Controls Over Leaf Litter Decomposition in Wet Tropical Forests

Tropical forests play a substantial role in the global carbon (C) cycle and are projected to experience significant changes in climate, highlighting the importance of understanding the factors that control organic matter decomposition in this biome. In the tropics, high temperature and rainfall lead to some of the highest rates of litter decomposition on earth, and given the near-optimal abiotic conditions, litter quality likely exerts disproportionate control over litter decomposition. Yet interactions between litter quality and abiotic variables, most notably precipitation, remain poorly resolved, especially for the wetter end of the tropical forest biome. We assessed the importance of variation in litter chemistry and precipitation in a lowland tropical rain forest in southwest Costa Rica that receives \u3e5000 mm of precipitation per year, using litter from 11 different canopy tree species in conjunction with a throughfall manipulation experiment. In general, despite the exceptionally high rainfall in this forest, simulated throughfall reductions consistently suppressed rates of litter decomposition. Overall, variation between species was greater than that induced by manipulating throughfall and was best explained by initial litter solubility and lignin:P ratios. Collectively, these results support a model of litter decomposition in which mass loss rates are positively correlated with rainfall up to very high rates of mean annual precipitation and highlight the importance of phosphorus availability in controlling microbial processes in many lowland tropical forests

Tree Species Control Rates of Free-Living Nitrogen Fixation in a Tropical Rain Forest

Tropical rain forests represent some of the most diverse ecosystems on earth, yet mechanistic links between tree species identity and ecosystem function in these forests remains poorly understood. Here, using free-living nitrogen (N) fixation as a model, we explore the idea that interspecies variation in canopy nutrient concentrations may drive significant local-scale variation in biogeochemical processes. Biological N fixation is the largest “natural” source of newly available N to terrestrial ecosystems, and estimates suggest the highest such inputs occur in tropical ecosystems. While patterns of and controls over N fixation in these systems remain poorly known, the data we do have suggest that chemical differences among tree species canopies could affect free-living N fixation rates. In a diverse lowland rain forest in Costa Rica, we established a series of vertical, canopy-to-soil profiles for six common canopy tree species, and we measured free-living N fixation rates and multiple aspects of chemistry of live canopy leaves, senesced canopy leaves, bulk leaf litter, and soil for eight individuals of each tree species. Free-living N fixation rates varied significantly among tree species for all four components, and independent of species identity, rates of N fixation ranged by orders of magnitude along the vertical profile. Our data suggest that variations in phosphorus (P) concentration drove a significant fraction of the observed species-specific variation in free-living N fixation rates within each layer of the vertical profile. Furthermore, our data suggest significant links between canopy and forest floor nutrient concentrations; canopy P was correlated with bulk leaf litter P below individual tree crowns. Thus, canopy chemistry may affect a suite of ecosystem processes not only within the canopy itself, but at and beneath the forest floor as well

Nutrient Regulation of Organic Matter Decomposition in a Tropical Rain Forest

errestrial biosphere–atmosphere CO2 exchange is dominated by tropical forests, so understanding how nutrient availability affects carbon (C) decomposition in these ecosystems is central to predicting the global C cycle\u27s response to environmental change. In tropical rain forests, phosphorus (P) limitation of primary production and decomposition is believed to be widespread, but direct evidence is rare. We assessed the effects of nitrogen (N) and P fertilization on litter-layer organic matter decomposition in two neighboring tropical rain forests in southwest Costa Rica that are similar in most ways, but that differ in soil P availability. The sites contain 100–200 tree species per hectare and between species foliar nutrient content is variable. To control for this heterogeneity, we decomposed leaves collected from a widespread neotropical species, Brosimum utile. Mass loss during decomposition was rapid in both forests, with B. utile leaves losing \u3e80% of their initial mass in (DOM) rather than direct CO2 mineralization. While P fertilization did not significantly affect mass loss in the litter layer, it did stimulate P immobilization in decomposing material, leading to increased P content and a lower C:P ratio in soluble DOM. In turn, increased P content of leached DOM stimulated significant increases in microbial mineralization of DOM in P-fertilized soil. These results show that, while nutrients may not affect mass loss during decomposition in nutrient-poor, wet ecosystems, they may ultimately regulate CO2 losses (and hence C storage) by limiting microbial mineralization of DOM leached from the litter layer to soil

Climatic, edaphic, and biotic controls over storage and turnover of carbon in soils

