Coarse woody debris mass and nutrients in forest ecosystems of Korea

Coarse woody debris (CWD) is an essential component of forests. However, quantification of both the mass and nutrient content of CWD within a given environment tends to be a fairly labor-intensive proposition that requires long-term studies to be conducted for viable data to be obtained. As a result, various aspects of CWD in forest ecosystems remain somewhat poorly understood. In this review, we have compiled all available estimates of CWD mass and nutrients from both coniferous and deciduous forests in Korea. The CWD mass data varied substantially by forest type, age, location, and sampling time, ranging from 1.5 to 24.5 Mg ha⁻¹, and for the amount (kg ha⁻¹) of nutrients in the CWD, ranging from 3.5 to 23.6 for nitrogen (N), 0.8 to 4.7 for phosphorus (P), 3.9 to 13.3 for potassium (K), 25.9 to 30.9 for calcium (Ca), 1.4 to 4.2 for magnesium (Mg), and 0.1 to 0.6 for sodium (Na). The mass of CWD transferred from live trees to the forest floor ranged between 0.1 and 4.9 Mg ha⁻¹ year⁻¹, and these values were roughly equivalent to 26–42% of the annual litterfall inputs (2.5–10.8 Mg ha⁻¹ year⁻¹) for mixed Quercus spp. forests within the relevant region. Annual nutrients inputs (kg ha⁻¹ year⁻¹) through CWD decomposition were 0.7–1.6 for N, 0.04–0.3 for P, 0.3–1.0 for K, 1.7–3.1 for Ca, and 0.1–0.3 for Mg. Consequently, these results revealed that the ecological value of CWD for C and nutrient cycling was relatively insignificant. However, only a limited number of studies have been conducted on CWD in different coniferous or mixed deciduous forests in the region. As a direct result of this paucity of data, further long-term studies on CWD mass and nutrients in a variety of forest types are required in order to be able to evaluate accurately the ecological value of CWD on biodiversity and physical properties in Korean forest ecosystems.     

Decomposition and Fragmentation of Coarse Woody Debris: Re-visiting a Boreal Black Spruce Chronosequence

We re-visited a seven-stand boreal chronosequence west of Thompson, Manitoba, Canada, in which coarse woody debris (CWD) and its instantaneous decomposition were measured in 2000. New CWD measurements were performed in 2007, and tree inventories updated to provide mortality and snag failure data. These data were used to model CWD changes, compare methods of estimating decomposition, and infer possible fragmentation rates. Measured CWD was between 9.7 (in both the 77- and 43-year-old stands) and 80.4 (in the 18-year-old stand) Mg ha⁻¹ in 2007. Spatial variability was high; at most stands CWD levels had not changed significantly from 2000 to 2007. Tree mortality was a significant flux only in older stands, whereas snag fall rate varied by an order of magnitude, from 2.9% y⁻¹ (0.2 Mg ha⁻¹ y⁻¹) in the 9-year-old stand to 9.8% y⁻¹ (2.3 Mg ha⁻¹ y⁻¹) in the 12-year-old stand. A one-pool model based on these inputs underestimated actual 2000–2007 CWD decomposition in the younger stands, suggesting that fragmentation could be an important part of the carbon flux exiting the CWD pool. We compared three independent measures of annual decomposition (k): direct measurements of CWD respiration, rates based on the 7-year re-sampling effort described here, and rates inferred from the chronosequence design itself. Mean k values arrived at via these techniques were 0.06 ± 0.03, 0.05 ± 0.04, and 0.05 ± 0.05 y⁻¹, respectively. The four-pool model suggested that the transition rate between decay classes was 0.14–0.19 y⁻¹; the model was most sensitive to initial CWD values. Although the computed k values implied a problem with chronosequence site selection for at least one site, the overall CWD trend was consistent with a larger number of sites surveyed in the region.     

