Experimental litterfall manipulation drives large and rapid changes in soil carbon cycling in a wet tropical forest

Global changes such as variations in plant net primary production are likely to drive shifts in leaf litterfall inputs to forest soils, but the effects of such changes on soil carbon (C) cycling and storage remain largely unknown, especially in C‐rich tropical forest ecosystems. We initiated a leaf litterfall manipulation experiment in a tropical rain forest in Costa Rica to test the sensitivity of surface soil C pools and fluxes to different litter inputs. After only 2 years of treatment, doubling litterfall inputs increased surface soil C concentrations by 31%, removing litter from the forest floor drove a 26% reduction over the same time period, and these changes in soil C concentrations were associated with variations in dissolved organic matter fluxes, fine root biomass, microbial biomass, soil moisture, and nutrient fluxes. However, the litter manipulations had only small effects on soil organic C (SOC) chemistry, suggesting that changes in C cycling, nutrient cycling, and microbial processes in response to litter manipulation reflect shifts in the quantity rather than quality of SOC. The manipulation also affected soil CO ₂ fluxes; the relative decline in CO ₂ production was greater in the litter removal plots (?22%) than the increase in the litter addition plots (+15%). Our analysis showed that variations in CO ₂ fluxes were strongly correlated with microbial biomass pools, soil C and nitrogen (N) pools, soil inorganic P fluxes, dissolved organic C fluxes, and fine root biomass. Together, our data suggest that shifts in leaf litter inputs in response to localized human disturbances and global environmental change could have rapid and important consequences for belowground C storage and fluxes in tropical rain forests, and highlight differences between tropical and temperate ecosystems, where belowground C cycling responses to changes in litterfall are generally slower and more subtle.     

Estimates of soil respiration and net primary production of three forests at different succession stages in South China

Soil respiration (heterotropic and autotropic respiration, R_g) and aboveground litter fall carbon were measured at three forests at different succession (early, middle and advanced) stages in Dinghushan Biosphere Reserve, Southern China. It was found that the soil respiration increases exponentially with soil temperature at 5 cm depth (T_s) according to the relation R_g=a exp(bT_s), and the more advanced forest community during succession has a higher value of a because of higher litter carbon input than the forests at early or middle succession stages. It was also found that the monthly soil respiration is linearly correlated with the aboveground litter carbon input of the previous month. Using measurements of aboveground litter and soil respiration, the net primary productions (NPPs) of three forests were estimated using nonlinear inversion. They are 475, 678 and 1148 g C m^?2 yr^?1 for the Masson pine forest (MPF), coniferous and broad‐leaf mixed forest (MF) and subtropical monsoon evergreen broad‐leaf forest (MEBF), respectively, in year 2003/2004, of which 54%, 37% and 62% are belowground NPP for those three respective forests if no change in live plant biomass is assumed. After taking account of the decrease in live plant biomass, we estimated the NPP of the subtropical MEBF is 970 g C m^?2 yr^?1 in year 2003/2004. Total amount of carbon allocated below ground for plant roots is 388 g C m^?2 yr^?1 for the MPF, 504 g C m^?2 yr^?1 for the coniferous and broad‐leaf MF and 1254 g C m^?2 yr^?1 for the subtropical MEBF in 2003/2004. Our results support the hypothesis that the amount of carbon allocation belowground increases during forest succession.     

