Seasonal and diurnal CO2 efflux from red wood ant (Formica aquilonia) mounds in boreal coniferous forests

Organic mounds of the red wood ants (Formica rufa group; RWA) have been shown to be “hot spots” of carbon dioxide (CO₂) efflux from the European forest soils. However, little information is available on the variability of CO₂ effluxes from RWA mounds and on the factors regulating CO₂ efflux. We assessed the seasonal and diurnal changes in CO₂ effluxes, temperatures and volumetric water contents from mounds of Formica aquilona, the important RWA of the boreal forests in Finland. The daily average CO₂ efflux from RWA mounds ranged 1.1-6.9 g CO₂ m⁻² h⁻¹ during the active ant season (May-September), and from 0.2 to 1.1 g CO₂ m⁻² h⁻¹ during their dormant period (October-April). Mound CO₂ efflux from May to September was 3.6-6.0 times higher than from the surrounding forest floors, and most likely came from RWA respiration. Seasonal changes in mound CO₂ effluxes were significantly correlated with mound temperature, but not with volumetric water content (7% on average). The high CO₂ efflux associated with increased volumetric water content (up to 34%) after a RWA mound was abandoned indicated that these dry mound conditions restrict microbial decomposition of mound organic matter. CO₂ effluxes were highest at night and lowest during the day, which is likely due to an increased ant activity or numbers in the mound at night. Diurnal changes in mound CO₂ efflux were negatively correlated with air temperature, and positively correlated with the difference between the mound and air temperature. This suggests that thermal convection of warmer mound air to the colder outside air at night might be also a cause of the diurnal changes. We conclude that seasonal and diurnal variations in mound CO₂ effluxes are dependent on RWA activities and fluctuation in RWA mound and outside temperatures.     

Parasitoids (Hymenoptera, Ichneumonoidea) of Saproxylic beetles are affected by forest successional stage and dead wood characteristics in boreal spruce forest

The habitat requirements and effects of forest management on insects belonging to higher trophic levels are relatively unknown in forest ecosystems. We tested the effect of forest successional stage and dead wood characteristics on the saproxylic parasitoid (Hymenoptera, Ichneumonoidea) assemblage in boreal spruce-dominated forests in northern Sweden. Within each of nine areas, we selected three sites with different management histories: (1) a clear-cut (2) a mature managed forest and (3) an old-growth forest. Parasitoids were collected in 2003 using eclector traps mounted on fresh logs, which were either untreated (control), burned, inoculated with fungi, or naturally shaded, and on artificially-created snags.Both forest type and dead wood characteristics had a significant effect on parasitoid assemblages. Grouped idiobionts and some species, such as Bracon obscurator and Ontsira antica, preferred clear-cuts, while others, such as Cosmophorus regius (Hym., Braconidae) and other koinobionts, were associated with older successional forest stages. No single dead wood substrate was sufficient to support the entire community of parasitoids in any forest type, even when the regular host was present. In particular, snags hosted a different assemblage of species from other types of dead wood, with parasitoids of Tetropium spp. such as Rhimphoctona spp. (Hym., Ichneumonidae) and Helconidea dentator (Hym., Braconidae) being abundant. These results indicate that a diversity of dead wood habitats is necessary to support complete assemblages of beetle-associated parasitoids from early successional stages of dead wood and that parasitoids may be more sensitive to habitat change than their hosts.     

Measuring forest floor CO2 fluxes in a Douglas-fir forest

CO₂ exchange was measured on the forest floor of a coastal temperate Douglas-fir forest located near Campbell River, British Columbia, Canada. Continuous measurements were obtained at six locations using an automated chamber system between April and December, 2000. Fluxes were measured every half hour by circulating chamber headspace air through a sampling manifold assembly and a closed-path infrared gas analyzer. Maximum CO₂ fluxes measured varied by a factor of almost 3 between the chamber locations, while the highest daily average fluxes observed at two chamber locations occasionally reached values near 15 μmol C m⁻² s⁻¹. Generally, fluxes ranged between 2 and 10 μmol C m⁻² s⁻¹ during the measurement period. CO₂ flux from the forest floor was strongly related to soil temperature with the highest correlation found with 5 cm depth temperature. A simple temperature dependent exponential model fit to the nighttime fluxes revealed Q₁₀ values in the normal range of 2–3 during the warmer parts of the year, but values of 4–5 during cooler periods. Moss photosynthesis was negligible in four of the six chambers, while at the other locations, it reduced daytime half-hourly net CO₂ flux by about 25%. Soil moisture had very little effect on forest floor CO₂ flux. Hysteresis in the annual relationship between chamber fluxes and soil temperatures was observed. Net exchange from the six chambers was estimated to be 1920±530 g C m⁻² per year, the higher estimates exceeding measurement of ecosystem respiration using year-round eddy correlation above the canopy at this site. This discrepancy is attributed to the inadequate number of chambers to obtain a reliable estimate of the spatial average soil CO₂ flux at the site and uncertainty in the eddy covariance respiration measurements.     