Woody debris stocks in different secondary and primary forests in the subtropical Ailao Mountains, southwest China

Woody debris (WD), including coarse woody debris (CWD) and fine woody debris (FWD), is an essential structural and functional component of many ecosystems, particularly in montane forests. CWD is considered to be the major part in forest WD and it is primarily composed of logs, snags, stumps and large branches, while FWD mainly consists of small twigs. Attributes of dead woody material may change in accordance with trends in stand dynamics. The primary forest (primary montane moist evergreen broad-leaved forest) in Ailao Mountain National Nature Reserve (NNR) preserves the largest tract of natural vegetation in China. The Alnus nepalensis (D. Don) association, Populus bonatii (Levl.) association and secondary Lithocarpus association represent the secondary and chronological types following human disturbance by fires and logging under different intensity. The mass and composition of coarse woody debris (CWD, ≥10 cm in diameter) and fine woody debris (FWD, 2.5–10 cm in diameter) were inventoried in a primary forest and its three secondary counterparts. Estimates of total mass of woody debris across secondary types to primary forest ranged from 2.4 to 74.9 Mg ha⁻¹. The lowest value was found in the A. nepalensis association and the highest values were in the primary forest of which logs are the considerable differences. The ratios of CWD to FWD were low in the secondary types (about 1–4) but high in the primary forest (above 15). Our results suggested that for the recovery of woody debris in the secondary forest, it might last longer than the age of the oldest successional stage studied. Yang Lipan and Ma Wenzhang contributed equally to this work.     

Dead wood biomass and turnover time,measured by radiocarbon,along a subalpine elevation gradient

Dead wood biomass can be a substantial fraction of stored carbon in forest ecosystems, and coarse woody debris (CWD) decay rates may be sensitive to climate warming. We used an elevation gradient in Colorado Rocky Mountain subalpine forest to examine climate and species effects on dead wood biomass, and on CWD decay rate. Using a new radiocarbon approach, we determined that the turnover time of lodgepole pine CWD (340±130 years) was roughly half as long in a site with 2.5–3°C warmer air temperature, as that of pine (630±400 years) or Engelmann spruce CWD (800±960 and 650±410 years) in cooler sites. Across all sites and both species, CWD age ranged from 2 to 600 years, and turnover time was 580±180 years. Total standing and fallen dead wood biomass ranged from 4.7±0.2 to 54±1 Mg ha^–1, and from 2.8 to 60% of aboveground live tree biomass. Dead wood biomass increased 75 kg ha^–1 per meter gain in elevation and decreased 13 Mg ha^–1 for every degree C increase in mean air temperature. Differences in biomass and decay rates along the elevation gradient suggest that climate warming will lead to a loss of dead wood carbon from subalpine forest.Electronic Supplementary Material Supplementary material is available for this article at     

The Imprint of Land-use History: Patterns of Carbon and Nitrogen in Downed Woody Debris at the Harvard Forest

Few data sets have characterized carbon (C) and nitrogen (N) pools in woody debris at sites where other aspects of C and N cycling are studied and histories of land use and disturbance are well documented. We quantified pools of mass, C, and N in fine and coarse woody debris (CWD) in two contrasting stands: a 73-year-old red pine plantation on abandoned agricultural land and a naturally regenerated deciduous forest that has experienced several disturbances in the past 150 years. Masses of downed woody debris amounted to 40.0 Mg ha⁻¹ in the coniferous stand and 26.9 Mg ha⁻¹ in the deciduous forest (20.4 and 13.8 Mg C ha⁻¹, respectively). Concentrations of N were higher and C:N ratios were lower in the deciduous forest compared to the coniferous. Pools of N amounted to 146 kg N ha⁻¹ in the coniferous stand and 155 kg N ha⁻¹ in the deciduous forest; both are larger than previously published pools of N in woody debris of temperate forests. Woody detritus buried in O horizons was minimal in these forests, contrary to previous findings in forests of New England. Differences in the patterns of mass, C, and N in size and decay classes of woody debris were related to stand histories. In the naturally regenerated deciduous forest, detritus was distributed across all size categories, and most CWD mass and N was present in the most advanced decay stages. In the coniferous plantation, nearly all of the CWD mass was present in the smallest size class (less than 25 cm diameter), and a recognizable cohort of decayed stems was evident from the stem-exclusion phase of this even-aged stand. These results indicate that heterogeneities in site histories should be explicitly included when biogeochemical process models are used to scale C and N stocks in woody debris to landscapes and regions. Received 27 April 2001; accepted 4 January 2002.     

