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.     

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.     

Carbon storage in post-mining forest soil,the role of tree biomass and soil bioturbation

Carbon storage in aboveground tree biomass and soil organic matter (in depth of A layer development i.e., up to 20 cm) was studied in 22–32 year-old post-mining sites in the northwest of the Czech Republic. Four replicated sites afforested with different tree species (spruce, pine, larch, oak, lime or alder) were compared with sites left to natural regeneration which were dominated by aspen, birch and willow. No topsoil was applied at the sites; hence carbon accumulation resulted from in situ soil development on alkaline tertiary clays that were dumped on the heaps. In aboveground tree biomass, carbon storage ranged from 17.0 ± 5.9 (mean ± SEM) to 67.6 ± 5.9 t ha⁻¹ and the rate of C accumulation increased from 0.60 ± 0.09 to 2.31 ± 0.23 t ha⁻¹ year⁻¹ (natural regeneration < pine < spruce < oak < lime < alder < larch). Carbon storage in soil organic matter varied from 4.5 ± 3.7 to 38.0 ± 7.1 t ha⁻¹ and the rate of C accumulation in soil organic matter increased from 0.15 ± 0.05 to 1.28 ± 0.34 t ha⁻¹ year⁻¹ at sites in the order: natural regeneration < spruce < pine, oak < larch < alder < lime. Carbon storage in the soil was positively correlated with aboveground tree biomass. Soil carbon was equivalent to 98.1% of the carbon found in aboveground tree biomass at lime dominated sites, but only 21.8% at sites with natural regeneration. No significant correlation was found between C storage in soil and aboveground litter input. Total soil carbon storage was correlated positively and significantly with earthworm density, and occurrence of earthworm cast in topsoil, which indicated that bioturbation could play an important role in soil carbon storage. Hence, not only restoring of wood production, but also restoring of soil community is critical for C storage in soil and whole ecosystem.     

Biomass and net production in a bambooPhyllostachys bambusoides stand

Biomass and net production were measured in aPhyllostachys bambusoides stand in Kyoto Prefecture, central Japan, which had carried out gregarious flowering in 1969 and has been recovering vegetatively. The culm density fluctuated around an average value of 12 040 ha⁻¹ during the research period (1985–91). Annual recruirment and mortality rates of culms were 1340 and 1133 ha⁻¹, respectively. The mean diameter at breast height increased from 7.28 cm in 1985 to 8.68 cm in 1991, and the biomass of culms increased from 71.3 to 111.6t ha⁻¹ over the same time period. Branch and leaf biomasses were almost constant, 10.0 and 9.4t ha⁻¹ on average, respectively. The leaf area index of the stand was 11.6 ha ha⁻¹, which is one of the largest values found in Japanese forests. The belowground biomass of 32.6t ha⁻¹ for rhizomes and 14.8t ha⁻¹ for fine roots resulted in the smaller ratio of aboveground parts to the root system (2.38) than those determined for forest stands. The amount of litterfall, excluding culms and large branches, was large (9.13t ha⁻¹ year⁻¹), corresponding to those measured in equatorial stands. The aboveground net production was 24.6t ha⁻¹ year⁻¹, larger than the average value reported for forest stands under similar weather conditions.     

Nitrogen dynamics and soil nitrate retention in a <Emphasis Type="Italic">Coffea arabica</Emphasis>—<Emphasis Type="Italic">Eucalyptus deglupta</Emphasis> agroforestry system in Southern Costa Rica