Decomposition of organic matter and nutrient mineralisation in wood ant (<Emphasis Type="Italic">Formica rufa</Emphasis> group) mounds in boreal coniferous forests of different age

Wood ants (Formica rufa group) are dominating ecosystem elements of the boreal region due to their wide and abundant occurrence. They collect and concentrate organic material from the surrounding forest floor by building large above-ground mounds. These mounds have higher temperature and lower water content than the surrounding forest floor. We studied how these peculiar environmental conditions affected mass loss and carbon (C), nitrogen (N), phosphorus (P) and potassium (K) mineralisation of organic matter in boreal Norway spruce (Picea abies L. Karst.)-dominated mixed forest stands of four different age classes (5-, 30-, 60-, and 100-year-old) situated in eastern Finland using the litter bag technique. Norway spruce needle litter was incubated in inhabited and abandoned wood ant mounds as well as on the surrounding forest floor. We expected decomposition to be extremely slow due to the dryness of the mounds. Mass losses inside inhabited mounds were lower compared to the surrounding forest floor (on average 30 vs 50% after 2 years) but not as low as we expected, which might be a result of ant and microbial activity in the mounds. Decomposition in the abandoned mounds proceeded similarly as on the forest floor. Nutrient mineralisation proceeded more slowly in the ant mounds than on the surrounding forest floor. Mineralisation occurred for all studied nutrients in the ant mounds, except for N, which net amount remained stable during the years of the experiment. When wood ant mounds are abandoned and their porous and dry structure is no longer maintained by the ants, their decomposition is accelerated, and nutrients may be available for uptake by plants, although the nutrient mineralisation seems still to remain lower compared to the surrounding forest floor. However, eventually the mound material will be decomposed and nutrients mineralised, thus providing a nutrient hot spot increasing the heterogeneity of forest floor nutrient availability.     

Biotic and abiotic factors regulating forest floor CO2 flux across a range of forest age classes in the southern Appalachians

We measured forest floor CO₂ flux in three age classes of forest in the southern Appalachians: 20-year-old, 85-year-old, and old-growth. Our objectives were to quantify differences in forest floor CO₂ flux among age classes, and determine the relative importance of abiotic and biotic driving variables. Forest floor CO₂ flux was measured using an openflow infrared gas analyzer measurement system for 24 h periods and samples were taken every 2 months over a 2-year period. Litter/soil interface, soil temperature (5 cm depth), soil moisture (%), forest floor moisture (%), forest floor mass, fine root (2 mm) mass, coarse root mass (>2 mm), forest floor C and N (%), fine root C and N, coarse root C and N, and soil N and C were co-measured during each sample period. Results showed significant nonlinear relationships (r²=0.68 to 0.81) between litter/soil interface temperature and forest floor CO₂ flux for all three forest age classes, but no differences in temperature response parameters. These results indicated no differences in forest floor CO₂ flux among age classes. Considerable temporal variation in abiotic and biotic variables was observed within and among forests. Biotic variables correlated with forest floor CO₂ flux included indices of litter and root quality. Differences in biotic variables correlated with forest floor CO₂ flux among forests may have been related to shifts in the relative importance of heterotrophic and autotrophic respiration components to overall forest floor CO₂ flux.     