Biomass dynamics in a logged forest: the role of wood density

Wood density (WD) is believed to be a key trait in driving growth strategies of tropical forest species, and as it entails the amount of mass per volume of wood, it also tends to correlate with forest carbon stocks. Yet there is relatively little information on how interspecific variation in WD correlates with biomass dynamics at the species and population level. We determined changes in biomass in permanent plots in a logged forest in Vietnam from 2004 to 2012, a period representing the last 8 years of a 30 years logging cycle. We measured diameter at breast height (DBH) and estimated aboveground biomass (AGB) growth, mortality, and net AGB increment (the difference between AGB gains and losses through growth and mortality) per species at the individual and population (i.e. corrected for species abundance) level, and correlated these with WD. At the population level, mean net AGB increment rates were 6.47 Mg ha^?1 year^?1 resulting from a mean AGB growth of 8.30 Mg ha^?1 year^?1, AGB recruitment of 0.67 Mg ha^?1 year^?1 and AGB losses through mortality of 2.50 Mg ha^?1 year^?1. Across species there was a negative relationship between WD and mortality rate, WD and DBH growth rate, and a positive relationship between WD and tree standing biomass. Standing biomass in turn was positively related to AGB growth, and net AGB increment both at the individual and population level. Our findings support the view that high wood density species contribute more to total biomass and indirectly to biomass increment than low wood density species in tropical forests. Maintaining high wood density species thus has potential to increase biomass recovery and carbon sequestration after logging.     

Coarse woody debris in undisturbed and logged forests in the eastern Brazilian Amazon

Coarse woody debris (CWD) is an important component of the carbon cycle in tropical forests. We measured the volume and density of fallen CWD at two sites, Cauaxi and Tapajós in the Eastern Amazon. At both sites we studied undisturbed forests (UFs) and logged forests 1 year after harvest. Conventional logging (CL) and reduced impact logging (RIL) were used for management on areas where the geometric volumes of logs harvested was about 25–30 m³ ha^?1. Density for five classes of fallen CWD for large material (>10 cm diameter) ranged from 0.71 to 0.28 Mg m^?3 depending upon the degree of decomposition. Density of wood within large fallen logs varied with position relative to the ground and with distance from the center of the log. Densities for materials with diameters from 2 to 5 and 5 to 10 cm were 0.36 and 0.45 Mg m^?3, respectively. The average mass (±SE) of fallen CWD at Cauaxi was 55.2 (4.7), 74.7 (0.6), and 107.8 (10.5) Mg ha^?1 for duplicate UF, RIL, and CL sites, respectively. At Tapajós, the average mass of fallen CWD was 50.7 (1.1) Mg ha^?1 for UF and 76.2 (10.2) Mg ha^?1 for RIL for duplicate sites compared with 282 Mg ha^?1 for live aboveground biomass. Small‐ and medium‐sized material (<10 cm dia.) accounted for 8–18% of the total fallen CWD mass. The large amount of fallen CWD at these UF sites relative to standing aboveground biomass suggests either that the forests have recently been subjected to a pulse of high mortality or that they normally suffer a high mortality rate in the range of 0.03 per year. Accounting for background CWD in UF, CL management produced 2.7 times as much CWD as RIL management. Excess CWD at logging sites would generate a substantial CO₂ emission given the high rates of decay in moist tropical forests.     

