Contemporary carbon balance and late Holocene carbon accumulation in a northern peatland

Northern peatlands contain up to 25% of the world's soil carbon (C) and have an estimated annual exchange of CO₂‐C with the atmosphere of 0.1–0.5 Pg yr⁻¹ and of CH₄‐C of 10–25 Tg yr⁻¹. Despite this overall importance to the global C cycle, there have been few, if any, complete multiyear annual C balances for these ecosystems. We report a 6‐year balance computed from continuous net ecosystem CO₂ exchange (NEE), regular instantaneous measurements of methane (CH₄) emissions, and export of dissolved organic C (DOC) from a northern ombrotrophic bog. From these observations, we have constructed complete seasonal and annual C balances, examined their seasonal and interannual variability, and compared the mean 6‐year contemporary C exchange with the apparent C accumulation for the last 3000 years obtained from C density and age‐depth profiles from two peat cores. The 6‐year mean NEE‐C and CH₄‐C exchange, and net DOC loss are −40.2±40.5 (±1 SD), 3.7±0.5, and 14.9±3.1 g m⁻² yr⁻¹, giving a 6‐year mean balance of −21.5±39.0 g m⁻² yr⁻¹ (where positive exchange is a loss of C from the ecosystem). NEE had the largest magnitude and variability of the components of the C balance, but DOC and CH₄ had similar proportional variabilities and their inclusion is essential to resolve the C balance. There are large interseasonal and interannual ranges to the exchanges due to variations in climatic conditions. We estimate from the largest and smallest seasonal exchanges, quasi‐maximum limits of the annual C balance between 50 and −105 g m⁻² yr⁻¹. The net C accumulation rate obtained from the two peatland cores for the interval 400–3000 bp (samples from the anoxic layer only) were 21.9±2.8 and 14.0±37.6 g m⁻² yr⁻¹, which are not significantly different from the 6‐year mean contemporary exchange.     

Carbon cycle responses of semi‐arid ecosystems to positive asymmetry in rainfall

Recent evidence shows that warm semi‐arid ecosystems are playing a disproportionate role in the interannual variability and greening trend of the global carbon cycle given their mean lower productivity when compared with other biomes (Ahlström et al. 2015 Science, 348, 895). Using multiple observations (land‐atmosphere fluxes, biomass, streamflow and remotely sensed vegetation cover) and two state‐of‐the‐art biospheric models, we show that climate variability and extremes lead to positive or negative responses in the biosphere, depending on vegetation type. We find Australia to be a global hot spot for variability, with semi‐arid ecosystems in that country exhibiting increased carbon uptake due to both asymmetry in the interannual distribution of rainfall (extrinsic forcing), and asymmetry in the response of gross primary production (GPP) to rainfall change (intrinsic response). The latter is attributable to the pulse‐response behaviour of the drought‐adapted biota of these systems, a response that is estimated to be as much as half of that from the CO₂ fertilization effect during 1990–2013. Mesic ecosystems, lacking drought‐adapted species, did not show an intrinsic asymmetric response. Our findings suggest that a future more variable climate will induce large but contrasting ecosystem responses, differing among biomes globally, independent of changes in mean precipitation alone. The most significant changes are occurring in the extensive arid and semi‐arid regions, and we suggest that the reported increased carbon uptake in response to asymmetric responses might be contributing to the observed greening trends there.     