Explaining temporal variation in soil CO2 efflux in a mature spruce forest in Southern Germany

An open dynamic chamber system was used to measure the soil CO₂ efflux intensively and continuously throughout a growing season in a mature spruce forest (Picea abies) in Southern Germany. The resulting data set contained a large amount of temporally highly resolved information on the variation in soil CO₂ efflux together with environmental variables. Based on this background, the dependencies of the soil CO₂ efflux rate on the controlling environmental factors were analysed in-depth. Of the abiotic factors, soil temperature alone explained 72% of the variation in the efflux rate, and including soil water content (SWC) as an additional variable increased the explained variance to about 83%. Between April and December, average rates ranged from 0.43 to 5.15 μmol CO₂ m⁻² s⁻¹ (in November and July, respectively) with diurnal variations of up to 50% throughout the experiment. The variability in wind speed above the forest floor influenced the CO₂ efflux rates for measuring locations with a litter layer of relatively low bulk density (and hence relatively high proportions of pore spaces). For the temporal integration of flux rates for time scales of hours to days, however, wind velocities were of no effect, reflecting the fact that wind forcing acts on the transport, but not the production of CO₂ in the soil. The variation in both the magnitude of the basal respiration rate and the temperature sensitivity throughout the growing season was only moderate (coefficient of variation of 15 and 25%, respectively). Soil water limitation of the CO₂ production in the soil could be best explained by a reduction in the temperature-insensitive basal respiration rate, with no discernible effect on the temperature sensitivity. Using a soil CO₂ efflux model with soil temperature and SWC as driving variables, it was possible to calculate the annual soil CO₂ efflux for four consecutive years for which meteorological data were available. These simulations indicate an average efflux sum of 560 g C m⁻² yr⁻¹ (SE=22 g C m⁻² yr⁻¹). An alternative model derived from the same data but using temperature alone as a driver over-estimated the annual flux sum by about 7% and showed less inter-annual variability. Given a likely shift in precipitation patterns alongside temperature changes under projected global change scenarios, these results demonstrate the necessity to include soil moisture in models that calculate the evolution of CO₂ from temperate forest soils.     

Fluxes of CO2, CH4 and N2O from drained organic soils in deciduous forests

We examined net greenhouse gas exchange at the soil surface in deciduous forests on soils with high organic contents. Fluxes of CO₂, CH₄ and N₂O were measured using dark static chambers for two consecutive years in three different forest types; (i) a drained and medium productivity site dominated by birch, (ii) a drained and highly productive site dominated by alder and (iii) an undrained and highly productive site dominated by alder. Although the drained sites had shallow mean groundwater tables (15 and 18 cm, respectively) their average annual rates of forest floor CO₂ release were almost twice as high compared to the undrained site (1.9±0.4 and 1.7±0.3, compared to 1.0±0.2 kg CO₂ m⁻² yr⁻¹). The average annual CH₄ emission was almost 10 times larger at the undrained site (7.6±3.1 compared to 0.9±0.5 g CH₄ m⁻² yr⁻¹ for the two drained sites). The average annual N₂O emissions at the undrained site (0.1±0.05 g N₂O m⁻² yr⁻¹) were lower than at the drained sites, and the emissions were almost five times higher at the drained alder site than at the drained birch site (0.9±0.35 compared to 0.2±0.11 g N₂O m⁻² yr⁻¹). The temporal variation in forest floor CO₂ release could be explained to a large extent by differences in groundwater table and air temperature, but little of the variation in the CH₄ and N₂O fluxes could be explained by these variables. The measured soil variables were only significant to explain for the within-site spatial variation in CH₄ and N₂O fluxes at the undrained swamp, and dark forest floor CO₂ release was not explained by these variables at any site. The between-site spatial variation was attributed to variations in drainage, groundwater level position, productivity and tree species for all three gases. The results indicate that N₂O emissions are of greater importance for the net greenhouse gas exchange at deciduous drained forest sites than at coniferous drained forest sites.     

Increasing net CO2 uptake by a Danish beech forest during the period from 1996 to 2009

The exchange of CO₂ between the atmosphere and a beech forest near Sorø, Denmark, was measured continuously over 14 years (1996-2009). The simultaneous measurement of many parameters that influence CO₂ uptake makes it possible to relate the CO₂ exchange to recent changes in e.g. temperature and atmospheric CO₂ concentration. The net CO₂ exchange (NEE) was measured by the eddy covariance method. Ecosystem respiration (RE) was estimated from nighttime values and gross ecosystem exchange (GEE) was calculated as the sum of RE and NEE. Over the years the beech forest acted as a sink of on average of 157 g C m⁻² yr⁻¹. In one of the years only, the forest acted as a small source. During 1996-2009 a significant increase in annual NEE was observed. A significant increase in GEE and a smaller and not significant increase in RE was also found. Thus the increased NEE was mainly attributed to an increase in GEE. The overall trend in NEE was significant with an average increase in uptake of 23 g C m⁻² yr⁻². The carbon uptake period (i.e. the period with daily net CO₂ gain) increased by 1.9 days per year, whereas there was a non significant tendency of increase of the leafed period. This means that the leaves stayed active longer. The analysis of CO₂ uptake by the forest by use of light response curves, revealed that the maximum rate of photosynthetic assimilation increased by 15% during the 14-year period. We conclude that the increase in the overall CO₂ uptake of the forest is due to a combination of increased growing season length and increased uptake capacity. We also conclude that long time series of flux measurements are necessary to reveal trends in the data because of the substantial inter-annual variation in the flux.     

