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Human domination of the biosphere includes changes to disturbance regimes, which push many ecosystems towards early-successional states. Ecological theory predicts that early-successional ecosystems are more sensitive to perturbations than mature systems, but little evidence supports this relationship for the perturbation of climate change. Here we show that vegetation (abundance, species richness and species composition) across seven European shrublands is quite resistant to moderate experimental warming and drought, and responsiveness is associated with the dynamic state of the ecosystem, with recently disturbed sites responding to treatments. Furthermore, most of these responses are not rapid (2–5 years) but emerge over a longer term (7–14 years). These results suggest that successional state influences the sensitivity of ecosystems to climate change, and that ecosystems recovering from disturbances may be sensitive to even modest climatic changes. A research bias towards undisturbed ecosystems might thus lead to an underestimation of the impacts of climate change.

As a key component of the carbon cycle, soil CO2 efflux (SCE) is being increasingly studied to improve our mechanistic understanding of this important carbon flux. Predicting ecosystem responses to climate change often depends on extrapolation of current relationships between ecosystem processes and their climatic drivers to conditions not yet experienced by the ecosystem. This raises the question of to what extent these relationships remain unaltered beyond the current climatic window for which observations are available to constrain the relationships. Here, we evaluate whether current responses of SCE to fluctuations in soil temperature and soil water content can be used to predict SCE under altered rainfall patterns. Of the 58 experiments for which we gathered SCE data, 20 were discarded because either too few data were available or inconsistencies precluded their incorporation in the analyses. The 38 remaining experiments were used to test the hypothesis that a model parameterized with data from the control plots (using soil temperature and water content as predictor variables) could adequately predict SCE measured in the manipulated treatment. Only for 7 of these 38 experiments was this hypothesis rejected. Importantly, these were the experiments with the most reliable data sets, i.e., those providing high-frequency measurements of SCE. Regression tree analysis demonstrated that our hypothesis could be rejected only for experiments with measurement intervals of less than 11 days, and was not rejected for any of the 24 experiments with larger measurement intervals. This highlights the importance of high-frequency measurements when studying effects of altered precipitation on SCE, probably because infrequent measurement schemes have insufficient capacity to detect shifts in the climate dependencies of SCE. Hence, the most justified answer to the question of whether current moisture responses of SCE can be extrapolated to predict SCE under altered

As a key component of the carbon cycle, soil CO2 efflux (SCE) is being increasingly studied to improve our mechanistic understanding of this important carbon flux. Predicting ecosystem responses to climate change often depends on extrapolation of current relationships between ecosystem processes and their climatic drivers to conditions not yet experienced by the ecosystem. This raises the question of to what extent these relationships remain unaltered beyond the current climatic window for which observations are available to constrain the relationships. Here, we evaluate whether current responses of SCE to fluctuations in soil temperature and soil water content can be used to predict SCE under altered rainfall patterns. Of the 58 experiments for which we gathered SCE data, 20 were discarded because either too few data were available or inconsistencies precluded their incorporation in the analyses. The 38 remaining experiments were used to test the hypothesis that a model parameterized with data from the control plots (using soil temperature and water content as predictor variables) could adequately predict SCE measured in the manipulated treatment. Only for 7 of these 38 experiments was this hypothesis rejected. Importantly, these were the experiments with the most reliable data sets, i.e., those providing high-frequency measurements of SCE. Regression tree analysis demonstrated that our hypothesis could be rejected only for experiments with measurement intervals of less than 11 days, and was not rejected for any of the 24 experiments with larger measurement intervals. This highlights the importance of high-frequency measurements when studying effects of altered precipitation on SCE, probably because infrequent measurement schemes have insufficient capacity to detect shifts in the climate dependencies of SCE. Hence, the most justified answer to the question of whether current moisture responses of SCE can be extrapolated to predict SCE under altered precipitation re

