Response of tree biomass and wood litter to disturbance in a Central Amazon forest

We developed an individual-based stochastic-empirical model to simulate the carbon dynamics of live and dead trees in a Central Amazon forest near Manaus, Brazil. The model is based on analyses of extensive field studies carried out on permanent forest inventory plots, and syntheses of published studies. New analyses included: (1) growth suppression of small trees, (2) maximum size (trunk base diameter) for 220 tree species, (3) the relationship between growth rate and wood density, and (4) the growth response of surviving trees to catastrophic mortality (from logging). The model simulates a forest inventory plot, and tracks recruitment, growth, and mortality of live trees, decomposition of dead trees (coarse litter), and how these processes vary with changing environmental conditions. Model predictions were tested against aggregated field data, and also compared with independent measurements including maximum tree age and coarse litter standing stocks. Spatial analyses demonstrated that a plot size of ~10 ha was required to accurately measure wood (live and dead) carbon balance. With the model accurately predicting relevant pools and fluxes, a number of model experiments were performed to predict forest carbon balance response to perturbations including: (1) increased productivity due to CO₂ fertilization, (2) a single semi-catastrophic (10%) mortality event, (3) increased recruitment and mortality (turnover) rates, and (4) the combined effects of increased turnover, increased tree growth rates, and decreased mean wood density of new recruits. Results demonstrated that carbon accumulation over the past few decades observed on tropical forest inventory plots (~0.5 Mg C ha^–1 year^–1) is not likely caused by CO₂ fertilization. A maximum 25% increase in woody tissue productivity with a doubling of atmospheric CO₂ only resulted in an accumulation rate of 0.05 Mg C ha^–1 year^–1 for the period 1980–2020 for a Central Amazon forest, or an order of magnitude less than observed on the inventory plots. In contrast, model parameterization based on extensive data from a logging experiment demonstrated a rapid increase in tree growth following disturbance, which could be misinterpreted as carbon sequestration if changes in coarse litter stocks were not considered. Combined results demonstrated that predictions of changes in forest carbon balance during the twenty-first century are highly dependent on assumptions of tree response to various perturbations, and underscores the importance of a close coupling of model and field investigations.     

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.     

Dynamics of aboveground big woody organs in a foothill dipterocarp forest,West Sumatra,Indonesia

The dynamics of aboveground big woody organs over 10 cm diameter was studied at a mature foothill dipterocarp forest in West Sumatra. The biomass of big woody organs was estimated to be 519 m³ ha⁻¹ or 408 metric ton ha⁻¹ by means of a pipe model theory. The diameter distribution showed a convex curve and the mode was found at a diameter of about 20 cm. The standing mass of big dead woody litter on the forest floor was 116 m³ ha⁻¹, which accounted for 22% by voume or 9.5% by weight of the biomass of living organs respectively. Thedbh observation with two 1-ha plots for 4 yr and 5 yr respectively revealed that the average net production rate was 9.5 ton ha⁻¹ yr⁻¹. The death rate (7.9 ton ha⁻¹ yr⁻¹) accounted for 83% of the net production rate and was nearly equivalent to the decay rate (7.5 ha⁻¹ yr⁻¹) of dead wood on the forest floor. The balance between the death and decay rates was confirmed for each diameter class. Average turnover periods for big woody organs and dead woody litter were estimated to be 43 and 8.1 yr, respectively. Standing masses of live anddead woody materials accumulated in the study forest were approximately equal to those obtained in a mature tropical lowland rainforest, whereas the flow rates were lower, being only 70% of the corresponding values.     