Nitrogen fertilization is a key factor for coffee production but creates a risk of water contamination through nitrate (NO₃⁻) leaching in heavily fertilized plantations under high rainfall. The inclusion of fast growing timber trees in these coffee plantations may increase total biomass and reduce nutrient leaching. Potential controls of N loss were measured in an unshaded coffee (Coffea arabica L.) plot and in an adjacent coffee plot shaded with the timber species Eucalyptus deglupta Blume (110 trees ha⁻¹), established on an Acrisol that received 180 kg N ha⁻¹ as ammonium-nitrate and 2,700 mm yr⁻¹ rainfall. Results of the one year study showed that these trees had little effect on the N budget although some N fluxes were modified. Soil N mineralization and nitrification rates in the 0–20 cm soil layer were similar in both systems (≈280 kg N ha⁻¹ yr⁻¹). N export in coffee harvest (2002) was 34 and 25 kg N ha⁻¹ yr⁻¹ in unshaded and shaded coffee, and N accumulation in permanent biomass and litter was 25 and 45 kg N ha⁻¹ yr⁻¹, respectively. The losses in surface runoff (≈0.8 kg mineral N ha⁻¹ yr⁻¹) and N₂O emissions (1.9 kg N ha⁻¹ yr⁻¹) were low in both cases. Lysimeters located at 60, 120, and 200 cm depths in shaded coffee, detected average concentrations of 12.9, 6.1 and 1.2 mg NO₃⁻-N l⁻¹, respectively. Drainage was slightly reduced in the coffee-timber plantation. NO₃⁻ leaching at 200 cm depth was about 27 ± 10 and 16 ± 7 kg N ha⁻¹ yr⁻¹ in unshaded and shaded coffee, respectively. In both plots, very low NO₃⁻ concentrations in soil solution at 200 cm depth (and in groundwater) were apparently due to NO₃⁻ adsorption in the subsoil but the duration of this process is not presently known. In these conventional coffee plantations, fertilization and agroforestry practices must be refined to match plant needs and limit potential NO₃⁻ contamination of subsoil and shallow soil water.     

Effects of soil chemistry on tropical forest biomass and productivity at different elevations in the equatorial Andes

The dependence of aboveground biomass and productivity of tropical forests on soil fertility is not fully understood, since previous studies yielded contrasting results. Here, we quantify aboveground biomass (AGB) and stem wood production, and examine the impact of soil chemistry on these parameters in mature tropical forest stands of the equatorial Andes in Ecuador. In 80 plots of 0.04 ha at four elevation levels (500, 1,000, 1,500 and 2,000 m a.s.l., total sample area = 3.2 ha), we measured ten important soil chemical parameters, inventoried all trees ≥10 cm dbh and monitored stem diameter growth with dendrometer tapes in 32 plots. Top canopy height and stem density significantly decreased from 500 to 2,000 m, while tree basal area increased and AGB remained invariant (344 ± 17 Mg DM ha^?1, mean ± SE) with elevation. Wood specific gravity (WSG) showed a significant, but small, decrease. Stem wood production decreased from 4.5 to 3.2 Mg DM ha^?1 year^?1 along the transect, indicating a higher biomass turnover at lower elevations. The only soil variable that covaried with AGB was exchangeable K in the topsoil. WSG increased with decreases in N mineralisation rate, soil pH and extractable Ca and P concentrations. Structural equation modelling (SEM) revealed that nitrogen availability acts on stem wood production only indirectly through a negative relation between N mineralisation rate and WSG, and a positive effect of a lowered WSG on stem growth. The SEM analysis showed neither direct nor indirect effects of resin-extractable P on wood production, but a negative P influence on AGB. We conclude that nitrogen availability significantly influences productivity in these Andean forests, but both N and P are affecting wood production mainly indirectly through alterations in WSG and stem density; the growth-promoting effect of N is apparently larger than that of P.     

Ecosystem Carbon Storage Across the Grassland–Forest Transition in the High Andes of Manu National Park, Peru

Improved management of carbon storage by terrestrial biomes has significant value for mitigating climate change. The carbon value of such management has the potential to provide additional income to rural communities and provide biodiversity and climate adaptation co-benefits. Here, we quantify the carbon stores in a 49,300-ha landscape centered on the cloud forest–grassland transition of the high Andes in Manu National Park, Peru. Aboveground carbon densities were measured across the landscape by field sampling of 70 sites above and below the treeline. The forest near the treeline contained 63.4 ± 5.2 Mg C ha⁻¹ aboveground, with an additional 13.9 ± 2.8 Mg C ha⁻¹ estimated to be stored in the coarse roots, using a root to shoot ratio of 0.26. Puna grasslands near the treeline were found to store 7.5 ± 0.7 Mg C ha⁻¹ in aboveground biomass. Comparing our result to soil data gathered by Zimmermann and others (Ecosystems 13:62–74, 2010), we found the ratio of belowground:aboveground carbon decreased from 15.8 on the puna to 8.6 in the transition zone and 2.1 in the forest. No significant relationships were found between carbon densities and slope, altitude or fire disturbance history, though grazing (for puna) was found to reduce aboveground carbon densities significantly. We scaled our study sites to the study region with remote sensing observations from Landsat. The carbon sequestration potential of improved grazing management and assisted upslope treeline migration was also estimated. Afforestation of puna at the treeline could generate revenues of US $1,374 per ha over the project lifetime via commercialization of the carbon credits from gains in aboveground carbon stocks. Uncertainties in the fate of the large soil carbon stocks under an afforestation scenario exist.     

