ecosystem functions depend on environmental conditions and the traits of species that comprise the ecological communities4. However, the axes of variation of ecosystem functions are largely unknown, which limits our understanding of how ecosystems respond as a whole to anthropogenic drivers, climate and environmental variability4,5. Here we derive a set of ecosystem functions6 from a dataset of surface gas exchange measurements across major terrestrial biomes. We find that most of the variability within ecosystem functions (71.8%) is captured by three key axes. The first axis reflects maximum ecosystem productivity and is mostly explained by vegetation structure. The second axis reflects ecosystem water-use strategies and is jointly explained by variation in vegetation height and climate. The third axis, which represents ecosystem carbon-use efficiency, features a gradient related to aridity, and is explained primarily by variation in vegetation structure. We show that two state-of-the-art land surface models reproduce the first and most important axis of ecosystem functions. However, the models tend to simulate more strongly correlated functions than those observed, which limits their ability to accurately predict the full range of responses to environmental changes in carbon, water and energy cycling in terrestrial ecosystems7,8., Three key axes of variation of ecosystem functional changes and their underlying causes are identified from a dataset of surface gas exchange measurements across major terrestrial biomes and climate zones.

Our study suggests that the global CO 2 fertilization effect (CFE) on vegetation photosynthesis has declined during the past four decades. The Comments suggest that the temporal inconsistency in AVHRR data and the attribution method undermine the results’ robustness. Here, we provide additional evidence that these arguments did not affect our finding and that the global decline in CFE is robust.

Kelly's commentary regarding our paper1, claiming that nutrient scarcity cannot select for masting behaviour in plants, initiated a fruitful discussion on traditionally settled hypotheses about the evolution of reproductive behaviour in plants. In his commentary, Prof. Kelly raises support for a contrasting hypothesis explaining our observation that temporally variable seed production is more pronounced under nutrient scarcity, namely that nutrient scarcity does not directly cause seed production variability, but instead increases variability induced by economies of scale. The commentary hinges mainly on the argument that an economy of scale (EOS) is necessary to select for highly variable seed production. It also points out that there are no mechanisms by which nutrient scarcity would select for that particular trait over generations. In reply to the stimulating comment, we 1) propose a mechanism by which nutrient scarcity may select for highly variable seed production, with larger-scale weather patterns inducing masting synchrony across populations; 2) we further discuss why wind-pollination and predator satiation, the EOS suggested by Kelly, cannot be the only selective pressures that select for highly variable reproduction.

Continuous atmospheric CO2 monitoring data indicate an increase in the amplitude of seasonal CO2-cycle exchange (SCANBP) in northern high latitudes. The major drivers of enhanced SCANBP remain unclear and intensely debated, with land-use change, CO2 fertilization and warming being identified as likely contributors. We integrated CO2-flux data from two atmospheric inversions (consistent with atmospheric records) and from 11 state-of-the-art land-surface models (LSMs) to evaluate the relative importance of individual contributors to trends and drivers of the SCANBP of CO2 fluxes for 1980–2015. The LSMs generally reproduce the latitudinal increase in SCANBP trends within the inversions range. Inversions and LSMs attribute SCANBP increase to boreal Asia and Europe due to enhanced vegetation productivity (in LSMs) and point to contrasting effects of CO2 fertilization (positive) and warming (negative) on SCANBP. Our results do not support land-use change as a key contributor to the increase in SCANBP. The sensitivity of simulated microbial respiration to temperature in LSMs explained biases in SCANBP trends, which suggests that SCANBP could help to constrain model turnover times.

Sea spray aerosol (SSA) is responsible for the large-scale transfer of particles from the sea to the land, leading to significant deposition of a range of ions, predominantly Na+, K+, Mg2+, Ca2+, and Cl−. Up to now, there has been little research into the effects of SSA on spring water chemistry. Therefore, we sampled 303 semi-natural springs across Catalonia (NE Iberian Peninsula) and analysed the concentrations of 20 different ions and elements, and determined the impact of SSA (using distance to the coast as a proxy) as well as climate, lithology and human disturbances. We found that distance to the coast had a clear effect on the water chemical composition of springs, while accounting for potentially confounding factors such as anthropogenic water pollution (nitrate, NO3−), differences in lithology and annual rainfall. Our results showed that springs located closer to the coast had higher Cl−, SO42−, Na+, Mg2+, K+ and Ca2+ concentrations than those of springs located further away. Precipitation was generally negatively correlated with the concentration of almost all elements analysed. The concentration of NO3− increased with distance to the coast, concurrently with farming activities, located mainly inland in the study area. These results demonstrate that SSA has an important effect on the groundwater of coastal zones, up to a distance of around 70 km from the coastline. This analysis reveals the main natural and human processes that influence spring water chemistry in this Mediterranean region, information that could be helpful in similar regions for ecological studies, water quality policies, and for the improvement of predictions in the current context of global change.

