Future climate warming is expected to enhance plant growth in temperate ecosystems and to increase carbon sequestration. But although severe regional heatwaves may become more frequent in a changing climate, their impact on terrestrial carbon cycling is unclear. Here we report measurements of ecosystem carbon dioxide fluxes, remotely sensed radiation absorbed by plants, and country-level crop yields taken during the European heatwave in 2003. We use a terrestrial biosphere simulation model to assess continental-scale changes in primary productivity during 2003, and their consequences for the net carbon balance. We estimate a 30 per cent reduction in gross primary productivity over Europe, which resulted in a strong anomalous net source of carbon dioxide (0.5 Pg C yr(-1)) to the atmosphere and reversed the effect of four years of net ecosystem carbon sequestration. Our results suggest that productivity reduction in eastern and western Europe can be explained by rainfall deficit and extreme summer heat, respectively. We also find that ecosystem respiration decreased together with gross primary productivity, rather than accelerating with the temperature rise. Model results, corroborated by historical records of crop yields, suggest that such a reduction in Europe's primary productivity is unprecedented during the last century. An increase in future drought events could turn temperate ecosystems into carbon sources, contributing to positive carbon-climate feedbacks already anticipated in the tropics and at high latitudes.

The European CARBOEUROPE/FLUXNET monitoring sites, spatial remote sensing observations via the EOS-MODIS sensor and ecosystem modelling provide independent and complementary views on the effect of the 2003 heatwave on the European biosphere's productivity and carbon balance. In our analysis, these data streams consistently demonstrate a strong negative anomaly of the primary productivity during the summer of 2003. FLUXNET eddy-covariance data indicate that the drop in productivity was not primarily caused by high temperatures ('heat stress') but rather by limitation of water (drought stress) and that, contrary to the classical expectation about a heat wave, not only gross primary productivity but also ecosystem respiration declined by up to more than to 80 gC m(-2) month(-1). Anomalies of carbon and water fluxes were strongly correlated. While there are large between-site differences in water-use efficiency (WUE, 1-6 kg C kg(-1) H₂O) here defined as gross carbon uptake divided by evapotranspiration (WUE=GPP/ET), the year-to-year changes in WUE were small (< 1 g kg(-1)) and quite similar for most sites (i.e. WUE decreased during the year of the heatwave). Remote sensing data from MODIS and AVHRR both indicate a strong negative anomaly of the fraction of absorbed photosynthetically active radiation in summer 2003, at more than five standard deviations of the previous years. The spatial differentiation of this anomaly follows climatic and land-use patterns: Largest anomalies occur in the centre of the meteorological anomaly (central Western Europe) and in areas dominated by crops or grassland. A preliminary model intercomparison along a gradient from data-oriented models to process-oriented models indicates that all approaches are similarly describing the spatial pattern of ecosystem sensitivity to the climatic 2003 event with major exceptions in the Alps and parts of Eastern Europe, but differed with respect to their interannual variability. [References: 46

The temperature sensitivity of soil respiration (SR) is often estimated from the seasonal changes in the flux relative to those in soil temperature, and subsequently used in models to interpolate or predict soil fluxes. However, temperature sensitivities derived from seasonal changes in SR (from here on denoted seasonal Q 10 ) may not solely reflect the temperature sensitivity of SR, because seasonal changes in SR can also be affected by other seasonally fluctuating conditions and processes. In this manuscript, we present a case study of how the seasonal Q 10 of SR can be decoupled from the temperature sensitivity of SR. In a mixed temperate forest, we measured SR under vegetations with different leaf strategies: pure evergreen, pure deciduous, and mixed. Seasonal Q 10 was much higher under deciduous than under evergreen canopies. However, at a shorter time scale, both vegetation types exhibited very similar Q 10 values, indicating that the large differences in seasonal Q 10 do not represent differences in the temperature sensitivity of the soil metabolism. The seasonal Q 10 depends strongly on the amplitude of the seasonal changes in SR (SR s ), which, under the particular climatic and edaphic conditions of our forest study site, were significantly larger in deciduous forest. In turn, SR s was positively correlated with the seasonal changes in leaf area index (LAI s ), a measure of the deciduousness of the vegetation. Thus, in this temperate maritime forest, seasonal Q 10 of SR was strongly influenced by the deciduousness of the vegetation. We conclude that the large differences in seasonal Q 10 were not entirely due to different temperature sensitivities, but also to different seasonal patterns of plant activity in the evergreen and deciduous plants of this site. Some coniferous forests may be more seasonal than the one we studied, and the deciduous-evergreen differences observed here may not be broadly applicable, but this case study demonstrates that variation of plant phenological process can significantly contribute to the seasonality of SR, and, hence, calculated Q 10 values. Where this occurs, the seasonal Q 10 value for SR does not accurately represent temperature sensitivity. Because the strong seasonal correlation between SR and temperature does not necessarily imply a causal relationship, Q 10 values derived form annual patterns of SR should be used with caution when predicting future responses of SR to climatic change.