Altitudinal changes in carbon storage of temperate forests on Mt Changbai, Northeast China

A number of studies have investigated regional and continental scale patterns of carbon (C) stocks in forest ecosystems; however, the altitudinal changes in C storage in different components (vegetation, detritus, and soil) of forest ecosystems remain poorly understood. In this study, we measured C stocks of vegetation, detritus, and soil of 22 forest plots along an altitudinal gradient of 700–2,000 m to quantify altitudinal changes in carbon storage of major forest ecosystems (Pinus koraiensis and broadleaf mixed forest, 700–1,100 m; Picea and Abies forest, 1,100–1,800 m; and Betula ermanii forest, 1,800–2,000 m) on Mt Changbai, Northeast China. Total ecosystem C density (carbon stock per hectare) averaged 237 t C ha⁻¹ (ranging from 112 to 338 t C ha⁻¹) across all the forest stands, of which 153 t C ha⁻¹ (52–245 t C ha⁻¹) was stored in vegetation biomass, 14 t C ha⁻¹ (2.2–48 t C ha⁻¹) in forest detritus (including standing dead trees, fallen trees, and floor material), and 70 t C ha⁻¹ (35–113 t C ha⁻¹) in soil organic matter (1-m depth). Among all the forest types, the lowest vegetation and total C density but the highest soil organic carbon (SOC) density occurred in Betula ermanii forest, whereas the highest detritus C density was observed in Picea and Abies forest. The C density of the three ecosystem components showed distinct altitudinal patterns: with increasing altitude, vegetation C density decreased significantly, detritus C density first increased and then decreased, and SOC density exhibited increasing but insignificant trends. The allocation of total ecosystem C to each component exhibited similar but more significant trends along the altitudinal gradient. Our results suggest that carbon storage and partitioning among different components in temperate forests on Mt Changbai vary greatly with forest type and altitude.     

Carbon storage,structure and composition of miombo woodlands in Tanzania’s Eastern Arc Mountains

We determine the aboveground biomass and carbon storage (ABGC) of trees and the herbaceous layer in miombo woodland in the Eastern Arc Mountains (EAM) of Tanzania. In four 1‐ha sample plots in Nyanganje and Kitonga Forests, we measured all trees ≥10 cm diameter alongside height and wood mass density. The plots contained an average of 20 tree species ha^?1 (range 11–29) and 344 stems ha^?1 (range 281–382) with Shannon diversity values of 1.05 and 1.25, respectively. We weighted nine previously published woody savannah allometric models based on whether: (i) the model was derived from the same geographical region; (ii) the model included tree height/wood mass density in addition to stem diameter; and (iii) sample size was used to fit the model. The weighted mean ABGC storage from the nine models range from 13.5 ± 2 to 29.8 ± 5 Mg ha^?1. Measured ABGC storage in the herbaceous layer, using the wet combustion method, adds 0.55 ± 0.02 Mg C ha^?1. Estimates suggest that EAM miombo woodlands store a range of 13–30 Mg ha^?1 of carbon. Although the estimates suggest that miombo woodlands store significant quantities of carbon, caution is required as this is the first estimate based on in situ data.     

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.     

