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.

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.

Revisiting the choice of the driving temperature for eddy covariance CO2 flux partitioning.

So-called CO2 flux partitioning algorithms are widely used to partition the net ecosystem CO2 exchange into the two component fluxes, gross primary productivity and ecosystem respiration. Common CO2 flux partitioning algorithms conceptualize ecosystem respiration to originate from a single source, requiring the choice of a corresponding driving temperature. Using a conceptual dual-source respiration model, consisting of an above- and a below-ground respiration source each driven by a corresponding temperature, we demonstrate that the typical phase shift between air and soil temperature gives rise to a hysteresis relationship between ecosystem respiration and temperature. The hysteresis proceeds in a clockwise fashion if soil temperature is used to drive ecosystem respiration, while a counter-clockwise response is observed when ecosystem respiration is related to air temperature. As a consequence, nighttime ecosystem respiration is smaller than daytime ecosystem respiration when referenced to soil temperature, while the reverse is true for air temperature. We confirm these qualitative modelling results using measurements of day and night ecosystem respiration made with opaque chambers in a short-statured mountain grassland. Inferring daytime from nighttime ecosystem respiration or vice versa, as attempted by CO2 flux partitioning algorithms, using a single-source respiration model is thus an oversimplification resulting in biased estimates of ecosystem respiration. We discuss the likely magnitude of the bias, options for minimizing it and conclude by emphasizing that the systematic uncertainty of gross primary productivity and ecosystem respiration inferred through CO2 flux partitioning needs to be better quantified and reported.

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.

Convergence of potential net ecosystem production among contrasting C3 grasslands.

Metabolic theory and body size constraints on biomass production and decomposition suggest that differences in the intrinsic potential net ecosystem production (NEPPOT ) should be small among contrasting C3 grasslands and therefore unable to explain the wide range in the annual apparent net ecosystem production (NEPAPP ) reported by previous studies. We estimated NEPPOT for nine C3 grasslands under contrasting climate and management regimes using multiyear eddy covariance data. NEPPOT converged within a narrow range, suggesting little difference in the net carbon dioxide uptake capacity among C3 grasslands. Our results indicate a unique feature of C3 grasslands compared with other terrestrial ecosystems and suggest a state of stability in NEPPOT due to tightly coupled production and respiration processes. Consequently, the annual NEPAPP of C3 grasslands is primarily a function of seasonal and short-term environmental and management constraints, and therefore especially susceptible to changes in future climate patterns and associated adaptation of management practices.

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.