Effects of management practices on the ecosystem-service multifunctionality of temperate grasslands.

Human wellbeing depends on ecosystem services, highlighting the need for improving the ecosystem-service multifunctionality of food and feed production systems. We study Swiss agricultural grasslands to assess how employing and combining three widespread aspects of grassland management and their interactions can enhance 22 plot-level ecosystem service indicators, as well as ecosystem-service multifunctionality. The three management aspects we assess are i) organic production system, ii) an eco-scheme prescribing extensive management (without fertilization), and iii) harvest type (pasture vs. meadow). While organic production system and interactions between the three management aspects play a minor role, the main effects of eco-scheme and harvest type considerably shape single services. Moreover, the eco-scheme 'extensive management' and the harvest type 'pasture' enhance plot-scale ecosystem-service multifunctionality, mostly through facilitating cultural services at the expense of provisioning services. These changes in ecosystem-service supply occur mainly via changes in land-use intensity, i.e., reduced fertilizer input and harvest frequency. In conclusion, diversifying grassland management where this is currently homogeneous across farms and landscapes depicts an important first step to improve landscape-scale multifunctionality for sustainable grassland systems. To meet societal ecosystem services demand, the three studied management aspects can be systematically combined to increase ecosystem services that are in short supply.

Contrasting impact of extreme soil and atmospheric dryness on the functioning of trees and forests.

Recent studies indicate an increase in the frequency of extreme compound dryness days (days with both extreme soil AND air dryness) across central Europe in the future, with little information on their impact on the functioning of trees and forests. This study aims to quantify and assess the impact of extreme soil dryness, extreme air dryness, and extreme compound dryness on the functioning of trees and forests. For this, >15 years of ecosystem-level (carbon dioxide and water vapor fluxes) and 6-10 years of tree-level measurements (transpiration and growth) each from a montane mixed deciduous forest (CH-Lae) and a subalpine evergreen coniferous forest (CH-Dav) in Switzerland, is used. The results showed extreme air dryness limitation on CO2 fluxes and extreme soil dryness limitations on water vapor fluxes. Additionally, CH-Dav was mainly affected by extreme air dryness whereas CH-Lae was affected by both extreme soil dryness and extreme air dryness. The impact of extreme compound dryness on net CO2 uptake (about 75 % decrease) was more due to higher increased ecosystem respiration (40 % and 70 % increase at CH-Dav and CH-Lae, respectively) than decreased gross primary productivity (10 % and 40 % decrease at CH-Dav and CH-Lae, respectively). A significant negative impact on evapotranspiration and transpiration was only observed at CH-Lae during extreme soil and compound dryness (about 25 % decrease). Furthermore, with some differences, the tree-level impact on tree water deficit, transpiration, and growth were consistent with the ecosystem-level impact on carbon uptake and evapotranspiration. Finally, the impact of extreme dryness showed no significant relationship with tree allometry (diameter and height) but across different tree species. The projected future is likely to expose these forest areas to more extreme and frequent dryness conditions, thus compromising the functioning of trees and forests, thereby calling for management interventions to increase the adaptive capacity and resistance of these forests.

Chronic warming and dry soils limit carbon uptake and growth despite a longer growing season in beech and oak.

Progressively warmer and drier climatic conditions impact tree phenology and carbon cycling with large consequences for forest carbon balance. However, it remains unclear how individual impacts of warming and drier soils differ from their combined effects and how species interactions modulate tree responses. Using mesocosms, we assessed the multiyear impact of continuous air warming and lower soil moisture alone or in combination on phenology, leaf-level photosynthesis, nonstructural carbohydrate concentrations, and aboveground growth of young European beech (Fagus sylvatica L.) and Downy oak (Quercus pubescens Willd.) trees. We further tested how species interactions (in monocultures and in mixtures) modulated these effects. Warming prolonged the growing season of both species but reduced growth in oak. In contrast, lower moisture did not impact phenology but reduced carbon assimilation and growth in both species. Combined impacts of warming and drier soils did not differ from their single effects. Under warmer and drier conditions, performances of both species were enhanced in mixtures compared to monocultures. Our work revealed that higher temperature and lower soil moisture have contrasting impacts on phenology vs. leaf-level assimilation and growth, with the former being driven by temperature and the latter by moisture. Furthermore, we showed a compensation in the negative impacts of chronic heat and drought by tree species interactions.