Seasonal changes in the spatial structures of N2O, CO2, and CH4 fluxes from Acacia mangium plantation soils in Indonesia

We evaluated the spatial structures of nitrous oxide (N₂O), carbon dioxide (CO₂), and methane (CH₄) fluxes in an Acacia mangium plantation stand in Sumatra, Indonesia, in drier (August) and wetter (March) seasons. A 60 × 100-m plot was established in an A. mangium plantation that included different topographical elements of the upper plateau, lower plateau, upper slope and foot slope. The plot was divided into 10 × 10-m grids and gas fluxes and soil properties were measured at 77 grid points at 10-m intervals within the plot. Spatial structures of the gas fluxes and soil properties were identified using geostatistical analyses. Averaged N₂O and CO₂ fluxes in the wetter season (1.85 mg N m⁻² d⁻¹ and 4.29 g C m⁻² d⁻¹, respectively) were significantly higher than those in the drier season (0.55 mg N m⁻² d⁻¹ and 2.73 g C m⁻² d⁻¹, respectively) and averaged CH₄ uptake rates in the drier season (−0.62 mg C m⁻² d⁻¹) were higher than those in the wetter season (−0.24 mg C m⁻² d⁻¹). These values of N₂O fluxes in A. mangium soils were higher than those reported for natural forest soils in Sumatra, while CO₂ and CH₄ fluxes were in the range of fluxes reported for natural forest soils. Seasonal differences in these gas fluxes appears to be controlled by soil water content and substrate availability due to differing precipitation and mineralization of litter between seasons. N₂O fluxes had strong spatial dependence with a range of about 18 m in both the drier and wetter seasons. Topography was associated with the N₂O fluxes in the wetter season with higher and lower fluxes on the foot slope and on the upper plateau, respectively, via controlling the anaerobic-aerobic conditions in the soils. In the drier season, however, we could not find obvious topographic influences on the spatial patterns of N₂O fluxes and they may have depended on litter amount distribution. CO₂ fluxes had no spatial dependence in both seasons, but the topographic influence was significant in the drier season with lowest fluxes on the foot slope, while there was no significant difference between topographic positions in the wetter season. The distributions of litter amount and soil organic matter were possibly associated with CO₂ fluxes through their effects on microbial activities and fine root distribution in this A. mangium plantation.     

CH4 oxidation and emissions of CH4 and N2O from Lolium perenne swards under elevated atmospheric CO2

Emissions of N₂O and CH₄ and CH₄ oxidation rates were measured from Lolium perenne swards in a short-term study under ambient (36 Pa) and elevated (60 Pa) atmospheric CO₂ at the Free Air Carbon dioxide Enrichment experiment, Eschikon, Switzerland. Elevated pCO₂ increased (P<0.05) N₂O emissions from high N fertilised (11.2 g N m⁻²) swards by 69%, but had no significant effect on net emissions of CH₄. Application of ¹³C-CH₄ (11 μl l⁻¹; 11 at.% excess ¹³C) to closed chamber headspaces in microplots enabled determination of rates of ¹³C-CH₄ oxidation even when net CH₄ fluxes from main plots were positive. We found a significant interaction between fertiliser application rate and atmospheric pCO₂ on ¹³C-CH₄ oxidation rates that was attributed to differences in gross nitrification rates and C and N availability. CH₄ oxidation was slower and thought to be temporarily inhibited in the high N ambient pCO₂ sward. The most rapid CH₄ oxidation of 14.6 μg ¹³C-CH₄ m⁻² h⁻¹ was measured in the high fertilised elevated pCO₂ sward, and we concluded that either elevated pCO₂ had a stimulatory effect on CH₄ oxidation or inhibition of oxidation following fertiliser application was lowered under elevated pCO₂. Application of ¹⁴NH₄¹⁵NO₃ and ¹⁵NH₄¹⁵NO₃ (10 at.% excess ¹⁵N) to different replicates enabled determination of the respective contributions of nitrification and denitrification to N₂O emissions. Inhibition of CH₄ oxidation in the high fertilised ambient pCO₂ sward, due to competition between NH₃ and CH₄ for methane monooxygenase enzymes or toxic effects of NH₂OH or NO₂⁻ produced during nitrification, was hypothesised to increase gross nitrification (12.0 mg N kg dry soil⁻¹) and N₂O emissions during nitrification (327 mg ¹⁵N-N₂O m⁻² over 11 d). Our results indicate that increasing atmospheric concentrations of CO₂ may increase emissions of N₂O by denitrification, lower nitrification rates and either increase or decrease the ability of soil to act as a sink for atmospheric CH₄ depending on fertiliser management.     