Carbon and nitrogen pools in a mangrove stand of <Emphasis Type="Italic">Kandelia obovata</Emphasis> (S., L.) Yong: vertical distribution in the soil–vegetation system

Attempts were made to quantify the carbon and nitrogen pools in a monospecific and pioneer mangrove stand of Kandelia obovata Sheue, Liu & Yong, Okinawa Island, Japan. The leaf C and N concentrations on a leaf area basis decreased with increasing PPFD (Photosysthetic Photon Flux Density). The total C and N stocks in foliage were estimated as 3.55 Mg ha^–1 and 0.105 Mg ha^–1, respectively. The bark (45.6–48.6% for C and 0.564–0.842% for N) contained significantly higher amount of C (P < 0.05) and N (P < 0.01) than wood (46.2–47.8% for C and 0.347–0.914% N). The total C stock of stem was 23.2 Mg ha^–1 in wood and 8.33 Mg ha^–1 in bark, and the total N stock was 0.222 Mg ha^–1 in wood and 0.116 Mg ha^–1 in bark. The root wood (37.1–45.0%) contained significantly higher amount of C than root bark (35.4–40.7%) (P < 0.01). The total C stock of root was 14.2 Mg ha^–1 in wood and 12.6 Mg ha^–1 in bark, and the total N stock of root was 0.157 Mg ha^–1 in wood and 0.155 Mg ha^–1 in bark. The soil organic C and total N stocks within 1 m soil depth were estimated as 57.3 Mg ha^–1 and 2.73 Mg ha^–1, respectively. The C pool in aboveground biomass (35.1 Mg ha^–1) was 1.3 times as large as that in belowground biomass (26.9 Mg ha^–1). However, the soil organic C pool (57.3 Mg ha^–1) was similar to the total C pool (62.0 Mg ha^–1) of vegetation, indicating that the mangrove stored a large part of production in the soil. About 50% of the C was in the soil. The N pool in aboveground biomass (0.442 Mg ha^–1) was 1.4 times as large as that in belowground biomass (0.312 Mg ha^–1). The soil N stock was 3.3 times as large as the biomass N stock (0.754 Mg ha^–1).     

Forest structure and carbon dynamics in Amazonian tropical rain forests

Living trees constitute one of the major stocks of carbon in tropical forests. A better understanding of variations in the dynamics and structure of tropical forests is necessary for predicting the potential for these ecosystems to lose or store carbon, and for understanding how they recover from disturbance. Amazonian tropical forests occur over a vast area that encompasses differences in topography, climate, and geologic substrate. We observed large differences in forest structure, biomass, and tree growth rates in permanent plots situated in the eastern (near Santarém, Pará), central (near Manaus, Amazonas) and southwestern (near Rio Branco, Acre) Amazon, which differed in dry season length, as well as other factors. Forests at the two sites experiencing longer dry seasons, near Rio Branco and Santarém, had lower stem frequencies (460 and 466 ha^–1 respectively), less biodiversity (Shannon–Wiener diversity index), and smaller aboveground C stocks (140.6 and 122.1 Mg C ha^–1) than the Manaus site (626 trees ha^–1, 180.1 Mg C ha^–1), which had less seasonal variation in rainfall. The forests experiencing longer dry seasons also stored a greater proportion of the total biomass in trees with >50 cm diameter (41–45 vs 30% in Manaus). Rates of annual addition of C to living trees calculated from monthly dendrometer band measurements were 1.9 (Manaus), 2.8 (Santarém), and 2.6 (Rio Branco) Mg C ha^–1 year^–1. At all sites, trees in the 10–30 cm diameter class accounted for the highest proportion of annual growth (38, 55 and 56% in Manaus, Rio Branco and Santarém, respectively). Growth showed marked seasonality, with largest stem diameter increment in the wet season and smallest in the dry season, though this may be confounded by seasonal variation in wood water content. Year-to-year variations in C allocated to stem growth ranged from nearly zero in Rio Branco, to 0.8 Mg C ha^–1 year^–1 in Manaus (40% of annual mean) and 0.9 Mg C ha^–1 year^–1 (33% of annual mean) in Santarém, though this variability showed no significant relation with precipitation among years. Initial estimates of the C balance of live wood including recruitment and mortality as well as growth suggests that live wood biomass is at near steady-state in Manaus, but accumulating at about 1.5 Mg C ha^–1 at the other two sites. The causes of C imbalance in living wood pools in Santarém and Rio Branco sites are unknown, but may be related to previous disturbance at these sites. Based on size distribution and growth rate differences in the three sites, we predict that trees in the Manaus forest have greater mean age (~240 years) than those of the other two forests (~140 years).     