Additive effects of two agri-environmental schemes on plant diversity but not on productivity indicators in permanent grasslands in Switzerland.

Different agri-environmental schemes (AES), such as ecological focus areas and organic farming, have been suggested to reduce the impact of intensive agriculture on the environment and to conserve or even restore farmland biodiversity. However, the effectiveness of such schemes, their ability to actually support biodiversity and associated trade-offs with agricultural production are still debated. We analysed a large dataset from the biodiversity monitoring in the Swiss agricultural landscape to assess the effects of two different grassland AES, i.e., extensively managed ecological focus areas (EFAs versus non-EFAs) and organic farming (versus conventional), on plant diversity, plant community composition and productivity indicators, i.e., weed abundance, forage value and nutrient availability. We also considered environmental factors, i.e., topography and soil conditions, which potentially modulate AES effects on biodiversity. We used in total 1170 plots in permanent grasslands, managed as meadows or pastures. Both AES had significant positive effects on plant diversity. However, EFAs increased plant richness considerably stronger (+6.6 species) than organic farming (+1.8 species). Effects of the two schemes were additive with organic EFA grasslands exhibiting highest plant diversity. Differences in topography partly explained AES effects on diversity as both AES were associated with differences in elevation and slope. Thus, future assessments of the effectiveness of AES need to consider the non-random placement of AES across heterogeneous landscapes. EFA grasslands revealed a considerably reduced agricultural productivity as shown by low forage values and low nutrient availability. Yet, the abundance of agricultural weeds, i.e., agriculturally undesired plant species, was lower in EFA compared to non-EFA grasslands. Productivity indicators were only weakly affected by organic farming and other than for plant diversity, productivity did not differ between organic and conventional EFA grasslands. The positive additive diversity effects of EFAs and organic grassland farming underline the potential of both AES to contribute to biodiversity conservation in agricultural landscapes, though to a different extent. Comparing the effects of the two AES revealed that the lower the reduction in agricultural productivity associated with an AES, the smaller the gains in plant diversity, highlighting the inevitable trade-off between productivity and plant diversity in semi-natural grasslands.

Long-term mitigation of drought changes the functional potential and life-strategies of the forest soil microbiome involved in organic matter decomposition.

Climate change can alter the flow of nutrients and energy through terrestrial ecosystems. Using an inverse climate change field experiment in the central European Alps, we explored how long-term irrigation of a naturally drought-stressed pine forest altered the metabolic potential of the soil microbiome and its ability to decompose lignocellulolytic compounds as a critical ecosystem function. Drought mitigation by a decade of irrigation stimulated profound changes in the functional capacity encoded in the soil microbiome, revealing alterations in carbon and nitrogen metabolism as well as regulatory processes protecting microorganisms from starvation and desiccation. Despite the structural and functional shifts from oligotrophic to copiotrophic microbial lifestyles under irrigation and the observation that different microbial taxa were involved in the degradation of cellulose and lignin as determined by a time-series stable-isotope probing incubation experiment with 13C-labeled substrates, degradation rates of these compounds were not affected by different water availabilities. These findings provide new insights into the impact of precipitation changes on the soil microbiome and associated ecosystem functioning in a drought-prone pine forest and will help to improve our understanding of alterations in biogeochemical cycling under a changing climate.

Monitoring of carbon-water fluxes at Eurasian meteorological stations using random forest and remote sensing.

Simulating the carbon-water fluxes at more widely distributed meteorological stations based on the sparsely and unevenly distributed eddy covariance flux stations is needed to accurately understand the carbon-water cycle of terrestrial ecosystems. We established a new framework consisting of machine learning, determination coefficient (R2), Euclidean distance, and remote sensing (RS), to simulate the daily net ecosystem carbon dioxide exchange (NEE) and water flux (WF) of the Eurasian meteorological stations using a random forest model or/and RS. The daily NEE and WF datasets with RS-based information (NEE-RS and WF-RS) for 3774 and 4427 meteorological stations during 2002-2020 were produced, respectively. And the daily NEE and WF datasets without RS-based information (NEE-WRS and WF-WRS) for 4667 and 6763 meteorological stations during 1983-2018 were generated, respectively. For each meteorological station, the carbon-water fluxes meet accuracy requirements and have quasi-observational properties. These four carbon-water flux datasets have great potential to improve the assessments of the ecosystem carbon-water dynamics.