FLUXNET and modelling the global carbon cycle

Measurements of the net CO₂ flux between terrestrial ecosystems and the atmosphere using the eddy covariance technique have the potential to underpin our interpretation of regional CO₂ source–sink patterns, CO₂ flux responses to forcings, and predictions of the future terrestrial C balance. Information contained in FLUXNET eddy covariance data has multiple uses for the development and application of global carbon models, including evaluation/validation, calibration, process parameterization, and data assimilation. This paper reviews examples of these uses, compares global estimates of the dynamics of the global carbon cycle, and suggests ways of improving the utility of such data for global carbon modelling. Net ecosystem exchange of CO₂ (NEE) predicted by different terrestrial biosphere models compares favourably with FLUXNET observations at diurnal and seasonal timescales. However, complete model validation, particularly over the full annual cycle, requires information on the balance between assimilation and decomposition processes, information not readily available for most FLUXNET sites. Site history, when known, can greatly help constrain the model‐data comparison. Flux measurements made over four vegetation types were used to calibrate the land‐surface scheme of the Goddard Institute for Space Studies global climate model, significantly improving simulated climate and demonstrating the utility of diurnal FLUXNET data for climate modelling. Land‐surface temperatures in many regions cool due to higher canopy conductances and latent heat fluxes, and the spatial distribution of CO₂ uptake provides a significant additional constraint on the realism of simulated surface fluxes. FLUXNET data are used to calibrate a global production efficiency model (PEM). This model is forced by satellite‐measured absorbed radiation and suggests that global net primary production (NPP) increased 6.2% over 1982–1999. Good agreement is found between global trends in NPP estimated by the PEM and a dynamic global vegetation model (DGVM), and between the DGVM and estimates of global NEE derived from a global inversion of atmospheric CO₂ measurements. Combining the PEM, DGVM, and inversion results suggests that CO₂ fertilization is playing a major role in current increases in NPP, with lesser impacts from increasing N deposition and growing season length. Both the PEM and the inversion identify the Amazon basin as a key region for the current net terrestrial CO₂ uptake (i.e. 33% of global NEE), as well as its interannual variability. The inversion's global NEE estimate of −1.2 Pg [C] yr⁻¹ for 1982–1995 is compatible with the PEM‐ and DGVM‐predicted trends in NPP. There is, thus, a convergence in understanding derived from process‐based models, remote‐sensing‐based observations, and inversion of atmospheric data. Future advances in field measurement techniques, including eddy covariance (particularly concerning the problem of night‐time fluxes in dense canopies and of advection or flow distortion over complex terrain), will result in improved constraints on land‐atmosphere CO₂ fluxes and the rigorous attribution of mechanisms to the current terrestrial net CO₂ uptake and its spatial and temporal heterogeneity. Global ecosystem models play a fundamental role in linking information derived from FLUXNET measurements to atmospheric CO₂ variability. A number of recommendations concerning FLUXNET data are made, including a request for more comprehensive site data (particularly historical information), more measurements in undisturbed ecosystems, and the systematic provision of error estimates. The greatest value of current FLUXNET data for global carbon cycle modelling is in evaluating process representations, rather than in providing an unbiased estimate of net CO₂ exchange.     

CO2 flux from soil in pastures and forests in southwestern Amazonia

Stocks of carbon in Amazonian forest biomass and soils have received considerable research attention because of their potential as sources and sinks of atmospheric CO₂. Fluxes of CO₂ from soil to the atmosphere, on the other hand, have not been addressed comprehensively in regard to temporal and spatial variations and to land cover change, and have been measured directly only in a few locations in Amazonia. Considerable variation exists across the Amazon Basin in soil properties, climate, and management practices in forests and cattle pastures that might affect soil CO₂ fluxes. Here we report soil CO₂ fluxes from an area of rapid deforestation in the southwestern Amazonian state of Acre. Specifically we addressed (1) the seasonal variation of soil CO₂ fluxes, soil moisture, and soil temperature; (2) the effects of land cover (pastures, mature, and secondary forests) on these fluxes; (3) annual estimates of soil respiration; and (4) the relative contributions of grass‐derived and forest‐derived C as indicated by δ¹³CO₂. Fluxes were greatest during the wet season and declined during the dry season in all land covers. Soil respiration was significantly correlated with soil water‐filled pore space but not correlated with temperature. Annual fluxes were higher in pastures compared with mature and secondary forests, and some of the pastures also had higher soil C stocks. The δ¹³C of CO₂ respired in pasture soils showed that high respiration rates in pastures were derived almost entirely from grass root respiration and decomposition of grass residues. These results indicate that the pastures are very productive and that the larger flux of C cycling through pasture soils compared with forest soils is probably due to greater allocation of C belowground. Secondary forests had soil respiration rates similar to mature forests, and there was no correlation between soil respiration and either forest age or forest biomass. Hence, belowground allocation of C does not appear to be directly related to the stature of vegetation in this region. Variation in seasonal and annual rates of soil respiration of these forests and pastures is more indicative of flux of C through the soil rather than major net changes in ecosystem C stocks.     

Aquatic carbon fluxes dampen the overall variation of net ecosystem productivity in the Amazon basin: An analysis of the interannual variability in the boundless carbon cycle