Carbon stock and cycling in a bambooPhyllostachys bambusoides stand

Gross production and carbon cycling in aPhyllostachys bambusoides stand in Kyoto Prefecture, central Japan, were determined, and then a compartment model showing the carbon stock and cycling within the ecosystem was developed. Aboveground carbon stock was 52.3 tC ha⁻¹, increasing at a rate of 3.6 tC ha⁻¹ year⁻¹. Belowground carbon stock was 20.8 tC ha⁻¹ in the root system and 92.0 tC ha⁻¹ in the soil. Aboveground net production was 11.2 tC ha⁻¹ year⁻¹. Belowground net production was crudely estimated at 4.5 tC ha⁻¹ year⁻¹. The gross production was estimated at 41.8 tC ha⁻¹ year⁻¹ by summing the amount of outflow to the environment and the increment in biomass. Leaves consumed 13.7 tC ha⁻¹ year⁻¹ by respiration; the rest (41.8−13.7=28.1 tC ha⁻¹ year⁻¹) was surplus production of the leaves and flowed into the other compartments. The amounts of construction and maintenance respiration of the aboveground compartments were 3.4 and 18.5 tC ha⁻¹ year⁻¹, respectively. The annual amount of soil respiration was 11.2 tC ha⁻¹ year⁻¹. Soil respiration levels of 4.3 and 3.1 tC ha⁻¹ year⁻¹ were estimated for the flow of root respiration and root detritus. The proportion of net to gross production was 37%, which fell within the range of young and mature forests. A shorter life span of culms, compared to tree trunks, resulted in smaller biomass accumulation ratio (biomass/net production) in the ecosystem, of 4.66.     

Low stocks of coarse woody debris in a southwest Amazonian forest

The stocks and dynamics of coarse woody debris (CWD) are significant components of the carbon cycle within tropical forests. However, to date, there have been no reports of CWD stocks and fluxes from the approximately 1.3 million km² of lowland western Amazonian forests. Here, we present estimates of CWD stocks and annual CWD inputs from forests in southern Peru. Total stocks were low compared to other tropical forest sites, whether estimated by line-intercept sampling (24.4 ± 5.3 Mg ha⁻¹) or by complete inventories within 11 permanent plots (17.7 ± 2.4 Mg ha⁻¹). However, annual inputs, estimated from long-term data on tree mortality rates in the same plots, were similar to other studies (3.8 ± 0.2 or 2.9 ± 0.2 Mg ha⁻¹ year⁻¹, depending on the equation used to estimate biomass). Assuming the CWD pool is at steady state, the turnover time of coarse woody debris is low (4.7 ± 2.6 or 6.1 ± 2.6 years). These results indicate that these sites have not experienced a recent, large-scale disturbance event and emphasise the distinctive, rapid nature of carbon cycling in these western Amazonian forests.     

Culm recruitment,dry matter dynamics and carbon flux in recently harvested and mature bamboo savannas in the Indian dry tropics

Culm recruitment, standing crop biomass, net production and carbon flux were estimated in mature (5 years after last harvest) and recently harvested bamboo (Dendrocalamus strictus (Roxb.) Nees) savanna sites in the dry tropics. During the 2 study years bamboo shoot recruitment was 1711–3182 and 1432–1510 shoots ha⁻¹ in harvested and mature sites, respectively. Corresponding shoot mortality was 66–93% and 62–69%, respectively. Total biomass was 34.9 t ha⁻¹ at the harvested site and 47.4 t ha⁻¹ at the mature site. Harvesting increased the relative contribution of belowground bamboo biomass. Annual litter input to soil was 2.7 and 5.9 t ha⁻¹ year⁻¹ at the harvested and mature sites, respectively. The bulk of the annual litterfall (78–88%) occurred in the cool dry season (November to February). The mean litter mass on the savanna floor ranged from 3.1 to 3.3 t ha⁻¹; at the harvested site wood litter contributed 70% of the litter mass and at the mature site leaves formed 77% of the litter mass. The mean total net production (TNP) for the two annual cycles was 15.8 t ha⁻¹ year⁻¹ at the harvested site and 19.3 t ha⁻¹ year⁻¹ at the mature site. Nearly half (46–57%) of the TNP was allocated to the belowground parts. Short lived components (leaves and fine roots) contributed about four-fifths of the net production of bamboo. Total carbon storage in the system was 64.4 t ha⁻¹ at the harvested site and 75.4 t ha⁻¹ at the mature site, of which 23–28% was distributed in vegetation, 2% in litter and 70–75% in soil. Annual net carbon deposition was 6.3 and 8.7 t ha⁻¹ year⁻¹ at harvested and mature sites, respectively.     

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.     