杉木林年龄序列地下碳分配变化

  森林地下碳分配在森林碳平衡和碳吸存中具有重要作用, 而揭示人工林生长过程中地下碳分配变化对于人工林碳汇估算和碳汇管理等有重要意义。通过采用年龄序列方法研究了杉木(Cunninghamia lanceolata)林生长过程中地下碳分配变化特点。年龄序列为福建省南平7 a生(幼龄林)、16 a生(中龄林)、21 a生(近熟林)、41 a生(成熟林)和88 a生(老龄林)的杉木林。细根净生产力测定采用连续土芯法, 根系呼吸测定采用壕沟法, 生物量增量测定采用异速生长方程, 地上年凋落物量采用凋落物收集框测定。结果表明: 杉木林细根净生产力在中龄林前没有显著差异, 维持在较高水平; 但此后则显著下降。细根净生产力/地上凋落物量比值随林龄增加而显著下降。老龄林的根系呼吸显著低于其它林龄林分, 根系呼吸与细根生物量间呈显著线性相关。中龄林和近成熟林的地下碳分配(Total belouground carbon allocation, TBCA)显著高于幼龄林和成熟林, 而老龄林的则最低。中龄林、近成熟林和成熟林的地上部分净生产力/TBCA比值显著高于幼龄林和老龄林, 而杉木林的根系碳利用效率(RCUE)则呈现出随林龄增加而降低的趋势。     

Postfire carbon pools and fluxes in semiarid ponderosa pine in Central Oregon

Forest fire dramatically affects the carbon storage and underlying mechanisms that control the carbon balance of recovering ecosystems. In western North America where fire extent has increased in recent years, we measured carbon pools and fluxes in moderately and severely burned forest stands 2 years after a fire to determine the controls on net ecosystem productivity (NEP) and make comparisons with unburned stands in the same region. Total ecosystem carbon in soil and live and dead pools in the burned stands was on average 66% that of unburned stands (11.0 and 16.5 kg C m⁻², respectively, P<0.01). Soil carbon accounted for 56% and 43% of the carbon pools in burned and unburned stands. NEP was significantly lower in severely burned compared with unburned stands (P<0.01) with an increasing trend from −125±44 g C m⁻² yr⁻¹ (±1 SD) in severely burned stands (stand replacing fire), to −38±96 and +50±47 g C m⁻² yr⁻¹ in moderately burned and unburned stands, respectively. Fire of moderate severity killed 82% of trees <20 cm in diameter (diameter at 1.3 m height, DBH); however, this size class only contributed 22% of prefire estimates of bole wood production. Larger trees (> 20 cm DBH) suffered only 34% mortality under moderate severity fire and contributed to 91% of postfire bole wood production. Growth rates of trees that survived the fire were comparable with their prefire rates. Net primary production NPP (g C m⁻² yr⁻¹, ±1 SD) of severely burned stands was 47% of unburned stands (167±76, 346±148, respectively, P<0.05), with forb and grass aboveground NPP accounting for 74% and 4% of total aboveground NPP, respectively. Based on continuous seasonal measurements of soil respiration in a severely burned stand, in areas kept free of ground vegetation, soil heterotrophic respiration accounted for 56% of total soil CO₂ efflux, comparable with the values of 54% and 49% previously reported for two of the unburned forest stands. Estimates of total ecosystem heterotrophic respiration (R_h) were not significantly different between stand types 2 years after fire. The ratio NPP/R_h averaged 0.55, 0.85 and 1.21 in the severely burned, moderately burned and unburned stands, respectively. Annual soil CO₂ efflux was linearly related to aboveground net primary productivity (ANPP) with an increase in soil CO₂ efflux of 1.48 g C yr⁻¹ for every 1 g increase in ANPP (P<0.01, r²= 0.76). There was no significant difference in this relationship between the recently burned and unburned stands. Contrary to expectations that the magnitude of NEP 2 years postfire would be principally driven by the sudden increase in detrital pools and increased rates of R_h, the data suggest NPP was more important in determining postfire NEP.     

Effects of nitrogen additions on above- and belowground carbon dynamics in two tropical forests