The river–floodplain network plays an important role in the carbon (C) cycle of the Amazon basin, as it transports and processes a significant fraction of the C fixed by terrestrial vegetation, most of which evades as CO₂ from rivers and floodplains back to the atmosphere. There is empirical evidence that exceptionally dry or wet years have an impact on the net C balance in the Amazon. While seasonal and interannual variations in hydrology have a direct impact on the amounts of C transferred through the river–floodplain system, it is not known how far the variation of these fluxes affects the overall Amazon C balance. Here, we introduce a new wetland forcing file for the ORCHILEAK model, which improves the representation of floodplain dynamics and allows us to closely reproduce data‐driven estimates of net C exports through the river–floodplain network. Based on this new wetland forcing and two climate forcing datasets, we show that across the Amazon, the percentage of net primary productivity lost to the river–floodplain system is highly variable at the interannual timescale, and wet years fuel aquatic CO₂ evasion. However, at the same time overall net ecosystem productivity (NEP) and C sequestration are highest during wet years, partly due to reduced decomposition rates in water‐logged floodplain soils. It is years with the lowest discharge and floodplain inundation, often associated with El Nino events, that have the lowest NEP and the highest total (terrestrial plus aquatic) CO₂ emissions back to atmosphere. Furthermore, we find that aquatic C fluxes display greater variation than terrestrial C fluxes, and that this variation significantly dampens the interannual variability in NEP of the Amazon basin. These results call for a more integrative view of the C fluxes through the vegetation‐soil‐river‐floodplain continuum, which directly places aquatic C fluxes into the overall C budget of the Amazon basin.     

Carbon emissions from fires in tropical and subtropical ecosystems

Global carbon emissions from fires are difficult to quantify and have the potential to influence interannual variability and long‐term trends in atmospheric CO₂ concentrations. We used 4 years of Tropical Rainfall Measuring Mission (TRMM) Visible and Infrared Scanner (VIRS) satellite data and a biogeochemical model to assess spatial and temporal variability of carbon emissions from tropical fires. The TRMM satellite data extended between 38°N and 38°S and covered the period from 1998 to 2001. A relationship between TRMM fire counts and burned area was derived using estimates of burned area from other satellite fire products in Africa and Australia and reported burned areas from the United States. We modified the Carnegie‐Ames‐Stanford‐Approach (CASA) biogeochemical model to account for both direct combustion losses and the decomposition from fire‐induced mortality, using both TRMM and Sea‐viewing Wide Field of view Sensor (SeaWiFS) satellite data as model drivers. Over the 1998–2001 period, we estimated that the sum of carbon emissions from tropical fires and fuel wood use was 2.6 Pg C yr^?1. An additional flux of 1.2 Pg C yr^?1 was released indirectly, as a result of decomposition of vegetation killed by fire but not combusted. The sum of direct and indirect carbon losses from fires represented 9% of tropical and subtropical net primary production (NPP). We found that including fire processes in the tropics substantially alters the seasonal cycle of net biome production by shifting carbon losses to months with low soil moisture and low rates of soil microbial respiration. Consequently, accounting for fires increases growing season net flux by ～12% between 38°N and 38°S, with the greatest effect occurring in highly productive savanna regions.     

CO2 balance of boreal, temperate, and tropical forests derived from a global database

Terrestrial ecosystems sequester 2.1 Pg of atmospheric carbon annually. A large amount of the terrestrial sink is realized by forests. However, considerable uncertainties remain regarding the fate of this carbon over both short and long timescales. Relevant data to address these uncertainties are being collected at many sites around the world, but syntheses of these data are still sparse. To facilitate future synthesis activities, we have assembled a comprehensive global database for forest ecosystems, which includes carbon budget variables (fluxes and stocks), ecosystem traits (e.g. leaf area index, age), as well as ancillary site information such as management regime, climate, and soil characteristics. This publicly available database can be used to quantify global, regional or biome‐specific carbon budgets; to re‐examine established relationships; to test emerging hypotheses about ecosystem functioning [e.g. a constant net ecosystem production (NEP) to gross primary production (GPP) ratio]; and as benchmarks for model evaluations. In this paper, we present the first analysis of this database. We discuss the climatic influences on GPP, net primary production (NPP) and NEP and present the CO₂ balances for boreal, temperate, and tropical forest biomes based on micrometeorological, ecophysiological, and biometric flux and inventory estimates. Globally, GPP of forests benefited from higher temperatures and precipitation whereas NPP saturated above either a threshold of 1500 mm precipitation or a mean annual temperature of 10 °C. The global pattern in NEP was insensitive to climate and is hypothesized to be mainly determined by nonclimatic conditions such as successional stage, management, site history, and site disturbance. In all biomes, closing the CO₂ balance required the introduction of substantial biome‐specific closure terms. Nonclosure was taken as an indication that respiratory processes, advection, and non‐CO₂ carbon fluxes are not presently being adequately accounted for.     