Differences in the elemental composition of plants, mainly C, N, and P, have been shown to be related to differences in their nutritional status, and their morphological and functional traits. The relationship between morphological traits and micronutrients and trace elements, however, has been much less studied. Additionally, in bryophytes, research devoted to investigating these relationships is still very scarce. Here, we analysed 80 samples from 29 aquatic and semi-aquatic (hygrophytic) moss species living in Mediterranean springs to investigate the relationship between moss nutrient concentrations and their micro- and macroscopic morphological traits and growth forms. We found that, across species, the elemental concentration of mosses was more tightly linked to macroscopic traits than to microscopic traits. Growth forms could also be successfully explained by the concentration of elements in mosses. Apart from macronutrients and their stoichiometric ratios (C:N, C:P, and N:P), micronutrients and trace elements were also important variables predicting moss morphological traits and growth forms. Additionally, our results showed that microscopic traits were well related to macroscopic traits. Overall, our results clearly indicate that the elemental composition of mosses can be used to infer their morphological traits, and that elements other than macronutrients should be taken into account to achieve a good representation of their morphological and, potentially, functional traits when comparing the elemental composition across species.

Altres ajuts: beca de la Fundació La Caixa LCF/BQ/PI21/11830010. The elemental composition of plants (the elementome) is a reliable indicator of their functional traits and the ecological strategies that they follow, and thus represents a good predictor of how ecosystems work. Biodiversity and, especially, functional diversity are also widely recognized as important drivers of ecosystem functioning, mainly because of niche partitioning amongst different species. Here, I review evidence indicating that plant elementomes relate to their ecological niches and how plant elemental concentrations may shift in response to abiotic and biotic drivers. I propose the use of ecosystem elementome diversity as a universal metric to compare ecosystems and investigate diversity-ecosystem functioning relationships. Future research using this promising novel approach will bring together elementomes, diversity, and ecosystem functioning.

Grasslands are key repositories of biodiversity and carbon storage and are heavily impacted by effects of global warming and changes in precipitation regimes. Patterns of grassland dynamics associated with variability in future climate conditions across spatiotemporal scales are yet to be adequately quantified. Here, we performed a global meta-analysis of year and growing season sensitivities of vegetation aboveground biomass (AGB), aboveground net primary productivity (ANPP), and species richness (SR) and diversity (Shannon index, H) to experimental climate warming and precipitation shifts. All four variables were sensitive to climate change. Their sensitivities to shifts in precipitation were correlated with local background water availability, such as mean annual precipitation (MAP) and aridity, and AGB and ANPP sensitivities were greater in dry habitats than in nonwater-limited habitats. There was no effect of duration of experiment (short vs long term) on sensitivities. Temporal trends in ANPP and SR sensitivity depended on local water availability; ANPP sensitivity to warming increased over time and SR sensitivity to irrigation decreased over time. Our results provide a global overview of the sensitivities of grassland function and diversity to climate change that will improve the understanding of ecological responses across spatiotemporal scales and inform policies for conservation in dry climates.

Ecological stoichiometry and studies of biogeochemical niches have mainly focused on plankton and vascular plants, but the phenotypically closest modern relatives of early plants, bryophytes, have been largely neglected. We analysed C:N:P stoichiometries and elemental compositions (K, Na, Mg, Ca, S, Fe) of 35 widely distributed bryophyte species inhabiting springs. We estimated bryophyte C:N:P ratios and their biogeochemical niches, investigated how elementomes respond to the environment and determined whether they tend to diverge more for coexisting than non-coexisting individuals and species. The median C:N:P was 145:8:1, intermediate between Redfield's ratio for marine plankton and those for vascular plants. Biogeochemical niches were differentiated amongst species and were phylogenetically conserved. Differences in individual and species-specific elementomes increased with coexistence between species. Our results provide an evolutionary bridge between the ecological stoichiometries of algae and vascular plants and suggest that differences in elementomes could be used to understand community assemblages and functional diversity.