Anthropogenic nitrogen (N) deposition is increasing rapidly in tropical regions, adding N to ecosystems that often have high background N availability. Tropical forests play an important role in the global carbon (C) cycle, yet the effects of N deposition on C cycling in these ecosystems are poorly understood. We used a field N-fertilization experiment in lower and upper elevation tropical rain forests in Puerto Rico to explore the responses of above- and belowground C pools to N addition. As expected, tree stem growth and litterfall productivity did not respond to N fertilization in either of these N-rich forests, indicating a lack of N limitation to net primary productivity (NPP). In contrast, soil C concentrations increased significantly with N fertilization in both forests, leading to larger C stocks in fertilized plots. However, different soil C pools responded to N fertilization differently. Labile (low density) soil C fractions and live fine roots declined with fertilization, while mineral-associated soil C increased in both forests. Decreased soil CO₂ fluxes in fertilized plots were correlated with smaller labile soil C pools in the lower elevation forest (R² = 0.65, p < 0.05), and with lower live fine root biomass in the upper elevation forest (R² = 0.90, p < 0.05). Our results indicate that soil C storage is sensitive to N deposition in tropical forests, even where plant productivity is not N-limited. The mineral-associated soil C pool has the potential to respond relatively quickly to N additions, and can drive increases in bulk soil C stocks in tropical forests.     

Biometric Based Carbon Flux Measurements and Net Ecosystem Production (NEP) in a Temperate Deciduous Broad-Leaved Forest Beneath a Flux Tower

Biometric based carbon flux measurements were conducted over 5 years (1999–2003) in a temperate deciduous broad-leaved forest of the AsiaFlux network to estimate net ecosystem production (NEP). Biometric based NEP, as measured by the balance between net primary production (including NPP of canopy trees and of forest floor dwarf bamboo) and heterotrophic respiration (RH), clarified the contribution of various biological processes to the ecosystem carbon budget, and also showed where and how the forest is storing C. The mean NPP of the trees was 5.4 ± 1.07 t C ha⁻¹ y⁻¹, including biomass increment (0.3 ± 0.82 t C ha⁻¹ y⁻¹), tree mortality (1.0 ± 0.61 t C ha⁻¹ y⁻¹), aboveground detritus production (2.3 ± 0.39 t C ha⁻¹ y⁻¹) and belowground fine root production (1.8 ± 0.31 t C ha⁻¹ y⁻¹). Annual biomass increment was rather small because of high tree mortality during the 5 years. Total NPP at the site was 6.5 ± 1.07 t C ha⁻¹ y⁻¹, including the NPP of the forest floor community (1.1 ± 0.06 t C ha⁻¹ y⁻¹). The soil surface CO₂ efflux (RS) was averaged across the 5 years of record using open-flow chambers. The mean estimated annual RS amounted to 7.1 ± 0.44 t C ha⁻¹, and the decomposition of soil organic matter (SOM) was estimated at 3.9 ± 0.24 t C ha⁻¹. RH was estimated at 4.4 ± 0.32 t C ha⁻¹ y⁻¹, which included decomposition of coarse woody debris. Biometric NEP in the forest was estimated at 2.1 ± 1.15 t C ha⁻¹ y⁻¹, which agreed well with the eddy-covariance based net ecosystem exchange (NEE). The contribution of woody increment (Δbiomass + mortality) of the canopy trees to NEP was rather small, and thus the SOM pool played an important role in carbon storage in the temperate forest. These results suggested that the dense forest floor of dwarf bamboo might have a critical role in soil carbon sequestration in temperate East Asian deciduous forests.     

Heterotrophic Soil Respiration in Relation to Environmental Factors and Microbial Biomass in Two Wet Tropical Forests

We examined the correlation between fungal and bacterial biomass, abiotic factors such as soil moisture, carbon in the light soil fraction and soil nitrogen to a depth of 0–25 cm and heterotrophic soil respiration using a trenching technique – in a secondary forest (Myrcia splendens, Miconia prasina and Casearia arborea) and a pine (Pinus caribeae) plantation in the Luquillo Experimental Forest in Puerto Rico. Soil respiration was significantly reduced where roots were excluded for 7 years in both the secondary forest and the pine plantation. Microbial biomass was also significantly reduced in the root exclusion plots. In root exclusion treatment, total fungal biomass was on average 31 and 65% lower than the control plots in the pine plantation and the secondary forest, respectively, but the total bacterial biomass was 24 and 8.3% lower than the control plots in the pine plantation and the secondary forest, respectively. Heterotrophic soil respiration was positively correlated with fungal biomass (R²=0.63, R²=0.39), bacterial biomass (R²=0.16, R²=0.45), soil moisture (R²=0.41, R²=0.56), carbon in light fraction (R²=0.45, R²=0.39) and total nitrogen (R²=0.69, R²=0.67) in the pine plantation and the secondary forest, respectively. The regression analysis suggested that fungal biomass might have a greater influence on heterotrophic soil respiration in the pine plantation, while the bacterial biomass might have a greater influence in the secondary forest. Heterotrophic soil respiration was more sensitive to total N than to carbon in the light fraction, and soil moisture was a major factor influencing heterotrophic soil respiration in these forests where temperature is high and relatively invariable.     

