Soil nutrients affect spatial patterns of aboveground biomass and emergent tree density in southwestern Borneo

Studies on the relationship between soil fertility and aboveground biomass in lowland tropical forests have yielded conflicting results, reporting positive, negative and no effect of soil nutrients on aboveground biomass. Here, we quantify the impact of soil variation on the stand structure of mature Bornean forest throughout a lowland watershed (8–196 m a.s.l.) with uniform climate and heterogeneous soils. Categorical and bivariate methods were used to quantify the effects of (1) parent material differing in nutrient content (alluvium > sedimentary > granite) and (2) 27 soil parameters on tree density, size distribution, basal area and aboveground biomass. Trees ≥10 cm (diameter at breast height, dbh) were enumerated in 30 (0.16 ha) plots (sample area = 4.8 ha). Six soil samples (0–20 cm) per plot were analyzed for physiochemical properties. Aboveground biomass was estimated using allometric equations. Across all plots, stem density averaged 521 ± 13 stems ha⁻¹, basal area 39.6 ± 1.4 m² ha⁻¹ and aboveground biomass 518 ± 28 Mg ha⁻¹ (mean ± SE). Adjusted forest-wide aboveground biomass to account for apparent overestimation of large tree density (based on 69 0.3-ha transects; sample area = 20.7 ha) was 430 ± 25 Mg ha⁻¹. Stand structure did not vary significantly among substrates, but it did show a clear trend toward larger stature on nutrient-rich alluvium, with a higher density and larger maximum size of emergent trees. Across all plots, surface soil phosphorus (P), potassium, magnesium and percentage sand content were significantly related to stem density and/or aboveground biomass (R _Pearson = 0.368–0.416). In multiple linear regression, extractable P and percentage sand combined explained 31% of the aboveground biomass variance. Regression analyses on size classes showed that the abundance of emergent trees >120 cm dbh was positively related to soil P and exchangeable bases, whereas trees 60–90 cm dbh were negatively related to these factors. Soil fertility thus had a significant effect on both total aboveground biomass and its distribution among size classes. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

不同林窗马尾松凋落叶与土壤养分变化研究

以现有42年生的马尾松（Pinus massoniana）人工纯林,经过采伐形成4种不同大小有效面积的林窗（100、400、900和1 600 m²）为研究对象,以未经采伐的42年生马尾松人工纯林为对照样地,采用凋落叶分解袋法,研究不同大小有效面积林窗对马尾松凋落叶、土壤C、N、P及化学计量比和养分损失率的影响。研究结果表明：（1）不同大小有效面积林窗下的马尾松凋落叶、土壤C、N、P含量及养分损失率除土壤P含量和马尾松凋落叶P养分损失率外,均存在显著差异。随着林窗有效面积G1~G4的增大,马尾松凋落叶C、N、P含量均呈降低趋势,三者均在G3林窗体现出较小值。马尾松凋落叶C、N、P养分损失率、土壤C、N、P养分含量多呈抛物线趋势,且均在G2或G3林窗体现出最大值。（2）不同大小有效面积林窗下的马尾松凋落叶、土壤C/N/P均存在显著差异。随着林窗有效面积G1~G4的增大,马尾松人工林土壤C/N/P基本呈抛物线变化趋势,土壤C/N在G3林窗出现最大值,土壤C/P、N/P均在G2林窗体现出最大值;土壤C/N、C/P、N/P变异系数分别为13.31%、16.51%、17.21%。马尾松凋落叶C/N、C/P均在G3体现出最小值。（3）马尾松凋落叶C、N含量与土壤C、C/N/P及环境因子的相关性较强,P含量与它们的相关性较弱;C/N与土壤P、C/N/P及环境因子的相关性较强,C/P、N/P与土壤C/P及环境因子的相关性较强;C、N养分损失率与土壤C、C/N、C/P及环境因子的相关性较强,P养分损失率与土壤C、N、P含量及其化学计量比和环境因子的相关性较弱。土壤C、N、P含量及其化学计量比与环境因子的相关性较强。     

Regional‐Scale Variation in Litter Production and Seasonality in Tropical Dry Forests of Southern Mexico1

Highly seasonal rainfall creates a pulse of litterfall in the southern Yucatan peninsula region, with cascading effects on the timing of essential nutrient fluxes, microbial dynamics, and vegetation growth. I investigated whether forest age or a regional environmental gradient related to rainfall has a greater effect on patterns of litterfall in this increasingly human‐dominated landscape. Litterfall was sampled in 10–13 stands in each of three locations spanning a rainfall gradient of ca 900–1400 mm/yr. Litter was collected monthly from November 1998 through January 2000 in mature forests and in secondary forests aged 2–25 yr. Despite a substantial precipitation gradient, age was the only significant predictor of annual litter mass. Two‐ to five‐yr‐old forests produced significantly less litter than 12–25‐yr‐old secondary forests (4.6 vs. 6.2 Mg/ha/yr), but the difference between older secondary forests and mature forests (9 percent) was not significant. Litter production increased with rainfall, but not significantly so. The pattern of litterfall was similar across locations and age classes, with a peak during late March or early April. However, litterfall seasonality was most pronounced in the old secondary and mature forests. Litterfall was more evenly distributed throughout the year in forests under 10 yr old. Seasonality of litterfall was also less pronounced at the wettest site, with less disparity between peak litterfall and off‐peak months. Seasonality was not related to soil texture. Forest age and rainfall are important drivers of litterfall dynamics; however, both litter mass and degree of seasonality depended more strongly on forest age. Thus, the impact of land‐use change on litter nutrient cycling is as great, if not greater, than the constraint imposed by the major natural environmental factor affecting tropical dry forests.     

