Semiempirical modeling of abiotic and biotic factors controlling ecosystem respiration across eddy covariance sites

In this study we examined ecosystem respiration (R_ECO) 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 R_ECO: 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 R_ECO 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 R_ECO. The maximum seasonal leaf area index (LAI_MAX) had an additional effect that explained the spatial variability of reference respiration (the respiration at reference temperature T_ref=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 LAI_MAX, 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 (N_depo) 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 (LAI_MAX) which performed well in predicting the spatio‐temporal variability of R_ECO, 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.     

Functional changes in the control of carbon fluxes after 3 years of increased drought in a Mediterranean evergreen forest?

Our objective was to test how a long‐term increased water limitation affects structural and functional properties of a Mediterranean ecosystem, and how these changes modify the response of the main carbon fluxes to climatic controls. In 2003, a 27% throughfall exclusion experiment was installed in a Quercus ilex L. forest in France. Gross primary production (GPP), ecosystem respiration (R_ECO) and net ecosystem exchange (NEE) were estimated in a control and a dry treatment. Decreasing throughfall decreased GPP by 14% and had a smaller effect on R_ECO (?12%), especially soil respiration R_S (?11%). Interannual variability of GPP (29%) was higher than for R_ECO (12%). Error propagation was used to estimates uncertainties in the NEE fluxes, which ranged from 3% to 10% in the control treatment but up to 167% for NEE in the dry treatment because more steps and data types were involved in the scaling. After 3 years of throughfall exclusion, we found no acclimation of R_S to climatic drivers. Functional properties of the response of R_S to soil water, temperature and rain pulse remained similar in the control and the dry treatments. A diurnal clockwise hysteresis in R_S was probably controlled by canopy photosynthesis with a 3 h lag. The proportion of diurnal variation of respiration due to photosynthesis was similar in all treatments (4–5%). Because of the characteristic of rain in Mediterranean climates, a continuous decrease of water input in these environments have an effect on topsoil water and consequently on R_S only during short periods when rainfall is characterized by infrequent and small events that does not allow the topsoil to reach field capacity and does not allow to dry completely. However, in the longer term, we expect a stronger decrease in R_S in the dry treatment driven by the decrease in GPP.     

Respiration acclimation contributes to high carbon-use efficiency in a seasonally dry pine forest

Predictions of warming and drying in the Mediterranean and other regions require quantifying of such effects on ecosystem carbon dynamics and respiration. Long‐term effects can only be obtained from forests in which seasonal drought is a regular feature. We carried out measurements in a semiarid Pinus halepensis (Aleppo pine) forest of aboveground respiration rates of foliage, R_f, and stem, R_t over 3 years. Component respiration combined with ongoing biometric, net CO₂ flux [net ecosystem productivity (NEP)] and soil respiration measurements were scaled to the ecosystem level to estimate gross and net primary productivity (GPP, NPP) and carbon‐use efficiency (CUE=NPP/GPP) using 6 years data. GPP, NPP and NEP were, on average, 880, 350 and 211 g C m^?2 yr^?1, respectively. The above ground respiration made up half of total ecosystem respiration but CUE remained high at 0.4. Large seasonal variations in both R_f and R_t were not consistently correlated with seasonal temperature trends. Seasonal adjustments of respiration were observed in both the normalized rate (R₂₀) and short‐term temperature sensitivity (Q₁₀), resulting in low respiration rates during the hot, dry period. R_f in fully developed needles was highest over winter–spring, and foliage R₂₀ was correlated with photosynthesis over the year. Needle growth occurred over summer, with respiration rates in developing needles higher than the fully developed foliage at most times. R_t showed a distinct seasonal maximum in May irrespective of year, which was not correlated to the winter stem growth, but could be associated with phenological drivers such as carbohydrate re‐mobilization and cambial activity. We show that in a semiarid pine forest photosynthesis and stem growth peak in (wet) winter and leaf growth in (dry) summer, and associated adjustments of component respiration, dominated by those in R₂₀, minimize annual respiratory losses. This is likely a key for maintaining high CUE and ecosystem productivity similar to much wetter sites, and could lead to different predictions of the effect of warming and drying climate on productivity of pine forests than based on short‐term droughts.     