The possibility of using the elemental compositions of species as a tool to identify species/genotype niche remains to be tested at a global scale. We investigated relationships between the foliar elemental compositions (elementomes) of trees at a global scale with phylogeny, climate, N deposition and soil traits. We analysed foliar N, P, K, Ca, Mg and S concentrations in 23,962 trees of 227 species. Shared ancestry explained 60-94% of the total variance in foliar nutrient concentrations and ratios whereas current climate, atmospheric N deposition and soil type together explained 1-7%, consistent with the biogeochemical niche hypothesis which predicts that each species will have a specific need for and use of each bio-element. The remaining variance was explained by the avoidance of nutritional competition with other species and natural variability within species. The biogeochemical niche hypothesis is thus able to quantify species-specific tree niches and their shifts in response to environmental changes. Based on a global-scale analysis of the leaf elemental composition of tree species, the authors show that shared ancestry is the major factor shaping plant elementomes, thus providing large-scale empirical support for the biogeochemical niche hypothesis.

Nutrient economy and plant nutrient concentrations have been suggested as important selective pressures of reproductive traits in vascular plants. However, evidence supporting that the reproduction of bryophytes and their reproductive traits have also been conditioned by nutrient availability is lacking. We here analysed the nutrient (N, P, K, and Fe) concentration of 35 aquatic and semi-aquatic bryophytes and the chemistry of the spring water in which they lived to determine whether bryophyte nutrient concentrations were correlated with the frequency of sexual reproduction, main reproductive mode (sexual vs. asexual), dioicy, and spore diameter, while controlling for climate and nutrient concentrations in the spring water. Water nutrient concentrations, except for K, had a small but positive influence on bryophyte nutrient concentrations. P-rich and K-poor species produced sporophytes more often and were more likely to reproduce sexually and be monoicous. Spore diameter was smaller in P- and Fe-rich mosses but larger in N-rich mosses living in humid climates. P-rich species thus produced more and smaller spores and were able to propagate more extensively. Overall, our results suggest that nutrient economy, especially P economy, has played an important role in shaping the reproductive traits of bryophytes.

Mast seeding is one of the most intriguing reproductive traits in nature. Despite its potential drawbacks in terms of fitness, the widespread existence of this phenomenon suggests that it should have evolutionary advantages under certain circumstances. Using a global dataset of seed production time series for 219 plant species from all of the continents, we tested whether masting behaviour appears predominantly in species with low foliar nitrogen and phosphorus concentrations when controlling for local climate and productivity. Here, we show that masting intensity is higher in species with low foliar N and P concentrations, and especially in those with imbalanced N/P ratios, and that the evolutionary history of masting behaviour has been linked to that of nutrient economy. Our results support the hypothesis that masting is stronger in species growing under limiting conditions and suggest that this reproductive behaviour might have evolved as an adaptation to nutrient limitations and imbalances.

Mast seeding, the extremely variable and synchronized production of fruits, is a common reproductive behavior in plants. Weather is centrally involved in driving masting. Yet, it is often claimed that it cannot be the sole proximate cause of masting because weather is less variable than fruit production and because the shape of their distributions differ. We used computer simulations to demonstrate that the assumption that weather cannot be the main driver of masting was only valid for linear relationships between weather and fruit production. Non-linear relationships between interannual variability in weather and crop size, however, can account for the differences in their variability and the shape of their distributions because of Jensen's inequality. Exponential relationships with weather can increase the variability of fruit production, and sigmoidal relationships can produce bimodal distributions. These results challenge the idea that meteorological variability cannot be the main proximate driver of mast seeding, returning meteorological variability to the forefront of masting research.

Bryophytes are some of the most sensitive biological indicators of environmental change. Springs have a significant presence of bryophytes and so are ideal habitats for studying their relationship with the environment. We tested whether bryophyte assemblages can be explained with macro-, meso- and micro-ecological variables (i.e. seasonal climate, altitude, water pH and conductivity) sampling bryophytes from 198 semi-natural springs distributed along montane regions in the north-eastern Iberian Peninsula. We tested the influence of environmental variables on bryophyte assemblages in springs using sparse Partial Least Squares. Our results show that variability in bryophyte assemblages is explained by seasonal climate (temperature and precipitation from winter, spring, summer and autumn and temperature and precipitation seasonality), altitude and water conductivity. The results obtained by the present study will be useful for predicting bryophyte diversity in springs using simple and easy to obtain variables such as climate, water pH and conductivity.

The drivers of global change, including increases in atmospheric CO2 concentrations, N and S deposition, and climate change, likely affect the nutritional status of forests. Here we show forest foliar concentrations of N, P, K, S and Mg decreased significantly in Europe by 5%, 11%, 8%, 6% and 7%, respectively during the last three decades. The decrease in nutritional status was especially large in Mediterranean and temperate forests. Increasing atmospheric CO2 concentration was well correlated with the decreases in N, P, K, Mg, S concentrations and the increase of N:P ratio. Regional analyses indicated that increases in some foliar nutrient concentrations such as N, S and Ca in northern Europe occurred associated with increasingly favourable conditions of mean annual precipitation and temperature. Crucial changes in forest health, structure, functioning and services, including negative feedbacks on C capture can be expected if these trends are not reversed., Penuelas et al analyse foliar concentrations of nutrients in the main tree species of European forests using data collected over 30 years, together with changes in atmospheric CO2 levels, N and S deposition and climate. They find that foliar concentrations of N, P, K, S and Mg have decreased in central and southern Europe, with implications for forest ecosystems.