Effects of root and litter exclusion on soil CO2 efflux and microbial biomass in wet tropical forests

We examined the effects of root and litter exclusion on the rate of soil CO₂ efflux and microbial biomass at a soil depth of 25 cm in a secondary forest (dominated by Tabebuia heterophylla) and a pine (Pinus caribaea) plantation in the Luquillo Experimental Forest in Puerto Rico. The experimental plots were initially established in 1990, when root, forest floor mass and new litterfall were excluded for 7 y since then. Soil respiration was significantly reduced in the litter and root exclusion plots in both the secondary forest and the pine plantation compared with the control. Root exclusion had a greater effect on soil CO₂ efflux than the litter exclusion in the plantation, whereas a reversed pattern was observed in the secondary forest. The reduction of microbial biomass in the root exclusion plot was greater in the secondary forest (59%) than in the plantation (31%), while there was no difference of the reduction in the litter exclusion plots between these forests. Our results suggest that above-ground input and roots (root litter and exudates) differentially affect soil CO₂ efflux under different vegetation types.     

Surface energy balance closure by the eddy-covariance method above three boreal forest stands and implications for the measurement of the CO2 flux

Closure of the surface energy balance provides an objective criterion for evaluating eddy-covariance (EC) flux measurements. This study analyses 5 years of EC carbon dioxide, water vapor, and sensible heat flux measurements from three mature boreal forest stands in central Saskatchewan, Canada. The EC sensible and latent heat fluxes, H and λE, underestimated the surface available energy by 11% (aspen), 15% (black spruce), and 14% (jack pine). At all sites, the energy-closure fraction CF responded similarly to the friction velocity u_*, atmospheric stability, and time of day. At night, CF increased from 0.3 at very low-u_* to an asymptotic maximum of 0.9 at u_* above 0.35 m s⁻¹. During unstable-daytime periods, CF varied linearly from 0.7 at low-u_* to 1.0 at high-u_*. The energy imbalance pattern was similar among sites and may be characteristic of the continental, boreal forest.EC measurements of net ecosystem exchange F_NEE have no objective, diagnostic parameter that is equivalent to CF. We therefore derived an analogous F_NEE “closure fraction” CF^NEE by normalizing measured F_NEE against estimates from an empirical model that was tuned to the high-u_* data. CF and CF^NEE responded similarly to u_*, atmospheric stability, and time of day. We discuss two implications for EC flux data post-processing. The results uphold the common practice of rejecting EC measurements during low-u_* periods. They also lend support to the application of energy-closure adjustments to H, λE, and F_NEE.     

Larger floods reduce soil CO2 efflux during the post-flooding phase in seasonally-flooded forests of Western Amazonia

Seasonally flooded várzea forests of Western Amazonia are one of the most productive and biodiverse wetland forests in the world.However,data on their soil CO₂emissions,soil organic matter decomposition rates,and soil C stocks are scarce.This is a concern because hydrological changes are predicted to lead to increases in the height,extent,and duration of seasonal floods,which are likely to have a significant effect on soil C stocks and fluxes.However,with no empirical data,the impact of altered flood regimes on várzea soil C cycles remains uncertain.This study quantified the effects of maximum annual flood height and soil moisture on soil CO₂efflux rate(R_s)and soil organic matter decomposition rate(k)in the várzea forests of Pacaya Samiria National Reserve,Peru.The study was conducted between May and August 2017.The results showed that R_s(10.6–182.7 mg C m^-2h^-1)and k(0.016–0.078)varied between and within sites,and were considerably lower than the values reported for other tropical forests.In addition,R_swas negatively affected by flood height(P<0.01)and soil moisture(P<0.001),and it decreased with decreasing river levels post flooding(P<0.001).In contrast,k was not affected by any of the above-mentioned factors.Soil moisture was the dominant factor influencing R_s,and it was significantly affected by maximum flood height,even after the floods had subsided(P<0.001).Consequently,we concluded that larger floods will likely lead to reduced R_s,whilst k could remain unchanged but with decomposition processes becoming more anaerobic.     