Seasonal variation in leaf properties and ecosystem carbon budget in a cool-temperate deciduous broad-leaved forest: simulation analysis at Takayama site, Japan

Seasonal changes in gross primary production (GPP) and net ecosystem production (NEP) in temperate deciduous forests are mostly driven by environmental conditions and the phenology of leaf demography. This study addresses another factor, temporal changes in leaf properties, i.e., leaf aging from emergence to senescence. A process-based model was used to link the ecosystem-scale carbon budget with leaf-level properties on the basis of field observation and scaling procedures; temporal variations in leaf thickness (leaf mass per area, LMA), photosynthetic rubisco (V_cmax) and electron-transport (J_max) capacity, and dark respiration (R_d) were empirically parameterized. The model was applied to a cool-temperate deciduous broad-leaved forest at Takayama, in central Japan, and validated with data of net ecosystem CO₂ exchange (NEE=–NEP) measured using the eddy-covariance method. NEP of the Takayama site varied seasonally from 3 g C m^–2 day^–1 net source in late winter to 5 g C m^–2 day^–1 net sink in early to mid-summer. A sensitivity experiment showed that removing the leaf-aging effect changed the seasonal CO₂ exchange pattern, and led to overestimation of annual GPP by 6% and annual NEP by 38%. We found that seasonal variation in V_cmax affected the seasonal pattern and annual budget of CO₂ exchange most strongly; LMA and R_d had moderate influences. The rapid change in V_cmax and R_d during leaf emergence and senescence was important in evaluating GPP and NEP of the temperate deciduous forest.     

Effects of seasonal and interannual variations in leaf photosynthesis and canopy leaf area index on gross primary production of a cool-temperate deciduous broadleaf forest in Takayama, Japan

Revealing the seasonal and interannual variations in forest canopy photosynthesis is a critical issue in understanding the ecological mechanisms underlying the dynamics of carbon dioxide exchange between the atmosphere and deciduous forests. This study examined the effects of temporal variations of canopy leaf area index (LAI) and leaf photosynthetic capacity [the maximum velocity of carboxylation (V _cmax)] on gross primary production (GPP) of a cool-temperate deciduous broadleaf forest for 5 years in Takayama AsiaFlux site, central Japan. We made two estimations to examine the effects of canopy properties on GPP; one is to incorporate the in situ observation of V _cmax and LAI throughout the growing season, and another considers seasonality of LAI but constantly high V _cmax. The simulations indicated that variation in V _cmax and LAI, especially in the leaf expansion period, had remarkable effects on GPP, and if V _cmax was assumed constant GPP will be overestimated by 15%. Monthly examination of air temperature, radiation, LAI and GPP suggested that spring temperature could affect canopy phenology, and also that GPP in summer was determined mainly by incoming radiation. However, the consequences among these factors responsible for interannual changes of GPP are not straightforward since leaf expansion and senescence patterns and summer meteorological conditions influence GPP independently. This simulation based on in situ ecophysiological research suggests the importance of intensive consideration and understanding of the phenology of leaf photosynthetic capacity and LAI to analyze and predict carbon fixation in forest ecosystems.     

Chlorophyll fluorescence tracks seasonal variations of photosynthesis from leaf to canopy in a temperate forest