Carbon respired by terrestrial ecosystems – recent progress and challenges

Net ecosystem production is the residual of two much larger fluxes: photosynthesis and respiration. While photosynthesis is a single process with a well‐established theoretical underpinning, respiration integrates the variety of plant and microbial processes by which CO₂ returns from ecosystems to the atmosphere. Limits to current capacity for predicting ecosystem respiration fluxes across biomes or years result from the mismatch between what is usually measured – bulk CO₂ fluxes – and what process‐based models can predict – fluxes of CO₂ from plant (autotrophic) or microbial (heterotrophic) respiration. Papers in this Thematic Issue and in the recent literature, document advances in methods for separating respiration into autotrophic and heterotrophic components using three approaches: (1) continuous measurements of CO₂ fluxes and assimilation of these data into process‐based models; (2) application of isotope measurements, particularly radiocarbon; and (3) manipulation experiments. They highlight the role of allocation of C fixed by plants to respiration, storage, growth or transfer to other organisms as a control of seasonal and interannual variability in soil respiration and the oxidation state of C in the terrestrial biosphere. A second theme is the potential for comparing C isotope signatures in organic matter, CO₂ evolved in incubations and microbial biomarkers to elucidate the pathways (respiration, recycling, or transformation) of C during decomposition. Together, these factors determine the continuum of timescales over which C is returned to the atmosphere by respiration and enable testing of theories of plant and microbial respiration that go beyond empirical models and allow predictions of future respiration responses to future change in climate, pollution and land use.     

Terrestrial biosphere model performance for inter‐annual variability of land‐atmosphere CO2 exchange

Interannual variability in biosphere‐atmosphere exchange of CO₂ is driven by a diverse range of biotic and abiotic factors. Replicating this variability thus represents the ‘acid test’ for terrestrial biosphere models. Although such models are commonly used to project responses to both normal and anomalous variability in climate, they are rarely tested explicitly against inter‐annual variability in observations. Herein, using standardized data from the North American Carbon Program, we assess the performance of 16 terrestrial biosphere models and 3 remote sensing products against long‐term measurements of biosphere‐atmosphere CO₂ exchange made with eddy‐covariance flux towers at 11 forested sites in North America. Instead of focusing on model‐data agreement we take a systematic, variability‐oriented approach and show that although the models tend to reproduce the mean magnitude of the observed annual flux variability, they fail to reproduce the timing. Large biases in modeled annual means are evident for all models. Observed interannual variability is found to commonly be on the order of magnitude of the mean fluxes. None of the models consistently reproduce observed interannual variability within measurement uncertainty. Underrepresentation of variability in spring phenology, soil thaw and snowpack melting, and difficulties in reproducing the lagged response to extreme climatic events are identified as systematic errors, common to all models included in this study.     

Modeling the effects of snowpack on heterotrophic respiration across northern temperate and high latitude regions: Comparison with measurements of atmospheric carbon dioxide in high latitudes

Simulations by global terrestrial biogeochemical models (TBMs) consistently underestimate the concentration of atmospheric carbon dioxide (CO₂ at high latitude monitoring stations during the non-growing season. We hypothesized that heterotrophic respiration is underestimated during the nongrowing season primarily because TBMs do not generally consider the insulative effects of snowpack on soil temperature. To evaluate this hypothesis, we compared the performance of baseline and modified versions of three TBMs in simulating the seasonal cycle of atmospheric CO₂ at high latitude CO₂ monitoring stations; the modified version maintained soil temperature at 0 °C when modeled snowpack was present. The three TBMs include the Carnegie-Ames-Stanford Approach (CASA), Century, and the Terrestrial Ecosystem Model (TEM). In comparison with the baseline simulation of each model, the snowpack simulations caused higher releases of CO₂ between November and March and greater uptake of CO₂ between June and August for latitudes north of 30° N. We coupled the monthly estimates of CO₂ exchange, the seasonal carbon dioxide flux fields generated by the HAMOCC3 seasonal ocean carbon cycle model, and fossil fuel source fields derived from standard sources to the three-dimensional atmospheric transport model TM2 forced by observed winds to simulate the seasonal cycle of atmospheric CO₂ at each of seven high latitude monitoring stations. In comparison to the CO₂ concentrations simulated with the baseline fluxes of each TBM, concentrations simulated using the snowpack fluxes are generally in better agreement with observed concentrations between August and March at each of the monitoring stations. Thus, representation of the insulative effects of snowpack in TBMs generally improves simulation of atmospheric CO₂ concentrations in high latitudes during both the late growing season and nongrowing season. These simulations highlight the global importance of biogeochemical processes during the nongrowing season in estimating carbon balance of ecosystems in northern high and temperate latitudes.     

