Soil CO2 efflux in contrasting boreal deciduous and coniferous stands and its contribution to the ecosystem carbon balance

Similar nonsteady‐state automated chamber systems were used to measure and partition soil CO₂ efflux in contrasting deciduous (trembling aspen) and coniferous (black spruce and jack pine) stands located within 100 km of each other near the southern edge of the Boreal forest in Canada. The stands were exposed to similar climate forcing in 2003, including marked seasonal variations in soil water availability, which provided a unique opportunity to investigate the influence of climate and stand characteristics on soil CO₂ efflux and to quantify its contribution to the net ecosystem CO₂ exchange (NEE) as measured with the eddy‐covariance technique. Partitioning of soil CO₂ efflux between soil respiration (including forest‐floor vegetation) and forest‐floor photosynthesis showed that short‐ and long‐term temporal variations of soil CO₂ efflux were related to the influence of (1) soil temperature and water content on soil respiration and (2) below‐canopy light availability, plant water status and forest‐floor plant species composition on forest‐floor photosynthesis. Overall, the three stands were weak to moderate sinks for CO₂ in 2003 (NEE of ?103, ?80 and ?28 g C m^?2 yr^?1 for aspen, black spruce and jack pine, respectively). Forest‐floor respiration accounted for 86%, 73% and 75% of annual ecosystem respiration, in the three respective stands, while forest‐floor photosynthesis contributed to 11% and 14% of annual gross ecosystem photosynthesis in the black spruce and jack pine stands, respectively. The results emphasize the need to perform concomitant measurements of NEE and soil CO₂ efflux at longer time scales in different ecosystems in order to better understand the impacts of future interannual climate variability and vegetation dynamics associated with climate change on each component of the carbon balance.     

Seasonal variations of CO2 and water vapour exchange rates over a temperate deciduous forest

Long-term and direct measurements of CO₂ and water vapour exchange are needed over forested ecosystems to determine their net annual fluxes of carbon dioxide and water. Such measurements are also needed to parameterize and test biogeochemical, ecological and hydrological assessment models. Responding to this need, eddy covariance measurements of CO₂ and water vapour were made ever a deciduous forest growing near Oak Ridge, TN, between April 1993 and April 1994. Periodic measurements were made of leaf area index, stomatal resistance, soil moisture and pre-dawn leaf water potential to characterize the gas exchange capacity of the canopy. Four factors had a disproportionate influence on the seasonal variation of CO₂ flux densities. These factors were photon flux densities (during the growing season), temperature (during the dormant season), leaf area index and the occurrence of drought The drought period occurred during the peak of the growing season and caused a significant decline in daily and hourly CO₂ flux densities, relative to observations over the stand when soil moisture was plentiful. The annual net uptake of carbon was calculated by integrating flux measurements and filling missing and spurious data with the relations obtained between measured CO₂ fluxes and environmental forcing variables. The net flux of carbon for the period between April 1993 and April 1994 was -525 g C m^?2 y^?1. This value represents a net flux of carbon from the atmosphere and into the forest. The net annual carbon exchange of this southern temperate broadleaved forest exceeded values measured over a northern temperate forest (which experiences a shorter growing season and has less leaf area) by 200 g C m^?2 y^?1 (cf. Wofsy et al 1993). The seasonal variation of canopy evaporation (latent heat flux) was controlled mostly by changes in leaf area and net radiation. A strong depression in evaporation rates was not observed during the drought Over a broadleaved forest large vapour pressure deficits promote evaporation and trees in a mixed stand are able to tap a variety of deep and shallow water sources.     

An initial intercomparison of micrometeorological and ecological inventory estimates of carbon exchange in a mid-latitude deciduous forest

The role of mid‐latitude forests in the sequestration of carbon (C) is of interest to an increasing number of scientists and policy‐makers alike. Net CO₂ exchange can be estimated on an annual basis, using eddy‐covariance techniques or from ecological inventories of C fluxes to and from a forest. Here we present an intercomparison of annual estimates of C exchange in a mixed hardwood forest in the Morgan‐Monroe State Forest, Indiana, USA for two years, 1998 and 1999. Based on eddy‐covariance measurements made at 1.8 times canopy height from a tower, C uptake by the forest was 237 and 287 g C m^?2 y^?1 for 1998 and 1999, respectively. For the same time period, biometric and ecophysiological measures and modelled estimates of all significant carbon fluxes within deciduous forests were made, including: change in living biomass, aboveground and belowground detritus production, foliage consumption, and forest floor and soil respiration. Using this ecological inventory method for these same two time periods, C uptake was estimated to be 271 and 377 g C m^?2 y^?1, which are 14.3% and 31.4% larger, respectively, than the tower‐based values. The relative change between this method's annual estimates is consistent with that of the eddy‐covariance based values. Our results indicate that the difference in annual C exchange rates was due to reduced heterotrophic soil respiration in 1999.     