Background Tropical rainforests are frequently limited by soil nutrient availability. However, the response of the metabolic phenotypic plasticity of trees to an increase of soil nutrient availabilities is poorly understood. We expected that increases in the ability of a nutrient that limits some plant processes should be detected by corresponding changes in plant metabolome profile related to such processes. Methodology/Principal findings We studied the foliar metabolome of saplings of three abundant tree species in a 15 year field NPK fertilization experiment in a Panamanian rainforest. The largest differences were among species and explained 75% of overall metabolome variation. The saplings of the large canopy species, Tetragastris panamensis, had the lowest concentrations of all identified amino acids and the highest concentrations of most identified secondary compounds. The saplings of the “mid canopy” species, Alseis blackiana, had the highest concentrations of amino acids coming from the biosynthesis pathways of glycerate-3P, oxaloacetate and α-ketoglutarate, and the saplings of the low canopy species, Heisteria concinna, had the highest concentrations of amino acids coming from the pyruvate synthesis pathways. Conclusions/Significance The changes in metabolome provided strong evidence that different nutrients limit different species in different ways. With increasing P availability, the two canopy species shifted their metabolome towards larger investment in protection mechanisms, whereas with increasing N availability, the sub-canopy species increased its primary metabolism. The results highlighted the proportional distinct use of different nutrients by different species and the resulting different metabolome profiles in this high diversity community are consistent with the ecological niche theory.

Plants acquire carbon through photosynthesis to sustain biomass production, autotrophic respiration and production of non-structural compounds for multiple purposes(1). The fraction of photosynthetic production used for biomass production, the biomass production efficiency(2), is a key determinant of the conversion of solar energy to biomass. In forest ecosystems, biomass production efficiency was suggested to be related to site fertility(2). Here we present a database of biomass production effciency from 131 sites compiled from individual studies using harvest, biometric, eddy covariance, or process-based model estimates of production. The database is global, but dominated by data from Europe and North America. We show that instead of site fertility, ecosystem management is the key factor that controls biomass production efficiency in terrestrial ecosystems. In addition, in natural forests, grasslands, tundra, boreal peatlands and marshes, biomass production efficiency is independent of vegetation, environmental and climatic drivers. This similarity of biomass production efficiency across natural ecosystem types suggests that the ratio of biomass production to gross primary productivity is constant across natural ecosystems. We suggest that plant adaptation results in similar growth efficiency in high- and low-fertility natural systems, but that nutrient influxes under managed conditions favour a shift to carbon investment from the belowground flux of non-structural compounds to aboveground biomass.

Elevated CO2 concentrations increase photosynthesis and, potentially, net ecosystem production (NEP), meaning a greater CO2 uptake. Climate, nutrients and ecosystem structure, however, influence the effect of increasing CO2. Here we analysed global NEP from MACC-II and Jena CarboScope atmospheric inversions and ten dynamic global vegetation models (TRENDY), using statistical models to attribute the trends in NEP to its potential drivers: CO2, climatic variables and land-use change. We found that an increased CO2 was consistently associated with an increased NEP (1995–2014). Conversely, increased temperatures were negatively associated with NEP. Using the two atmospheric inversions and TRENDY, the estimated global sensitivities for CO2 were 6.0 ± 0.1, 8.1 ± 0.3 and 3.1 ± 0.1 PgC per 100 ppm (~1 °C increase), and −0.5 ± 0.2, −0.9 ± 0.4 and −1.1 ± 0.1 PgC °C−1 for temperature. These results indicate a positive CO2 effect on terrestrial C sinks that is constrained by climate warming. Global net ecosystem production (NEP) from a number of atmospheric inversions and dynamic global vegetation models is analysed to attribute trends to potential drivers. CO2 is found to have a positive effect on NEP that is constrained by climate warming.