Soil respiration (SR) was monitored periodically throughout 2001 in a Scots pine (Pinus sylvestris L.) stand located in the Belgian Campine region. As expected for a temperate maritime forest, temperature was the dominant control over SR during most of the year. However, during late spring and summer, when soil water content (SWC) was limiting, SR was insensitive to temperature (Q(10) = 1.24). We observed that during prolonged rain-free periods, when SWC was less than 15% (v/v), SR decreased dramatically (up to 50%) and SWC took over control of SR. During such drought periods, however, rain events sometimes stimulated SR and restored temperature control over SR, even though SWC in the mineral soil was low. We hypothesize that restoration of temperature control occurred only when rain events adequately rewetted the uppermost soil layers, where most of the respiratory activity occurred. To quantify the rewetting capacity of rain events, an index (I(w)) was designed that incorporated rainfall intensity, time elapsed since the last rain event, and atmospheric vapor pressure deficit (a proxy for evaporative water losses). To simulate SR fluxes, a model was developed that included the effects of soil temperature and, under drought and non-rewetting conditions (I(w) and SWC < threshold), an SWC response function. The model explained 95% of the temporal variability in SR observed during summer, whereas the temperature function alone explained only 73% of this variability. Our results revealed that, in addition to temperature and SWC, rain plays a role in determining the total amount of carbon released from soils, even in a maritime climate.

Abstract. We applied a site evaluation approach combining Lagrangian Stochastic footprint modeling with a quality assessment approach for eddy-covariance data to 25 forested sites of the CarboEurope-IP network. The analysis addresses the spatial representativeness of the flux measurements, instrumental effects on data quality, spatial patterns in the data quality, and the performance of the coordinate rotation method. Our findings demonstrate that application of a footprint filter could strengthen the CarboEurope-IP flux database, since only one third of the sites is situated in truly homogeneous terrain. Almost half of the sites experience a significant reduction in eddy-covariance data quality under certain conditions, though these effects are mostly constricted to a small portion of the dataset. Reductions in data quality of the sensible heat flux are mostly induced by characteristics of the surrounding terrain, while the latent heat flux is subject to instrumentation-related problems. The Planar-Fit coordinate rotation proved to be a reliable tool for the majority of the sites using only a single set of rotation angles. Overall, we found a high average data quality for the CarboEurope-IP network, with good representativeness of the measurement data for the specified target land cover types.

