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Soil respiration is a major source of atmospheric CO2. If it increases with warming, it will counteract efforts to minimize climate change. To improve understanding of environmental controls over soil CO2 emission, we applied generalized linear modeling to a large dataset of in situ measurements of short-term soil respiration rate, with associated environmental attributes, which was gathered over multiple years from four locations that varied in climate, soil type, and vegetation. Soil respiration includes many CO2-producing processes: we theorized that different environmental factors could limit each process distinctly, thereby diminishing overall CO2 emissions. A baseline model that included soil temperature, soil volumetric water content, and their interaction was effective in estimating soil respiration at all four locations (p < 0.0001). Model fits, based on model log likelihoods, improved continuously as additional covariates were added, including mean daily air temperature, enhanced vegetation index (EVI), and quadratic terms for soil temperature and water content, and their interactions. The addition of land cover and its direct interactions with environmental variables further improved model fits. Significant interactions between covariates were observed at each location and at every stage of analysis, but the interaction terms varied among sites and models, and did not consistently maintain importance in more complex models. A main-effects model was therefore tested, which included soil temperature and water content, their quadratic effects, EVI, and air temperature, but no interactions. In that case all six covariates were significant (p < 0.0001) when applied across sites. We infer that local-scale soil-CO2 emissions are commonly co-limited by EVI and air temperature, in addition to soil temperature and water content. Importantly, the quadratic soil temperature and moisture terms were significantly negative: estimated soil-CO2 emissions declined when soil temperature exceeded 22.5°C, and as soil moisture differed from the optimum of 0.27 m3 m-3., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2023 Raich et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Young secondary forests and plantations in the moist tropics often have rapid rates of biomass accumulation and thus sequester large amounts of carbon. Here, we compare results from mature forest and nearby 15-20 year old tree plantations in lowland Costa Rica to evaluate differences in allocation of carbon to aboveground production and root systems. We found that the tree plantations, which had fully developed, closed canopies, allocated more carbon belowground - to their root systems - than did mature forest. This increase in belowground carbon allocation correlated significantly with aboveground tree growth but not with canopy production (i.e., leaf fall or fine litter production). In contrast, there were no correlations between canopy production and either tree growth or belowground carbon allocation. Enhanced allocation of carbon to root systems can enhance plant nutrient uptake, providing nutrients beyond those required for the production of short-lived tissues such as leaves and fine roots, and thus enabling biomass accumulation. Our analyses support this deduction at our site, showing that enhanced allocation of carbon to root systems can be an important mechanism promoting biomass accumulation during forest growth in the moist tropics. Identifying factors that control when, where and for how long this occurs would help us to improve models of forest growth and nutrient cycling, and to ascertain the role that young forests play in mitigating increased atmospheric carbon dioxide.

Fast-growing forests such as tropical secondary forests can accumulate large amounts of carbon (C), and thereby play an important role in the atmospheric CO(2) balance. Because nitrogen (N) cycling is inextricably linked with C cycling, the question becomes: Where does the N come from to match high rates of C accumulation? In unique experimental 16-y-old plantations established in abandoned pasture in lowland Costa Rica, we used a mass-balance approach to quantify N accumulation in vegetation, identify sources of N, and evaluate differences among tree species in N cycling. The replicated design contained four broad-leaved evergreen tree species growing under similar environmental conditions. Nitrogen uptake was rapid, reaching 409 (± 30) kg · ha(-1) · y(-1), double the rate reported from a Puerto Rican forest and greater than four times that observed at Hubbard Brook Forest (New Hampshire, USA). Nitrogen amassed in vegetation was 874 (± 176) kg · ha(-1), whereas net losses of soil N (0-100 cm) varied from 217 (±146) to 3,354 (± 915) kg · ha(-1) (P = 0.018) over 16 y. Soil C:N, δ(13)C values, and N budgets indicated that soil was the main source of biomass N. In Vochysia guatemalensis, however, N fixation contributed >60 kg · ha(-1) · y(-1). All species apparently promoted soil N turnover, such that the soil N mean residence time was 32-54 y, an order of magnitude lower than the global mean. High rates of N uptake were associated with substantial N losses in three of the species, in which an average of 1.6 g N was lost for every gram of N accumulated in biomass.