Anthropogenic activities and intensive farming arc causing nitrate pollution in groundwater bodies. These aquifers are drained by springs which, in the Mediterranean region, act as refugia for preserving biodiversity of species that need continuous water. Some springs are also used for drinking water for wild animals, livestock and humans, so if their water quality is compromised it can become a threat to public health. However, the impact of nitrate pollution on these biotic communities remains unknown. We sampled 338 assemblages of aquatic and semi-aquatic bryophytes (i.e., hygrophytic mosses and liverworts) growing in springs in a gradient of water conductivity, nitrate concentration and climate and distributed across the north-east of the Iberian Peninsula to investigate the impact of nitrate pollution on the diversity of bryophytes and moss functional traits in Mediterranean springs. Based on previous literature suggesting that increased nitrogen load decreases biodiversity in grasslands and freshwater ecosystems, we hypothesised that water nitrate pollution in springs decreases bryophyte diversity at the local and regional scales. Our results indicated that, at the local scale (spring), nitrate pollution reduced the number and the likelihood of finding a rare species in springs. Rare species were found in 4% of the springs with nitrate above 50 mg but in 32% of the springs with nitrate below 50 mg L-1. Moss, liv- erwort and overall bryophyte diversity were not directly affected by nitrate at the local scale but nitrate consistently decreased diversity of mosses, liverworts and rare bryophyte species at the regional scale. We also found that warmer and drier springs presented fewer bryophyte species. Our results show that the combination of nitrate pollution, increasing temperature and drought could severely threaten bryophyte diversity Mediterranean springs. Our results indicate that the absence of rare bryophytes could be used as a bioindicator of nitrate pollution in springs. (C) 2018 Elsevier B.V. All rights reserved.

A decline in the carbon fertilization effect One source of uncertainty in climate science is how the carbon fertilization effect (CFE) will contribute to mitigation of anthropogenic climate change. Wang et al. explored the temporal dynamics of CFE on vegetation photosynthesis at the global scale. There has been a decline over recent decades in the contribution of CFE to vegetation photosynthesis, perhaps owing to the limiting effects of plant nutrients such as nitrogen and phosphorus. This declining trend has not been adequately accounted for in carbon cycle models. CFE thus has limitations for long-term mitigation of climate change, and future warming might currently be underestimated. Science , this issue p. 1295

Global change is affecting terrestrial carbon balances. The effect of climate on ecosystem carbon balance has been largely explored, but the roles of other concurrently changing factors, such as diversity and nutrient availability, remain elusive. We used eddy‐covariance carbon‐flux measurements from 62 ecosystems from which we compiled information on climate, ecosystem type, stand age, species abundance and foliar concentrations of N and P of the main species, to assess their importance in the ecosystem carbon balance. Climate and productivity were the main determinants of ecosystem C balance and its stability. In P‐rich sites, increasing N was related with increased gross primary production and respiration and vice versa, but reduced net C uptake. Our analyses did not provide evidence for a strong relation between ecosystem diversity and their productivity and stability. Nonetheless, these results suggest that nutrient imbalances and, potentially, diversity loss may alter future global carbon balance.

Unidad de excelencia María de Maeztu CEX2019-000940-M Annually variable and synchronous seed production, or masting, is often correlated with environmental factors and in oaks involves differential pollination success that depends on phenological synchrony in flowering. The synchronization of phenology of flowering was thought to be driven by temperature during flowering (microclimatic hypothesis). We tested an alternative, whereby phenological synchronization is driven by the timing of the onset of flowering (photoperiod-sensitivity hypothesis). This hypothesis assumes that flowering synchrony is driven by interaction between daylength and temperature, and individual variation in sensitivity to daylength as a phenological cue. We used long-term (23-26 years) records of airborne pollen in Quercus robur, Q. petraea, Q. ilex, and Q. humilis. Late pollen seasons were short, as predicted by photoperiod-sensitivity hypothesis. The onset of pollen seasons was delayed as preseason temperatures cooled over the last three decades at our Mediterranean sites, which was paralleled by shortening in pollen seasons, providing additional support for the photoperiod-sensitivity hypothesis. Global warming under the microclimatic hypothesis is predicted to lead to less frequent reproductive failures and thus decreased variability and synchrony of mast seeding. In contrast, warming under the photoperiod-sensitivity hypothesis should advance the onset of and desynchronize flowering, a pattern supported by our data. This pattern suggests that global warming will lead to more frequent vetoes and more stochastic and variable patterns of oak reproduction.

Forest production efficiency (FPE) metric describes how efficiently the assimilated carbon is partitioned into plants organs (biomass production, BP) or—more generally—for the production of organic matter (net primary production, NPP). We present a global analysis of the relationship of FPE to stand-age and climate, based on a large compilation of data on gross primary production and either BP or NPP. FPE is important for both forest production and atmospheric carbon dioxide uptake. We find that FPE increases with absolute latitude, precipitation and (all else equal) with temperature. Earlier findings—FPE declining with age—are also supported by this analysis. However, the temperature effect is opposite to what would be expected based on the short-term physiological response of respiration rates to temperature, implying a top-down regulation of carbon loss, perhaps reflecting the higher carbon costs of nutrient acquisition in colder climates. Current ecosystem models do not reproduce this phenomenon. They consistently predict lower FPE in warmer climates, and are therefore likely to overestimate carbon losses in a warming climate., Many models assume a universal carbon use efficiency across forest biomes, in contrast to assumptions of other process-based models. Here the authors analyse forest production efficiency across a wide range of climates to show a positive relationship with annual temperature and precipitation, indicating that ecosystem models are overestimating forest carbon losses under warming.