Long-term carbon exchange in a sparse, seasonally dry tussock grassland

Rainfall and its seasonal distribution can alter carbon dioxide (CO₂) exchange and the sustainability of grassland ecosystems. Using eddy covariance, CO₂ exchange between the atmosphere and a sparse grassland was measured for 2 years at Twizel, New Zealand. The years had contrasting distributions of rain and falls (446 mm followed by 933 mm; long‐term mean=646 mm). The vegetation was sparse with total above‐ground biomass of only 1410 g m^?2. During the dry year, leaf area index peaked in spring (November) at 0.7, but it was <0.2 by early summer. The maximum daily net CO₂ uptake rate was only 1.5 g C m^?2 day^?1, and it occurred before mid‐summer in both years. On an annual basis, for the dry year, 9 g C m^?2 was lost to the atmosphere. During the wet year, 41 g C m^?2 was sequestered from the atmosphere. The net exchange rates were determined mostly by the timing and intensity of spring rainfall. The components of ecosystem respiration were measured using chambers. Combining scaled‐up measurements with the eddy CO₂ effluxes, it was estimated that 85% of ecosystem respiration emanated from the soil surface. Under well‐watered conditions, 26% of the soil surface CO₂ efflux came from soil microbial activity. Rates of soil microbial CO₂ production and net mineral‐N production were low and indicative of substrate limitation. Soil respiration declined by a factor of four as the soil water content declined from field capacity (0.21 m³ m^?3) to the driest value obtained (0.04 m³ m^?3). Rainfall after periods of drought resulted in large, but short‐lived, respiration pulses that were curvilinearly related to the increase in root‐zone water content. Coupled with the low leaf area and high root : shoot ratio, this sparse grassland had a limited capacity to sequester and store carbon. Assuming a proportionality between carbon gain and rainfall during the summer, rainfall distribution statistics suggest that the ecosystem is sustainable in the long term.     

Carbon exchange of grazed pasture on a drained peat soil

Land‐use changes have contributed to increased atmospheric CO₂ concentrations. Conversion from natural peatlands to agricultural land has led to widespread subsidence of the peat surface caused by soil compaction and mineralization. To study the net ecosystem exchange of carbon (C) and the contribution of respiration to peat subsidence, eddy covariance measurements were made over pasture on a well‐developed, drained peat soil from 22 May 2002 to 21 May 2003. The depth to the water table fluctuated between 0.02 m in winter 2002 to 0.75 m during late summer and early autumn 2003. Peat soil moisture content varied between 0.6 and 0.7 m³ m^?3 until the water table dropped below 0.5 m, when moisture content reached 0.38 m³ m^?3. Neither depth to water table nor soil moisture was found to have an effect on the rate of night‐time respiration (ranging from 0.4–8.0 μmol CO₂ m^?2 s^?1 in winter and summer, respectively). Most of the variance in night‐time respiration was explained by changes in the 0.1 m soil temperature (r²=0.93). The highest values for daytime net ecosystem exchange were measured in September 2002, with a maximum of ?17.2 μmol CO₂ m^?2 s^?1. Grazing events and soil moisture deficiencies during a short period in summer reduced net CO₂ exchange. To establish an annual C balance for this ecosystem, non‐linear regression was used to model missing data. Annually integrated (CO₂) C exchange for this peat–pasture ecosystem was 45±500 kg C ha^?1 yr^?1. After including other C exchanges (methane emissions from cows and production of milk), the net annual C loss was 1061±500 kg C ha^?1 yr^?1.     