In this study, we compare annual fluxes of methane (CH4), nitrous oxide (N2O) and soil respiratory carbon dioxide (CO2) measured at nine European peatlands (n = 4) and shrublands (n = 5). The sites range from northern Sweden to Spain, covering a span in mean annual air temperature from 0 to 16 degrees C, and in annual precipitation from 300 to 1300 mm yr(-1). The effects of climate change, including temperature increase and prolonged drought, were tested at five shrubland sites. At one peatland site, the long-term (> 30 yr) effect of drainage was assessed, while increased nitrogen deposition was investigated at three peatland sites. The shrublands were generally sinks for atmospheric CH4, whereas the peatlands were CH4 sources, with fluxes ranging from -519 to + 6890 mg CH4-Cm-2 yr(-1) across the studied ecosystems. At the peatland sites, annual CH4 emission increased with mean annual air temperature, while a negative relationship was found between net CH4 uptake and the soil carbon stock at the shrubland sites. Annual N2O fluxes were generally small ranging from -14 to 42 mg N2O-Nm(-2) yr(-1). Highest N2O emission occurred at the sites that had highest nitrate (NO3-) concentration in the soil water. Furthermore, experimentally increased NO3- deposition led to increased N2O efflux, whereas prolonged drought and long-term drainage reduced the N2O efflux. Soil CO2 emissions in control plots ranged from 310 to 732 g CO2-C m(-2) yr(-1). Drought and long-term drainage from -519 to + 6890 mg CH4-C m(-2) yr(-1) across the studied ecosystems. At the peatland sites, annual CH4 emission increased with mean annual air temperature, while a negative relationship was found between net CH4 uptake and the soil carbon stock at the shrubland sites. Annual N2O fluxes were generally small ranging from -14 to 42 mg N2O-N m(-2) yr(-1). Highest N2O emission occurred at the sites that had highest nitrate (NO3-) concentration in the soil water. Furthermore, experimentally increased NO3

This study investigated the impact of predicted future climatic and atmospheric conditions on soil respiration (RS) in a Danish Calluna-Deschampsia-heathland. A fully factorial in situ experiment with treatments of elevated atmospheric CO2 (+130 ppm), raised soil temperature (+0.4 degrees C) and extended summer drought (58% precipitation exclusion) was established in 2005. The average RS, observed in the control over 3 years of measurements (1.7 mu mol CO2 m-2 sec-1), increased 38% under elevated CO2, irrespective of combination with the drought or temperature treatments. In contrast, extended summer drought decreased RS by 14%, while elevated soil temperature did not affect RS overall. A significant interaction between elevated temperature and drought resulted in further reduction of RS when these treatments were combined. A detailed analysis of short-term RS dynamics associated with drought periods showed that RS was reduced by 50% and was strongly correlated with soil moisture during these events. Recovery of RS to pre-drought levels occurred within 2 weeks of rewetting; however, unexpected drought effects were observed several months after summer drought treatment in 2 of the 3 years, possibly due to reduced plant growth or changes in soil water holding capacity. An empirical model that predicts RS from soil temperature, soil moisture and plant biomass was developed and accounted for 55% of the observed variability in RS. The model predicted annual sums of RS in 2006 and 2007, in the control, were 672 and 719 g C m-2 y-1, respectively. For the full treatment combination, i.e. the future climate scenario, the model predicted that soil respiratory C losses would increase by 21% (140150 g C m-2 y-1). Therefore, in the future climate, stimulation of C storage in plant biomass and litter must be in excess of 21% for this ecosystem to not suffer a reduction in net ecosystem exchange.

In recent years, increased awareness of the potential interactions between rising atmospheric CO2 concentrations ([ CO2 ]) and temperature has illustrated the importance of multifactorial ecosystem manipulation experiments for validating Earth System models. To address the urgent need for increased understanding of responses in multifactorial experiments, this article synthesizes how ecosystem productivity and soil processes respond to combined warming and [ CO2 ] manipulation, and compares it with those obtained in single factor [ CO2 ] and temperature manipulation experiments. Across all combined elevated [ CO2 ] and warming experiments, biomass production and soil respiration were typically enhanced. Responses to the combined treatment were more similar to those in the [ CO2 ]-only treatment than to those in the warming-only treatment. In contrast to warming-only experiments, both the combined and the [ CO2 ]-only treatments elicited larger stimulation of fine root biomass than of aboveground biomass, consistently stimulated soil respiration, and decreased foliar nitrogen (N) concentration. Nonetheless, mineral N availability declined less in the combined treatment than in the [ CO2 ]-only treatment, possibly due to the warming-induced acceleration of decomposition, implying that progressive nitrogen limitation (PNL) may not occur as commonly as anticipated from single factor [ CO2 ] treatment studies. Responses of total plant biomass, especially of aboveground biomass, revealed antagonistic interactions between elevated [ CO2 ] and warming, i.e. the response to the combined treatment was usually less-than-additive. This implies that productivity projections might be overestimated when models are parameterized based on single factor responses. Our results highlight the need for more (and especially more long-term) multifactor manipulation experiments. Because single factor CO2 responses often dominated over warming responses in the combined treatments, our result