Agricultural ecosystems are globally important sinks of carbon and other nutrient elements. In China, acid rain events affect about 0.62 million km2, representing about 6.4% of total land area; however, the impacts of acid rain mediated nitrogen (N) and sulfur (S) depositions on soil organic carbon (SOC) fractions and nutrient stocks in paddy soils and implications for yield production under climate change are unclear. We conducted a field experiment during two annual crop seasons to determine the effects of simulated acid rain on SOC fractions and nutrients in a subtropical paddy in China. Acid rain treatments comprised solutions of HNO3 + H2SO4 to simulate N and S deposition at pH levels of 4.5, 3.5, and 2.5. The results showed that content of soil C fractions varied with acid rain pH. Acid rain led to increased SOC concentrations and decreased the ratios of soil labile organic carbon (LOC): SOC and dissolved organic carbon (DOC): SOC concentrations, independently of crop season and growth stage. Soil salinity was positively associated with SOC suggesting that higher levels of salinity inhibit C decomposition favoring SOC accumulation. Treatment effects of acid rain on soil microbial C and N depended on crop growth stage. Concentration of Fe2+ was positively correlated with DOC in early and late paddy soils under acid rain, possibly as a result of Fe2+ retention by DOC, when Fe3+ is reduced to Fe2+. Acid rain led to increases in soil TN, TP, available N, NH4+ concentration, and SOC, and decreases in ratios of DOC: SOC, indicating decreases in soil biological activity and mineralization processes. Increases in dead rice plant biomass under acid rain were consistent with the increase in soil N and P concentrations, due to reduced nutrient uptake, and higher levels of total SOC.

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Continuous atmospheric CO2 monitoring data indicate an increase in seasonal-cycle amplitude (SCA) of CO2 exchange in northern high latitudes. The major drivers of enhanced SCA remain unclear and intensely debated with land-use change, CO2 fertilization and warming identified as likely contributors. We integrated CO2-flux data from two atmospheric inversions (consistent with atmospheric records) and from and 11 state-of-the-art land-surface models (LSMs) to evaluate the relative importance of individual contributors to trends and drivers of the SCA of CO2-fluxes for 1980−2015. The LSMs generally reproduce the latitudinal increase in SCA trends within the inversions range. Inversions and LSMs attribute SCA increase to boreal Asia and Europe due to enhanced vegetation productivity (in LSMs) and point to contrasting effects of CO2 fertilisation (positive) and warming (negative) on SCA. Our results do not support land-use change as a key contributor to the increase in SCA. The sensitivity of simulated microbial respiration to temperature in LSMs explained biases in SCA trends, which suggests SCA could help to constrain model turnover times.

Concentrations of nutrient elements in organisms and in the abiotic environment are key factors influencing ecosystem structure and function. We studied how concentrations and stoichiometries of nitrogen (N), phosphorus (P) and potassium (K) in leaves of forest trees are related to phylogeny and to environmental factors (mean annual precipitation, mean annual temperature, forest type, and nitrogen deposition). Using data for 4691 forest plots from across Spain, we tested the following hypotheses: (i) that foliar stoichiometries of forest trees are strongly influenced by phylogeny, (ii) that climate, as an important driver of plant uptake and nutrient use efficiency, affects foliar stoichiometry, (iii) that long-term loads of N influence N, P and K concentrations and ratios in natural vegetation, and (iv) that sympatric species are differentiated according to their foliar stoichiometry, thereby reducing the intensity of resource competition. Our analyses revealed that several factors contributed to interspecific variation in elemental composition and stoichiometry. These included phylogeny, forest type, climate, N deposition, and competitive neighborhood relationships (probably related to niche segregation effect). These findings support the notion that foliar elemental composition reflects adaptation both to regional factors such as climate and to local factors such as competition with co-occurring species.