Primary production and carbon allocation in relation to nutrient supply in a tropical experimental forest

Nutrient supply commonly limits aboveground plant productivity in forests, but the effects of an altered nutrient supply on gross primary production (GPP) and patterns of carbon (C) allocation remain poorly characterized. Increased nutrient supply may lead to a higher aboveground net primary production (ANPP), but a lower total belowground carbon allocation (TBCA), with little change in either aboveground plant respiration (APR) or GPP. Alternatively, increases in nutrient supply may increase GPP, with the quantity of GPP allocated aboveground increasing more steeply than the quantity of GPP allocated belowground. To examine the effects of an elevated nutrient supply on the C allocation patterns in forests, we determined whole‐ecosystem C budgets in unfertilized plots of Eucalyptus saligna and in adjacent plots receiving regular additions of 65 kg N ha^?1, 31 kg P ha^?1, 46 kg K ha^?1, and macro‐ and micronutrients. We measured the absolute flux of C allocated to the components of GPP (ANPP, TBCA and APR), as well as the fraction of GPP allocated to these components. Fertilization dramatically increased GPP. Averaged over 3 years, GPP in the fertilized plots was 34% higher than that in the unfertilized controls (3.95 vs. 2.95 kg C m^?2 yr^?1). Fertilization‐related increases in GPP were allocated entirely aboveground – ANPP was 85% higher and APR was 57% higher in the fertilized than in the control plots, while TBCA did not differ significantly between treatments. Carbon use efficiency (NPP/GPP) was slightly higher in the fertilized (0.53) compared with the control plots (0.51). Overall, fertilization increased ANPP and APR, and these increases were related to a greater GPP and an increase in the fraction of GPP allocated aboveground.     

Soil respiration in six temperate forests in China

Scaling soil respiration (R_S), the major CO₂ source to the atmosphere from terrestrial ecosystems, from chamber‐based measurements to ecosystems requires studies on variations and correlations of R_S from various biomes and across geographic regions. However, few studies on R_S are available for Chinese temperate forest despite the importance of this forest in the national and global carbon budgets. In this study, we conducted 18‐month R_S measurements during 2004–2005 in six temperate forest types, representing the typical secondary forest ecosystems across various site conditions in northeastern China: Mongolian oak (Quercus mongolica Fisch.), aspen‐birch (Populous davidiana Dode and Betula platyphylla Suk.), mixed deciduous (no dominant tree species), hardwood (dominated by Fraxinus mandshurica Rupr., Juglans mandshurica Maxim., and Phellodendron amurense Rupr.) forests, Korean pine (Pinus koraiensis Sieb. et Zucc.) and Dahurian larch (Larix gmelinii Rupr.) plantations. Our specific objectives were to: (1) explore relationships of R_S against soil temperature and water content for the six forest ecosystems, (2) quantify annual soil surface CO₂ flux and its relations to belowground carbon storage, (3) examine seasonal variations in R_S and related environmental factors, and (4) quantify among‐ and within‐ecosystem variations in R_S. The R_S was positively correlated to soil temperature in all forest types, and was significantly influenced by the interactions of soil temperature and water content in the pine, larch, and mixed deciduous forests. The sensitivity of R_S to soil temperature at 10 cm depth (Q₁₀) ranged from 2.61 in the oak forest to 3.75 in the aspen‐birch forests. The Q₁₀ tended to increase with soil water content until reaching a threshold, and then decline. The annual R_S for the larch, pine, hardwood, oak, mixed deciduous, and aspen‐birch forests averaged 403, 514, 781, 785, 786, and 813 g C m^?2 yr^?1, respectively. The annual R_S of the broadleaved forests was 72% greater than that of the coniferous forests. The annual R_S was positively correlated to soil organic carbon (SOC) concentration at O horizon (R²=0.868) and total biomass of roots <0.5 cm in diameter (R²=0.748). The coefficient of variation (CV) of R_S among forest types averaged 25% across the 18‐month measurements. The CV of R_S within plots varied from 20% to 27%, significantly (P<0.001) greater than those among plots (9–15%), indicating the importance of the fine‐scaled heterogeneity in R_S. This study emphasized that variations in soil respiration and potential sampling bias should be appropriately tackled for accurate soil CO₂ flux estimates.     