Accurate estimation of terrestrial gross primary productivity (GPP) remains a challenge despite its importance in the global carbon cycle. Chlorophyll fluorescence (ChlF) has been recently adopted to understand photosynthesis and its response to the environment, particularly with remote sensing data. However, it remains unclear how ChlF and photosynthesis are linked at different spatial scales across the growing season. We examined seasonal relationships between ChlF and photosynthesis at the leaf, canopy, and ecosystem scales and explored how leaf‐level ChlF was linked with canopy‐scale solar‐induced chlorophyll fluorescence (SIF) in a temperate deciduous forest at Harvard Forest, Massachusetts, USA. Our results show that ChlF captured the seasonal variations of photosynthesis with significant linear relationships between ChlF and photosynthesis across the growing season over different spatial scales (R² = 0.73, 0.77, and 0.86 at leaf, canopy, and satellite scales, respectively; P < 0.0001). We developed a model to estimate GPP from the tower‐based measurement of SIF and leaf‐level ChlF parameters. The estimation of GPP from this model agreed well with flux tower observations of GPP (R² = 0.68; P < 0.0001), demonstrating the potential of SIF for modeling GPP. At the leaf scale, we found that leaf F_q’/F_m’, the fraction of absorbed photons that are used for photochemistry for a light‐adapted measurement from a pulse amplitude modulation fluorometer, was the best leaf fluorescence parameter to correlate with canopy SIF yield (SIF/APAR, R² = 0.79; P < 0.0001). We also found that canopy SIF and SIF‐derived GPP (GPP_SIF) were strongly correlated to leaf‐level biochemistry and canopy structure, including chlorophyll content (R² = 0.65 for canopy GPP_SIF and chlorophyll content; P < 0.0001), leaf area index (LAI) (R² = 0.35 for canopy GPP_SIF and LAI; P < 0.0001), and normalized difference vegetation index (NDVI) (R² = 0.36 for canopy GPP_SIF and NDVI; P < 0.0001). Our results suggest that ChlF can be a powerful tool to track photosynthetic rates at leaf, canopy, and ecosystem scales.     

Climatic controls on the carbon and water balances of a boreal aspen forest, 1994–2003

The carbon and water budgets of boreal and temperate broadleaf forests are sensitive to interannual climatic variability and are likely to respond to climate change. This study analyses 9 years of eddy‐covariance data from the Boreal Ecosystem Research and Monitoring Sites (BERMS) Southern Old Aspen site in central Saskatchewan, Canada and characterizes the primary climatic controls on evapotranspiration, net ecosystem production (F_NEP), gross ecosystem photosynthesis (P) and ecosystem respiration (R). The study period was dominated by two climatic extremes: extreme warm and cool springs, which produced marked contrasts in the canopy duration, and a severe, 3‐year drought. Annual F_NEP varied among years from 55 to 367 g C m⁻² (mean 172, SD 94). Interannual variability in F_NEP was controlled primarily by factors that affected the R/P ratio, which varied between 0.74 and 0.96 (mean 0.87, SD 0.06). Canopy duration enhanced P and F_NEP with no apparent effect on R. The fraction of annual photosynthetically active radiation (PAR) that was absorbed by the canopy foliage varied from 38% in late leaf‐emergence years to 51% in early leaf‐emergence years. Photosynthetic light‐use efficiency (mean 0.0275, SD 0.026 mol C mol⁻¹ photons) was relatively constant during nondrought years but declined with drought intensity to a minimum of 0.0228 mol C mol⁻¹ photons during the most severe drought year. The impact of drought on F_NEP varied with drought intensity. Years of mild‐to‐moderate drought suppressed R while having little effect on P, so that F_NEP was enhanced. Years of severe drought suppressed both R and P, causing either little change or a subtle reduction in F_NEP. The analysis produced new insights into the dominance of canopy duration as the most important biophysical control on F_NEP. The results suggested a simple conceptual model for annual F_NEP in boreal deciduous forests. When water is not limiting, annual P is controlled by canopy duration via its influence on absorbed PAR at constant light‐use efficiency. Water stress suppresses P, by reducing light‐use efficiency, and R, by limiting growth and/or suppressing microbial respiration. The high photosynthetic light‐use efficiency showed this site to be a highly productive boreal deciduous forest, with properties similar to many temperate deciduous forests.     

Estimates of CO2 uptake and release among European forests based on eddy covariance data