Tree species and wood ash affect soil in Michigan’s Upper Peninsula

Tree species and wood ash application in plantations of short-rotation woody crops (SRWC) may have important effects on the soil productive capacity through their influence on soil organic matter (SOM) and exchangeable cations. An experiment was conducted to assess changes in soil C and N contents and pH within the 0–50 cm depth, and exchangeable cation (Ca²⁺, Mg²⁺, K⁺, and Na⁺) and extractable acidity concentrations within the 0–10 cm depth. The effects of different species (European larch [Larix decidua P. Mill.], aspen [Populus tremula L. × Populus tremuloides Michx.], and four poplar [Populus spp.] clones) and wood ash applications (0, 9, and 18 Mg ha⁻¹) on soil properties were evaluated, using a common garden experiment (N = 70 stands) over 7 years of management in Michigan’s Upper Peninsula. Soils were of the Onaway series (fine-loamy, mixed, active, frigid Inceptic Hapludalfs). The NM-6 poplar clone had the greatest soil C and N contents in almost all ash treatment levels. Soil C contents were 7.5, 19.4, and 10.7 Mg C ha⁻¹ greater under the NM-6 poplar than under larch in the ash-free, medium-, and high-level plots, respectively. Within the surface layer, ash application increased soil C and N contents (P < 0.05) through the addition of about 0.7 Mg C ha⁻¹ and 3 kg N ha⁻¹ with the 9 Mg ha⁻¹ ash application (twofold greater C and N amounts were added with the 18 Mg ha⁻¹ application). During a decadal time scale, tree species had no effects—except for K⁺—on the concentrations of the exchangeable cations, pH, and extractable acidity. In contrast, ash application increased soil pH and the concentration of Ca²⁺ (P < 0.05), from 5.2 ± 0.4 cmol_c kg⁻¹ (ash-free plots) to 8.6 ± 0.4 cmol_c kg⁻¹ (high-level ash plots), and tended to increase the concentration of Mg²⁺ (P < 0.1), while extractable acidity was reduced (P < 0.05) from 5.6 ± 0.2 cmol_c kg⁻¹ (ash-free plots) to 3.7 ± 0.2 cmol_c kg⁻¹ (high-level plots). Wood ash application, within certain limits, not only had a beneficial effect on soil properties important to the long-term productivity of fast-growing plantations but also enhanced long-term soil C sequestration.     

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.     

Decomposition and carbon cycling of dead trees in tropical forests of the central Amazon

Decomposition rate constants were measured for boles of 155 large dead trees (>10 cm diameter) in central Amazon forests. Mortality data from 21 ha of permanent inventory plots, monitored for 10–15 years, were used to select dead trees for sampling. Measured rate constants varied by over 1.5 orders of magnitude (0.015–0.67 year^–1), averaging 0.19 year^–1 with predicted error of 0.026 year. Wood density and bole diameter were significantly and inversely correlated with rate constants. A tree of average biomass was predicted to decompose at 0.17 year^–1. Based on mortality data, an average of 7.0 trees ha^–1 year^–1 died producing 3.6 Mg ha^–1 year^–1 of coarse litter (>10 cm diameter). Mean coarse litter standing-stocks were predicted to be 21 Mg ha^–1, with a mean residence time of 5.9 years, and a maximum mean carbon flux to the atmosphere of 1.8 Mg C ha^–1 year^–1. Total litter is estimated to be partitioned into 16% fine wood, 30% coarse wood, and 54% non-woody litter (e.g., leaves, fruits, flowers). Decomposition rate constants for coarse litter were compiled from 20 globally distributed studies. Rates were highly correlated with mean annual temperature, giving a respiration quotient (Q ₁₀) of 2.4 (10°C^–1). Received: 14 June 1999 / Accepted: 31 August 1999     

Fire in the Brazilian Amazon 2. Biomass, nutrient pools and losses in cattle pastures