Elevated CO2 alters carbon fluxes in early successional Mediterranean ecosystems

Annual carbon budgets of ecosystems are central to our understanding of the biotic control of atmospheric composition, but they are not available under elevated CO₂ for most vegetation types. Using gas exchange techniques, we assessed carbon fluxes of four early successional Mediterranean model communities, consisting of grasses, legumes and composites. The assemblages were grown on the same monoliths for three consecutive years in greenhouses tracking field conditions except for CO₂ maintained at ambient (370 μmol mol^?1) or elevated (700 μmol mol^?1) concentration. During the third year of study, CO₂ enrichment consistently shifted the annual carbon balance towards lower efflux, with displacements between 4.3 and 26.2 mol m^?2 y^?1 (one assemblage became a net CO₂ sink, another just reached equilibrium, and the remaining two remained as a CO₂ source). At least 50% of the shift under elevated CO₂ originated from a decrease in belowground respiration. This indicates that, during this year, CO₂ enrichment did not predominantly enhance C‐cycling, but on the contrary inhibited root respiration or microbial C‐utilization. Although elevated‐CO₂‐grown systems acted as a net CO₂ sink during a longer period of the year (4–7 months) compared with ambient‐CO₂‐grown systems (3–3.5 months), gross canopy photosynthesis was modified only to a limited extent (between ?5.9 and + 14.8%). Interaction between the carbon and the water cycle was apparently responsible for this weak stimulation. In particular, reduced evapotranspiration under elevated CO₂ coincided with inhibited canopy photosynthesis in early spring, most likely resulting from water saturation of the soil. In addition, only the earliest‐planted assemblages had an increased gross canopy photosynthesis during late autumn and early winter. This suggests that a longer summer drought, by delaying the establishment of such an annual type of vegetation, would reduce the positive impact of elevated CO₂ on productivity. Water regime appears to strongly govern the influence of CO₂ on the carbon fluxes in Mediterranean ecosystems with annual herbaceous vegetation.     

Do the energy fluxes and surface conductance of boreal coniferous forests in Europe scale with leaf area?

Earth observing systems are now routinely used to infer leaf area index (LAI) given its significance in spatial aggregation of land surface fluxes. Whether LAI is an appropriate scaling parameter for daytime growing season energy budget, surface conductance (G_s), water‐ and light‐use efficiency and surface–atmosphere coupling of European boreal coniferous forests was explored using eddy‐covariance (EC) energy and CO₂ fluxes. The observed scaling relations were then explained using a biophysical multilayer soil–vegetation–atmosphere transfer model as well as by a bulk G_s representation. The LAI variations significantly alter radiation regime, within‐canopy microclimate, sink/source distributions of CO₂, H₂O and heat, and forest floor fluxes. The contribution of forest floor to ecosystem‐scale energy exchange is shown to decrease asymptotically with increased LAI, as expected. Compared with other energy budget components, dry‐canopy evapotranspiration (ET) was reasonably ‘conservative’ over the studied LAI range 0.5–7.0 m² m^?2. Both ET and G_s experienced a minimum in the LAI range 1–2 m² m^?2 caused by opposing nonproportional response of stomatally controlled transpiration and ‘free’ forest floor evaporation to changes in canopy density. The young forests had strongest coupling with the atmosphere while stomatal control of energy partitioning was strongest in relatively sparse (LAI ~2 m² m^?2) pine stands growing on mineral soils. The data analysis and model results suggest that LAI may be an effective scaling parameter for net radiation and its partitioning but only in sparse stands (LAI <3 m² m^?2). This finding emphasizes the significance of stand‐replacing disturbances on the controls of surface energy exchange. In denser forests, any LAI dependency varies with physiological traits such as light‐saturated water‐use efficiency. The results suggest that incorporating species traits and site conditions are necessary when LAI is used in upscaling energy exchanges of boreal coniferous forests.     

Modelling night‐time ecosystem respiration by a constrained source optimization method

One of the main challenges to quantifying ecosystem carbon budgets is properly quantifying the magnitude of night‐time ecosystem respiration. Inverse Lagrangian dispersion analysis provides a promising approach to addressing such a problem when measured mean CO₂ concentration profiles and nocturnal velocity statistics are available. An inverse method, termed ‘Constrained Source Optimization’ or CSO, which couples a localized near‐field theory (LNF) of turbulent dispersion to respiratory sources, is developed to estimate seasonal and annual components of ecosystem respiration. A key advantage to the proposed method is that the effects of variable leaf area density on flow statistics are explicitly resolved via higher‐order closure principles. In CSO, the source distribution was computed after optimizing key physiological parameters to recover the measured mean concentration profile in a least‐square fashion. The proposed method was field‐tested using 1 year of 30‐min mean CO₂ concentration and CO₂ flux measurements collected within a 17‐year‐old (in 1999) even‐aged loblolly pine (Pinus taeda L.) stand in central North Carolina. Eddy‐covariance flux measurements conditioned on large friction velocity, leaf‐level porometry and forest‐floor respiration chamber measurements were used to assess the performance of the CSO model. The CSO approach produced reasonable estimates of ecosystem respiration, which permits estimation of ecosystem gross primary production when combined with daytime net ecosystem exchange (NEE) measurements. We employed the CSO approach in modelling annual respiration of above‐ground plant components (c. 214 g C m^?2 year^?1) and forest floor (c. 989 g C m^?2 year^?1) for estimating gross primary production (c. 1800 g C m^?2 year^?1) with a NEE of c. 605 g C m^?2 year^?1 for this pine forest ecosystem. We conclude that the CSO approach can utilise routine CO₂ concentration profile measurements to corroborate forest carbon balance estimates from eddy‐covariance NEE and chamber‐based component flux measurements.     