The net ecosystem exchange (NEE) of forests represents the balance of gross primary productivity (GPP) and respiration (R). Methods to estimate these two components from eddy covariance flux measurements are usually based on a functional relationship between respiration and temperature that is calibrated for night‐time (respiration) fluxes and subsequently extrapolated using daytime temperature measurements. However, respiration fluxes originate from different parts of the ecosystem, each of which experiences its own course of temperature. Moreover, if the temperature–respiration function is fitted to combined data from different stages of biological development or seasons, a spurious temperature effect may be included that will lead to overestimation of the direct effect of temperature and therefore to overestimates of daytime respiration. We used the EUROFLUX eddy covariance data set for 15 European forests and pooled data per site, month and for conditions of low and sufficient soil moisture, respectively. We found that using air temperature (measured above the canopy) rather than soil temperature (measured 5 cm below the surface) yielded the most reliable and consistent exponential (Q₁₀) temperature–respiration relationship. A fundamental difference in air temperature‐based Q₁₀ values for different sites, times of year or soil moisture conditions could not be established; all were in the range 1.6–2.5. However, base respiration (R₀, i.e. respiration rate scaled to 0°C) did vary significantly among sites and over the course of the year, with increased base respiration rates during the growing season. We used the overall mean Q₁₀ of 2.0 to estimate annual GPP and R. Testing suggested that the uncertainty in total GPP and R associated with the method of separation was generally well within 15%. For the sites investigated, we found a positive relationship between GPP and R, indicating that there is a latitudinal trend in NEE because the absolute decrease in GPP towards the pole is greater than in R.     

Simulation and scaling of temporal variation in gross primary production for coniferous and deciduous temperate forests

Observations of ecosystem net carbon dioxide exchange obtained with eddy covariance techniques over a 4‐year period at spruce, beech and pine forest sites were used to derive time series data for gross primary production (GPP) and ecosystem respiration (R_eco). A detailed canopy gas exchange model (PROXEL_NEE) was inverted at half‐hour time step to estimate seasonal changes in carboxylation capacity and light utilization efficiency of the forest canopies. The parameter estimates were then used further to develop a time‐dependent modifier of physiological activity in the daily time step gas exchange model of Chen et al. (1999) , previously used for regional simulations in BOREAS. The daily model was run under a variety of assumptions and the results emphasize the need in future analyses: (1) to focus on time‐dependent seasonal changes in canopy physiology as well as in leaf area index, (2) to compare time courses of physiological change in different habitats in terms of recognizable cardinal points in the seasonal course, and (3) to develop methods for utilizing information on seasonal changes in physiology in regional and continental carbon budget simulations. The results suggest that the daily model with appropriate seasonal adjustments for physiological process regulation should be an efficient tool for use in conjunction with remote sensing for regional evaluation of global change scenarios.     

表征亚热带常绿林光合作用季节变化特征的多种植被指数

利用遥感方法可以在区域尺度反演地表植被的光合生理状况和生产力变化,但亚热带常绿林冠层结构季节变化较小,传统的光谱植被指数对植被光合作用难以准确捕捉。利用2014—2015年中国科学院广东省鼎湖山森林生态试验站多角度自动光谱观测系统的光谱反射数据,分别反演传统冠层结构型植被指数(NDVI)、光合生理生化型植被指数(CCI)和叶绿素荧光型植被指数(NDFI_(685)和NDFI_(760)),并利用不同类型植被指数的组合,构建多元线性回归模型。结果表明:亚热带常绿针阔混交林三种类型植被指数均与GPP的动态变化有显著的相关性,其中,NDVI是表征GPP较优的植被指数(R~2=0.60,P0.01),其次为CCI(R~2=0.55,P0.01),而NDFI能够作为辅助指数,有效提高NDVI(R~2=0.68,P0.001)和CCI(R~2=0.67,P0.001)表征GPP的程度。多个植被指数参与构建的多元回归模型能够有效提高亚热带地区常绿林GPP季节动态变化的拟合精度,提升遥感精确评估亚热带森林生产力的能力。     

Carbon dynamics of mature and regrowth tropical forests derived from a pantropical database (TropForC‐db)