An analysis of soil respiration across northern hemisphere temperate ecosystems

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

The influence of fire on carbon distribution and net primary production of boreal Larix gmelinii forests in north-eastern China

The boreal larch forest of Eurasia is a widespread forest ecosystem and plays an important role in the carbon budget of boreal forests. However, few carbon budgets exist for these forests, and the effects of wildfire, the dominant natural disturbance in this region, on carbon budgets are poorly understood. The objective of this study was to quantify the effects of wildfire on carbon distribution and net primary production (NPP) for three major Dahurian larch (Larix gmelinii Rupr.) forest ecosystems in Tahe, Daxing'anling, north‐eastern China: Larix gmelinii–Ledum palustre, Larix gmelinii–grass and Larix gmelinii–Rhododendron dahurica forests. The experimental design included mature forests (unburned), and lightly and heavily burned forests from the 1.3‐million‐ha 1987 wildfire. We measured carbon distribution and above‐ground NPP, and estimated fine root production from literature values. Total ecosystem carbon content for the mature forests was greatest for Larix–Ledum forests (251.4 t C ha^?1) and smallest for Larix–grass forests (123.8 t C ha^?1). Larix–Ledum forests contained the smallest vegetation carbon (13.5%), while Larix–Rhododendron contained the largest vegetation carbon (63.1%). Fires tended to transfer carbon from vegetation to detritus and soil. Total NPP did not differ significantly between the lightly burned and unburned stands, and averaged 1.58, 1.29 and 1.01 t C ha^?1 year^?1 for Larix–grass, Larix–Rhododendron and Larix–Ledum lightly burned stands, respectively. Above‐ground net primary production (ANPP) of heavily burned stands was 92–95% less than unburned and lightly burned stands. The estimated carbon loss during the 1987 fire showed substantial variability among forest types and fire severity levels. Depending upon the assumptions made about the fraction of the landscape occupied by the three larch forest types, the 1987 conflagration in north‐east China released 2.5 × 10⁷?4.9 × 10⁷ t C to the atmosphere. This study illustrates the need to distinguish between the different larch forests for developing general carbon budgets.     

Fine Root Production across a Primary Successional Ecosystem Chronosequence at Mt. Shasta,California

Estimating changes in belowground biomass and production is essential for understanding fundamental patterns and processes during ecosystem development. We examined patterns of fine root production, aboveground litterfall, and forest floor accumulation during forest primary succession at the Mt. Shasta Mudflows ecosystem chronosequence. Fine root production was measured using the root ingrowth cores method over 1 year, and aboveground litterfall was collected over 2 years. Fine root production increased significantly with ecosystem age, but only the youngest ecosystem was significantly different from all of the older ecosystems. Root production was 44.5 ± 13.3, 168.3 ± 20.6, 190.5 ± 33.8, and 236.3 ± 65.4 g m⁻² y⁻¹ in the 77, 255, 616, and >850-year-old ecosystems, respectively. Generally, aboveground litterfall and forest floor accumulation did not follow the same pattern as root production. The relative contribution of fine root production to total fine detrital production increased significantly with ecosystem age, from 14 to 49%, but only the youngest ecosystem was significantly different from all of the older ecosystems. Fine root production was significantly correlated with some measures of soil fertility but was not correlated with leaf or total litterfall, or forest floor accumulation. It was best predicted by soil N concentration alone, but this relationship may not be causal, as soil N concentration was also correlated with ecosystem age. For the oldest ecosystem, fine root production was also measured using the sequential intact cores/compartment-flow model method, and the difference between the two estimates was not significant. Our study suggests that the relative contribution of fine roots to fine detrital production, and hence to soil organic matter accumulation, may increase during forest primary succession.     