Temperature and biomass influences on interannual changes in CO2 exchange in an alpine meadow on the Qinghai-Tibetan Plateau

Three years of eddy covariance measurements were used to characterize the seasonal and interannual variability of the CO₂ fluxes above an alpine meadow (3250 m a.s.l.) on the Qinghai‐Tibetan Plateau, China. This alpine meadow was a weak sink for atmospheric CO₂, with a net ecosystem production (NEP) of 78.5, 91.7, and 192.5 g C m^?2 yr^?1 in 2002, 2003, and 2004, respectively. The prominent, high NEP in 2004 resulted from the combination of high gross primary production (GPP) and low ecosystem respiration (R_e) during the growing season. The period of net absorption of CO₂ in 2004, 179 days, was 10 days longer than that in 2002 and 5 days longer than that in 2003. Moreover, the date on which the mean air temperature first exceeded 5.0°C was 10 days earlier in 2004 (DOY110) than in 2002 or 2003. This date agrees well with that on which the green aboveground biomass (Green AGB) started to increase. The relationship between light‐use efficiency and Green AGB was similar among the three years. In 2002, however, earlier senescence possibly caused low autumn GPP, and thus the annual NEP, to be lower. The low summertime R_e in 2004 was apparently caused by lower soil temperatures and the relatively lower temperature dependence of R_e in comparison with the other years. These results suggest that (1) the Qinghai‐Tibetan Plateau plays a potentially significant role in global carbon sequestration, because alpine meadow covers about one‐third of this vast plateau, and (2) the annual NEP in the alpine meadow was comprehensively controlled by the temperature environment, including its effect on biomass growth.     

Biogeographic variation in temperature sensitivity of decomposition in forest soils

Determining soil carbon (C) responses to rising temperature is critical for projections of the feedbacks between terrestrial ecosystems, C cycle, and climate change. However, the direction and magnitude of this feedback remain highly uncertain due largely to our limited understanding of the spatial heterogeneity of soil C decomposition and its temperature sensitivity. Here we quantified C decomposition and its response to temperature change with an incubation study of soils from 203 sites across tropical to boreal forests in China spanning a wide range of latitudes (18°16′ to 51°37′N) and longitudes (81°01′ to 129°28′E). Mean annual temperature (MAT) and mean annual precipitation primarily explained the biogeographic variation in the decomposition rate and temperature sensitivity of soils: soil C decomposition rate decreased from warm and wet forests to cold and dry forests, while Q_10‐MAT (standardized to the MAT of each site) values displayed the opposite pattern. In contrast, biological factors (i.e. plant productivity and soil bacterial diversity) and soil factors (e.g. clay, pH, and C availability of microbial biomass C and dissolved organic C) played relatively small roles in the biogeographic patterns. Moreover, no significant relationship was found between Q_10‐MAT and soil C quality, challenging the current C quality–temperature hypothesis. Using a single, fixed Q_10‐MAT value (the mean across all forests), as is usually done in model predictions, would bias the estimated soil CO₂ emissions at a temperature increase of 3.0°C. This would lead to overestimation of emissions in warm biomes, underestimation in cold biomes, and likely significant overestimation of overall C release from soil to the atmosphere. Our results highlight that climate‐related biogeographic variation in soil C responses to temperature needs to be included in next‐generation C cycle models to improve predictions of C‐climate feedbacks.     

Process-level controls on CO<Subscript>2</Subscript> fluxes from a seasonally snow-covered subalpine meadow soil,Niwot Ridge,Colorado

Fluxes of CO₂ during the snow-covered season contribute to annual carbon budgets, but our understanding of the mechanisms controlling the seasonal pattern and magnitude of carbon emissions in seasonally snow-covered areas is still developing. In a subalpine meadow on Niwot Ridge, Colorado, soil CO₂ fluxes were quantified with the gradient method through the snowpack in winter 2006 and 2007 and with chamber measurements during summer 2007. The CO₂ fluxes of 0.71 μmol m⁻² s⁻¹ in 2006 and 0.86 μmol m⁻² s⁻¹ in 2007 are among the highest reported for snow-covered ecosystems in the literature. These fluxes resulted in 156 and 189 g C m⁻² emitted over the winter, ~30% of the annual soil CO₂ efflux at this site. In general, the CO₂ flux increased during the winter as soil moisture increased. A conceptual model was developed with distinct snow cover zones to describe this as well as the three other reported temporal patterns in CO₂ flux from seasonally snow-covered soils. As snow depth and duration increase, the factor controlling the CO₂ flux shifts from freeze–thaw cycles (zone I) to soil temperature (zone II) to soil moisture (zone III) to carbon availability (zone IV). The temporal pattern in CO₂ flux in each zone changes from periodic pulses of CO₂ during thaw events (zone I), to CO₂ fluxes reaching a minimum when soil temperatures are lowest in mid-winter (zone II), to CO₂ fluxes increasing gradually as soil moisture increases (zone III), to CO₂ fluxes decreasing as available carbon is consumed. This model predicts that interannual variability in snow cover or directional shifts in climate may result in dramatically different seasonal patterns of CO₂ flux from seasonally snow-covered soils.     