Water Relations and CO2 Exchange of Tropical Bryophytes in a Lower Montane Rain Forest in Panama

For 6 tropical bryophytes, measurements of the diel courses in water status and net CO₂ exchange were made in a submontane tropical rain forest in Panama. In addition, the response of gas exchange to changes in photon flux density (PPFD) and thallus water content (WC) was studied under controlled conditions. Diel variation of WC was pronounced, and both low and high WC limited carbon gain considerably. Low PPFD, e.g. during rain storms, was less important in limiting CO₂ exchange. More than half of the mean diurnal carbon gain of 2.9 mg C per g thallus carbon was lost during the night as respiration. Assuming that the average 24-h carbon gain was representative for the entire year, we estimated the net annual primary productivity of the mosses and liverworts to be 45% of the initial plant carbon content.     

Reduction of forest floor respiration by fertilization on both carbon dioxide-enriched and reference 17-year-old loblolly pine stands

Elevated atmospheric carbon dioxide (CO₂^e) increases soil respiration rates in forest, grassland, agricultural and wetland systems as a result of increased growth, root biomass and enhanced biological activity of soil microorganisms. Less is known about how forest floor fluxes respond to the combined effects of elevated CO₂ and nutrient amendments; until now no experiments have been in place with large forest trees to allow even preliminary investigations. We investigated changes in forest floor respiration (S_ff) in a Pinus taeda L. plantation fumigated with CO₂ by using free‐air CO₂ enrichment (FACE) technology and given nutrient amendments. The prototype FACE apparatus (FACEp; 707 m²) was constructed in 1993, 10 years after planting, on a moderate fertility site in Duke Forest, North Carolina, USA, enriching the stand to 55 Pa (CO₂^e). A nearby ambient CO₂ (CO₂^a) plot (117 m²) was designated at the inception of the study as a reference (Ref). Both FACEp and Ref plot were divided in half and urea fertilizer was applied to one half at an annual rate of 11.2 g N m^?2 in the spring of 1998, 1999 and 2000. Forest floor respiration was monitored continuously for 220 days – March through November 2000 – by using two Automated Carbon Efflux Systems. Thirty locations (491 cm² each) were sampled in both FACEp and Ref, about half in each fertility treatment. Forest floor respiration was strongly correlated with soil temperature at 5 cm. Rates of S_ff were greater in CO₂^e relative to CO₂^a (an enhancement of ～178 g C m^?2) during the measurement period. Application of fertilizer resulted in a statistically significant depression of respiration rates in both the CO₂^a and CO₂^e plots (a reduction of ～186 g C m^?2). The results suggest that closed canopy forests on moderate fertility sites cycle back to the atmosphere more assimilated carbon (C) than similar forests on sites of high fertility. We recognize the limitations of this non‐replicated study, but its clear results offer strong testable hypotheses for future research in this important area.     

Carbon storage and fluxes in ponderosa pine forests at different developmental stages