The above-ground coarse wood productivity of 104 Neotropical forest plots

The net primary production of tropical forests and its partitioning between long‐lived carbon pools (wood) and shorter‐lived pools (leaves, fine roots) are of considerable importance in the global carbon cycle. However, these terms have only been studied at a handful of field sites, and with no consistent calculation methodology. Here we calculate above‐ground coarse wood carbon productivity for 104 forest plots in lowland New World humid tropical forests, using a consistent calculation methodology that incorporates corrections for spatial variations in tree‐size distributions and wood density, and for census interval length. Mean wood density is found to be lower in more productive forests. We estimate that above‐ground coarse wood productivity varies by more than a factor of three (between 1.5 and 5.5 Mg C ha^?1 a^?1) across the Neotropical plots, with a mean value of 3.1 Mg C ha^?1 a^?1. There appear to be no obvious relationships between wood productivity and rainfall, dry season length or sunshine, but there is some hint of increased productivity at lower temperatures. There is, however, also strong evidence for a positive relationship between wood productivity and soil fertility. Fertile soils tend to become more common towards the Andes and at slightly higher than average elevations, so the apparent temperature/productivity relationship is probably not a direct one. Coarse wood productivity accounts for only a fraction of overall tropical forest net primary productivity, but the available data indicate that it is approximately proportional to total above‐ground productivity. We speculate that the large variation in wood productivity is unlikely to directly imply an equivalent variation in gross primary production. Instead a shifting balance in carbon allocation between respiration, wood carbon and fine root production seems the more likely explanation.     

Carbon pools in a boreal mixedwood logging chronosequence

Mixedwood forests are an ecologically and economically important forest type in central Canada, but the ecology of these forests is not as well studied as that of single-species dominated stands in the boreal forest. Northern boreal mixedwood forests have only recently been harvested and the effects of harvesting on carbon content in these stands are unknown. We quantified the carbon content and aboveground net primary production (NPP) for four different-aged mixedwood boreal forest stands in northern Manitoba, Canada. The stands included 11-, 18-, and 30-year-old stands that originated from harvesting and a 65-year-old fire-originated stand that typifies the origin of all northern boreal mixed-wood forests that are coming under management. Trees included black spruce (Picea mariana (Mill.) B.S.P.), jack pine (Pinus banksiana Lamb.), balsam poplar (Populus balsamifera L.), and quaking aspen (Populus tremuloides Michx.). Overstory biomass was estimated using species-specific allometric models that generally explained greater than 95% of the observed variation in biomass. Carbon content of the overstory vegetation was greatest in the 65-year-old stand and was 74% larger than the 11-year-old stand and showed a positive relationship with stand age (F_{1, 2}=122.62, P=0.0081 R²=0.99). The slope of mineral soil carbon did not differ significantly among stands (F_{1, 2}=0.39, P=0.5956, R²=0.16). Coarse woody debris carbon content followed a U-shaped pattern among stands. Aboveground NPP differed by 24% between the youngest and oldest stand. Mean annual carbon accumulation and aboveground NPP rates of the mixedwood forests were on average two times greater than nearby relatively pure stands studied during the BOREAS (BOReal Ecosystem Atmospheric Study) project. The trends in the results, along with other field studies, suggest that harvesting does not significantly affect the total soil carbon content. The results of this study suggest that scientists should be cautious about extrapolating results from BOREAS stands to a broader region until more data on other forest types and regions are available.     