Tropical forests play a critical role in the global carbon (C) cycle, storing ~45% of terrestrial C and constituting the largest component of the terrestrial C sink. Despite their central importance to the global C cycle, their ecosystem‐level C cycles are not as well‐characterized as those of extra‐tropical forests, and knowledge gaps hamper efforts to quantify C budgets across the tropics and to model tropical forest‐climate interactions. To advance understanding of C dynamics of pantropical forests, we compiled a new database, the Tropical Forest C database (TropForC‐db), which contains data on ground‐based measurements of ecosystem‐level C stocks and annual fluxes along with disturbance history. This database currently contains 3568 records from 845 plots in 178 geographically distinct areas, making it the largest and most comprehensive database of its type. Using TropForC‐db, we characterized C stocks and fluxes for young, intermediate‐aged, and mature forests. Relative to existing C budgets of extra‐tropical forests, mature tropical broadleaf evergreen forests had substantially higher gross primary productivity (GPP) and ecosystem respiration (R_eco), their autotropic respiration (R_a) consumed a larger proportion (~67%) of GPP, and their woody stem growth (ANPP_stem) represented a smaller proportion of net primary productivity (NPP, ~32%) or GPP (~9%). In regrowth stands, aboveground biomass increased rapidly during the first 20 years following stand‐clearing disturbance, with slower accumulation following agriculture and in deciduous forests, and continued to accumulate at a slower pace in forests aged 20–100 years. Most other C stocks likewise increased with stand age, while potential to describe age trends in C fluxes was generally data‐limited. We expect that TropForC‐db will prove useful for model evaluation and for quantifying the contribution of forests to the global C cycle. The database version associated with this publication is archived in Dryad (DOI: 10.5061/dryad.t516f ) and a dynamic version is maintained at https://github.com/forc-db .     

Sensitivity of CO<Subscript>2</Subscript> Exchange of Fen Ecosystem Components to Water Level Variation

Abstract Climate change is predicted to bring about a water level (WL) draw-down in boreal peatlands. This study aimed to assess the effect of WL on the carbon dioxide (CO₂) dynamics of a boreal oligotrophic fen ecosystem and its components; Sphagnum mosses, sedges, dwarf shrubs and the underlying peat. We measured CO₂ exchange with closed chambers during four growing seasons in a study site that comprised different vegetation treatments. WL gradient in the site was broadened by surrounding half of the site with a shallow ditch that lowered the WL by 10–25 cm. We modeled gross photosynthesis (P _G) and ecosystem respiration (R _ECO) and simulated the CO₂ exchange in three WL conditions: prevailing and WL draw-down scenarios of 14 and 22 cm. WL draw-down both reduced the P _G and increased the R _ECO, thus leading to a smaller net CO₂ uptake in the ecosystem. Of the different components, Sphagnum mosses were most sensitive to WL draw-down and their physiological activities almost ceased. Vascular plant CO₂ exchange, en bloc, hardly changed but whereas sedges contributed twice as much to the CO₂ exchange as shrubs in the prevailing conditions, the situation was reversed in the WL draw-down scenarios. Peat respiration was the biggest component in R _ECO in all WL conditions and the increase in R _ECO following the WL draw-down was due to the increase in peat respiration. The results imply that functional diversity buffers the ecosystem against environmental variability and that in the long term, WL draw-down changes the vegetation composition of boreal fens.     

Partitioning controls on Amazon forest photosynthesis between environmental and biotic factors at hourly to interannual timescales

Gross ecosystem productivity (GEP) in tropical forests varies both with the environment and with biotic changes in photosynthetic infrastructure, but our understanding of the relative effects of these factors across timescales is limited. Here, we used a statistical model to partition the variability of seven years of eddy covariance‐derived GEP in a central Amazon evergreen forest into two main causes: variation in environmental drivers (solar radiation, diffuse light fraction, and vapor pressure deficit) that interact with model parameters that govern photosynthesis and biotic variation in canopy photosynthetic light‐use efficiency associated with changes in the parameters themselves. Our fitted model was able to explain most of the variability in GEP at hourly (R² = 0.77) to interannual (R² = 0.80) timescales. At hourly timescales, we found that 75% of observed GEP variability could be attributed to environmental variability. When aggregating GEP to the longer timescales (daily, monthly, and yearly), however, environmental variation explained progressively less GEP variability: At monthly timescales, it explained only 3%, much less than biotic variation in canopy photosynthetic light‐use efficiency, which accounted for 63%. These results challenge modeling approaches that assume GEP is primarily controlled by the environment at both short and long timescales. Our approach distinguishing biotic from environmental variability can help to resolve debates about environmental limitations to tropical forest photosynthesis. For example, we found that biotically regulated canopy photosynthetic light‐use efficiency (associated with leaf phenology) increased with sunlight during dry seasons (consistent with light but not water limitation of canopy development) but that realized GEP was nonetheless lower relative to its potential efficiency during dry than wet seasons (consistent with water limitation of photosynthesis in given assemblages of leaves). This work highlights the importance of accounting for differential regulation of GEP at different timescales and of identifying the underlying feedbacks and adaptive mechanisms.     