Soil carbon, a major component of the global carbon inventory, has significant potential for change with changing climate and human land use. We applied the Century ecosystem model to a series of forest and grassland sites distributed globally to examine large-scale controls over soil carbon. Key site-specific parameters influencing soil carbon dynamics are soil texture and foliar lignin content; accordingly, we perturbed these variables at each site to establish a range of carbon concentrations and turnover times. We examined the simulated soil carbon stores, turnover times, and C:N ratios for correlations with patterns of independent variables. Results showed that soil carbon is related linearly to soil texture, increasing as clay content increases, that soil carbon stores and turnover time are related to mean annual temperature by negative exponential functions, and that heterotrophic respiration originates from recent detritus (∼50%), microbial turnover (∼30%), and soil organic matter (∼20%) with modest variations between forest and grassland ecosystems. The effect of changing temperature on soil organic carbon (SOC) estimated by Century is dSOC/dT= 183e−0.034T. Global extrapolation of this relationship leads to an estimated sensitivity of soil C storage to a temperature of −11.1 Pg° C−1, excluding extreme arid and organic soils. In Century, net primary production (NPP) and soil carbon are closely coupled through the N cycle, so that as temperatures increase, accelerated N release first results in fertilization responses, increasing C inputs. The Century-predicted effect of temperature on carbon storage is modified by as much as 100% by the N cycle feedback. Century-estimated soil C sensitivity (−11.1 Pg° C−1) is similar to losses predicted with a simple data-based calculation (−14.1 Pg° C−1). Inclusion of the N cycle is important for even first-order predictions of terrestrial carbon balance. If the NPP-SOC feedback is disrupted by land use or other disturbances, then SOC sensitivity can greatly exceed that estimated in our simulations. Century results further suggest that if climate change results in drying of organic soils (peats), soil carbon loss rates can be high

Experimental Drought in a Tropical Rain Forest Increases Soil Carbon Dioxide Losses to the Atmosphere

Climate models predict precipitation changes for much of the humid tropics, yet few studies have investigated the potential consequences of drought on soil carbon (C) cycling in this important biome. In wet tropical forests, drought could stimulate soil respiration via overall reductions in soil anoxia, but previous research suggests that litter decomposition is positively correlated with high rainfall fluxes that move large quantities of dissolved organic matter (DOM) from the litter layer to the soil surface. Thus, reduced rainfall could also limit C delivery to the soil surface, reducing respiration rates. We conducted a throughfall manipulation experiment to investigate how 25% and 50% reductions in rainfall altered both C movement into soils and the effects of those DOM fluxes on soil respiration rates. In response to the experimental drought, soil respiration rates increased in both the −25% and −50% treatments. Throughfall fluxes were reduced by 26% and 55% in the −25% and −50% treatments, respectively. However, total DOM fluxes leached from the litter did not vary between treatments, because the concentrations of leached DOM reaching the soil surface increased in response to the simulated drought. Annual DOM concentrations averaged 7.7 ± 0.8, 11.2 ± 0.9, and 15.8 ± 1.2 mg C/L in the control, −25%, and −50% plots, respectively, and DOM concentrations were positively correlated with soil respiration rates. A laboratory incubation experiment confirmed the potential importance of DOM concentration on soil respiration rates, suggesting that this mechanism could contribute to the increase in CO2 fluxes observed in the reduced rainfall plots. Across all plots, the data suggested that soil CO2 fluxes were partially regulated by the magnitude and concentration of soluble C delivered to the soil, but also by soil moisture and soil oxygen availability. Together, our data suggest that declines in precipitation in tropical rain forests could drive higher CO2 fluxes to the atmosphere both via increased soil O2 availability and through responses to elevated DOM concentrations

Soil Microbial Dynamics and Biogeochemistry in Tropical Forests and Pastures, Southwestern Costa Rica

Tropical rain forest ecosystems are currently undergoing unprecedented rates of land conversion and land use change. Recent research suggests these activities profoundly influence nutrient cycling, but the principal mechanisms driving variation in nutrient status following land conversion are still not well understood. In this study, we used soils of varying fertility (oxisols and mollisols) in Costa Rica to investigate how conversion of tropical rain forest to cattle pasture affects the size and function of the microbial community, and to explore possible relationships between microbial dynamics and biogeochemistry. Our pasture sites are relatively lightly managed, and total pools of carbon (C), nitrogen (N), and phosphorus (P) were not significantly different from their forest counterparts. However, pools of available elements were different; most notably, plant available forms of P were significantly lower in the oxisol pasture than in the oxisol forest site. In addition, we found that land conversion led to fundamental changes in the size and activity of the soil microbial community. Microbial biomass was consistently higher in forests than in pastures, particularly in the oxisol sites, where it was more than twice the pasture value. Forest sites were also characterized by a microbial community that was more active, responded more rapidly to carbon substrate additions, and showed strong seasonal variation. Our results provide evidence that changes in biogeochemical cycling following land conversion observed here and elsewhere may be directly related to changes in microbial community structure and function