We compared carbon storage and fluxes in young and old ponderosa pine stands in Oregon, including plant and soil storage, net primary productivity, respiration fluxes, eddy flux estimates of net ecosystem exchange (NEE), and Biome‐BGC simulations of fluxes. The young forest (Y site) was previously an old‐growth ponderosa pine forest that had been clearcut in 1978, and the old forest (O site), which has never been logged, consists of two primary age classes (50 and 250 years old). Total ecosystem carbon content (vegetation, detritus and soil) of the O forest was about twice that of the Y site (21 vs. 10 kg C m^?2 ground), and significantly more of the total is stored in living vegetation at the O site (61% vs. 15%). Ecosystem respiration (R_e) was higher at the O site (1014 vs. 835 g C m^?2 year^?1), and it was largely from soils at both sites (77% of R_e). The biological data show that above‐ground net primary productivity (ANPP), NPP and net ecosystem production (NEP) were greater at the O site than the Y site. Monte Carlo estimates of NEP show that the young site is a source of CO₂ to the atmosphere, and is significantly lower than NEP(O) by c. 100 g C m^?2 year^?1. Eddy covariance measurements also show that the O site was a stronger sink for CO₂ than the Y site. Across a 15‐km swath in the region, ANPP ranged from 76 g C m^?2 year^?1 at the Y site to 236 g C m^?2 year^?1 (overall mean 158 ± 14 g C m^?2 year^?1). The lowest ANPP values were for the youngest and oldest stands, but there was a large range of ANPP for mature stands. Carbon, water and nitrogen cycle simulations with the Biome‐BGC model suggest that disturbance type and frequency, time since disturbance, age‐dependent changes in below‐ground allocation, and increasing atmospheric concentration of CO₂ all exert significant control on the net ecosystem exchange of carbon at the two sites. Model estimates of major carbon flux components agree with budget‐based observations to within ± 20%, with larger differences for NEP and for several storage terms. Simulations showed the period of regrowth required to replace carbon lost during and after a stand‐replacing fire (O) or a clearcut (Y) to be between 50 and 100 years. In both cases, simulations showed a shift from net carbon source to net sink (on an annual basis) 10–20 years after disturbance. These results suggest that the net ecosystem production of young stands may be low because heterotrophic respiration, particularly from soils, is higher than the NPP of the regrowth. The amount of carbon stored in long‐term pools (biomass and soils) in addition to short‐term fluxes has important implications for management of forests in the Pacific North‐west for carbon sequestration.     

Growth,Net Production,Litter Decomposition,and Net Nitrogen Accumulation by Epiphytic Bryophytes in a Tropical Montane Forest1

To understand the ecological roles of epiphytic bryophytes in the carbon (C) and nitrogen (N) cycles of a tropical montane forest, we used samples in enclosures to estimate rates of growth, net production, and N accumulation by shoots in the canopy, and litterbags, to estimate rates of decomposition and N dynamics of epiphytic bryophyte litter in the canopy and on the forest floor in Monteverde, Costa Rica. Growth of epiphytic bryophytes was estimated at 30.0–49.9 percent/yr, net production at 122–203 g/m²/yr, and N accumulation at 1.8–3.0 g N/m²/yr. Cumulative mass loss from litterbags after one and two years in the canopy was 17 ± 2 and 19 ± 2 percent (mean ± 1 SE) of initial sample mass, respectively, and mass loss from litter and green shoots in litterbags after one year on the forest floor was 29 ± 2 and 45 ± 3 percent, respectively. Approximately 30 percent of the initial N mass was released rapidly from litter in both locations. Nitrogen loss from green shoots on the forest floor was greater; about 47 percent of the initial N mass was lost within the first three months. There was no evidence for net N immobilization by litter or green shoots, but the remaining N in litter was apparently recalcitrant. Annual net accumulation of C and N by epiphytic bryophytes was estimated at 37–64 g C/m²/yr and 0.8–1.3 g N/m²/yr, respectively. Previous research at this site indicated that epiphytic bryophytes retain inorganic N from atmospheric deposition to the canopy. Therefore, they play a major role in transforming N from mobile to highly recalcitrant forms in this ecosystem.     

A multiyear synthesis of soil respiration responses to elevated atmospheric CO2 from four forest FACE experiments

The rapidly rising concentration of atmospheric CO₂ has the potential to alter forest and global carbon cycles by altering important processes that occur in soil. Forest soils contain the largest and longest lived carbon pools in terrestrial ecosystems and are therefore extremely important to the land–atmosphere exchange of carbon and future climate. Soil respiration is a sensitive integrator of many soil processes that control carbon storage in soil, and is therefore a good metric of changes to soil carbon cycling. Here, we summarize soil respiration data from four forest free‐air carbon dioxide enrichment (FACE) experiments in developing and established forests that have been exposed to elevated atmospheric [CO₂] (168 μL L^?1 average enrichment) for 2–6 years. The sites have similar experimental design and use similar methodology (closed‐path infrared gas analyzers) to measure soil respiration, but differ in species composition of the respective forest communities. We found that elevated atmospheric [CO₂] stimulated soil respiration at all sites, and this response persisted for up to 6 years. Young developing stands experienced greater stimulation than did more established stands, increasing 39% and 16%, respectively, averaged over all years and communities. Further, at sites that had more than one community, we found that species composition of the dominant trees was a major controller of the absolute soil CO₂ efflux and the degree of stimulation from CO₂ enrichment. Interestingly, we found that the temperature sensitivity of bulk soil respiration appeared to be unaffected by elevated atmospheric CO₂. These findings suggest that stage of stand development and species composition should be explicitly accounted for when extrapolating results from elevated CO₂ experiments or modeling forest and global carbon cycles.     