Conversion to cattle pasture is the most common fate of the ≈426,000 km² of tropical forest that has been deforested in the Brazilian Amazon. Yet little is known about the biomass, C, nutrient pools, or their responses to the frequent fires occurring in these pastures. We sampled biomass, nutrient pools and their losses or transformation during fire in three Amazonian cattle pastures with typical, but different, land-use histories. Total aboveground biomass (TAGB) ranged from to 53 to 119 Mg ha⁻¹. Residual wood debris from the forests that formally occupied the sites composed the majority of TAGB (47–87%). Biomass of fine fuels, principally pasture grasses, was ≈16–29 Mg ha⁻¹. Grasses contained as much as 52% of the aboveground K pool and the grass and litter components combined composed as much as 88% of the aboveground P pool. Fires consumed 21–84% of the TAGB. Losses of C to the atmosphere ranged from 11 to 21 Mg ha⁻¹ and N losses ranged from 205 to 261 kg ha⁻¹. Losses of S, P, Ca, and K were <33 kg ha⁻¹. There were no changes in surface soil (0–10 cm) nutrient concentration in pastures compared to adjacent primary forests. Fires occur frequently in cattle pastures (i.e., about every 2 years) and pastures are now likely the most common type of land burned in Amazonia. The first 6 years of a pastures existence would likely include the primary forest slash fire and three pasture fires. Based upon our results, the cumulative losses of N from these fires would be 1935 kg ha⁻¹ (equivalent to 94% of the aboveground pool of primary forest). Postfire aboveground C pools in old pastures are as low as 3% of those in adjacent primary forest. The initial primary forest slash fire and the repeated fires occurring in the pastures result in the majority of aboveground C and nutrient pools being released via combustion processes rather than decomposition processes. Received: 6 January 1997 / Accepted: 2 September 1997     

Invasion by an exotic tree alters above and belowground ecosystem components

With the widespread introduction and invasion of exotic plants there is a need for studies that quantify alterations of basic ecosystem structure and function. Ecosystem invasion by Melaleuca quinquenervia significantly altered both above- and belowground ecosystem components in this study. We measured the quantity and nutrient concentration of the litterfall, litter layer, and soil; microbial biomass pools; and rates of potentially mineralizable nitrogen and soil oxygen demand. Annual litterfall was 4.9 times higher in the non-invaded sites and contained 1.9 times more phosphorus than invaded sites. Non-invaded plots contained a larger litter layer compared to invaded plots: 2.4 ± 1.2 kg m⁻² and 0.62 ± 0.3 kg m⁻² , respectively. Lower nutrient concentration and quantity of the litter layer in the invaded plots led to changes in the aboveground storage of nutrients. In the invaded plots there was four times less carbon, seven times less nitrogen, and ten times less phosphorus stored in the organic litter layer compared to the non-invaded plots. Microbial biomass nutrient pools were consistently lower at both the 0–5 cm and 5–15 cm depth in the invaded soils compared to non-invaded soils, indicating a plant mediated change. Although M. quinquenervia altered microbial community structure, microbial activities were not different between invaded and non-invaded plots at either depth as measured by rates of soil oxygen demand and potentially mineralizable nitrogen. These changes may affect both native plant growth and water quality, and may act to promote and maintain site dominance by M. quinquenervia.     

Mechanisms of plant regeneration during succession after shifting cultivation in eastern Amazonia

The role of seed bank, seed rain, and regeneration from seedlings and sprouts after swidden agriculture was compared in 5-, 10- and 20-year-old secondary forest and in a primary forest in Bragantina, Pará, Brazil. The seed bank (0–5 cm soil depth) was largest in the 5-year-old forest (1190 ± 284 seeds m⁻²) and decreased nearly ten-fold with age to 137 ± 19 seeds m⁻² in the primary forest. The highest seed rain was in the 5-year-old forest (883 ± 230 seeds m⁻² year⁻¹) and the least in the primary forest (220 ± 80 seeds m⁻² year⁻¹). Large plants (≥5 cm dbh) had more individuals and species that regenerated from sprouts than from seeds and the most abundant tree species in the secondary forest stands of all ages appear to be maintained by sprouting. The smaller individuals (≥1 m tall, <5 cm dbh) in the 5-year-old forest were mainly from sprouts, but those in the older secondary forests originated mainly from seeds. These results show that at the beginning of succession, although many species can be introduced to swidden fallow from seed bank and seed rain, it is the sprout that is the main source of recruits of primary forest species in secondary forests in Bragantina.     

