The three major axes of terrestrial ecosystem function.

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.

Uncovering the critical soil moisture thresholds of plant water stress for European ecosystems.

Understanding the critical soil moisture (SM) threshold (θcrit ) of plant water stress and land surface energy partitioning is a basis to evaluate drought impacts and improve models for predicting future ecosystem condition and climate. Quantifying the θcrit across biomes and climates is challenging because observations of surface energy fluxes and SM remain sparse. Here, we used the latest database of eddy covariance measurements to estimate θcrit across Europe by evaluating evaporative fraction (EF)-SM relationships and investigating the covariance between vapor pressure deficit (VPD) and gross primary production (GPP) during SM dry-down periods. We found that the θcrit and soil matric potential threshold in Europe are 16.5% and -0.7 MPa, respectively. Surface energy partitioning characteristics varied among different vegetation types; EF in savannas had the highest sensitivities to SM in water-limited stage, and the lowest in forests. The sign of the covariance between daily VPD and GPP consistently changed from positive to negative during dry-down across all sites when EF shifted from relatively high to low values. This sign of the covariance changed after longer period of SM decline in forests than in grasslands and savannas. Estimated θcrit from the VPD-GPP covariance method match well with the EF-SM method, showing this covariance method can be used to detect the θcrit . We further found that soil texture dominates the spatial variability of θcrit while shortwave radiation and VPD are the major drivers in determining the spatial pattern of EF sensitivities. Our results highlight for the first time that the sign change of the covariance between daily VPD and GPP can be used as an indicator of how ecosystems transition from energy to SM limitation. We also characterized the corresponding θcrit and its drivers across diverse ecosystems in Europe, an essential variable to improve the representation of water stress in land surface models.

The tempo of greening in the European Alps: Spatial variations on a common theme.

The long-term increase in satellite-based proxies of vegetation cover is a well-documented response of seasonally snow-covered ecosystems to climate warming. However, observed greening trends are far from uniform, and substantial uncertainty remains concerning the underlying causes of this spatial variability. Here, we processed surface reflectance of the moderate resolution imaging spectroradiometer (MODIS) to investigate trends and drivers of changes in the annual peak values of the Normalized Difference Vegetation Index (NDVI). Our study focuses on above-treeline ecosystems in the European Alps. NDVI changes in these ecosystems are highly sensitive to land cover and biomass changes and are marginally affected by anthropogenic disturbances. We observed widespread greening for the 2000-2020 period, a pattern that is consistent with the overall increase in summer temperature. At the local scale, the spatial variability of greening was mainly due to the preferential response of north-facing slopes between 1900 and 2400 m. Using high-resolution imagery, we noticed that the presence of screes and outcrops locally magnified this response. At the regional scale, we identified hotspots of greening where vegetation cover is sparser than expected given the elevation and exposure. Most of these hotspots experienced delayed snow melt and green-up dates in recent years. We conclude that the ongoing greening in the Alps primarily reflects the high responsiveness of sparsely vegetated ecosystems that are able to benefit the most from temperature and water-related habitat amelioration above treeline.

Nutrients and water availability constrain the seasonality of vegetation activity in a Mediterranean ecosystem.

Anthropogenic nitrogen (N) deposition and resulting differences in ecosystem N and phosphorus (P) ratios are expected to impact photosynthetic capacity, that is, maximum gross primary productivity (GPPmax ). However, the interplay between N and P availability with other critical resources on seasonal dynamics of ecosystem productivity remains largely unknown. In a Mediterranean tree-grass ecosystem, we established three landscape-level (24 ha) nutrient addition treatments: N addition (NT), N and P addition (NPT), and a control site (CT). We analyzed the response of ecosystem to altered nutrient stoichiometry using eddy covariance fluxes measurements, satellite observations, and digital repeat photography. A set of metrics, including phenological transition dates (PTDs; timing of green-up and dry-down), slopes during green-up and dry-down period, and seasonal amplitude, were extracted from time series of GPPmax and used to represent the seasonality of vegetation activity. The seasonal amplitude of GPPmax was higher for NT and NPT than CT, which was attributed to changes in structure and physiology induced by fertilization. PTDs were mainly driven by rainfall and exhibited no significant differences among treatments during the green-up period. Yet, both fertilized sites senesced earlier during the dry-down period (17-19 days), which was more pronounced in the NT due to larger evapotranspiration and water usage. Fertilization also resulted in a faster increase in GPPmax during the green-up period and a sharper decline in GPPmax during the dry-down period, with less prominent decline response in NPT. Overall, we demonstrated seasonality of vegetation activity was altered after fertilization and the importance of nutrient-water interaction in such water-limited ecosystems. With the projected warming-drying trend, the positive effects of N fertilization induced by N deposition on GPPmax may be counteracted by an earlier and faster dry-down in particular in areas where the N:P ratio increases, with potential impact on the carbon cycle of water-limited ecosystems.