International audience; Soil fauna is a key control of the decomposition rate of leaf litter, yet its interactions with litter quality and the soil environment remain elusive. We conducted a litter decomposition experiment across different topographic levels within the landscape replicated in two rainforest sites providing natural gradients in soil fertility to test the hypothesis that low nutrient availability in litter and soil increases the strength of fauna control over litter decomposition. We crossed these data with a large dataset of 44 variables characterizing the biotic and abiotic microenvironment of each sampling point and found that microbe-driven carbon (C) and nitrogen (N) losses from leaf litter were 10.1 and 17.9% lower, respectively, in the nutrient-poorest site, but this among-site difference was equalized when meso- and macrofauna had access to the litterbags. Further, on average, soil fauna enhanced the rate of litter decomposition by 22.6%, and this contribution consistently increased as nutrient availability in the microenvironment declined. Our results indicate that nutrient scarcity increases the importance of soil fauna on C and N cycling in tropical rainforests. Further, soil fauna is able to equalize differences in microbial decomposition potential, thus buffering to a remarkable extent nutrient shortages at an ecosystem level.

Mosses are amongst the oldest and simplest plants, they can be found almost everywhere in the world, and they condition the structure and function of many ecosystems. Their sensitivity to environmental changes makes them very interesting subjects of study in ecology, and understanding them can provide insights into the evolutionary history of plants. However, the study of moss traits and their relationship with their environment is far behind that of vascular plants. We sampled 303 assemblages of aquatic and semi‐aquatic (hygrophytic) mosses growing in semi‐natural springs distributed around the northeast of the Iberian Peninsula to study how moss traits vary depending on their evolutionary history, climate and water chemistry. To do so, we analysed 30 moss species and 17 traits using phylogenetic comparative methods and an extended RLQ analysis, accounting for spatial and phylogenetic information. We hypothesized that there is a sclerophylly continuum in mosses living across a gradient of high and low water conductivity springs that may mimic sclerophylly in vascular plants that live in stressful environments. Results indicated that life‐forms and, especially, morphological traits were well preserved phylogenetically and responsive to water chemistry and climate. That combined with spatial autocorrelation in environmental variables resulted in a clustered distribution of phylogenetically closely related mosses in space. Mosses living in springs with a warm and dry climate that discharge hard water mainly presented species with needle‐like leaves, were denser, and had lower water absorption capacity. The opposite was found in cold, humid and soft water springs. Synthesis. Our results suggest that climate and water chemistry are main determinants of traits of hygrophytic mosses and of species distributions. We found evidence of a potential sclerophylly continuum in moss traits, which we hypothesize may be mainly related to physical and physiological constraints produced by water chemistry. Our findings describe moss sclerophylly in a gradient of water conductivity similar to that found in vascular plants with water availability and temperature. Further experimental studies will be required to confirm the observations found in this study. A free Plain Language Summary can be found within the Supporting Information of this article.

Some tree species have a highly variable year-to-year pattern of reproduction which has repercussions for the entire ecosystem. Links between meteorological variability, fruit production and crown cover, and trade-offs between reproduction and vegetative growth, remain elusive, despite a long history of research. We explored how meteorological conditions determined variations in fruit production and crown cover and how remotely sensed vegetation indices, such as the enhanced vegetation index (EVI), may be used to characterize the fluctuations in fruit production. We used data for fruit production from six European tree species (Abies alba, Picea abies, Pseudotsuga menziesii, Fagus sylvatica, Quercus petraea and Q. robur) growing in monospecific stands, EVI and seasonal meteorological variables (precipitation and temperature) for 2002–2010. Weather accounted for fruit production better than EVI. Deciduous trees were more responsive to weather than evergreens, most notably to different seasonal temperatures, which were positively correlated mainly with crown cover and fruit production in deciduous species. Our results also suggested different patterns of relationships between fruit production, crown cover and weather, indicating different strategies of resource management. These patterns indicated a possible internal trade-off in evergreens, with resources allocated to either growth or reproduction. In contrast, in deciduous species we found no evidence for such a trade-off between vegetative growth and reproduction.

Synchronized and variable reproduction by perennial plants, called mast seeding, is a major reproductive strategy of trees. The need to accumulate sufficient resources after depletion following fruiting (resource budget), the efficiency of mass flowering for outcross pollination (pollen coupling), or the external factors preventing reproduction (environmental veto) could all synchronize masting. We used seed production data for four species (Quercus ilex, Quercus humilis, Sorbus aucuparia, and Pinus albicaulis) to parametrize resource budget models of masting. Based on species life-history characteristics, we hypothesized that pollen coupling should synchronize reproduction in S. aucuparia and P. albicaulis, while in Q. ilex and Q. humilis, environmental veto should be a major factor. Pollen coupling was stronger in S. aucuparia and P. albicaulis than in oaks, while veto was more frequent in the latter. Yet in all species, costs of reproduction were too small to impose a replenishment period. A synchronous environmental veto, in the presence of environmental stochasticity, was sufficient to produce observed variability and synchrony in reproduction. In the past, vetoes like frost events that prevent reproduction have been perceived as negative for plants. In fact, they could be selectively favored as a way to create mast seeding.