Nitrogen (N) is a critical resource for plant growth in tundra ecosystems, and species differences in the timing of N uptake may be an important feature regulating community composition and ecosystem productivity. We added N-15-labelled glycine to a subarctic heath tundra dominated by dwarf shrubs, mosses and graminoids in fall, and investigated its partitioning among ecosystem components at several time points (October, November, April, May, June) through to the following spring/early summer. Soil microbes had acquired 65 +/- A 7% of the N-15 tracer by October, but this pool decreased through winter to 37 +/- A 7% by April indicating significant microbial N turnover prior to spring thaw. Only the evergreen dwarf shrubs showed active N-15 acquisition before early May indicating that they had the highest potential of all functional groups for acquiring nutrients that became available in early spring. The faster-growing deciduous shrubs did not resume N-15 acquisition until after early May indicating that they relied more on nitrogen made available later during the spring/early summer. The graminoids and mosses had no significant increases in N-15 tracer recovery or tissue N-15 tracer concentrations after the first harvest in October. However, the graminoids had the highest root N-15 tracer concentrations of all functional groups in October indicating that they primarily relied on N made available during summer and fall. Our results suggest a temporal differentiation among plant functional groups in the post-winter resumption of N uptake with evergreen dwarf shrubs having the highest potential for early N uptake, followed by deciduous dwarf shrubs and graminoids.

In this study, we compare annual fluxes of methane (CH4), nitrous oxide (N2O) and soil respiratory carbon dioxide (CO2) measured at nine European peatlands (n = 4) and shrublands (n = 5). The sites range from northern Sweden to Spain, covering a span in mean annual air temperature from 0 to 16 °C, and in annual precipitation from 300 to 1300 mm yr−1. The effects of climate change, including temperature increase and prolonged drought, were tested at five shrubland sites. At one peatland site, the long-term (>30 yr) effect of drainage was assessed, while increased nitrogen deposition was investigated at three peatland sites. The shrublands were generally sinks for atmospheric CH4 whereas the peatlands were CH4 sources, with fluxes ranging from −519 to +6890 mg CH4-C m−2 yr−1 across the studied ecosystems. At the peatland sites, annual CH4 emission increased with mean annual air temperature, while a negative relationship was found between net CH4 uptake and the soil carbon stock at the shrubland sites. Annual N2O fluxes were generally small ranging from –14 to 42 mg N2O-N m−2 yr−1. Highest N2O emission occurred at the sites that had highest concentration of nitrate (NO3−) in soil water. Furthermore, experimentally increased NO3− deposition led to increased N2O efflux, whereas prolonged drought and long-term drainage reduced the N2O efflux. Soil CO2 emissions in control plots ranged from 310 to 732 g CO2-C m−2 yr−1. Drought and long-term drainage generally reduced the soil CO2 efflux, except at a~hydric shrubland where drought tended to increase soil respiration. When comparing the fractional importance of each greenhouse gas to the total n