Controls Over Foliar N:P Ratios in Tropical Rain Forests

Correlations between foliar nutrient concentrations and soil nutrient availability have been found in multiple ecosystems. These relationships have led to the use of foliar nutrients as an index of nutrient status and to the prediction of broadscale patterns in ecosystem processes. More recently, a growing interest in ecological stoichiometry has fueled multiple analyses of foliar nitrogen : phosphorus (N:P) ratios within and across ecosystems. These studies have observed that N:P values are generally elevated in tropical forests when compared to higher latitude ecosystems, adding weight to a common belief that tropical forests are generally N rich and P poor. However, while these broad generalizations may have merit, their simplicity masks the enormous environmental heterogeneity that exists within the tropics; such variation includes large ranges in soil fertility and climate, as well as the highest plant species diversity of any biome. Here we present original data on foliar N and P concentrations from 150 mature canopy tree species in Costa Rica and Brazil, and combine those data with a comprehensive new literature synthesis to explore the major sources of variation in foliar N:P values within the tropics. We found no relationship between N:P ratios and either latitude or mean annual precipitation within the tropics alone. There is, however, evidence of seasonal controls; in our Costa Rica sites, foliar N:P values differed by 25% between wet and dry seasons. The N:P ratios do vary with soil P availability and/or soil order, but there is substantial overlap across coarse divisions in soil type, and perhaps the most striking feature of the data set is variation at the species level. Taken as a whole, our results imply that the dominant influence on foliar N:P ratios in the tropics is species variability and that, unlike marine systems and perhaps many other terrestrial biomes, the N:P stoichiometry of tropical forests is not well constrained. Thus any use of N:P ratios in the tropics to infer larger-scale ecosystem processes must comprehensively account for the diversity of any given site and recognize the broad range in nutrient requirements, even at the local scale

Topographic controls on soil nitrogen availability in a lowland tropical forest

Geomorphic position often correlates with nutrient cycling across landscapes. In tropical forests, topography is known to influence phosphorus (P) availability, but its effect on nitrogen (N) cycling has received less exploration, especially in lowland forests where widespread N richness is frequently assumed. Here, we report significant effects of topographic slope and landscape position on multiple aspects of the N cycle across a highly dissected lowland tropical forest on the Osa Peninsula, Costa Rica. A suite of N cycle metrics measured along a topographic sequence revealed a distinct gradient in N availability. Values of soil d15N, inorganic N pools, net nitrification rates, and nitrification potentials were all substantially lower on a flanking steep hillslope (;288) compared to a relatively flat ridge top (;68), indicating lower N availability and a less open N cycle in steep parts of the landscape. Slope soils also hosted smaller total carbon and nitrogen stocks and notably less weathered soil minerals than did ridge soils. These latter findings suggest that elevated N loss resulting from high rates of soil and particulate organic matter erosion could underpin the spatial variation in N cycling and availability. Expanding our analysis to the larger study landscape, a strong negative linear relationship between soil d15N values and surface slope angles was observed. N isotope mass balance models suggest that this pattern is most plausibly explained by an increase in N loss via erosive, non-fractioning pathways from steep zones, as most other variables commonly assumed to affect soil d15N values (such as temperature, precipitation, and vegetation type) did not vary across the sampled region. Together, these results reveal notable hillslope-scale variation in N richness and suggest an important role for nonfractionating N loss in the maintenance of this pattern. Such findings highlight the importance of geomorphology and the significant capacity of erosion to influence N availability in steepland ecosystems

Unexpected Changes in Soil Phosphorus Dynamics Along Pasture Chronosequences in the Humid Tropics

Phosphorus (P) is widely believed to limit plant growth and organic matter storage in a large fraction of the world\u27s lowland tropical rain forests. We investigated how the most common land use change in such forests, conversion to cattle pasture, affects soil P fractions along pasture chronosequences in the central Brazilian Amazon and in southwestern Costa Rica. Our sites represent a broad range in rainfall, soil type, management strategies, and total soil P (45.2–1228.0 μg P/g soil), yet we found some unexpected and at times quite similar changes in soil P in all sites. In the Brazilian sites, where rainfall is relatively low and pasture management is more intense than in the Costa Rican sites, significant losses in total soil P and soil organic carbon (SOC) were seen with pasture age on already P-deficient Oxisol and Entisol soils. However, P losses were from inorganic soil P fractions, while organic forms of soil P remained constant or increased with pasture age, despite the declines in SOC. In Costa Rica, SOC remained constant across the Oxisol sites and increased from forest to pasture on the Mollisols, while soil organic P increased with pasture age in both sequences