'Hearing' alpine plants growing after snowmelt: ultrasonic snow sensors provide long-term series of alpine plant phenology.

In alpine environments, the growing season is severely constrained by low temperature and snow. Here, we aim at determining the climatic factors that best explain the interannual variation in spring growth onset of alpine plants, and at examining whether photoperiod might limit their phenological response during exceptionally warm springs and early snowmelts. We analysed 17 years of data (1998-2014) from 35 automatic weather stations located in subalpine and alpine zones ranging from 1560 to 2450 m asl in the Swiss Alps. These stations are equipped with ultrasonic sensors for snow depth measurements that are also able to detect plant growth in spring and summer, giving a unique opportunity to analyse snow and climate effects on alpine plant phenology. Our analysis showed high phenological variation among years, with one exceptionally early and late spring, namely 2011 and 2013. Overall, the timing of snowmelt and the beginning of plant growth were tightly linked irrespective of the elevation of the station. Snowmelt date was the best predictor of plant growth onset with air temperature after snowmelt modulating the plants' development rate. This multiple series of alpine plant phenology suggests that currently alpine plants are directly tracking climate change with no major photoperiod limitation.

Assessing Crop Coefficients for Natural Vegetated Areas Using Satellite Data and Eddy Covariance Stations.

The Food and Agricultural Organization (FAO) method for potential evapotranspiration assessment is based on the crop coefficient, which allows one to relate the reference evapotranspiration of well irrigated grass to the potential evapotranspiration of specific crops. The method was originally developed for cultivated species based on lysimeter measurements of potential evapotranspiration. Not many applications to natural vegetated areas exist due to the lack of available data for these species. In this paper we investigate the potential of using evapotranspiration measurements acquired by micrometeorological stations for the definition of crop coefficient functions of natural vegetated areas and extrapolation to ungauged sites through remotely sensed data. Pastures, deciduous and evergreen forests have been considered and lower crop coefficient values are found with respect to FAO data.

Influence of physiological phenology on the seasonal pattern of ecosystem respiration in deciduous forests.

Understanding the environmental and biotic drivers of respiration at the ecosystem level is a prerequisite to further improve scenarios of the global carbon cycle. In this study we investigated the relevance of physiological phenology, defined as seasonal changes in plant physiological properties, for explaining the temporal dynamics of ecosystem respiration (RECO) in deciduous forests. Previous studies showed that empirical RECO models can be substantially improved by considering the biotic dependency of RECO on the short-term productivity (e.g., daily gross primary production, GPP) in addition to the well-known environmental controls of temperature and water availability. Here, we use a model-data integration approach to investigate the added value of physiological phenology, represented by the first temporal derivative of GPP, or alternatively of the fraction of absorbed photosynthetically active radiation, for modeling RECO at 19 deciduous broadleaved forests in the FLUXNET La Thuile database. The new data-oriented semiempirical model leads to an 8% decrease in root mean square error (RMSE) and a 6% increase in the modeling efficiency (EF) of modeled RECO when compared to a version of the model that does not consider the physiological phenology. The reduction of the model-observation bias occurred mainly at the monthly time scale, and in spring and summer, while a smaller reduction was observed at the annual time scale. The proposed approach did not improve the model performance at several sites, and we identified as potential causes the plant canopy heterogeneity and the use of air temperature as a driver of ecosystem respiration instead of soil temperature. However, in the majority of sites the model-error remained unchanged regardless of the driving temperature. Overall, our results point toward the potential for improving current approaches for modeling RECO in deciduous forests by including the phenological cycle of the canopy.

Five years of phenological monitoring in a mountain grassland: inter-annual patterns and evaluation of the sampling protocol.

The increasingly important effect of climate change and extremes on alpine phenology highlights the need to establish accurate monitoring methods to track inter-annual variation (IAV) and long-term trends in plant phenology. We evaluated four different indices of phenological development (two for plant productivity, i.e., green biomass and leaf area index; two for plant greenness, i.e., greenness from visual inspection and from digital images) from a 5-year monitoring of ecosystem phenology, here defined as the seasonal development of the grassland canopy, in a subalpine grassland site (NW Alps). Our aim was to establish an effective observation strategy that enables the detection of shifts in grassland phenology in response to climate trends and meteorological extremes. The seasonal development of the vegetation at this site appears strongly controlled by snowmelt mostly in its first stages and to a lesser extent in the overall development trajectory. All indices were able to detect an anomalous beginning of the growing season in 2011 due to an exceptionally early snowmelt, whereas only some of them revealed a later beginning of the growing season in 2013 due to a late snowmelt. A method is developed to derive the number of samples that maximise the trade-off between sampling effort and accuracy in IAV detection in the context of long-term phenology monitoring programmes. Results show that spring phenology requires a smaller number of samples than autumn phenology to track a given target of IAV. Additionally, productivity indices (leaf area index and green biomass) have a higher sampling requirement than greenness derived from visual estimation and from the analysis of digital images. Of the latter two, the analysis of digital images stands out as the more effective, rapid and objective method to detect IAV in vegetation development.