In this study, we compare annual fluxes of methane (CH4), nitrous oxide (N2O) and soil respiratory carbon dioxide (CO2) measured at nine European peatlands (n = 4) and shrublands (n = 5). The sites range from northern Sweden to Spain, covering a span in mean annual air temperature from 0 to 16 C, and in annual precipitation from 300 to 1300mmyr-1. The effects of climate change, including temperature increase and prolonged drought, were tested at five shrubland sites. At one peatland site, the long-term (>30 yr) effect of drainage was assessed, while increased nitrogen deposition was investigated at three peatland sites. The shrublands were generally sinks for atmospheric CH4, whereas the peatlands were CH4 sources, with fluxes ranging from -519 to +6890 mgCH4-Cm-2 yr-1 across the studied ecosystems. At the peatland sites, annual CH4 emission increased with mean annual air temperature, while a negative relationship was found between net CH4 uptake and the soil carbon stock at the shrubland sites. Annual N2O fluxes were generally small ranging from -14 to 42 mgN2O-Nm-2 yr-1. Highest N2O emission occurred at the sites that had highest nitrate (NO- 3 ) concentration in the soil water. Furthermore, experimentally increased NO- 3 deposition led to increased N2O efflux, whereas prolonged drought and long-term drainage reduced the N2O efflux. Soil CO2 emissions in control plots ranged from 310 to 732 gCO2-Cm-2 yr-1. Drought and long-term drainage generally reduced the soil CO2 efflux, except at a hydric shrubland where drought tended to increase soil respiration. In terms of fractional importance of each greenhouse gas to the total numerical global warming response, the change in CO2 efflux dominated the response in all treatments (ranging 71–96 %), except for NO- 3 addition where 89% was due to change in CH4 emissions. Thus, in European peatlan

Laboratory experiments show that rates of CH4 emission from plant material depend exponentially on temperature and linearly on UV irradiance. The UV irradiance shall be spectrally weighted and shorter wavelengths results in higher CH4 emissions. Global upscaling models for estimating aerobic CH4, based on lab results, have be conducted with varying results, but until now field measurements based on profile and eddy covariance measurements have failed to show CH4 emissions from forest canopies. To detect CH4 production or consumption in the canopy of a beech stand we connected a CH4 analyzer to a canopy air profile system that samples air below and above the canopy from seven different heights. A profile system with many vertical sample points can detect gas concentration gradients with a high sensitivity only under conditions with no or little air movements. Under these conditions we found indications of periodic CH4 emissions in the canopy, but more data need to be analyzed before the magnitude of the canopy source of CH4 can be established.

CLIMMANI/INTERFACE Joint Workshop: Nutrient Constraints on the Net Carbon Balance; Keflavik, Iceland, 15-17 June 2011 A challenge to improving predictions of the consequences of climate change is bringing empiricists and theoreticians together to work toward common goals. Critical to this challenge is ensuring that novel, process-based information is incorporated into the land surface portion of Earth system models and, at the same time, that field-based experiments are designed to explore model weaknesses. The Integrated Network for Terrestrial Ecosystem Research on Feedbacks to the Atmosphere and Climate (INTERFACE) and Climate Change: Manipulation Experiments in Terrestrial Ecosystems (ClimMani) joint meeting in Iceland fostered interdisciplinary interaction among land modelers, biogeochemists, and ecologists while focusing on nutrient constraints on the net carbon budget. Discussions at the meeting aimed to identify the status of current knowledge, major knowledge gaps, and the steps needed to move forward.

Field-scale experiments simulating realistic future climate scenarios are important tools for investigating the effects of current and future climate changes on ecosystem functioning and biogeochemical cycling. We exposed a seminatural Danish heathland ecosystem to elevated atmospheric carbon dioxide (CO2), warming, and extended summer drought in all combinations. Here, we report on the short-term responses of the nitrogen (N) cycle after 2 years of treatments. Elevated CO2 significantly affected aboveground stoichiometry by increasing the carbon to nitrogen (C/N) ratios in the leaves of both co-dominant species (Calluna vulgaris and Deschampsia flexuosa), as well as the C/N ratios of Calluna flowers and by reducing the N concentration of Deschampsia litter. Belowground, elevated CO2 had only minor effects, whereas warming increased N turnover, as indicated by increased rates of microbial NH4+ consumption, gross mineralization, potential nitrification, denitrification and N2O emissions. Drought reduced belowground gross N mineralization and decreased fauna N mass and fauna N mineralization. Leaching was unaffected by treatments but was significantly higher across all treatments in the second year than in the much drier first year indicating that ecosystem N loss is highly sensitive to changes and variability in amount and timing of precipitation. Interactions between treatments were common and although some synergistic effects were observed, antagonism dominated the interactive responses in treatment combinations, i.e. responses were smaller in combinations than in single treatments. Nonetheless, increased C/N ratios of photosynthetic tissue in response to elevated CO2, as well as drought-induced decreases in litter N production and fauna N mineralization prevailed in the full treatment combination. Overall, the simulated future climate scenario therefore lead to reduced N turnover, which could act to reduce the potential growth response of plants to elevated atmosp