Temperature sensitivity of respiration differs among forest floor layers in a Pinus resinosa plantation

Decomposer microorganisms contribute to carbon loss from the forest floor as they metabolize organic substances and respire CO₂. In temperate and boreal forest ecosystems, the temperature of the forest floor can fluctuate significantly on a day-to-night or day-to-day basis. In order to estimate total respiratory CO₂ loss over even relatively short durations, therefore, we need to know the temperature sensitivity (Q₁₀) of microbial respiration. Temperature sensitivity has been calculated for microbes in different soil horizons, soil fractions, and at different depths, but we would suggest that for some forests, other ecologically relative soil portions should be considered to accurately predict the contribution of soil to respiration under warming. The floor of many forests is heterogeneous, consisting of an organic horizon comprising a few more-or-less distinct layers varying in decomposition status. We therefore determined at various measurement temperatures the respiration rates of litter, F-layer, and H-layer collected from a Pinus resinosa plantation, and calculated Q₁₀ values for each layer. Q₁₀ depended on measurement temperature, and was significantly greater in H-layer than in litter or F-layer between 5 and 17 °C. Our results indicate, therefore, that as the temperature of the forest floor rises, the increase in respiration by the H-layer will be disproportionate to the increase by other layers. However, change in respiration by the H-layer associated with change in temperature may contribute minimally or significantly to changes of total forest floor respiration in response to changes in temperature depending on the depth and thickness of the layer in different forest ecosystems.     

Factors controlling soil CO2 effluxes and the effects of rewetting on effluxes in adjacent deciduous, coniferous, and mixed forests in Korea

To better understand the factors that control forest soil CO₂ efflux and the effects of rewetting on efflux, we measured soil CO₂ efflux in adjacent deciduous, coniferous, and mixed forests in the central part of the Korean Peninsula over the course of one year. We also conducted laboratory rewetting experiments with soil collected from the three sites using three different incubation temperatures (4 °C, 10 °C, and 20 °C). Soil moisture (SM), soil organic matter (SOM), and total root mass values of the three sites were significantly different from one another; however, soil temperature (ST), observed soil CO₂ efflux and sensitivity of soil CO₂ efflux to ST (i.e., Q₁₀ = 3.7 ± 0.1) were not significantly different among the three sites. Soil temperature was a dominant control factor regulating soil CO₂ efflux during most of the year. We infer that soil CO₂ efflux was not significantly different among the sites due to similar ST and Q₁₀. Though a significant increase in soil CO₂ efflux following rewetting of dry soil was observed both in the field observations (60-170%) and laboratory incubation experiments (100-1000%), both the increased rates of soil CO₂ efflux and the magnitude of change in SM were not significantly different among the sites. The increased rates of soil CO₂ efflux following rewetting depended on the initial SM before rewetting. During drying phase after rewetting, a significant correlation between SM and soil CO₂ efflux was found, but the effect of ST on increased soil CO₂ efflux was not clear. Cumulative peak soil CO₂ efflux (11.3 ± 0.7 g CO₂ m⁻²) following rewetting in the field was not significantly different among the sites. Those evidences indicate that the observed similar rewetting effects on soil CO₂ efflux can be explained by the similar magnitude of change in SM after rewetting at the sites. We conclude that regardless of vegetation type, soil CO₂ efflux and the effect of rewetting on soil CO₂ efflux do not differ among the sites, and ST is a primary control factor for soil CO₂ efflux while SM modulates the effect of rewetting on soil CO₂ efflux. Further studies are needed to quantify and incorporate relationship of initial dryness of the soil and the frequency of the dry-wet cycle on soil CO₂ efflux into models describing carbon (C) processes in forested ecosystems.     