Land-use conversion effects on CO2 emissions: from agricultural to hybrid poplar plantation

Land-use changes such as deforestation have been considered one of the main contributors to increased greenhouse gas emissions, while verifiable C sequestration through afforestation projects is eligible to receive C credits under the Kyoto Protocol. We studied the short-term effects on CO₂ emissions of converting agricultural land-use (planted to barley) to a hybrid poplar (Populus deltoids × Populus × petrowskyana var. Walker) plantation in the Parkland region in northern Alberta, where large areas are being planted to hybrid poplars. CO₂ emissions were measured using a static gas chamber method. No differences were found in soil temperature, volumetric moisture content, or soil respiration rates between the barley and Walker plots. The mean soil respiration rate in 2005 was 1.83 ± 0.19 (mean ± 1 SE) and 1.89 ± 0.13 μmol CO₂ m⁻² s⁻¹ in the barley and Walker plots, respectively. However, biomass production was higher in the barley plots, indicating that the agricultural land-use system had a greater ability to fix atmospheric CO₂. The C balance in the land-use systems were estimated to be a small net gain (before considering straw and grain removal through harvesting) of 0.03 ± 0.187 Mg C ha⁻¹ year⁻¹ in the barley plots and a net loss of 3.35 ± 0.080 Mg C ha⁻¹ year⁻¹ from the Walker poplar plots. Over the long-term, we expect the hybrid poplar plantation to become a net C sink as the trees grow bigger and net primary productivity increases.     

Coarse Woody Debris following Fire and Logging in Wyoming Lodgepole Pine Forests

The accumulation and decomposition of coarse woody debris (CWD) are processes that affect habitat, soil structure and organic matter inputs, and energy and nutrient flows in forest ecosystems. Natural disturbances such as fires typically produce large quantities of CWD as trees fall and break, whereas human disturbances such as timber harvesting remove much of the CWD. Our objective was to compare the amount of CWD removed and left behind after clear-cutting to the amount consumed and left behind after natural fires in Rocky Mountain lodgepole pine. The masses of fallen logs, dead-standing trees, stumps, and root crowns more than 7.5 cm in diameter were estimated in clear-cut and intact lodgepole pine forests in Wyoming and compared to estimates made in burned and unburned stands in Yellowstone National Park (YNP), where no timber harvesting has occurred. Estimates of downed CWD consumed or converted to charcoal during an intense crown fire were also made in YNP. No significant differences in biomass of downed CWD more than 7.5 cm in diameter were detected between burned stands and those following a single clear-cut. However, the total mass of downed CWD plus the mass of snags that will become CWD was nearly twice as high in burned stands than in clear-cuts. In YNP, approximately 8% of the downed CWD was consumed by fire and an additional 8% was converted to charcoal, for an estimated loss of about 16%. In contrast, approximately four times more wood (70%) was removed by clear-cutting. Considering all CWD more than 7.5 cm in diameter that was either still present in the stand or removed by harvesting, slash treatment, or burning, clear-cut stands lost an average of 80 Mg ha⁻¹ whereas stands that burned gained an average of 95 Mg ha⁻¹. Some CWD remains as slash and stumps left behind after harvesting, but stands subjected to repeated harvesting will have forest floor and surface soil characteristics that are beyond the historic range of variability of naturally developing stands. Received 16 November 1999; Accepted 31 May 2000.     

Deadwood stocks increase with selective logging and large tree frequency in Gabon

Deadwood is a major component of aboveground biomass (AGB) in tropical forests and is important as habitat and for nutrient cycling and carbon storage. With deforestation and degradation taking place throughout the tropics, improved understanding of the magnitude and spatial variation in deadwood is vital for the development of regional and global carbon budgets. However, this potentially important carbon pool is poorly quantified in Afrotropical forests and the regional drivers of deadwood stocks are unknown. In the first large‐scale study of deadwood in Central Africa, we quantified stocks in 47 forest sites across Gabon and evaluated the effects of disturbance (logging), forest structure variables (live AGB, wood density, abundance of large trees), and abiotic variables (temperature, precipitation, seasonality). Average deadwood stocks (measured as necromass, the biomass of deadwood) were 65 Mg ha^?1 or 23% of live AGB. Deadwood stocks varied spatially with disturbance and forest structure, but not abiotic variables. Deadwood stocks increased significantly with logging (+38 Mg ha^?1) and the abundance of large trees (+2.4 Mg ha^?1 for every tree >60 cm dbh). Gabon holds 0.74 Pg C, or 21% of total aboveground carbon in deadwood, a threefold increase over previous estimates. Importantly, deadwood densities in Gabon are comparable to those in the Neotropics and respond similarly to logging, but represent a lower proportion of live AGB (median of 18% in Gabon compared to 26% in the Neotropics). In forest carbon accounting, necromass is often assumed to be a constant proportion (9%) of biomass, but in humid tropical forests this ratio varies from 2% in undisturbed forest to 300% in logged forest. Because logging significantly increases the deadwood carbon pool, estimates of tropical forest carbon should at a minimum use different ratios for logged (mean of 30%) and unlogged forests (mean of 18%).     