An analysis of soil respiration across northern hemisphere temperate ecosystems

Over two-thirds of terrestrial carbon is stored belowground and a significant amount of atmospheric CO₂ is respired by roots and microbes in soils. For this analysis, soil respiration (Rs) data were assembled from 31 AmeriFlux and CarboEurope sites representing deciduous broadleaf, evergreen needleleaf, grasslands, mixed deciduous/evergreen and woodland/savanna ecosystem types. Lowest to highest rates of soil respiration averaged over the growing season were grassland and woodland/savanna < deciduous broadleaf forests < evergreen needleleaf, mixed deciduous/evergreen forests with growing season soil respiration significantly different between forested and non-forested biomes (p < 0.001). Timing of peak respiration rates during the growing season varied from March/April in grasslands to July–September for all other biomes. Biomes with overall strongest relationship between soil respiration and soil temperature were from the deciduous and mixed forests (R² ≥ 0.65). Maximum soil respiration was weakly related to maximum fine root biomass (R² = 0.28) and positively related to the previous years’ annual litterfall (R² = 0.46). Published rates of annual soil respiration were linearly related to LAI and fine root carbon (R² = 0.48, 0.47), as well as net primary production (NPP) (R² = 0.44). At 10 sites, maximum growing season Rs was weakly correlated with annual GPP estimated from eddy covariance towersites (R² = 0.29; p < 0.05), and annual soil respiration and total growing season Rs were not correlated with annual GPP (p > 0.1). Yet, previous studies indicate correlations on shorter time scales within site (e.g., weekly, monthly). Estimates of annual GPP from the Biome-BGC model were strongly correlated with observed annual estimates of soil respiration for six sites (R² = 0.84; p < 0.01). Correlations from observations of Rs with NPP, LAI, fine root biomass and litterfall relate above and belowground inputs to labile pools that are available for decomposition. Our results suggest that simple empirical relationships with temperature and/or moisture that may be robust at individual sites may not be adequate to characterize soil CO₂ effluxes across space and time, agreeing with other multi-site studies. Information is needed on the timing and phenological controls of substrate availability (e.g., fine roots, LAI) and inputs (e.g., root turnover, litterfall) to improve our ability to accurately quantify the relationships between soil CO₂ effluxes and carbon substrate storage.For this study, these authors received significant contributions from: M. Aubinet, D. Baldocchi, C. Bernhofer, P. Bolstad, A. Bosc, J.L. Campbell, Y. Cheng, J. Curiel Yuste, P. Curtis, E.A. Davidson, D. Epron, A. Granier, T. Grünwald, D. Hollinger, I.A. Janssens, B. Longdoz, D. Loustau, J. Martin, R. Monson, W. Oechel, J. Pippen, F. Ponti, R. Ryel, K. Savage, L. Scott-Denton, J.-A. Subke, J. Tang, J. Tenhunen, V. Turcu, C. S. Vogel.     

Assessment of ecosystem CO2 efflux and its components in a Pinus canariensis forest at the treeline

We estimated respiratory fluxes in a treeline-associated Pinus canariensis forest in Tenerife, Canary Islands, an ecotone with strong seasonal changes in soil water availability. CO₂ efflux rates from the foliage, above ground woody tissue and the soil were measured by chamber techniques. Site-specific models obtained from these chamber measurements were then combined with half-hourly measurements of canopy, stem and soil temperature as well as with soil water potential, leaf area and stem sapwood volume data for scaling up component-specific CO₂ efflux to ecosystem respiration (R _ECO). Integrated over an entire year R _ECO was 550 g C m^?2 ground surface area (average of 2008 and 2009) and comprised the following component fluxes: 57 % from the soil, 10 % from above ground woody tissue and 33 % from the foliage. Between year differences in R _ECO and its components were <3 %. R _ECO varied markedly throughout an entire year generally following the seasonal trends in temperature during most of the year. During the dry summer, however, R _ECO was significantly reduced due to limited soil water availability in the rooting horizon. Thus, in treeline-associated forests under predicted scenarios of increasing aridity in Mediterranean regions, it is likely that there will be a shift in the contribution of CO₂ efflux from the soil (R _S), the foliage (R _F) and above ground woody tissue (R _W) to R _ECO from predominately below ground to increasingly above ground. Such changes should be taken into account for predicting the response of treeline forests to a changing climate at their upper distribution limit.     