Severe drought effects on ecosystem CO2 and H2O fluxes at three Mediterranean evergreen sites: revision of current hypotheses?

Eddy covariance and sapflow data from three Mediterranean ecosystems were analysed via top‐down approaches in conjunction with a mechanistic ecosystem gas‐exchange model to test current assumptions about drought effects on ecosystem respiration and canopy CO₂/H₂O exchange. The three sites include two nearly monospecific Quercus ilex L. forests – one on karstic limestone (Puéchabon), the other on fluvial sand with access to ground water (Castelporziano) – and a typical mixed macchia on limestone (Arca di Noè). Estimates of ecosystem respiration were derived from light response curves of net ecosystem CO₂ exchange. Subsequently, values of ecosystem gross carbon uptake were computed from eddy covariance CO₂ fluxes and estimates of ecosystem respiration as a function of soil temperature and moisture. Bulk canopy conductance was calculated by inversion of the Penman‐Monteith equation. In a top‐down analysis, it was shown that all three sites exhibit similar behaviour in terms of their overall response to drought. In contrast to common assumptions, at all sites ecosystem respiration revealed a decreasing temperature sensitivity ( Q ₁₀) in response to drought. Soil temperature and soil water content explained 70–80% of the seasonal variability of ecosystem respiration. During the drought, light‐saturated ecosystem gross carbon uptake and day‐time averaged canopy conductance declined by up to 90%. These changes were closely related to soil water content. Ecosystem water‐use efficiency of gross carbon uptake decreased during the drought, regardless whether evapotranspiration from eddy covariance or transpiration from sapflow had been used for the calculation. We evidence that this clearly contrasts current models of canopy function which predict increasing ecosystem water‐use efficiency (WUE) during the drought. Four potential explanations to those results were identified (patchy stomatal closure, changes in physiological capacities of photosynthesis, decreases in mesophyll conductance for CO₂, and photoinhibition), which will be tested in a forthcoming paper. It is suggested to incorporate the new findings into current biogeochemical models after further testing as this will improve estimates of climate change effects on (semi)arid ecosystems' carbon balances.     

Differential responses of production and respiration to temperature and moisture drive the carbon balance across a climatic gradient in New Mexico

Southwestern North America faces an imminent transition to a warmer, more arid climate, and it is critical to understand how these changes will affect the carbon balance of southwest ecosystems. In order to test our hypothesis that differential responses of production and respiration to temperature and moisture shape the carbon balance across a range of spatio‐temporal scales, we quantified net ecosystem exchange (NEE) of CO₂ and carbon storage across the New Mexico Elevational Gradient, which consists of six eddy‐covariance sites representing biomes ranging from desert to subalpine conifer forest. Within sites, hotter and drier conditions were associated with an increasing advantage of respiration relative to production such that daily carbon uptake peaked at intermediate temperatures – with carbon release often occurring on the hottest days – and increased with soil moisture. Across sites, biotic adaptations modified but did not override the dominant effects of climate. Carbon uptake increased with decreasing temperature and increasing precipitation across the elevational gradient; NEE ranged from a source of ～30 g C m^?2 yr^?1 in the desert grassland to a sink of ～350 g C m^?2 yr^?1 in the subalpine conifer forest. Total aboveground carbon storage increased dramatically with elevation, ranging from 186 g C m^?2 in the desert grassland to 26 600 g C m^?2 in the subalpine conifer forest. These results make sense in the context of global patterns in NEE and biomass storage, and support that increasing temperature and decreasing moisture shift the carbon balance of ecosystems in favor of respiration, such that the potential for ecosystems to sequester and store carbon is reduced under hot and/or dry conditions. This implies that projected climate change will trigger a substantial net release of carbon in these New Mexico ecosystems (～3 Gt CO₂ statewide by the end of the century), thereby acting as a positive feedback to climate change.     