Detection and attribution of an anomaly in terrestrial photosynthesis in Europe during the COVID-19 lockdown.

Carbon dioxide (CO2) uptake by plant photosynthesis, referred to as gross primary production (GPP) at the ecosystem level, is sensitive to environmental factors, including pollutant exposure, pollutant uptake, and changes in the scattering of solar shortwave irradiance (SWin) - the energy source for photosynthesis. The 2020 spring lockdown due to COVID-19 resulted in improved air quality and atmospheric transparency, providing a unique opportunity to assess the impact of air pollutants on terrestrial ecosystem functioning. However, detecting these effects can be challenging as GPP is influenced by other meteorological drivers and management practices. Based on data collected from 44 European ecosystem-scale CO2 flux monitoring stations, we observed significant changes in spring GPP at 34 sites during 2020 compared to 2015-2019. Among these, 14 sites showed an increase in GPP associated with higher SWin, 10 sites had lower GPP linked to atmospheric and soil dryness, and seven sites were subjected to management practices. The remaining three sites exhibited varying dynamics, with one experiencing colder and rainier weather resulting in lower GPP, and two showing higher GPP associated with earlier spring melts. Analysis using the regional atmospheric chemical transport model (LOTOS-EUROS) indicated that the ozone (O3) concentration remained relatively unchanged at the research sites, making it unlikely that O3 exposure was the dominant factor driving the primary production anomaly. In contrast, SWin increased by 9.4 % at 36 sites, suggesting enhanced GPP possibly due to reduced aerosol optical depth and cloudiness. Our findings indicate that air pollution and cloudiness may weaken the terrestrial carbon sink by up to 16 %. Accurate and continuous ground-based observations are crucial for detecting and attributing subtle changes in terrestrial ecosystem functioning in response to environmental and anthropogenic drivers.

N2O and CH4 fluxes from intensively managed grassland: The importance of biological and environmental drivers vs. management.

Agriculture is the main contributor to anthropogenic nitrous oxide (N2O) and methane (CH4) emissions. Therefore, mitigation options are urgently needed. In contrast to carbon dioxide, eddy covariance measurements of N2O and CH4 fluxes are still scarce, and thus little is known how environmental and biotic drivers as well as management affect the net N2O and CH4 exchange in grasslands. Thus, we investigated the most important drivers of net ecosystem N2O and CH4 fluxes in a temperate grassland, and continued a N2O mitigation experiment (increased clover proportion vs. fertilization with slurry). Random forest gap-filling models were able to capture intermittent emission peaks, performing better for half-hourly N2O than for CH4 fluxes. The unfertilized clover parcel (parcel B) continued to show lower N2O emissions (4.4 and 2.7 kg N2O-N ha-1 yr-1) compared to the fertilized parcel (parcel A; 6.9 and 5.9 kg N2O-N ha-1 yr-1) for 2019 and 2020, respectively. Tier 1 nitrogen (N) emission factors of 2.6 % and 1.9 % were observed at the fertilized parcel during the study period. Lower soil N concentrations indicated a lower N leaching risk at the clover than at the fertilized parcel. Annual CH4 emissions (including periods with sheep grazing) were similar from both parcels, and ranged from 25 to 38.5 kg CH4-C ha-1. The most important drivers of both N2O and CH4 fluxes were lagged precipitation and water filled pore space, but also management (for N2O from parcel B; CH4 from parcel A). Biotic variables such as vegetation height and leaf area index were important predictors for the N2O exchange, while grazing temporarily increased CH4 emissions. Overall, reducing N fertilization and increasing the legume proportion were effective N2O reduction measures. In particular, adjusting N fertilization to plant N demands can help to avoid high N2O emissions from grasslands.

Leaf-level coordination principles propagate to the ecosystem scale.