Towards physiologically meaningful water‐use efficiency estimates from eddy covariance data

Intrinsic water‐use efficiency (iWUE) characterizes the physiological control on the simultaneous exchange of water and carbon dioxide in terrestrial ecosystems. Knowledge of iWUE is commonly gained from leaf‐level gas exchange measurements, which are inevitably restricted in their spatial and temporal coverage. Flux measurements based on the eddy covariance (EC) technique can overcome these limitations, as they provide continuous and long‐term records of carbon and water fluxes at the ecosystem scale. However, vegetation gas exchange parameters derived from EC data are subject to scale‐dependent and method‐specific uncertainties that compromise their ecophysiological interpretation as well as their comparability among ecosystems and across spatial scales. Here, we use estimates of canopy conductance and gross primary productivity (GPP) derived from EC data to calculate a measure of iWUE (G₁, “stomatal slope”) at the ecosystem level at six sites comprising tropical, Mediterranean, temperate, and boreal forests. We assess the following six mechanisms potentially causing discrepancies between leaf and ecosystem‐level estimates of G₁: (i) non‐transpirational water fluxes; (ii) aerodynamic conductance; (iii) meteorological deviations between measurement height and canopy surface; (iv) energy balance non‐closure; (v) uncertainties in net ecosystem exchange partitioning; and (vi) physiological within‐canopy gradients. Our results demonstrate that an unclosed energy balance caused the largest uncertainties, in particular if it was associated with erroneous latent heat flux estimates. The effect of aerodynamic conductance on G₁ was sufficiently captured with a simple representation. G₁ was found to be less sensitive to meteorological deviations between canopy surface and measurement height and, given that data are appropriately filtered, to non‐transpirational water fluxes. Uncertainties in the derived GPP and physiological within‐canopy gradients and their implications for parameter estimates at leaf and ecosystem level are discussed. Our results highlight the importance of adequately considering the sources of uncertainty outlined here when EC‐derived water‐use efficiency is interpreted in an ecophysiological context.     

Multiscale modeling of spring phenology across Deciduous Forests in the Eastern United States

Phenological events, such as bud burst, are strongly linked to ecosystem processes in temperate deciduous forests. However, the exact nature and magnitude of how seasonal and interannual variation in air temperatures influence phenology is poorly understood, and model‐based phenology representations fail to capture local‐ to regional‐scale variability arising from differences in species composition. In this paper, we use a combination of surface meteorological data, species composition maps, remote sensing, and ground‐based observations to estimate models that better represent how community‐level species composition affects the phenological response of deciduous broadleaf forests to climate forcing at spatial scales that are typically used in ecosystem models. Using time series of canopy greenness from repeat digital photography, citizen science data from the USA National Phenology Network, and satellite remote sensing‐based observations of phenology, we estimated and tested models that predict the timing of spring leaf emergence across five different deciduous broadleaf forest types in the eastern United States. Specifically, we evaluated two different approaches: (i) using species‐specific models in combination with species composition information to ‘upscale’ model predictions and (ii) using repeat digital photography of forest canopies that observe and integrate the phenological behavior of multiple representative species at each camera site to calibrate a single model for all deciduous broadleaf forests. Our results demonstrate variability in cumulative forcing requirements and photoperiod cues across species and forest types, and show how community composition influences phenological dynamics over large areas. At the same time, the response of different species to spatial and interannual variation in weather is, under the current climate regime, sufficiently similar that the generic deciduous forest model based on repeat digital photography performed comparably to the upscaled species‐specific models. More generally, results from this analysis demonstrate how in situ observation networks and remote sensing data can be used to synergistically calibrate and assess regional parameterizations of phenology in models.     