Carbon sequestration in a high-elevation, subalpine forest

We studied net ecosystem CO₂ exchange (NEE) dynamics in a high‐elevation, subalpine forest in Colorado, USA, over a two‐year period. Annual carbon sequestration for the forest was 6.71 mol C m^?2 (80.5 g C m^?2) for the year between November 1, 1998 and October 31, 1999, and 4.80 mol C m^?2 (57.6 g C m^?2) for the year between November 1, 1999 and October 31, 2000. Despite its evergreen nature, the forest did not exhibit net CO₂ uptake during the winter, even during periods of favourable weather. The largest fraction of annual carbon sequestration occurred in the early growing‐season; during the first 30 days of both years. Reductions in the rate of carbon sequestration after the first 30 days were due to higher ecosystem respiration rates when mid‐summer moisture was adequate (as in the first year of the study) or lower mid‐day photosynthesis rates when mid‐summer moisture was not adequate (as in the second year of the study). The lower annual rate of carbon sequestration during the second year of the study was due to lower rates of CO₂ uptake during both the first 30 days of the growing season and the mid‐summer months. The reduction in CO₂ uptake during the first 30 days of the second year was due to an earlier‐than‐normal spring warm‐up, which caused snow melt during a period when air temperatures were lower and atmospheric vapour pressure deficits were higher, compared to the first 30 days of the first year. The reduction in CO₂ uptake during the mid‐summer of the second year was due to an extended drought, which was accompanied by reduced latent heat exchange and increased sensible heat exchange. Day‐to‐day variation in the daily integrated NEE during the summers of both years was high, and was correlated with frequent convective storm clouds and concomitant variation in the photosynthetic photon flux density (PPFD). Carbon sequestration rates were highest when some cloud cover was present, which tended to diffuse the photosynthetic photon flux, compared to periods with completely clear weather. The results of this study are in contrast to those of other studies that have reported increased annual NEE during years with earlier‐than‐normal spring warming. In the current study, the lower annual NEE during 2000, the year with the earlier spring warm‐up, was due to (1) coupling of the highest seasonal rates of carbon sequestration to the spring climate, rather than the summer climate as in other forest ecosystems that have been studied, and (2) delivery of snow melt water to the soil when the spring climate was cooler and the atmosphere drier than in years with a later spring warm‐up. Furthermore, the strong influence of mid‐summer precipitation on CO₂ uptake rates make it clear that water supplied by the spring snow melt is a seasonally limited resource, and summer rains are critical for sustaining high rates of annual carbon sequestration.     

Spring warming and carbon dioxide exchange over low Arctic tundra in central Canada

Tundra‐atmosphere exchanges of carbon dioxide (CO₂) and water vapour were measured near Daring Lake, Northwest Territories in the Canadian Low Arctic for 3 years, 2004–2006. The measurement period spanned late‐winter until the end of the growing period. Mean temperatures during the measurement period varied from about 2 °C less than historical average in 2004 and 2005 to 2 °C greater in 2006. Much of the added warmth in 2006 occurred at the beginning of the study, when snow melt occurred 3 weeks earlier than in the other years. Total precipitation in 2006 (163 mm) was more than double that of the driest year, 2004 (71 mm). The tundra was a net sink for CO₂ carbon in all years. Mid‐summer net ecosystem exchange of CO₂ (NEE) achieved maximum values of ?1.3 g C m^?2 day^?1 (2004) to ?1.8 g C m^?2 day^?1 (2006). Accumulated NEE values over the 109‐day period were ?32,?51 and ?61 g C m^?2 in 2004, 2005 and 2006, respectively. The larger CO₂ uptake in 2006 was attributed to the early spring coupled with warmer air and soil conditions. In 2004, CO₂ uptake was limited by the shorter growing season and mid‐summer dryness, which likely reduced ecosystem productivity. Seasonal total evapotranspiration (ET) ranged from 130 mm (2004) to 181 mm (2006) and varied in accordance with the precipitation received and with the timing of snow melt. Maximum daily ET rates ranged from 2.3 to 2.7 mm day^?1, occurring in mid July. Ecosystem water use efficiency (WUE_eco) varied slightly between years, ranging from 2.2 in the driest year to 2.5 in the year with intermediate rainfall amounts. In the wettest year, increased soil evaporation may have contributed to a lower WUE_eco (2.3). We speculate that most, if not all, of the modest growing season CO₂ sink measured at this site could be lost due to fall and winter respiration leading to the tundra being a net CO₂ source or CO₂ neutral on an annual basis. However, this hypothesis is untested as yet.     