Root turnover in a beech and a spruce stand of the Belgian Ardennes

The theoretical basis of fine root turnover estimation in forest soils is discussed, in relation to appropriate experimental techniques of measurement. After sequential coring, the correct expression is the sum of significant positive increments of live and dead roots of the various diameter categories, to which the transfer of dead roots to organic matter derived from roots, OMDR, has to be added. This should not be confounded with dead root mineralization. The transfer rates should first be estimated in root dimensions and not in weight of dry matter. The measurements were carried out in a 120 year old beech (Fagus sylvatica L.) stand and a 35 year old Norway spruce (Picea abies Karst) stand, in the Eastern Ardennes, Belgium. The turnover rate of fine roots (diam. <5 mm) was 4393 kg ha⁻¹ year⁻¹ (root dry weight), including 711.2 kg ha⁻¹ year⁻¹ for dead root transfer to OMDR, for beech. For spruce, turnover rate was 7011 kg ha⁻¹ year⁻¹ (root dry weight), including 1498 kg ha⁻¹ year⁻¹ for dead root transfer to OMDR. Under beech, there was a slight root density increase in spring. No seasonal fluctuations were observed under spruce, but a strong irreversible drop in live root growth was found in the later season 1980–1981, corresponding to a decrease of tree height growth and trunk radius increment. Turnover rates were further expressed in dry weight and in amounts of elements (kg ha⁻¹ year⁻¹) (Ca, Mg, K, Na, Al, N, P, S). Correlative relations between root dimensions and dry weight and element concentrations show that the derived values, and in particular root specific density (dry weight volume⁻¹) vary according to species, root category, and seasonal sampling. Various schemes of seasonal variations of root growth, described in Europe, show that the major dependance on general climate is obscured by environmental factors (soil, exposure, species). It is suggested that root density fluctuation approach the steady state on an annual basis under mild Atlantic conditions.     

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.     

Nutrient transfer by leaf litterfall during a sugar maple decline episode at Lake Clair watershed,Québec,Canada

In a declining sugar maple (SM) stand, we tested the hypothesis that an increasing relative abundance of American beech (AB) and yellow birch (YB) would improve litter quality by providing a higher proportion of litterfall richer in base cations and lower in acidity. From 1989 to 2006, SM leaf fall diminished from 59% (1,718 kg ha⁻¹ year⁻¹) to 36% (915 kg ha⁻¹ year⁻¹) of the total leaf fall biomass. Overall, the increase in AB and YB litterfall compensated for the SM decrease, resulting in constant annual leaf litterfall fluxes (2,803 kg ha⁻¹ year⁻¹) over the period studied. However, because the leaf litter for AB and YB had Ca and Mg concentrations 2–3 times higher than did SM, Ca and Mg concentrations and fluxes in leaf litterfall significantly increased between 1989 and 2006. Leaf litterfall of AB and YB also has a higher base/acid ratio than SM. Consequently, changes in forest composition following SM decline led to a clear improvement in litterfall quality in terms of base cations content and fluxes and acid–base properties.     

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.     

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.     