Gaps and Soil C Dynamics in Old Growth Northern Hardwood–Hemlock Forests

Old growth forest soils are large C reservoirs, but the impacts of tree-fall gaps on soil C in these forests are not well understood. The effects of forest gaps on soil C dynamics in old growth northern hardwood–hemlock forests in the upper Great Lakes region, USA, were assessed from measurements of litter and soil C stocks, surface C efflux, and soil microbial indices over two consecutive growing seasons. Forest floor C was significantly less in gaps (19.0 Mg C ha⁻¹) compared to gap-edges (39.5 Mg C ha⁻¹) and the closed forest (38.0 Mg C ha⁻¹). Labile soil C (coarse particulate organic matter, cPOM) was significantly less in gaps and edges (11.1 and 11.2 Mg C ha⁻¹) compared to forest plots (15.3 Mg C ha⁻¹). In situ surface C efflux was significantly greater in gaps (12.0 Mg C ha⁻¹ y⁻¹) compared to edges and the closed forest (9.2 and 8.9 Mg C ha⁻¹ y⁻¹). Microbial biomass N (MBN) was significantly greater in edges (0.14 Mg N ha⁻¹) than in the contiguous forest (0.09 Mg N ha⁻¹). The metabolic quotient (qCO₂) was significantly greater in the forest (0.0031 mg CO₂ h⁻¹ g⁻¹/mg MBC g⁻¹) relative to gaps or edges (0.0014 mg CO₂ h⁻¹ g⁻¹/mg MBC g⁻¹). A case is made for gaps as alleviators of old growth forest soil C saturation. Relative to the undisturbed closed forest, gaps have significantly less labile C, significantly greater in situ surface C efflux, and significantly lower decreased qCO₂ values.     

Simulating root responses to grazing of a Mongolian grassland ecosystem

A new Sim-CYCLE grazing model has been obtained by combining a grazing model (Seligman et al. 1992, Ecol. Model. 60: 45–61) with the Sim-CYCLE model (Ito and Oikawa 2002, Ecol. Model. 151: 143–176). The new model has been validated against a set of field data obtained at Kherlen Bayaan-Ulaan (KBU) grassland. On the basis of the model, the root responses to grazing of KBU grassland have been studied under different conditions of stocking rates and precipitation. Model results indicate that both below-ground biomass (BB) and below-ground net primary production (BNPP) generally decrease with increasing stocking rate. However, if stocking rate is not higher than 0.7 sheep ha⁻¹, a sustainable state of the grassland ecosystem can be achieved after about 100 years, which suggests that the maximum sustainable stocking rate at KBU should be 0.7 sheep ha⁻¹. At the sustainable state, the maximum BB in a year is about 11 Mg DM ha⁻¹ under non-grazing condition, 5 Mg DM ha⁻¹ under 0.4 sheep ha⁻¹ stocking rate, and 4 Mg DM ha⁻¹ under 0.7 sheep ha⁻¹ stocking rate; the BNPP is 1.3 Mg DM ha⁻¹ year⁻¹ under non-grazing condition, and 0.6 Mg DM ha⁻¹ year⁻¹ under 0.4 sheep ha⁻¹ stocking rate, and 0.4 Mg DM ha⁻¹ year⁻¹ under 0.7 sheep ha⁻¹ stocking rate. Ratio of non-assimilation organ to assimilation organ (C/F) increases with increasing stocking rate. The C/F ratio is 10.99 under non-grazing conditions, and 12.11 under 0.7 sheep ha⁻¹ stocking rate. Root turnover rate decreases with increasing stocking rate. The rate is 12% each year under non-grazing conditions, and 11% each year under 0.7 sheep ha⁻¹ stocking rate. In addition, the effect of grazing on the grassland ecosystem under different scenarios of precipitation is also analyzed. Both BB and BNPP increase with increased precipitation, and vice versa. When precipitation is set to be 10% higher than the averaged from 1993 to 2002, the maximum sustainable stocking rate is 0.8 sheep ha⁻¹, and when the precipitation is set to be 15% lower than the averaged, the maximum sustainable stocking rate is 0.6 sheep ha⁻¹.     