The magnitude and variability of soil-surface CO2 efflux increase with mean annual temperature in Hawaiian tropical montane wet forests

Soil-surface CO₂ efflux (F_S; ‘soil respiration’) accounts for ≥50% of the CO₂ released annually by the terrestrial biosphere to the atmosphere, and the magnitude and variability of this flux are likely to be sensitive to climate change. We measured F_S in nine permanent plots along a 5.2 °C mean annual temperature (MAT) gradient (13-18.2 °C) in Hawaiian tropical montane wet forests where substrate type and age, soil type, soil water balance, disturbance history, and canopy vegetation are constant. The objectives of this study were to quantify how the (i) magnitude, (ii) plot-level spatial variability, and (iii) plot-level diel variability of F_S vary with MAT. To address the first objective, annual F_S budgets were constructed by measuring instantaneous F_S monthly in all plots for one year. For the second objective, we compared plot-level mean instantaneous F_S in six plots derived from 8 versus 16 measurements, and conducted a power analysis to determine adequate sample sizes. For the third objective, we measured instantaneous F_S hourly for 24 h in three plots (cool, intermediate and warm MATs). The magnitude of annual F_S and the spatial variability of plot-level instantaneous F_S increased linearly with MAT, likely due to concomitant increases in stand productivity. Mean plot-level instantaneous F_S from 8 versus 16 measurements per plot yielded statistically similar patterns. The number of samples required to estimate plot-level instantaneous F_S within 10% and 20% of the actual mean increased with MAT. In two of three plots examined, diel variability in instantaneous F_S was significantly correlated with soil temperature but minimal diel fluctuations in soil temperature (<0.6 °C) resulted in minimal diel variability in F_S. Our results suggest that as MAT increases in tropical montane wet forests, F_S will increase and become more spatially variable if ecosystem characteristics and functioning undergo concurrent changes as measured along this gradient. However, diel variation in F_S will remain a minor component of overall plot-level variation.     

Sources of CO2 efflux from soil and review of partitioning methods

Five main biogenic sources of CO₂ efflux from soils have been distinguished and described according to their turnover rates and the mean residence time of carbon. They are root respiration, rhizomicrobial respiration, decomposition of plant residues, the priming effect induced by root exudation or by addition of plant residues, and basal respiration by microbial decomposition of soil organic matter (SOM). These sources can be grouped in several combinations to summarize CO₂ efflux from the soil including: root-derived CO₂, plant-derived CO₂, SOM-derived CO₂, rhizosphere respiration, heterotrophic microbial respiration (respiration by heterotrophs), and respiration by autotrophs. These distinctions are important because without separation of SOM-derived CO₂ from plant-derived CO₂, measurements of total soil respiration have very limited value for evaluation of the soil as a source or sink of atmospheric CO₂ and for interpreting the sources of CO₂ and the fate of carbon within soils and ecosystems. Additionally, the processes linked to the five sources of CO₂ efflux from soil have various responses to environmental variables and consequently to global warming. This review describes the basic principles and assumptions of the following methods which allow SOM-derived and root-derived CO₂ efflux to be separated under laboratory and field conditions: root exclusion techniques, shading and clipping, tree girdling, regression, component integration, excised roots and insitu root respiration; continuous and pulse labeling, ¹³C natural abundance and FACE, and radiocarbon dating and bomb-¹⁴C. A short sections cover the separation of the respiration of autotrophs and that of heterotrophs, i.e. the separation of actual root respiration from microbial respiration, as well as methods allowing the amount of CO₂ evolved by decomposition of plant residues and by priming effects to be estimated. All these methods have been evaluated according to their inherent disturbance of the ecosystem and C fluxes, and their versatility under various conditions. The shortfalls of existing approaches and the need for further development and standardization of methods are highlighted.     

Can a mixed stand of N2-fixing and non-fixing plants restrict N2O emissions with increasing CO2 concentration?

Initial effects of elevated atmospheric CO₂ concentration on N₂O fluxes and biomass production of timothy/red clover were studied in the laboratory. The experimental design consisted of two levels of atmospheric CO₂ (ca. 360 and 720 μmol CO₂ mol⁻¹) and two N fertilisation levels (5 and 10 g N m⁻²). There was a total of 36 mesocosms comprising sandy loam soil, which were equally distributed in four thermo-controlled greenhouses. In two of the greenhouses, the CO₂ concentration was kept at ambient concentration and in the other two at doubled concentration. Forage was harvested and the plants fertilised three times during the basic experiment, followed by harvest, a fertilisation with the double amount of nitrogen and rise of water level. Under elevated CO₂, harvestable and total aboveground dry biomass production of a mixed Trifolium/Phleum stand was increased at both N treatments compared to ambient CO₂. The N₂O flux rates under ambient CO₂ were significantly higher at both N treatments during the early growth of mixed Phleum/Trifolium mesocosms compared to the N₂O flux rate under elevated CO₂. However, when the conditions were favourable for denitrification at the end of the experiment, i.e. N availability and soil moisture were high enough, the elevated CO₂ concentration enhanced the N₂O efflux.     