An analysis of flow tilt angles from a fetch-limited beech forest site with clearings is presented in the context of vertical advection of carbon dioxide. Flow angles and vertical velocities from two sonic anemometers by different manufacturers were analyzed. Instead of using rotations, where zero-flow angles were assumed for neutral flow, the data was interpreted in relation to upstream and downstream forest edges. Uncertainties caused by flow distortion, vertical misalignment and limited sampling time (statistical uncertainty) were evaluated and found to be highly significant. Since the attack angle distribution of the wind on the sonic anemometer is a function of atmospheric stratification, an instrumental error caused by imperfect flow distortion correction is also a function of the atmospheric stratification. In addition, it is discussed that the sonic anemometers have temperature dependent off-sets. These features of the investigated sonic anemometers make them unsuitable for measuring vertical velocities over highly turbulent forested terrain. By comparing the sonic anemometer results to that of a conically scanning Doppler lidar (Dellwik et al., 2010b), sonic anemometer accuracy for measuring mean flow tilt angles was estimated to between 2° and 3°. Use of planar fit algorithms, where the mean vertical velocity is calculated as the difference between the neutral and non-neutral flow, does not solve this problem of low accuracy and is not recommended. Because of the large uncertainties caused by flow distortion and vertical alignment, it was only possible to a limited extent to relate sonic anemometer flow tilt angles to upwind forest edges, but the results by the lidar indicated that an internal boundary layer affect flow tilt angles at 21m above the forest. This is in accordance with earlier studies at the site. Since the mean flow tilt angles do not follow the terrain, an estimate of the vertical advection term for near-neutral conditions was calculated us

1. Recent findings indicate that the interactions among CO2, temperature and water can be substantial, and that the combined effects on the biological systems of several factors may not be predicted from experiments with one or a few factors. Therefore realistic multifactorial experiments involving a larger set of main factors are needed.2. We describe a new Danish climate change-related field scale experiment, CLIMAITE, in a heath/grassland ecosystem. CLIMAITE is a full factorial combination of elevated CO2, elevated temperature and prolonged summer drought. The manipulations are intended to mimic anticipated major environmental changes at the site by year 2075 as closely as possible. The impacts on ecosystem processes and functioning (at ecophysiological levels, through responses by individuals and communities to ecosystem-level responses) are investigated simultaneously.3. The increase of [CO2] closely corresponds with the scenarios for year 2075, while the warming treatment is at the lower end of the predictions and seems to be the most difficult treatment to increase without unwanted side effects on the other variables. The drought treatment follows predictions of increased frequency of drought periods in summer. The combination of the treatments does not create new unwanted side effects on the treatments relative to the treatments alone.

We measured net ecosystem CO2 flux (F-n) and ecosystem respiration (R-E), and estimated gross ecosystem photosynthesis (P-g) by difference, for two years in a temperate heath ecosystem using a chamber method. The exchange rates of carbon were high and of similar magnitude as for productive forest ecosystems with a net ecosystem carbon gain during the second year of 293 +/- 11 g C m(-2) year(-1) showing that the carbon sink strength of heather-dominated ecosystems may be considerable when C. vulgaris is in the building phase of its life cycle. The estimated gross ecosystem photosynthesis and ecosystem respiration from October to March was 22% and 30% of annual flux, respectively, suggesting that both cold-season carbon gain and loss were important in the annual carbon cycle of the ecosystem. Model fit of R-E of a classic, first-order exponential equation related to temperature ( second year; R-2 = 0.65) was improved when the P-g rate was incorporated into the model (second year; R-2 = 0.79), suggesting that daytime R-E increased with increasing photosynthesis. Furthermore, the temperature sensitivity of R-E decreased from apparent Q(10) values of 3.3 to 3.9 by the classic equation to a more realistic Q(10) of 2.5 by the modified model. The model introduces R-photo, which describes the part of respiration being tightly coupled to the photosynthetic rate. It makes up 5% of the assimilated carbon dioxide flux at 0 degrees C and 35% at 20 degrees C implying a high sensitivity of respiration to photosynthesis during summer. The simple model provides an easily applied, non-intrusive tool for investigating seasonal trends in the relationship between ecosystem carbon sequestration and respiration.