Biometric-based estimation of net ecosystem production in a mature Japanese cedar (<Emphasis Type="Italic">Cryptomeria japonica</Emphasis>) plantation beneath a flux tower

Quantification of carbon budgets and cycling in Japanese cedar (Cryptomeria japonica D. Don) plantations is essential for understanding forest functions in Japan because these plantations occupy about 20% of the total forested area. We conducted a biometric estimate of net ecosystem production (NEP) in a mature Japanese cedar plantation beneath a flux tower over a 4-year period. Net primary production (NPP) was 7.9 Mg C ha⁻¹ year⁻¹ and consisted mainly of tree biomass increment and aboveground litter production. Respiration was calculated as 6.8 (soil) and 3.3 (root) Mg C ha⁻¹ year⁻¹. Thus, NEP in the plantation was 4.3 Mg C ha⁻¹ year⁻¹. In agreement with the tower-based flux findings, this result suggests that the Japanese cedar plantation was a strong carbon sink. The biometric-based NEP was higher among most other types of Japanese forests studied. Carbon sequestration in the mature plantation was characterized by a larger increment in tree biomass and lower mortality than in natural forests. Land-use change from natural forest to Japanese cedar plantation might, therefore, stimulate carbon sequestration and change the carbon allocation of NPP from an increment in coarse woody debris to an increase in tree biomass.     

Carbon, nitrogen and phosphorus in volcanic soils following afforestation with native birch (Betula pubescens) and introduced larch (Larix sibirica) in Iceland

Afforestation has become an important tool for soil protection and land reclamation in Iceland. Nevertheless, the harsh climate and degraded soils are growth-limiting for trees, and little is know about changes in soil nutrients in maturing forests planted on the volcanic soils. In the present chronosequence study, changes in C, N and total P in soil (0–10 and 10–20 cm depth) and C and N in foliar tissue were investigated in stands of native Downy birch (Betula pubescens Enrh.) and the in Iceland introduced Siberian larch (Larix sibirica Ledeb.). The forest stands were between 14 and 97 years old and were established on heath land that had been treeless for centuries. Soils were Andosols derived from basaltic material and rhyolitic volcanic ash. A significant effect of tree species was only found for the N content in foliar tissue. Foliar N concentrations were significantly higher and foliar C/N ratios significantly lower in larch needles than in birch leaves. There was no effect of stand age. Changes in soil C and the soil nutrient status with time after afforestation were little significant. Soil C concentrations in 0–10 cm depth in forest stands older than 30 years were significantly higher than in heath land and forest stands younger than 30 years. This was attributed to a slow accumulation of organic matter. Soil N concentrations and soil P_tot were not affected by stand age. Nutrient pools in the two soil layers were calculated for an average weight of soil material (400 Mg soil ha⁻¹ in 0–10 cm depth and 600 Mg soil ha⁻¹ in 10–20 cm depth, respectively). Soil nutrient pools did not change significantly with time. Soil C pools were in average 23.6 Mg ha⁻¹ in the upper soil layer and 16.9 Mg ha⁻¹ in the lower soil layer. The highest annual increase in soil C under forest compared to heath land was 0.23 Mg C ha⁻¹ year⁻¹ in 0–10 cm depth calculated for the 53-year-old larch stand. Soil N pools were in average 1.0 Mg N ha⁻¹ in both soil layers and did not decrease with time despite a low N deposition and the uptake and accumulation of N in biomass of the growing trees. Soil P_tot pools were in average 220 and 320 kg P ha⁻¹ in the upper and lower soil layer, respectively. It was assumed that mycorrhizal fungi present in the stands had an influence on the availability of N and P to the trees. Responsible Editor: Hans Lambers.     