Factors controlling the water quality of rock glacier springs in European and American mountain ranges.

Rock glaciers (RGs) provide significant water resources in mountain areas under climate change. Recent research has highlighted high concentrations of solutes including trace elements in RG-fed waters, with negative implications on water quality. Yet, sparse studies from a few locations hinder conclusions about the main drivers of solute export from RGs. Here, in an unprecedented effort, we collected published and unpublished data on rock glacier hydrochemistry around the globe. We considered 201 RG springs from mountain ranges across Europe, North and South America, using a combination of machine learning, multivariate and univariate analyses, and geochemical modeling. We found that 35 % of springs issuing from intact RGs (containing internal ice) have water quality below drinking water standards, compared to 5 % of springs connected to relict RGs (without internal ice). The interaction of ice and bedrock lithology is responsible for solute concentrations in RG springs. Indeed, we found higher concentrations of sulfate and trace elements in springs sourcing from intact RGs compared to water originating from relict RGs, mostly in specific lithological settings. Enhanced sulfide oxidation in intact RGs is responsible for the elevated trace element concentrations. Challenges for water management may arise in mountain catchments rich in intact RGs, and where the predisposing geology would make these areas geochemical RG hotspots. Our work represents a first comprehensive attempt to identify the main drivers of solute concentrations in RG waters.

Tradeoffs between global warming and day length on the start of the carbon uptake period in seasonally cold ecosystems.

It is well established that warming leads to longer growing seasons in seasonally cold ecosystems. Whether this goes along with an increase in the net ecosystem carbon dioxide (CO2) uptake is much more controversial. We studied the effects of warming on the start of the carbon uptake period (CUP) of three mountain grasslands situated along an elevational gradient in the Alps. To this end we used a simple empirical model of the net ecosystem CO2 exchange, calibrated and forced with multi-year empirical data from each site. We show that reductions in the quantity and duration of daylight associated with earlier snowmelts were responsible for diminishing returns, in terms of carbon gain, from longer growing seasons caused by reductions in daytime photosynthetic uptake and increases in nighttime losses of CO2. This effect was less pronounced at high, compared to low, elevations, where the start of the CUP occurred closer to the summer solstice when changes in day length and incident radiation are minimal.

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.

Sensitivity of gross primary productivity to climatic drivers during the summer drought of 2018 in Europe.

In summer 2018, Europe experienced a record drought, but it remains unknown how the drought affected ecosystem carbon dynamics. Using observations from 34 eddy covariance sites in different biomes across Europe, we studied the sensitivity of gross primary productivity (GPP) to environmental drivers during the summer drought of 2018 versus the reference summer of 2016. We found a greater drought-induced decline of summer GPP in grasslands (-38%) than in forests (-10%), which coincided with reduced evapotranspiration and soil water content (SWC). As compared to the 'normal year' of 2016, GPP in different ecosystems exhibited more negative sensitivity to summer air temperature (Ta) but stronger positive sensitivity to SWC during summer drought in 2018, that is, a stronger reduction of GPP with soil moisture deficit. We found larger negative effects of Ta and vapour pressure deficit (VPD) but a lower positive effect of photosynthetic photon flux density on GPP in 2018 compared to 2016, which contributed to reduced summer GPP in 2018. Our results demonstrate that high temperature-induced increases in VPD and decreases in SWC aggravated drought impacts on GPP. This article is part of the theme issue 'Impacts of the 2018 severe drought and heatwave in Europe: from site to continental scale'.

Altered energy partitioning across terrestrial ecosystems in the European drought year 2018.

Drought and heat events, such as the 2018 European drought, interact with the exchange of energy between the land surface and the atmosphere, potentially affecting albedo, sensible and latent heat fluxes, as well as CO2 exchange. Each of these quantities may aggravate or mitigate the drought, heat, their side effects on productivity, water scarcity and global warming. We used measurements of 56 eddy covariance sites across Europe to examine the response of fluxes to extreme drought prevailing most of the year 2018 and how the response differed across various ecosystem types (forests, grasslands, croplands and peatlands). Each component of the surface radiation and energy balance observed in 2018 was compared to available data per site during a reference period 2004-2017. Based on anomalies in precipitation and reference evapotranspiration, we classified 46 sites as drought affected. These received on average 9% more solar radiation and released 32% more sensible heat to the atmosphere compared to the mean of the reference period. In general, drought decreased net CO2 uptake by 17.8%, but did not significantly change net evapotranspiration. The response of these fluxes differed characteristically between ecosystems; in particular, the general increase in the evaporative index was strongest in peatlands and weakest in croplands. This article is part of the theme issue 'Impacts of the 2018 severe drought and heatwave in Europe: from site to continental scale'.