Production and Resource Use Efficiencies in N- and P-Limited Tropical Forests: A Comparison of Responses to Long-term Fertilization

At two sites at the extreme ends of a soil development chronosequence in Hawaii, we investigated whether forest responses to fertilization on young soils were similar to those on highly weathered soils and whether the initial responses were maintained after 6–11 years of fertilization. Aboveground net primary production (ANPP) was increased by nitrogen (N) application at the 300-year-old site and phosphorus (P) application at the 4.1-million-year-old site, thus confirming earlier results and their designations as N- and P-limited forests. Along with ANPP, application of the limiting element consistently increased leaf area index (LAI), radiation conversion efficiency (RCE), and foliar and litter nutrient concentrations. Fertilization did not consistently alter N or P retranslocation from senescent leaves at either site, but a comparison with other sites on the chronosequence and with a common-garden study suggests that there is a genetic basis for low foliar and litter nutrients and higher retranslocation at infertile sites vs more fertile sites. N limitation appears to be expressed as limitation to carbon gain, with long leaf lifespans and high leaf mass per area. P limitation results in high P-use efficiency and disproportionally large increases in P uptake after fertilization; a comparison with other studies indicates large investments in acquiring and storing P. Although the general responses of ANPP, LAI, and RCE were similar for the two sites, other aspects of nutrient use differ in relation to the physiological and biogeochemical roles of the two elements. Received 2 June 2000; Accepted 4 April 2001.     

Dynamics of soil carbon following destruction of tropical rainforest and the subsequent establishment of Imperata grassland in Indonesian Borneo using stable carbon isotopes

Southeast Asia has the highest rate of tropical rainforest deforestation worldwide, and large deforested areas have been replaced ultimately by the highly invasive grass Imperata cylindrica. However, information on the carbon (C) budget with such land transition is very scarce. This study presents the dynamics of soil C following rainforest destruction and the subsequent establishment of Imperata grassland in the lowland humid tropics of Indonesian Borneo using stable C isotopes. To evaluate the relative contribution of organic matter originating from primary forest (C₃) and grasslands (C₄), we compared soil C stock and natural ¹³C abundance from six sites to a depth of 100 cm using samples with a wide range of soil textures. Twelve years after the first soil sampling in the grasslands, we re‐sampled to examine temporal changes in soil organic matter. The grassland topsoil (0–5 cm) is an active layer with rapid decomposition and incorporation of fresh C (mean residence time: 7.5 year) and a substantial proportion of the stable C pool (37%). The decline in forest‐derived C was slight, even at 5–10 cm depths, and subsoil (20–100 cm depth) forest‐derived C did not change along the forest‐to‐grassland chronosequence. Grassland‐derived C stock increased significantly in the subsurface and subsoils (5–100 cm). Simulation indicated that total soil C stock (0–100 cm) increased by 18.6 Mg ha^?1 from initial primary forest (58.0 Mg ha^?1) to a new equilibrium state of the grassland (76.6 Mg ha^?1) after 30–50 years of grassland establishment. This research indicates that the soil did not function as a CO₂ source when the deforested area was replaced by Imperata grassland on the Ultisols of the Asian humid tropics. Instead, increased soil C stocks offset CO₂ emissions, with the C offset accounting for 6.6–7.4% of the loss of biomass C stock.     

Effects of Plant Growth Characteristics on Biogeochemistry and Community Composition in a Changing Climate

Vegetation growth characteristics influence ecosystem biogeochemistry and must be incorporated in models used to project biogeochemical responses to climate variations. We used a multiple-element limitation model (MEL) to examine how variations in nutrient use efficiency (NUE) and net primary production to biomass ratio (nPBR) affect changes in ecosystem C stocks after an increase in temperature and atmospheric CO₂. nPBR influences the initial rates of response, but the magnitude and direction of long-term responses are determined by NUE. MEL was used to simulate responses to climate change in communities composed of two species differing in nPBR and/or NUE. When only nPBR differed between the species, the high-nPBR species outgrew the low-nPBR species early in the simulations, but the shift in dominance was transitory because of secondary N limitations. High-NUE species were less affected by secondary N limitations and were therefore favored under elevated CO₂. Increased temperature stimulated N release from soil organic matter (SOM) and therefore favored low-NUE species. The combined release from C and N limitation under the combination of increased temperature and elevated CO₂ favored high-NUE species. High C:N litter from high-NUE species limited the N-supply rate from SOM, which favors the dominance of the high-NUE species in the short term. However, in the long term increased litter production resulted in SOM accumulation, which reestablished a N supply rate favorable to the reestablishment and dominance of the low-NUE species. Conditions then reverted to a state favorable to the high-NUE species. Received 8 October 1998; accepted 9 April 1999.