Fine Root Dynamics and Forest Production Across a Calcium Gradient in Northern Hardwood and Conifer Ecosystems

Losses of soil base cations due to acid rain have been implicated in declines of red spruce and sugar maple in the northeastern USA. We studied fine root and aboveground biomass and production in five northern hardwood and three conifer stands differing in soil Ca status at Sleepers River, VT; Hubbard Brook, NH; and Cone Pond, NH. Neither aboveground biomass and production nor belowground biomass were related to soil Ca or Ca:Al ratios across this gradient. Hardwood stands had 37% higher aboveground biomass (P = 0.03) and 44% higher leaf litter production (P < 0.01) than the conifer stands, on average. Fine root biomass (<2 mm in diameter) in the upper 35 cm of the soil, including the forest floor, was very similar in hardwoods and conifers (5.92 and 5.93 Mg ha⁻¹). The turnover coefficient (TC) of fine roots smaller than 1 mm ranged from 0.62 to 1.86 y⁻¹ and increased significantly with soil exchangeable Ca (P = 0.03). As a result, calculated fine root production was clearly higher in sites with higher soil Ca (P = 0.02). Fine root production (biomass times turnover) ranged from 1.2 to 3.7 Mg ha⁻¹ y⁻¹ for hardwood stands and from 0.9 to 2.3 Mg ha⁻¹ y⁻¹ for conifer stands. The relationship we observed between soil Ca availability and root production suggests that cation depletion might lead to reduced carbon allocation to roots in these ecosystems.     

Aboveground net production and dry matter allocation ofPinus pumila forests in the Kiso mountain range,central Japan

Aboveground net production rates of the subalpine stone pine (Pinus pumila) forests in central Japan were estimated by the summation method; net production was defined as the sum of annual biomass increment and annual loss due to death. In the two pine stands of different scrub heights, P1 (200 cm) and P2 (140 cm), aboveground biomass reached 177 and 126 ton ha⁻¹, respectively. Leaf biomass was about 14 ton ha⁻¹ in each stand. The estimates of aboveground net production during the 2 year period (1987–1989) averaged 4.1 and 3.7 ton ha⁻¹ y⁻¹ in P1 and P2, respectively, which were at the lowest among the pine forests in the world. Two indices of efficiency of energy fixation, that is, the ratio of net production to the total radiation during a growing season and the ratio of net production to total radiation per unit of leaf weight, were evaluated. Both efficiency indices for the twoP. pumila stands fell in the range obtained for other Japanese evergreen conifer forests. This suggested that the low annual net production of the stone pine stands were mainly due to a limitation in the length of the growing season. The pine forests were also characterized by a small allocation (about 17%) of aboveground net production into biomass increment, in comparison with other evergreen conifer forest types. Annual net carbon gain in theP. pumila forests was suggested to be largely invested in leaf production at the expense of the growth of woody parts.     

Seasonal dynamics of fine root biomass, root length density, specific root length, and soil resource availability in a Larix gmelinii plantation