International audience; Every living creature on Earth is made of atoms of the various bioelements that are harnessed in the construction of molecules, tissues, organisms, and communities, as we know them. Organisms need these bioelements in specific quantities and proportions to survive and grow. Distinct species have different functions and life strategies, and have therefore developed distinct structures and adopted a certain combination of metabolic and physiological processes. Each species is thus also expected to have different requirements for each bioelement. We therefore propose that a “biogeochemical niche” can be associated with the classical ecological niche of each species. We show from field data examples that a biogeochemical niche is characterized by a particular elementome defined as the content of all (or at least most) bioelements. The differences in elementome among species are a function of taxonomy and phylogenetic distance, sympatry (the bioelemental compositions should differ more among coexisting than among non-coexisting species to avoid competitive pressure), and homeostasis with a continuum between high homeostasis/low plasticity and low homeostasis/high plasticity. This proposed biogeochemical niche hypothesis has the advantage relative to other associated theoretical niche hypotheses that it can be easily characterized by actual quantification of a measurable trait: the elementome of a given organism or a community, being potentially applicable across taxa and habitats. The changes in bioelemental availability can determine genotypic selection and therefore have a feedback on ecosystem function and organization, and, at the end, become another driving factor of the evolution of life and the environment.

Unidad de excelencia María de Maeztu CEX2019-000940-M Nutrient availability influences virtually every aspect of an ecosystem, and is a critical modifier of ecosystem responses to global change. Although this crucial role of nutrient availability in regulating ecosystem structure and functioning has been widely acknowledged, nutrients are still often neglected in observational and experimental synthesis studies due to difficulties in comparing the nutrient status across sites. In the current study, we explain different nutrient-related concepts and discuss the potential of soil-, plant- and remote sensing-based metrics to compare the nutrient status across space. Based on our review and additional analyses on a dataset of European, managed temperate and boreal forests (ICP [International Co-operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests] Forests dataset), we conclude that the use of plant- and remote sensing-based metrics that rely on tissue stoichiometry is limited due to their strong dependence on species identity. The potential use of other plant-based metrics such as Ellenberg indicator values and plant-functional traits is also discussed. We conclude from our analyses and review that soil-based metrics have the highest potential for successful intersite comparison of the nutrient status. As an example, we used and adjusted a soil-based metric, previously developed for conifer forests across Sweden, against the same ICP Forests data. We suggest that this adjusted and further adaptable metric, which included the organic carbon concentration in the upper 20 cm of the soil (including the organic fermentation-humus [FH] layer), the C:N ratio and of the FH layer, can be used as a complementary tool along with other indicators of nutrient availability, to compare the background nutrient status across temperate and boreal forests dominated by spruce, pine or beech. Future collection and provision of harmonized soil data from observational and experimental sites is crucial for further testing and adjusting the metric.

The quantification of isoprene and monoterpene emissions at the ecosystem level with available models and field measurements is not entirely satisfactory. Remote-sensing techniques can extend the spatial and temporal assessment of isoprenoid fluxes. Detecting the exchange of biogenic volatile organic compounds (BVOCs) using these techniques is, however, a very challenging goal. Recent evidence suggests that a simple remotely sensed index, the photochemical reflectance index (PRI), which is indicative of light-use efficiency, relative pigment levels and excess reducing power, is a good indirect estimator of foliar isoprenoid emissions. We tested the ability of PRI to assess isoprenoid fluxes in a temperate deciduous forest in central USA throughout the entire growing season and under moderate and extreme drought conditions. We compared PRI time series calculated with MODIS bands to isoprene emissions measured with eddy covariance. MODIS PRI was correlated with isoprene emissions for most of the season, until emissions peaked. MODIS PRI was also able to detect the timing of the annual peak of emissions, even when it was advanced in response to drought conditions. PRI is thus a promising index to estimate isoprene emissions when it is complemented by information on potential emission. It may also be used to further improve models of isoprene emission under drought and other stress conditions. Direct estimation of isoprene emission by PRI is, however, limited, because PRI estimates LUE, and the relationship between LUE and isoprene emissions can be modified by severe stress conditions.

1. 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 analyzing data from 2245 forest stands. 2. 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 toward foliar P deficiency were thus more marked when high N deposition coincided with climatic conditions favorable to plant production. 3. 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 negatively correlated with soil Olsen P concentration. 4. 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 were related with relative annual BD growth. In contrast, needle P concentration was positively related with both absolute and relative annual BD growth. 5. 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 toward increased nutrient losses in runoff 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 center 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.