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.     

Changes in carbon storage and fluxes in a chronosequence of ponderosa pine

Forest development following stand‐replacing disturbance influences a variety of ecosystem processes including carbon exchange with the atmosphere. On a series of ponderosa pine (Pinius ponderosa var. Laws.) stands ranging from 9 to> 300 years in central Oregon, USA, we used biological measurements to estimate carbon storage in vegetation and soil pools, net primary productivity (NPP) and net ecosystem productivity (NEP) to examine variation with stand age. Measurements were made on plots representing four age classes with three replications: initiation (I, 9–23 years), young (Y, 56–89 years), mature (M, 95–106 years), and old (O, 190–316 years) stands typical of the forest type in the region. Net ecosystem productivity was lowest in the I stands (?124 g C m^?2 yr^?1), moderate in Y stands (118 g C m^?2 yr^?1), highest in M stands (170 g C m^?2 yr^?1), and low in the O stands (35 g C m^?2 yr^?1). Net primary productivity followed similar trends, but did not decline as much in the O stands. The ratio of fine root to foliage carbon was highest in the I stands, which is likely necessary for establishment in the semiarid environment, where forests are subject to drought during the growing season (300–800 mm precipitation per year). Carbon storage in live mass was the highest in the O stands (mean 17.6 kg C m^?2). Total ecosystem carbon storage and the fraction of ecosystem carbon in aboveground wood mass increased rapidly until 150–200 years, and did not decline in older stands. Forest inventory data on 950 ponderosa pine plots in Oregon show that the greatest proportion of plots exist in stands ～ 100 years old, indicating that a majority of stands are approaching maximum carbon storage and net carbon uptake. Our data suggests that NEP averages ～ 70 g C m^?2 year^?1 for ponderosa pine forests in Oregon. About 85% of the total carbon storage in biomass on the survey plots exists in stands greater than 100 years, which has implications for managing forests for carbon sequestration. To investigate variation in carbon storage and fluxes with disturbance, simulation with process models requires a dynamic parameterization for biomass allocation that depends on stand age, and should include a representation of competition between multiple plant functional types for space, water, and nutrients.     

CO2 flux from soil in pastures and forests in southwestern Amazonia

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

Seasonal and annual respiration of a ponderosa pine ecosystem

The net ecosystem exchange of CO₂ between forests and the atmosphere, measured by eddy covariance, is the small difference between two large fluxes of photosynthesis and respiration. Chamber measurements of soil surface CO₂ efflux (F_s), wood respiration (F_w) and foliage respiration (F_f) help identify the contributions of these individual components to net ecosystem exchange. Models developed from the chamber data also provide independent estimates of respiration costs. We measured CO₂ efflux with chambers periodically in 1996–97 in a ponderosa pine forest in Oregon, scaled these measurements to the ecosystem, and computed annual totals for respiration by component. We also compared estimated half-hourly ecosystem respiration at night (F_nc) with eddy covariance measurements. Mean foliage respiration normalized to 10 °C was 0.20 μmol m^–2 (hemi-leaf surface area) s^–1, and reached a maximum of 0.24 μmol m^–2 HSA s^–1 between days 162 and 208. Mean wood respiration normalized to 10 °C was 5.9 μmol m^–3 sapwood s^–1, with slightly higher rates in mid-summer, when growth occurs. There was no significant difference (P > 0.10) between wood respiration of young (45 years) and old trees (250 years). Soil surface respiration normalized to 10 °C ranged from 0.7 to 3.0 μmol m^–2 (ground) s^–1 from days 23 to 329, with the lowest rates in winter and highest rates in late spring. Annual CO₂ flux from soil surface, foliage and wood was 683, 157, and 54 g C m^–2 y^–1, with soil fluxes responsible for 76% of ecosystem respiration. The ratio of net primary production to gross primary production was 0.45, consistent with values for conifer sites in Oregon and Australia, but higher than values reported for boreal coniferous forests. Below-ground carbon allocation (root turnover and respiration, estimated as F_s– litterfall carbon) consumed 61% of GPP; high ratios such as this are typical of sites with more water and nutrient constraints. The chamber estimates were moderately correlated with change in CO₂ storage in the canopy (F_stor) on calm nights (friction velocity u* < 0.25 m s^–1; R² = 0.60); F_stor was not significantly different from summed chamber estimates. On windy nights (u* > 0.25 m s^–1), the sum of turbulent flux measured above the canopy by eddy covariance and F_stor was only weakly correlated with summed chamber estimates (R² = 0.14); the eddy covariance estimates were lower than chamber estimates by 50%.     