Fundamental axes of variation in plant traits result from trade-offs between costs and benefits of resource-use strategies at the leaf scale. However, it is unclear whether similar trade-offs propagate to the ecosystem level. Here, we test whether trait correlation patterns predicted by three well-known leaf- and plant-level coordination theories - the leaf economics spectrum, the global spectrum of plant form and function, and the least-cost hypothesis - are also observed between community mean traits and ecosystem processes. We combined ecosystem functional properties from FLUXNET sites, vegetation properties, and community mean plant traits into three corresponding principal component analyses. We find that the leaf economics spectrum (90 sites), the global spectrum of plant form and function (89 sites), and the least-cost hypothesis (82 sites) all propagate at the ecosystem level. However, we also find evidence of additional scale-emergent properties. Evaluating the coordination of ecosystem functional properties may aid the development of more realistic global dynamic vegetation models with critical empirical data, reducing the uncertainty of climate change projections.

Long-term changes in forest response to extreme atmospheric dryness.

Atmospheric dryness, as indicated by vapor pressure deficit (VPD), has a strong influence on forest greenhouse gas exchange with the atmosphere. In this study, we used long-term (10-30 years) net ecosystem productivity (NEP) measurements from 60 forest sites across the world (1003 site-years) to quantify long-term changes in forest NEP resistance and NEP recovery in response to extreme atmospheric dryness. We tested two hypotheses: first, across sites differences in NEP resistance and NEP recovery of forests will depend on both the biophysical characteristics (i.e., leaf area index [LAI] and forest type) of the forest as well as on the local meteorological conditions of the site (i.e., mean VPD of the site), and second, forests experiencing an increasing trend in frequency and intensity of extreme dryness will show an increasing trend in NEP resistance and NEP recovery over time due to emergence of long-term ecological stress memory. We used a data-driven statistical learning approach to quantify NEP resistance and NEP recovery over multiple years. Our results showed that forest types, LAI, and median local VPD conditions explained over 50% of variance in both NEP resistance and NEP recovery, with drier sites showing higher NEP resistance and NEP recovery compared to sites with less atmospheric dryness. The impact of extreme atmospheric dryness events on NEP lasted for up to 3 days following most severe extreme events in most forests, indicated by an NEP recovery of less than 100%. We rejected our second hypothesis as we found no consistent relationship between trends of extreme VPD with trends in NEP resistance and NEP recovery across different forest sites, thus an increase in atmospheric dryness as it is predicted might not increase the resistance or recovery of forests in terms of NEP.

Dew benefits on alpine grasslands are cancelled out by combined heatwave and drought stress.

Increasing frequencies of heatwaves combined with simultaneous drought stress in Europe threaten the ecosystem water and carbon budgets of alpine grasslands. Dew as an additional water source can promote ecosystem carbon assimilation. It is known that grassland ecosystems keep high evapotranspiration as long as soil water is available. However, it is rarely being investigated whether dew can mitigate the impact of such extreme climatic events on grassland ecosystem carbon and water exchange. Here we use stable isotopes in meteoric waters and leaf sugars, eddy covariance fluxes for H2O vapor and CO2, in combination with meteorological and plant physiological measurements, to investigate the combined effect of dew and heat-drought stress on plant water status and net ecosystem production (NEP) in an alpine grassland (2000 m elevation) during the June 2019 European heatwave. Before the heatwave, enhanced NEP in the early morning hours can be attributed to leaf wetting by dew. However, dew benefits on NEP were cancelled out by the heatwave, due to the minor contribution of dew in leaf water. Heat-induced reduction in NEP was intensified by the combined effect of drought stress. The recovery of NEP after the peak of the heatwave could be linked to the refilling of plant tissues during nighttime. Among-genera differences of plant water status affected by dew and heat-drought stress can be attributed to differences in their foliar dew water uptake, and their reliance on soil moisture or the impact of the atmospheric evaporative demand. Our results indicate that dew influence on alpine grassland ecosystems varies according to the environmental stress and plant physiology.

Daytime stomatal regulation in mature temperate trees prioritizes stem rehydration at night.