Respiration from coarse wood litter in central Amazon forests

Respiration from coarse litter (trunks and large branches >10 cm diameter) was studied in central Amazon forests. Respiration ratesvaried over almost two orders of magnitude (1.003–0.014 µg Cg^–1 C min^–1, n = 61), and weresignificantly correlated with wood density (r² _adj= 0.42), and moisture content (r² _adj= 0.39). Additional samples taken from a nearby pasture indicatedthat wood moisture content was the most important factor controllingrespiration rates across sites (r² _adj =0.65). Based on average coarse litter wood density and moisture content,the mean long-term carbon loss rate due to respiration was estimated tobe 0.13 yr^–1 (range of 95% prediction interval(PI) = 0.11–0.15 yr^–1). Comparing meanrespiration rate with mean mass loss (decomposition) rate from aprevious study, respiratory emissions to the atmosphere from coarselitter were predicted to be 76% (95% PI =65–88%) of total carbon loss, or about 1.9 (95% PI= 1.6–2.2) Mg C ha^–1yr^–1. Optimum respiration activity corresponded toabout 2.5 g H₂O g^–1 dry wood, and severelyrestricted respiration to < 0.5 g H₂O g^–1dry wood. Respiration from coarse litter in central Amazon forests iscomparable in magnitude to decomposing fine surface litter (e.g. leaves,twigs) and is an important carbon cycling component when characterizingheterotrophic respiration budgets and net ecosystem exchange(NEE).     

Symptoms of nitrogen saturation in an aggrading forested watershed in western Maryland

The objective of this study was to evaluate the nitrogen (N) biogeochemistry of an 18–22 year old forested watershed in western Maryland. We hypothesized that this watershed should not exhibit symptoms of N saturation. This watershed was a strong source of nitrate (NO₃ ⁻) to the stream in all years, with a mean annual export of 9.5 kg N ha⁻¹ year⁻¹ and a range of 4.4–18.4 kg N ha⁻¹ year⁻¹. During the 2001 and 2002 water years, wet deposition of inorganic N was 9.0 kg N ha⁻¹ year⁻¹ and 6.3 kg N ha⁻¹ year⁻¹, respectively. Watershed N export rates in 2001 and 2002 water years were 4.2 kg N ha⁻¹ year⁻¹ and 5.3 kg N ha⁻¹ year⁻¹, respectively. During the wetter water years of 2003 and 2004, the watershed exported 15.0 kg N ha⁻¹ year⁻¹ and 18.4 kg N ha⁻¹ year⁻¹, rates that exceeded annual wet deposition of N by a factor of two (7.5 kg N ha⁻¹ year⁻¹ in 2003) and three (5.5 kg N ha⁻¹ year⁻¹ in 2004). Consistent with the high rates of N export, were high concentrations (2.1–3.3%) of N in foliage, wood (0.3%) and fine roots, low C:N ratios in the forest floor (17–24) and mineral soil (14), high percentages (83–96%) of the amount of mineralized N that was nitrified and elevated N concentrations (up to 3 mg N l⁻¹) in soil solution. Although this watershed contained a young aggrading forest, it exhibited several symptoms of N saturation commonly observed in more mature forests.     

Woody debris contribution to the carbon budget of selectively logged and maturing mid-latitude forests

Woody debris (WD) is an important component of forest C budgets, both as a C reservoir and source of CO₂ to the atmosphere. We used an infrared gas analyzer and closed dynamic chamber to measure CO₂ efflux from downed coarse WD (CWD; diameter≥7.5 cm) and fine WD (FWD; 7.5 cm>diameter≥2 cm) to assess respiration in a selectively logged forest and a maturing forest (control site) in the northeastern USA. We developed two linear regression models to predict WD respiration: one based on WD temperature, moisture, and size (R ²=0.57), and the other on decay class and air temperature (R ²=0.32). WD respiration (0.28±0.09 Mg C ha⁻¹ year⁻¹) contributed only ≈2% of total ecosystem respiration (12.3±0.7 Mg C ha⁻¹ year⁻¹, 1999–2003), but net C flux from CWD accounted for up to 30% of net ecosystem exchange in the maturing forest. C flux from CWD on the logged site increased modestly, from 0.61±0.29 Mg C ha⁻¹ year⁻¹ prior to logging to 0.77±0.23 Mg C ha⁻¹ year⁻¹ after logging, reflecting increased CWD stocks. FWD biomass and associated respiration flux were ≈7 times and ≈5 times greater, respectively, in the logged site than the control site. The net C flux associated with CWD, including inputs and respiratory outputs, was 0.35±0.19 Mg C ha⁻¹ year⁻¹ (net C sink) in the control site and −0.30±0.30 Mg C ha⁻¹ year⁻¹ (net C source) in the logged site. We infer that accumulation of WD may represent a small net C sink in maturing northern hardwood forests. Disturbance, such as selective logging, can enlarge the WD pool, increasing the net C flux from the WD pool to the atmosphere and potentially causing it to become a net C source.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Climatically induced interannual variability in aboveground production in forest-tundra and northern taiga of central Siberia