The legacy of harvest and fire on ecosystem carbon storage in a north temperate forest

Forest harvesting and wildfire were widespread in the upper Great Lakes region of North America during the early 20th century. We examined how long this legacy of disturbance constrains forest carbon (C) storage rates by quantifying C pools and fluxes after harvest and fire in a mixed deciduous forest chronosequence in northern lower Michigan, USA. Study plots ranged in age from 6 to 68 years and were created following experimental clear‐cut harvesting and fire disturbance. Annual C storage was estimated biometrically from measurements of wood, leaf, fine root, and woody debris mass, mass losses to herbivory, soil C content, and soil respiration. Maximum annual C storage in stands that were disturbed by harvest and fire twice was 26% less than a reference stand receiving the same disturbance only once. The mechanism for this reduction in annual C storage was a long‐lasting decrease in site quality that endured over the 62‐year timeframe examined. However, during regrowth the harvested and burned forest rapidly became a net C sink, storing 0.53 Mg C ha⁻¹ yr⁻¹ after 6 years. Maximum net ecosystem production (1.35 Mg C ha⁻¹ yr⁻¹) and annual C increment (0.95 Mg C ha⁻¹ yr⁻¹) were recorded in the 24‐ and 50‐year‐old stands, respectively. Net primary production averaged 5.19 Mg C ha⁻¹ yr⁻¹ in experimental stands, increasing by < 10% from 6 to 50 years. Soil heterotrophic respiration was more variable across stand ages, ranging from 3.85 Mg C ha⁻¹ yr⁻¹ in the 6‐year‐old stand to 4.56 Mg C ha⁻¹ yr⁻¹ in the 68‐year‐old stand. These results suggest that harvesting and fire disturbances broadly distributed across the region decades ago caused changes in site quality and successional status that continue to limit forest C storage rates.     

The role of environmental, root, and microbial biomass characteristics in soil respiration in temperate secondary forests of Northeast China

For secondary forests, the major forest resources in China (accounting for more than 50% of the national total), soil respiration (R _S) and the relationship between R _S and various biotic/abiotic factors are poorly understood. The objectives of the present study were to examine seasonal variations in soil respiration during the growing season, and to explore the factors affecting the variation in soil respiration rates for three forest types (Mongolian oak, Manchurian walnut and mixed forests) of temperate secondary forest in Northeast China. The results showed that (1) the maximum total R _S rate occurred in July, following a bell-shaped curve with season, (2) for all forest types, the total R _S was significantly influenced by soil temperature (P < 0.01), and did not significantly correlate with soil moisture, (3) compared with fine root biomass, coarse root biomass was more closely related with the root respiration in mixed forest (R ² = 0.711, P = 0.017) and in Manchurian walnut forest (R ² = 0.768, P = 0.010), and (4) microbial biomass carbon (MBC) and nitrogen were significantly correlated with heterotrophic R _S in Mongolian oak forest (R ² = 0.664, P = 0.026; R ² = 0.784, P = 0.008, respectively) and in mixed forest (R ² = 0.918, P = 0.001; R ² = 0.967, P = 0.001, respectively). We can conclude that in temperate secondary forests: (1) the R _S rate and the relationships between R _S and abiotic/biotic factors change greatly with forest types, and (2) R _S is strongly influenced by soil temperature, MBC, microbial biomass nitrogen and coarse root biomass in temperate secondary forests.