Trees remain sufficiently hydrated during drought by closing stomata and reducing canopy conductance (Gc ) in response to variations in atmospheric water demand and soil water availability. Thresholds that control the reduction of Gc are proposed to optimize hydraulic safety against carbon assimilation efficiency. However, the link between Gc and the ability of stem tissues to rehydrate at night remains unclear. We investigated whether species-specific Gc responses aim to prevent branch embolisms, or enable night-time stem rehydration, which is critical for turgor-dependent growth. For this, we used a unique combination of concurrent dendrometer, sap flow and leaf water potential measurements and collected branch-vulnerability curves of six common European tree species. Species-specific Gc reduction was weakly related to the water potentials at which 50% of branch xylem conductivity is lost (P50 ). Instead, we found a stronger relationship with stem rehydration. Species with a stronger Gc control were less effective at refilling stem-water storage as the soil dries, which appeared related to their xylem architecture. Our findings highlight the importance of stem rehydration for water-use regulation in mature trees, which likely relates to the maintenance of adequate stem turgor. We thus conclude that stem rehydration must complement the widely accepted safety-efficiency stomatal control paradigm.

ForestClim-Bioclimatic variables for microclimate temperatures of European forests.

Microclimate research gained renewed interest over the last decade and its importance for many ecological processes is increasingly being recognized. Consequently, the call for high-resolution microclimatic temperature grids across broad spatial extents is becoming more pressing to improve ecological models. Here, we provide a new set of open-access bioclimatic variables for microclimate temperatures of European forests at 25 × 25 m2 resolution.

Hydrogen isotope fractionation in carbohydrates of leaves and xylem tissues follows distinct phylogenetic patterns: a common garden experiment with 73 tree and shrub species.

Recent methodological advancements in determining the nonexchangeable hydrogen isotopic composition (δ2 Hne ) of plant carbohydrates make it possible to disentangle the drivers of hydrogen isotope (2 H) fractionation processes in plants. Here, we investigated the influence of phylogeny on the δ2 Hne of twig xylem cellulose and xylem water, as well as leaf sugars and leaf water, across 73 Northern Hemisphere tree and shrub species growing in a common garden. 2 H fractionation in plant carbohydrates followed distinct phylogenetic patterns, with phylogeny reflected more in the δ2 Hne of leaf sugars than in that of twig xylem cellulose. Phylogeny had no detectable influence on the δ2 Hne of twig or leaf water, showing that biochemistry, not isotopic differences in plant water, caused the observed phylogenetic pattern in carbohydrates. Angiosperms were more 2 H-enriched than gymnosperms, but substantial δ2 Hne variations also occurred at the order, family, and species levels within both clades. Differences in the strength of the phylogenetic signals in δ2 Hne of leaf sugars and twig xylem cellulose suggest that the original phylogenetic signal of autotrophic processes was altered by subsequent species-specific metabolism. Our results will help improve 2 H fractionation models for plant carbohydrates and have important consequences for dendrochronological and ecophysiological studies.

Networking the forest infrastructure towards near real-time monitoring - A white paper.

Forests account for nearly 90 % of the world's terrestrial biomass in the form of carbon and they support 80 % of the global biodiversity. To understand the underlying forest dynamics, we need a long-term but also relatively high-frequency, networked monitoring system, as traditionally used in meteorology or hydrology. While there are numerous existing forest monitoring sites, particularly in temperate regions, the resulting data streams are rarely connected and do not provide information promptly, which hampers real-time assessments of forest responses to extreme climate events. The technology to build a better global forest monitoring network now exists. This white paper addresses the key structural components needed to achieve a novel meta-network. We propose to complement - rather than replace or unify - the existing heterogeneous infrastructure with standardized, quality-assured linking methods and interacting data processing centers to create an integrated forest monitoring network. These automated (research topic-dependent) linking methods in atmosphere, biosphere, and pedosphere play a key role in scaling site-specific results and processing them in a timely manner. To ensure broad participation from existing monitoring sites and to establish new sites, these linking methods must be as informative, reliable, affordable, and maintainable as possible, and should be supplemented by near real-time remote sensing data. The proposed novel meta-network will enable the detection of emergent patterns that would not be visible from isolated analyses of individual sites. In addition, the near real-time availability of data will facilitate predictions of current forest conditions (nowcasts), which are urgently needed for research and decision making in the face of rapid climate change. We call for international and interdisciplinary efforts in this direction.