To investigate the variability of primary production of boreal forest ecosystems under the current climatic changes, we compared the dynamics of annual increments and productivity of the main components of plant community (trees, shrubs, mosses) at three sites in the north of Siberia (Russia). Annual radial growth of trees and shrubs was mostly defined by summer temperature regime (positive correlation), but climatic response of woody plants was species specific and depends on local conditions. Dynamics of annual increments of mosses were opposite to tree growth. The difference in climatic response of the different vegetation components of the forest ecosystems indicates that these components seem to be adapted to use climatic conditions during the short and severe northern summer, and decreasing in annual production of one component is usually combined with the increase of other component productivity. Average productivity in the northern forest ecosystems varies from 0.05 to 0.14 t ha⁻¹ year⁻¹ for trees, from 0.05 to 0.18 t ha⁻¹ year⁻¹ for shrubs and from 0.54 to 0.66 t ha⁻¹ year⁻¹ for mosses. Higher values of tree productivity combined with lower annual moss productivity were found in sites in northern taiga in comparison with forest-tundra. Different tendencies in the productivity of the dominant species from each vegetation level (trees, shrubs, mosses) were indicated for the last 10 years studied (1990–1999): while productivity of mosses is increasing, productivity of trees is decreasing, but there is no obvious trend in the productivity of shrubs. Our results show that in the long term, the main contribution to changes in annual biomass productivity in forest-tundra and northern taiga ecosystems under the predicted climatic changes will be determined by living ground cover.     

Litterfall dynamics in carbonate and deltaic mangrove ecosystems in the Gulf of Mexico

From 1996 to 2002, we measured litterfall, standing litter crop, and litter turnover rates in scrub, basin, fringe and riverine forests in two contrasting mangrove ecosystems: a carbonate-dominated system in the Southeastern Everglades and a terrigenous-dominated system in Laguna de Terminos (LT), Mexico. We hypothesized that litter dynamics is driven by latitude, geomorphology, hydrology, soil fertility and soil salinity stress. There were significant temporal patterns in LT with litterfall rates higher during the rainy season (2.4 g m⁻² day⁻¹) than during the dry season (1.8 g m⁻² day⁻¹). Total annual litterfall was significantly higher in the riverine forest (12.8 Mg ha⁻² year⁻¹) than in the fringe and basin forests (9.7 and 5.2 Mg ha⁻² year⁻¹, respectively). In Southeastern Everglades, total annual litterfall was also significantly higher during the rainy season than during the dry season. Spatially, the scrub forest had the lowest annual litterfall (2.5 Mg ha⁻² year⁻¹), while the fringe and basin had the highest (9.1 and 6.5 Mg ha⁻² year⁻¹, respectively). In LT, annual standing litter crop was 3.3 Mg ha⁻¹ in the fringe and 2.2 Mg ha⁻¹ in the basin. Litter turnover rates were significantly higher in the fringe mangrove forest (4.1 year⁻¹) relative to the basin forests (2.2 year⁻¹). At Southeastern Everglades there were significant differences in annual standing litter crop: 1.9, 3.3 and 4.5 Mg ha⁻¹ at scrub, basin and fringe mangrove sites, respectively. Furthermore, turnover rates were similar at both basin and fringe mangrove types (2.1 and 2.0 year⁻¹, respectively) but significantly higher than scrub mangrove forest (1.3 year⁻¹). These findings suggest that litter export is important in regulating litter turnover rates in frequently flooded riverine and fringe forests, while in infrequently flooded basin forests, in situ litter decomposition controls litter turnover rates.     