Fine root turnover is a major pathway for carbon and nutrient cycling in terrestrial ecosystems and is most likely sensitive to many global change factors. Despite the importance of fine root turnover in plant C allocation and nutrient cycling dynamics and the tremendous research efforts in the past, our understanding of it remains limited. This is because the dynamics processes associated with soil resources availability are still poorly understood. Soil moisture, temperature, and available nitrogen are the most important soil characteristics that impact fine root growth and mortality at both the individual root branch and at the ecosystem level. In temperate forest ecosystems, seasonal changes of soil resource availability will alter the pattern of carbon allocation to belowground. Therefore, fine root biomass, root length density (RLD) and specific root length (SRL) vary during the growing season. Studying seasonal changes of fine root biomass, RLD, and SRL associated with soil resource availability will help us understand the mechanistic controls of carbon to fine root longevity and turnover. The objective of this study was to understand whether seasonal variations of fine root biomass, RLD and SRL were associated with soil resource availability, such as moisture, temperature, and nitrogen, and to understand how these soil components impact fine root dynamics in Larix gmelinii plantation. We used a soil coring method to obtain fine root samples (⩽2 mm in diameter) every month from May to October in 2002 from a 17-year-old L. gmelinii plantation in Maoershan Experiment Station, Northeast Forestry University, China. Seventy-two soil cores (inside diameter 60 mm; depth intervals: 0–10 cm, 10–20 cm, 20–30 cm) were sampled randomly from three replicates 25 m × 30 m plots to estimate fine root biomass (live and dead), and calculate RLD and SRL. Soil moisture, temperature, and nitrogen (ammonia and nitrates) at three depth intervals were also analyzed in these plots. Results showed that the average standing fine root biomass (live and dead) was 189.1 g·m⁻²·a⁻¹, 50% (95.4 g·m⁻²·a⁻¹) in the surface soil layer (0–10 cm), 33% (61.5 g·m⁻²·a⁻¹), 17% (32.2 g·m⁻²·a⁻¹) in the middle (10–20 cm) and deep layer (20–30cm), respectively. Live and dead fine root biomass was the highest from May to July and in September, but lower in August and October. The live fine root biomass decreased and dead biomass increased during the growing season. Mean RLD (7,411.56 m·m⁻³·a⁻¹) and SRL (10.83 m·g⁻¹·a⁻¹) in the surface layer were higher than RLD (1 474.68 m·m⁻³·a⁻¹) and SRL (8.56 m·g⁻¹·a⁻¹) in the deep soil layer. RLD and SRL in May were the highest (10 621.45 m·m⁻³ and 14.83m·g⁻¹) compared with those in the other months, and RLD was the lowest in September (2 198.20 m·m⁻³) and SRL in October (3.77 m·g⁻¹). Seasonal dynamics of fine root biomass, RLD, and SRL showed a close relationship with changes in soil moisture, temperature, and nitrogen availability. To a lesser extent, the temperature could be determined by regression analysis. Fine roots in the upper soil layer have a function of absorbing moisture and nutrients, while the main function of deeper soil may be moisture uptake rather than nutrient acquisition. Therefore, carbon allocation to roots in the upper soil layer and deeper soil layer was different. Multiple regression analysis showed that variation in soil resource availability could explain 71–73% of the seasonal variation of RLD and SRL and 58% of the variation in fine root biomass. These results suggested a greater metabolic activity of fine roots living in soil with higher resource availability, which resulted in an increased allocation of carbohydrate to these roots, but a lower allocation of carbohydrate to those in soil with lower resource availability. __________ Translated from Acta Phytoecologica Sinica, 2005, 29(3): 403–410 [译自: 植物生态学报, 2005, 29(3): 403–410]     

No changes in soil organic carbon and nitrogen following long-term prescribed burning and livestock exclusion in the Sudan-savanna woodlands of Burkina Faso

Fire and overgrazing reduce aboveground biomass, leading to land degradation and potential impacts on soil organic carbon (SOC) and total nitrogen (TN) dynamics. However, empirical data are lacking on how prescribed burning and livestock exclusion impact SOC in the long-term. Here we analyse the effects of 19 years of prescribed annual burning and livestock exclusion on tree density, SOC and TN concentrations in the Sudanian savanna ecoregion at two sites (Tiogo and Laba) in Burkina Faso. Results revealed that neither livestock exclusion nor prescribed burning had significant impact on SOC and TN concentrations. The results at both sites indicate that 19 years of livestock and fire exclusion did not result in a significant increase in tree density compared to grazing and annual prescribed burning. The overall mean (± SEM) of SOC stocks in the 0–50 cm depth increment in the unburnt (53.5 ± 4.7 Mg C ha⁻¹) and annually burnt (56.4 ± 4.3 Mg C ha⁻¹) plots at Tiogo were not statistically different. Similarly, at Laba there was no significant difference between the corresponding figures in the unburnt (37.9 ± 2.6 Mg ha⁻¹) and in the annually burnt plots (38.6 ± 1.9 Mg ha⁻¹). Increases in belowground inputs from root turnover may have countered changes in aboveground biomass, resulting in no net change in SOC and TN. We conclude that, contrary to our expectation and current policy recommendations, restricting burning or grazing did not result in increase in SOC stocks in this dry savanna ecosystem.