Biodiversity-stability relationships strengthen over time in a long-term grassland experiment.

Numerous studies have demonstrated that biodiversity drives ecosystem functioning, yet how biodiversity loss alters ecosystems functioning and stability in the long-term lacks experimental evidence. We report temporal effects of species richness on community productivity, stability, species asynchrony, and complementarity, and how the relationships among them change over 17 years in a grassland biodiversity experiment. Productivity declined more rapidly in less diverse communities resulting in temporally strengthening positive effects of richness on productivity, complementarity, and stability. In later years asynchrony played a more important role in increasing community stability as the negative effect of richness on population stability diminished. Only during later years did species complementarity relate to species asynchrony. These results show that species complementarity and asynchrony can take more than a decade to develop strong stabilizing effects on ecosystem functioning in diverse plant communities. Thus, the mechanisms stabilizing ecosystem functioning change with community age.

Greenhouse gas fluxes (CO2, N2O and CH4) of pea and maize during two cropping seasons: Drivers, budgets, and emission factors for nitrous oxide.

Agriculture contributes considerably to the increase of global greenhouse gas (GHG) emissions. Hence, magnitude and drivers of temporal variations in carbon dioxide (CO2), nitrous oxide (N2O) and methane (CH4) fluxes in croplands are urgently needed to develop sustainable, climate-smart agricultural practices. However, our knowledge of GHG fluxes from croplands is still very limited. The eddy covariance technique was used to quantify GHG budgets and N2O emission factors (EF) for pea and maize in Switzerland. The random forest technique was applied for gap-filling N2O and CH4 fluxes as well as to determine the relevance of environmental, vegetation vs. management drivers of the GHG fluxes during two cropping seasons. Environmental (i.e., net radiation, soil water content, soil temperature) and vegetation drivers (i.e., vegetation height) were more important drivers for GHG fluxes at field scale than time since management for the two crop species. Both crops acted as GHG sinks between sowing and harvest, clearly dominated by net CO2 fluxes, while CH4 emissions were negligible. However, considerable N2O emissions occurred in both crop fields early in the season when crops were still establishing. N2O fluxes in both crops were small later in the season when vegetation was tall, despite high soil water contents and temperatures. Results clearly show a strong and highly dynamic microbial-plant competition for N driving N2O fluxes at the field scale. The total loss was 1.4 kg N2O-N ha-1 over 55 days for pea and 4.8 kg N2O-N ha-1 over 127 days for maize. EFs of N2O were 1.5 % (pea) and 4.4 % (maize) during the cropping seasons, clearly exceeding the IPCC Tier 1 EF for N2O. Thus, sustainable, climate-smart agriculture needs to consider crop phenology and better adapt N supply to crop N demand for growth, particularly during the early cropping season when competition for N between establishing crops and soil microorganisms modulates N2O losses.

Impact of warmer and drier conditions on tree photosynthetic properties and the role of species interactions.

Increased temperature and prolonged soil moisture reduction have distinct impacts on tree photosynthetic properties. Yet, our knowledge of their combined effect is limited. Moreover, how species interactions alter photosynthetic responses to warming and drought remains unclear. Using mesocosms, we studied how photosynthetic properties of European beech and downy oak were impacted by multi-year warming and soil moisture reduction alone or combined, and how species interactions (intra- vs inter-specific interactions) modulated these effects. Warming of +5°C enhanced photosynthetic properties in oak but not beech, while moisture reduction decreased them in both species. Combined warming and moisture reduction reduced photosynthetic properties for both species, but no exacerbated effects were observed. Oak was less impacted by combined warming and limited moisture when interacting with beech than in intra-specific stands. For beech, species interactions had no impact on the photosynthetic responses to warming and moisture reduction, alone or combined. Warming had either no or beneficial effects on the photosynthetic properties, while moisture reduction and their combined effects strongly reduced photosynthetic responses. However, inter-specific interactions mitigated the adverse impacts of combined warming and drought in oak, thereby highlighting the need to deepen our understanding of the role of species interactions under climate change.