Evaluation of terrestrial carbon cycle models for their response to climate variability and to CO2 trends

The purpose of this study was to evaluate 10 process‐based terrestrial biosphere models that were used for the IPCC fifth Assessment Report. The simulated gross primary productivity (GPP) is compared with flux‐tower‐based estimates by Jung et al. [Journal of Geophysical Research 116 (2011) G00J07] (JU11). The net primary productivity (NPP) apparent sensitivity to climate variability and atmospheric CO₂ trends is diagnosed from each model output, using statistical functions. The temperature sensitivity is compared against ecosystem field warming experiments results. The CO₂ sensitivity of NPP is compared to the results from four Free‐Air CO₂ Enrichment (FACE) experiments. The simulated global net biome productivity (NBP) is compared with the residual land sink (RLS) of the global carbon budget from Friedlingstein et al. [Nature Geoscience 3 (2010) 811] (FR10). We found that models produce a higher GPP (133 ± 15 Pg C yr^?1) than JU11 (118 ± 6 Pg C yr^?1). In response to rising atmospheric CO₂ concentration, modeled NPP increases on average by 16% (5–20%) per 100 ppm, a slightly larger apparent sensitivity of NPP to CO₂ than that measured at the FACE experiment locations (13% per 100 ppm). Global NBP differs markedly among individual models, although the mean value of 2.0 ± 0.8 Pg C yr^?1 is remarkably close to the mean value of RLS (2.1 ± 1.2 Pg C yr^?1). The interannual variability in modeled NBP is significantly correlated with that of RLS for the period 1980–2009. Both model‐to‐model and interannual variation in model GPP is larger than that in model NBP due to the strong coupling causing a positive correlation between ecosystem respiration and GPP in the model. The average linear regression slope of global NBP vs. temperature across the 10 models is ?3.0 ± 1.5 Pg C yr^?1 °C^?1, within the uncertainty of what derived from RLS (?3.9 ± 1.1 Pg C yr^?1 °C^?1). However, 9 of 10 models overestimate the regression slope of NBP vs. precipitation, compared with the slope of the observed RLS vs. precipitation. With most models lacking processes that control GPP and NBP in addition to CO₂ and climate, the agreement between modeled and observation‐based GPP and NBP can be fortuitous. Carbon–nitrogen interactions (only separable in one model) significantly influence the simulated response of carbon cycle to temperature and atmospheric CO₂ concentration, suggesting that nutrients limitations should be included in the next generation of terrestrial biosphere models.     

Terrestrial nitrogen cycle simulation with a dynamic global vegetation model

A global scale Dynamic Nitrogen scheme (DyN) has been developed and incorporated into the Lund–Posdam–Jena (LPJ) dynamic global vegetation model (DGVM). The DyN is a comprehensive process‐based model of the cycling of N through and within terrestrial ecosystems, with fully interactive coupling to vegetation and C dynamics. The model represents the uptake, allocation and turnover of N in plants, and soil N transformations including mineralization, N₂ fixation, nitrification and denitrification, NH₃ volatilization, N leaching, and N₂, N₂O and NO production and emission. Modelled global patterns of site‐scale nitrogen fluxes and reservoirs are highly correlated to observations reported from different biomes. The simulation of site‐scale net primary production and soil carbon content was improved relative to the original LPJ, which lacked an interactive N cycle, especially in the temporal and boreal regions. Annual N uptake by global natural vegetation was simulated as 1.084 Pg N yr⁻¹, with lowest values <1 g N m⁻² yr⁻¹ (polar desert) and highest values in the range 24–36.5 g N m⁻² yr⁻¹ (tropical forests). Simulated global patterns of annual N uptake are consistent with previous model results by Melillo et al. The model estimates global total nitrogen storage potentials in vegetation (5.3 Pg N), litter (4.6 Pg N) and soil (≥67 Pg as organic N and 0.94 Pg as inorganic N). Simulated global patterns of soil N storage are consistent with the analysis by Post et al. although total simulated N storage is less. Deserts were simulated to store 460 Tg N (up to 0.262 kg N m⁻²) as NO₃⁻, contributing 80% of the global total NO₃⁻ inventory of 580 Tg N. This model result is in agreement with the findings of a large NO₃⁻ pool beneath deserts. Globally, inorganic soil N is a small reservoir, comprising only 1.6% of the global soil N content to 1.5 m soil depth, but the ratio has a very high spatial variability and in hot desert regions, inorganic NO₃⁻ is estimated to be the dominant form of stored N in the soil.     