In this study we examined ecosystem respiration (RECO) data from 104 sites belonging to FLUXNET, the global network of eddy covariance flux measurements. The goal was to identify the main factors involved in the variability of RECO: temporally and between sites as affected by climate, vegetation structure and plant functional type (PFT) (evergreen needleleaf, grasslands, etc.). We demonstrated that a model using only climate drivers as predictors of RECO failed to describe part of the temporal variability in the data and that the dependency on gross primary production (GPP) needed to be included as an additional driver of RECO. The maximum seasonal leaf area index (LAIMAX) had an additional effect that explained the spatial variability of reference respiration (the respiration at reference temperature Tref=15 °C, without stimulation introduced by photosynthetic activity and without water limitations), with a statistically significant linear relationship (r2=0.52, P<0.001, n=104) even within each PFT. Besides LAIMAX, we found that reference respiration may be explained partially by total soil carbon content (SoilC). For undisturbed temperate and boreal forests a negative control of total nitrogen deposition (Ndepo) on reference respiration was also identified. We developed a new semiempirical model incorporating abiotic factors (climate), recent productivity (daily GPP), general site productivity and canopy structure (LAIMAX) which performed well in predicting the spatio‐temporal variability of RECO, explaining >70% of the variance for most vegetation types. Exceptions include tropical and Mediterranean broadleaf forests and deciduous broadleaf forests. Part of the variability in respiration that could not be described by our model may be attributed to a series of factors, including phenology in deciduous broadleaf forests and management practices in grasslands and croplands.

[1] We analyze the impacts of the steady state assumption on inverse model parameter retrieval from biogeochemical models. An inverse model parameterization study using eddy covariance CO 2 flux data was performed with the Carnegie Ames Stanford Approach (CASA) model under conditions of strict and relaxed carbon cycle steady state assumption (CCSSA) in order to evaluate both the robustness of the model's structure for the simulation of net ecosystem carbon fluxes and the assessment of the CCSSA effects on simulations and parameter estimation. Net ecosystem production (NEP) measurements from several eddy covariance sites were compared with NEP estimates from the CASA model driven by local weather station climate inputs as well as by remotely sensed fraction of photosynthetically active radiation absorbed by vegetation and leaf area index. The parameters considered for optimization are directly related to aboveground and belowground modeled responses to temperature and water availability, as well as a parameter (h) that relaxed the CCSSA in the model, allowing for site level simulations to be initialized either as net sinks or sources. A robust relationship was observed between NEP observations and predictions for most of the sites through the range of temporal scales considered (daily, weekly, biweekly, and monthly), supporting the conclusion that the model structure is able to capture the main processes explaining NEP variability. Overall, relaxing CCSSA increased model efficiency (21%) and decreased normalized average error (À92%). Intersite variability was a major source of variance in model performance differences between fixed (CCSSA f ) and relaxed (CCSSA r ) CCSSA conditions. These differences were correlated with mean annual NEP observations, where an average increase in modeling efficiency of 0.06 per 100 g C m À2 a À1(where a is years) of NEP is observed (a < 0.003). The parameter h was found to be a key parameter in the optimization exercise, generating significant model efficiency losses when removed from the initial parameter set and parameter uncertainties were significantly lower under CCSSA r . Moreover, modeled soil carbon stocks were generally closer to observations once the steady state assumption was relaxed. Finally, we also show that estimates of individual parameters are affected by the steady state assumption. For example, estimates of radiation-use efficiency were strongly affected by the CCSSA f indicating compensation effects for the inadequate steady state assumption, leading to GLOBAL

Sun-induced fluorescence (SIF) in the far-red region provides a new noninvasive measurement approach that has the potential to quantify dynamic changes in light-use efficiency and gross primary production (GPP). However, the mechanistic link between GPP and SIF is not completely understood. We analyzed the structural and functional factors controlling the emission of SIF at 760 nm (F ) in a Mediterranean grassland manipulated with nutrient addition of nitrogen (N), phosphorous (P) or nitrogen-phosphorous (NP). Using the soil-canopy observation of photosynthesis and energy (SCOPE) model, we investigated how nutrient-induced changes in canopy structure (i.e. changes in plant forms abundance that influence leaf inclination distribution function, LIDF) and functional traits (e.g. N content in dry mass of leaves, N%, Chlorophyll a+b concentration (Cab) and maximum carboxylation capacity (V )) affected the observed linear relationship between F and GPP. We conclude that the addition of nutrients imposed a change in the abundance of different plant forms and biochemistry of the canopy that controls F . Changes in canopy structure mainly control the GPP-F relationship, with a secondary effect of Cab and V . In order to exploit F data to model GPP at the global/regional scale, canopy structural variability, biodiversity and functional traits are important factors that have to be considered.