Ecosystem CO2 exchange and plant biomass in the littoral zone of a boreal eutrophic lake

• 1 In order to study the dynamics of primary production and decomposition in the lake littoral, an interface zone between the pelagial, the catchment and the atmosphere, we measured ecosystem/atmosphere carbon dioxide (CO₂) exchange in the littoral zone of an eutrophic boreal lake in Finland during two open water periods (1998–1999). We reconstructed the seasonal net CO₂ exchange and identified the key factors controlling CO₂ dynamics. The seasonal net ecosystem exchange (NEE) was related to the amount of carbon accumulated in plant biomass.
• 2 In the continuously inundated zones, spatial and temporal variation in the density of aerial shoots controlled CO₂ fluxes, but seasonal net exchange was in most cases close to zero. The lower flooded zone had a net CO₂ uptake of 1.8–6.2 mol m^?2 per open water period, but the upper flooded zone with the highest photosynthetic capacity and above‐ground plant biomass, had a net CO₂ loss of 1.1–7.1 mol m^?2 per open water period as a result of the high respiration rate. The excess of respiration can be explained by decomposition of organic matter produced on site in previous years or leached from the catchment.
• 3 Our results from the two study years suggest that changes in phenology and water level were the prime cause of the large interannual difference in NEE in the littoral zone. Thus, the littoral is a dynamic buffer and source for the load of allochthonous and autochthonous carbon to small lakes.

     

High rates of net ecosystem carbon assimilation by Brachiara pasture in the Brazilian Cerrado

To investigate the consequences of land use on carbon and energy exchanges between the ecosystem and atmosphere, we measured CO₂ and water vapour fluxes over an introduced Brachiara brizantha pasture located in the Cerrado region of Central Brazil. Measurements using eddy covariance technique were carried out in field campaigns during the wet and dry seasons. Midday CO₂ net ecosystem exchange rates during the wet season were ?40 μmol m^?2 s^?1, which is more than twice the rate found in the dry season (?15 μmol m^?2 s^?1). This was observed despite similar magnitudes of irradiance, air and soil temperatures. During the wet season, inferred rates of canopy photosynthesis did not show any tendency to saturate at high solar radiation levels, with rates of around 50 μmol m^?2 s^?1 being observed at the maximum incoming photon flux densities of 2200 μmol m^?2 s^?1. This contrasted strongly to the dry period when light saturation occurred with 1500 μmol m^?2 s^?1 and with maximum canopy photosynthetic rates of only 20 μmol m^?2 s^?1. Both canopy photosynthetic rates and night‐time ecosystem CO₂ efflux rates were much greater than has been observed for cerrado native vegetation in both the wet and dry seasons. Indeed, observed CO₂ exchange rates were also much greater than has previously been reported for C₄ pastures in the tropics. The high rates in the wet season may have been attributable, at least in part, to the pasture not being grazed. Higher than expected net rates of carbon acquisition during the dry season may also have been attributable to some early rain events. Nevertheless, the present study demonstrates that well‐managed, productive tropical pastures can attain ecosystem gas exchange rates equivalent to fertilized C₄ crops growing in the temperate zone.     

Old trees contribute bio-available nitrogen through canopy bryophytes

Symbiotic cyanobacteria??bryophyte associations on the forest floor are shown to contribute significantly to stand-level nitrogen budgets through the process of biological nitrogen fixation (BNF), but few studies have considered the role of canopy bryophytes. Given the high biomass of epiphytic bryophytes in many tree species of the North American temperate rain forest, we suggest that canopy bryophytes may contribute substantially to stand-level N dynamics. We confirm the presence of cyanobacteria and measure rates of BNF at three heights (0, 15 and 30 m) in Sitka spruce trees across three watershed estuaries of Clayoquot Sound, British Columbia, Canada. This study is the first to report BNF by cyanobacteria associated with epiphytic and forest floor bryophytes in the coastal temperate rain forest of North America. Cyanobacteria density was significantly greater in epiphytic bryophytes compared to mosses on the forest floor, and rates of BNF were highest at 30 m in the canopy. The majority of total stand-level BNF (0.76 kg N · ha^-1 · yr^-1) occurs in the canopy, rather than on the forest floor (0.26 kg N · ha^-1 · yr^-1). We suggest that BNF by cyanobacterial-bryophyte associations in the canopy of coastal temperate rain forests is a unique source of ecosystem N, which is dependent on large, old trees with high epiphytic bryophyte biomass.