Greenhouse gas fluxes from tropical peatlands in south‐east Asia

The lowland peatlands of south‐east Asia represent an immense reservoir of fossil carbon and are reportedly responsible for 30% of the global carbon dioxide (CO₂) emissions from Land Use, Land Use Change and Forestry. This paper provides a review and meta‐analysis of available literature on greenhouse gas fluxes from tropical peat soils in south‐east Asia. As in other parts of the world, water level is the main control on greenhouse gas fluxes from south‐east Asian peat soils. Based on subsidence data we calculate emissions of at least 900 g CO₂ m^?2 a^?1 (～250 g C m^?2 a^?1) for each 10 cm of additional drainage depth. This is a conservative estimate as the role of oxidation in subsidence and the increased bulk density of the uppermost drained peat layers are yet insufficiently quantified. The majority of published CO₂ flux measurements from south‐east Asian peat soils concerns undifferentiated respiration at floor level, providing inadequate insight on the peat carbon balance. In contrast to previous assumptions, regular peat oxidation after drainage might contribute more to the regional long‐term annual CO₂ emissions than peat fires. Methane fluxes are negligible at low water levels and amount to up to 3 mg CH₄ m^?2 h^?1 at high water levels, which is low compared with emissions from boreal and temperate peatlands. The latter emissions may be exceeded by fluxes from rice paddies on tropical peat soil, however. N₂O fluxes are erratic with extremely high values upon application of fertilizer to wet peat soils. Current data on CO₂ and CH₄ fluxes indicate that peatland rewetting in south‐east Asia will lead to substantial reductions of net greenhouse gas emissions. There is, however, an urgent need for further quantitative research on carbon exchange to support the development of consistent policies for climate change mitigation.     

The contributions of land-use change, CO2 fertilization, and climate variability to the Eastern US carbon sink

Atmospheric measurements and land‐based inventories imply that terrestrial ecosystems in the northern hemisphere are taking up significant amounts of anthropogenic cabon dioxide (CO₂) emissions; however, there is considerable disagreement about the causes of this uptake, and its expected future trajectory. In this paper, we use the ecosystem demography (ED) model to quantify the contributions of disturbance history, CO₂ fertilization and climate variability to the past, current, and future terrestrial carbon fluxes in the Eastern United States. The simulations indicate that forest regrowth following agricultural abandonment accounts for uptake of 0.11 Pg C yr^?1 in the 1980s and 0.15 Pg C yr^?1 in the 1990s, and regrowth following forest harvesting accounts for an additional 0.1 Pg C yr^?1 of uptake during both these decades. The addition of CO₂ fertilization into the model simulations increases carbon uptake rates to 0.38 Pg C yr^?1 in the 1980s and 0.47 Pg C yr^?1 in the 1990s. Comparisons of predicted aboveground carbon uptake to regional‐scale forest inventory measurements indicate that the model's predictions in the absence of CO₂ fertilization are 14% lower than observed, while in the presence of CO₂ fertilization, predicted uptake rates are 28% larger than observed. Comparable results are obtained from comparisons of predicted total Net Ecosystem Productivity to the carbon fluxes observed at the Harvard Forest flux tower site and in model simulations free‐air CO₂ enrichment (FACE) experiments. These results imply that disturbance history is the principal mechanism responsible for current carbon uptake in the Eastern United States, and that conventional biogeochemical formulations of plant growth overestimate the response of plants to rising CO₂ levels. Model projections out to 2100 imply that the carbon uptake arising from forest regrowth will increasingly be dominated by forest regrowth following harvesting. Consequently, actual carbon storage declines to near zero by the end of the 21st century as the forest regrowth that has occurred since agricultural abandonment comes into equilibrium with the landscape's new disturbance regime. Incorporating interannual climate variability into the model simulations gives rise to large interannual variation in regional carbon fluxes, indicating that long‐term measurements are necessary to detect the signature of processes that give rise to long‐term uptake and storage.     