Seasonal and annual variation of carbon exchange in an evergreen Mediterranean forest in southern France

We present 9 years of eddy covariance measurements made over an evergreen Mediterranean forest in southern France. The goal of this study was to quantify the different components of the carbon (C) cycle, gross primary production (GPP) and ecosystem respiration (R_eco), and to assess the effects of climatic variables on these fluxes and on the net ecosystem exchange of carbon dioxide. The Puéchabon forest acted as a net C sink of ?254 g C m^?2 yr^?1, with a GPP of 1275 g C m^?2 yr^?1 and a R_eco of 1021 g C m^?2 yr^?1. On average, 83% of the net annual C sink occurred between March and June. The effects of exceptional events such the insect‐induced partial canopy defoliation that occurred in spring 2005, and the spring droughts of 2005 and 2006 are discussed. A high interannual variability of ecosystem C fluxes during summer and autumn was observed but the resulting effect on the annual net C budget was moderate. Increased severity and/or duration of summer drought under climate change do not appear to have the potential to negatively impact the average C budget of this ecosystem. On the contrary, factors affecting ecosystem functioning (drought and/or defoliation) during March–June period may reduce dramatically the annual C balance of evergreen Mediterranean forests.     

Seasonal and interannual variations in carbon dioxide exchange over a cropland in the North China Plain

In China, croplands account for a relatively large form of vegetation cover. Quantifying carbon dioxide exchange and understanding the environmental controls on carbon fluxes over croplands are critical in understanding regional carbon budgets and ecosystem behaviors. In this study, the net ecosystem exchange (NEE) at a winter wheat/summer maize rotation cropping site, representative of the main cropping system in the North China Plain, was continuously measured using the eddy covariance technique from 2005 to 2009. In order to interpret the abiotic factors regulating NEE, NEE was partitioned into gross primary production (GPP) and ecosystem respiration (R_eco). Daytime R_eco was extrapolated from the relationship between nighttime NEE and soil temperature under high turbulent conditions. GPP was then estimated by subtracting daytime NEE from the daytime estimates of R_eco. Results show that the seasonal patterns of the temperature responses of R_eco and light‐response parameters are closely related to the crop phenology. Daily R_eco was highly dependent on both daily GPP and air temperature. Interannual variability showed that GPP and R_eco were mainly controlled by temperature. Water availability also exerted a limit on R_eco. The annual NEE was ?585 and ?533 g C m^?2 for two seasons of 2006–2007 and 2007–2008, respectively, and the wheat field absorbed more carbon than the maize field. Thus, we concluded that this cropland was a strong carbon sink. However, when the grain harvest was taken into account, the wheat field was diminished into a weak carbon sink, whereas the maize field was converted into a weak carbon source. The observations showed that severe drought occurring during winter did not reduce wheat yield (or integrated NEE) when sufficient irrigation was carried out during spring.     

Impacts of Hurricane Frances on Florida scrub-oak ecosystem processes: defoliation, net CO2 exchange and interactions with elevated CO2

Hurricane disturbances have profound impacts on ecosystem structure and function, yet their effects on ecosystem CO₂ exchange have not been reported. In September 2004, our research site on a fire‐regenerated scrub‐oak ecosystem in central Florida was struck by Hurricane Frances with sustained winds of 113 km h⁻¹ and wind gusts as high as 152 km h⁻¹. We quantified the hurricane damage on this ecosystem resulting from defoliation: we measured net ecosystem CO₂ exchange, the damage and recovery of leaf area, and determined whether growth in elevated carbon dioxide concentration in the atmosphere (C_a) altered this disturbance. The hurricane decreased leaf area index (LAI) by 21%, which was equal to 60% of seasonal variation in canopy growth during the previous 3 years, but stem damage was negligible. The reduction in LAI led to a 22% decline in gross primary production (GPP) and a 25% decline in ecosystem respiration (R_e). The compensatory declines in GPP and R_e resulted in no significant change in net ecosystem production (NEP). Refoliation began within a month after the hurricane, although this period was out of phase with the regular foliation period, and recovered 20% of the defoliation loss within 2.5 months. Full recovery of LAI, ecosystem CO₂ assimilation, and ecosystem respiration did not occur until the next growing season. Plants exposed to elevated C_a did not sustain greater damage, nor did they recover faster than plants grown under ambient C_a. Thus, our results indicate that hurricanes capable of causing significant defoliation with negligible damage to stems have negligible effects on NEP under current or future CO₂‐enriched environment.