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.     

Zonal distribution of coarse woody debris and its contribution to net primary production in a secondary mangrove forest

Coarse woody debris (CWD) plays an important role in long-term carbon storage in forest ecosystems. However, few studies have examined CWD in mangrove forests. A secondary mangrove forest on an estuary of the Trat River showed different structures along vegetation zones ranging from the river’s edge to inland parts of the forest (the Sonneratia–Avicennia, Avicennia, Rhizophora, and Xylocarpus zones, respectively). The mass distribution of CWD stock in downed wood and standing dead trees along these vegetation zones was evaluated. Most of the CWD stock in the Sonneratia–Avicennia and Avicennia zones was found in downed wood, while it mainly accumulated in standing dead trees in the Rhizophora and Xylocarpus zones. The total mass of CWD stock that accumulated in each zone ranged from 1.56–8.39 t ha^?1, depending on the forest structure and inundation regimes. The annual woody debris flux in each zone was calculated by summing the necromass (excluding foliage) of dead trees and coarse litter from 2010 to 2013. The average woody debris flux was 5.4 t ha^?1 year^?1, and its zonal variation principally depended on the necromass production that resulted from forest succession, high tree-density, and lightning. Over all the zones, the above- and below-ground net primary production (ANPP and BNPP, respectively) was estimated at 18.0 and 3.6 t ha^?1 year^?1, respectively. The magnitude of BNPP and its contribution to the NPP was markedly increased when fine root production was taken into consideration. The contribution of the woody debris flux without root necromass to the ANPP ranged from 12 to 28%.     

Litter and fine-root production in three types of tropical premontane rain forest in SE Venezuela

We investigated the productivity of three premontane primary forest sites in an extremely nutrient-poor environment on the Guyana shield in SE Venezuela. Aboveground litter production (total 5.58; leaves 4.30 Mg ha^–1 year^–1) was low, but comparable to other tropical forests. Due to the low nutrient status, net production of fine roots was among the highest ever reported (11.14 Mg ha^–1 year^–1). Only 20% of fine root stock was alive, 80% consisted of necromass. Similar values were obtained for dead and living root tips. Element concentrations in fine roots (including bio- and necromass), especially Ca, were low, whereas Al concentrations were relatively high.The upper limit of average fine root lifetime was 253 days. We conclude that the high proportion of necromass was mainly caused by slow mineralisation of nutrient-poor fine roots.     

The change of soil carbon stocks and fine root dynamics after land use change from a native pasture to a pine plantation

A published meta-analysis of worldwide data showed soil carbon decreasing following land use change from pasture to conifer plantation. A paired site (a native pasture with Themeda triandra dominant, and an adjacent Pinus radiata plantation planted onto the pasture 16 years ago) was set up as a case study to assess the soil carbon reduction and the possible reason for the reduction under pine, including the change in fine root (diameter <2 mm) dynamics (production and mortality). Soil analysis confirmed that soil carbon and nitrogen stocks to 100 cm under the plantation were significantly less than under the pasture by 20 and 15%, respectively. A 36% greater mass of fine root was found in the soil under the pasture than under the plantation and the length of fine root was about nine times greater in the pasture. Much less fine root length was produced and roots died more slowly under the plantation than under the pasture based on observations of fine root dynamics in minirhizotrons. The annual inputs of fine root litter to the top 100 cm soil, estimated from soil coring and minirhizotron observations, were 6.3 Mg dry matter ha⁻¹ year⁻¹ (containing 2.7 Mg C and 38.9 kg N) under the plantation, and 9.7 Mg ha⁻¹ year⁻¹ (containing 3.6 Mg C and 81.4 kg N) under the pasture. The reduced amount of carbon, following afforestation of the pasture, in each depth-layer of the soil profile correlated with the lower length of dead fine roots in the layer under the plantation compared with the pasture. This correlation was consistent with the hypothesis that the soil carbon reduction after land use change from pasture to conifer plantation might be related to change of fine root dynamics, at least in part.     

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.