Soil Nutrients Limit Fine Litter Production and Tree Growth in Mature Lowland Forest of Southwestern Borneo

Efforts to improve models of terrestrial productivity and to understand the function of tropical forests in global carbon cycles require a mechanistic understanding of spatial variation in aboveground net primary productivity (ANPP) across tropical landscapes. To help derive such an understanding for Borneo, we monitored aboveground fine litterfall, woody biomass increment and ANPP (their sum) in mature forest over 29 months across a soil nutrient gradient in southwestern Kalimantan. In 30 (0.07 ha) plots stratified throughout the watershed (∼340 ha, 8–190 m a.s.l.), we measured productivity and tested its relationship with 27 soil parameters. ANPP across the study area was among the highest reported for mature lowland tropical forests. Aboveground fine litterfall ranged from 5.1 to 11.0 Mg ha⁻¹ year⁻¹ and averaged 7.7 ± 0.4 (mean ± 95 C.I.). Woody biomass increment ranged from 5.8 to 23.6 Mg ha⁻¹ year⁻¹ and averaged 12.0 ± 2.0. Growth of large trees (≥60 cm dbh) contributed 38–82% of plot-wide biomass increment and explained 92% of variation among plots. ANPP, the sum of these parameters, ranged from 11.1 to 32.3 Mg ha⁻¹ year⁻¹ and averaged 19.7 ± 2.2. ANPP was weakly related to fine litterfall (r ² = 0.176), but strongly related to growth of large trees at least 60 cm dbh (r ² = 0.848). Adjusted ANPP after accounting for apparent “mature forest bias” in our sampling method was 17.5 ± 1.2 Mg ha⁻¹ year⁻¹.Relating productivity measures to soil parameters showed that spatial patterning in productivity was significantly related to soil nutrients, especially phosphorus (P). Fine litterfall increased strongly with extractable P (r ² = 0.646), but reached an asymptote at moderate P levels, whereas biomass increment (r ² = 0.473) and ANPP (r ² = 0.603) increased linearly across the gradient. Biomass increment of large trees was more frequently and strongly related to nutrients than small trees, suggesting size dependency of tree growth on nutrients. Multiple linear regression confirmed the leading importance of soil P, and identified Ca as a potential co-limiting factor. Our findings strongly suggest that (1) soil nutrients, especially P, limit aboveground productivity in lowland Bornean forests, and (2) these forests play an important, but changing role in carbon cycles, as canopy tree logging alters these terrestrial carbon sinks. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Carbon and nitrogen storage in an age-sequence of <Emphasis Type="Italic">Pinus densiflora</Emphasis> stands in Korea

The carbon (C) and nitrogen (N) storage capabilities of Pinus densiflora in six different stand ages (10, 27, 30, 32, 44, and 71 years old) were investigated in Korea. Thirty sample trees were destructively harvested and 12 were excavated. Samples from the above and belowground tree components, coarse woody debris (CWD), forest floor, and mineral soil (0–30 cm) were collected. Tree biomass was highest in the 71-year-old stand (202.8 t ha⁻¹) and lowest in the 10-year-old stand (18.4 t ha⁻¹). C and N storage in the mineral soil was higher in the 71-year-old stand than in the other stands, mainly due to higher soil C and N concentrations. Consequently, the total ecosystem C and N storage (tree+forest floor+CWD+soil) was positively correlated with stand age: increasing from a minimum in the 10 year old stand (18.8 t C ha⁻¹ and 1.3 t N ha⁻¹) to a maximum in the 71-year-old stand (201.4 t C ha⁻¹ and 8.5 t N ha⁻¹). The total ecosystem C storage showed a similar sigmoidal pattern to that of tree C storage as a function of the age-sequence, while N storage in the CWD, forest floor and mineral soil showed no significant temporal trends. Our results provide important insights that will increase our understanding of C and N storage in P. densiflora stands and our ability to predict changes according to stand age in the region.