Concentration of sugars, phenolic acids, and amino acids in forest soils exposed to elevated atmospheric CO2 and O3

Concentrations of soluble soil sugars, soluble phenolic acids, and free amino acids were measured in three forest communities at the FACTS-II Aspen FACE Site near Rhinelander, WI, in order to better understand how elevated atmospheric CO₂ and O₃ are influencing soil nutrient availability and cycling. Sugars, phenolic acids, and amino acids are mostly derived from plant and microbial processes, and have the potential to be influenced by changes in carbon inputs. We hypothesized that concentrations in the soil would parallel increases seen in biological activity, due to greater net primary productivity under elevated CO₂ and seasonal patterns of root growth. Chemical analysis of soils revealed marginally significant increases of total soluble sugars and total soluble phenolic acids in the elevated CO₂ treatment (+27 mg kg⁻¹, +0.02 μmol g⁻¹), but there were no significant differences in concentrations due to elevated O₃ or CO₂+O₃. Total free amino acid concentrations were not affected by any of the treatments, but significant shifts in individual amino acids were observed. Elevated CO₂ and the interaction treatment (elevated CO₂+O₃) increased aspartic acid concentrations, while elevated O₃ treatment decreased the concentration of valine. Concentrations of sugars increased throughout the growing season, while phenolic acids were constant and amino acids decreased. The birch-aspen community had the highest concentration of phenolic acids and sugars overall, while maple-aspen had the lowest. These findings suggest that concentrations of soluble sugars, soluble phenolic acids, and free amino acids in the soil are strongly influenced by soil properties, plant and microbial activity, plant community composition, and to a lesser degree, changes in atmospheric CO₂ and O₃.     

Soil CO2 efflux in a temperate deciduous forest: Environmental drivers and component contributions

Soil CO₂ efflux is a large component of total respiration in many ecosystems. It is important to understand the environmental controls on soil CO₂ efflux, in order to evaluate potential responses of ecosystems to climate change. This study investigated the relationship between total soil CO₂ efflux and soil temperature, soil moisture and solar radiation on an interannual basis for a plot of temperate deciduous ancient semi-natural woodland at Wytham Woods in central southern England. We also aimed to quantify the contribution of soil organic matter decomposition (SOM), root-and-rhizosphere respiration, and mycorrhizal respiration components to total soil CO₂ efflux, and determine their environmental correlates. Total soil CO₂ efflux was measured regularly from April 2006 to December 2008 and found to average 4.1 Mg C ha⁻¹ yr⁻¹ in both 2007 and 2008. In addition, we applied a recently developed approach to partition the efflux into SOM, root-and-rhizosphere, and mycorrhizal components in situ using mesh bags. SOM decomposition, root-and-rhizosphere, and mycorrhizal respiration were estimated to contribute 70 ± 6%, 22 ± 6% and 8 ± 3% of total soil CO₂ efflux respectively, equating to 3.0 ± 0.3, 0.9 ± 0.2 and 0.3 ± 0.1 Mg C ha⁻¹ yr⁻¹. In order to avoid the effect of temporal correlation between variables caused by seasonality, we investigated interannual variability by examining the relationship between CO₂ flux anomalies and anomalies in environmental variables. Variation in soil temperature explained 50% of the interannual variance in soil CO₂ efflux, and soil moisture a further 18% of the residual variance. Solar radiation, as a proxy for plant photosynthesis, had no significant effect on total soil CO₂ efflux, but was positively correlated with root-and-rhizosphere respiration, and mycorrhizal respiration. The relationship between anomalies in soil CO₂ efflux and soil temperature was highly significant, with a sensitivity of 0.164 ± 0.023 μmol CO₂ m⁻² s⁻¹ °C⁻¹. For mean peak summer efflux rates (2.03 μmol CO₂ m² s⁻¹), this is equivalent to 8% per °C, or a Q₁₀ temperature sensitivity of 2.2 ± 0.2. We demonstrate the utility of an anomaly analysis approach and conclude that soil temperature is the key driver of total soil CO₂ efflux primarily through its positive relationship with SOM-decomposition rate.