Tundra ecosystems observed to be CO2 sources due to differential amplification of the carbon cycle

Are tundra ecosystems currently a carbon source or sink? What is the future trajectory of tundra carbon fluxes in response to climate change? These questions are of global importance because of the vast quantities of organic carbon stored in permafrost soils. In this meta‐analysis, we compile 40 years of CO₂ flux observations from 54 studies spanning 32 sites across northern high latitudes. Using time‐series analysis, we investigated if seasonal or annual CO₂ fluxes have changed over time, and whether spatial differences in mean annual temperature could help explain temporal changes in CO₂ flux. Growing season net CO₂ uptake has definitely increased since the 1990s; the data also suggest (albeit less definitively) an increase in winter CO₂ emissions, especially in the last decade. In spite of the uncertainty in the winter trend, we estimate that tundra sites were annual CO₂ sources from the mid‐1980s until the 2000s, and data from the last 7 years show that tundra continue to emit CO₂ annually. CO₂ emissions exceed CO₂ uptake across the range of temperatures that occur in the tundra biome. Taken together, these data suggest that despite increases in growing season uptake, tundra ecosystems are currently CO₂ sources on an annual basis.     

Soil water content and temperature as independent or confounded factors controlling soil respiration in a temperate mixed hardwood forest

Variation in soil temperature can account for most of the seasonal and diel variation in soil CO₂ efflux, but the temperature effect is not always consistent, and other factors such as soil water content are known to influence soil respiration. The objectives of this research were to study the spatial and temporal variation in soil respiration in a temperate forested landscape and to evaluate temperature and soil water functions as predictors of soil respiration. Soil CO₂ fluxes were measured with chambers throughout an annual cycle in six study areas at the Harvard Forest in central Massachusetts that include soil drainage classes from well drained to very poorly drained. The mean annual estimate of soil CO₂ efflux was 7.2 Mg ha^–1, but ranged from 5.3 in the swamp site to 8.5 in a well-drained site, indicating that landscape heterogeneity is related to soil drainage class. An exponential function relating CO₂ fluxes to soil temperature accounted for 80% of the seasonal variation in fluxes across all sites (Q₁₀ = 3.9), but the Q₁₀ ranged from 3.4 to 5.6 for the individual study sites. A significant drought in 1995 caused rapid declines in soil respiration rates in August and September in five of the six sites (a swamp site was the exception). This decline in CO₂ fluxes correlated exponentially with decreasing soil matric potential, indicating a mechanistic effect of drought stress. At moderate to high water contents, however, soil water content was negatively correlated with soil temperature, which precluded distinguishing between the effects of these two confounded factors on CO₂ flux. Occurrence of high Q₁₀ values and variation in Q₁₀ values among sites may be related to: (i) confounding effects of high soil water content; (ii) seasonal and diel patterns in root respiration and turnover of fine roots that are linked to above ground phenology and metabolism; and (iii) variation in the depth where CO₂ is produced. The Q₁₀ function can yield reasonably good predictions of annual fluxes of CO₂, but it is a simplification that masks responses of root and microbial processes to variation in temperature and water content throughout the soil.     

Eutrophication effects on greenhouse gas fluxes from shallow‐lake mesocosms override those of climate warming

Fresh waters make a disproportionately large contribution to greenhouse gas (GHG) emissions, with shallow lakes being particular hot spots. Given their global prevalence, how GHG fluxes from shallow lakes are altered by climate change may have profound implications for the global carbon cycle. Empirical evidence for the temperature dependence of the processes controlling GHG production in natural systems is largely based on the correlation between seasonal temperature variation and seasonal change in GHG fluxes. However, ecosystem‐level GHG fluxes could be influenced by factors, which while varying seasonally with temperature are actually either indirectly related (e.g. primary producer biomass) or largely unrelated to temperature, for instance nutrient loading. Here, we present results from the longest running shallow‐lake mesocosm experiment which demonstrate that nutrient concentrations override temperature as a control of both the total and individual GHG flux. Furthermore, testing for temperature treatment effects at low and high nutrient levels separately showed only one, rather weak, positive effect of temperature (CH₄ flux at high nutrients). In contrast, at low nutrients, the CO₂ efflux was lower in the elevated temperature treatments, with no significant effect on CH₄ or N₂O fluxes. Further analysis identified possible indirect effects of temperature treatment. For example, at low nutrient levels, increased macrophyte abundance was associated with significantly reduced fluxes of both CH₄ and CO₂ for both total annual flux and monthly observation data. As macrophyte abundance was positively related to temperature treatment, this suggests the possibility of indirect temperature effects, via macrophyte abundance, on CH₄ and CO₂ flux. These findings indicate that fluxes of GHGs from shallow lakes may be controlled more by factors indirectly related to temperature, in this case nutrient concentration and the abundance of primary producers. Thus, at ecosystem scale, response to climate change may not follow predictions based on the temperature dependence of metabolic processes.