In the moist tropical forest biome, which cycles carbon (C) rapidly and stores huge amounts of C, the impacts of individual species on C balances are not well known. In one of the earliest replicated experimental sites for investigating growth of native tropical trees, we examined traits of tree species in relation to their effects on forest C balances, mechanisms of influence, and consequences for C sequestration. The monodominant stands, established in abandoned pasture in 1988 at La Selva Biological Station, Costa Rica, contained five species in a complete randomized block design. Native species were: Hieronyma alchorneoides, Pentaclethra macroloba, Virola koschnyi, and Vochysia guatemalensis. The exotic species was Pinus patula. By 16 years, the lack of differences among species in some attributes suggested strong abiotic control in this environment, where conditions are very favorable for growth, These attributes included aboveground net primary productivity (ANPP), averaging 11.7 Mg C x ha(-1) x yr(-1) across species, and soil organic C (0-100 cm, 167 Mg C/ha). Other traits differed significantly, however, indicating some degree of biological control. In Vochysia plots, both aboveground biomass of 99 Mg C/ha, and belowground biomass of 20 Mg C/ha were 1.8 times that of Virola (P = 0.02 and 0.03, respectively). Differences among species in overstory biomass were not compensated by understory vegetation. Belowground NPP of 4.6 Mg C x ha(-1) yr(-1) in Hieronyma was 2.4 times that of Pinus (P < 0.01). Partitioning of NPP to belowground components in Hieronyma was more than double that of Pinus (P = 0.03). The canopy turnover rate in Hieronyma was 42% faster than that of Virola (P < 0.01). Carbon sequestration, highest in Vochysia (7.4 Mg C x ha(-1) x yr(-1), P = 0.02), averaged 5.2 Mg C x ha(-1) x yr(-1), close to the annual per capita fossil fuel use in the United States of 5.3 Mg C. Our results indicated that differences in species effects on forest C balances were related primarily to differences in growth rates, partitioning of C among biomass components, tissue turnover rates, and tissue chemistry. Inclusion of those biological attributes may be critical for robust modeling of C cycling across the moist tropical forest biome.

Prolific fine root growth coupled with small accumulations of dead fine roots indicate rapid rates of fine root production, mortality and decay in young tree plantations in lowland Costa Rica. However, published studies indicate that fine roots decay relatively slowly in tropical forests. To resolve this discrepancy, we used the intact-core technique to quantify first-year decay rates of fine roots in four single-species plantations of native tree species. We tested three hypotheses: first, that fine roots from different tree species would decay at different rates; second, that species having rapid fine root growth rates would also have rapid rates of fine root decay; and third, that differences in fine root decay among species could be explained by fine root chemistry variables previously identified as influencing decay rates. Fine roots in Virola koschnyi plantations decayed very slowly (k = 0.29 +/- 0.15 year(-1)); those of Vochysia guatemalensis decayed seven times faster (k = 2.00 +/- 0.13 year(-1)). Decay rates of the remaining two species, Hieronyma alchorneoides and Pentaclethra macroloba, were 1.36 and 1.28 year(-1), respectively. We found a positive, marginally significant correlation between fine root decay rates and the relative growth rates of live fine roots (R = 0.93, n = 4, P = 0.072). There was a highly significant negative correlation between fine root decay and fine root lignin:N (R = 0.99, P = 0.01), which supports the use of lignin:N as a decay-controlling factor within terrestrial ecosystem models. The decay rates that we observed in this single study location encompassed the entire range of fine root decay rates previously observed in moist tropical forests, and thus suggest great potential for individual tree species to alter belowground organic matter and nutrient dynamics within a biotically rich rainforest environment.

Evergreen broad-leaved tropical forests can have high rates of productivity and large accumulations of carbon in plant biomass and soils. They can therefore play an important role in the global carbon cycle, influencing atmospheric CO2 concentrations if climate warms. We applied meta-analyses to published data to evaluate the apparent effects of temperature on carbon fluxes and storages in mature, moist tropical evergreen forest ecosystems. Among forests, litter production, tree growth, and belowground carbon allocation all increased significantly with site mean annual temperature (MAT); total net primary productivity (NPP) increased by an estimated 0.2-0.7 Mg C x ha(-1) x yr(-1) x degrees C(-1). Temperature had no discernible effect on the turnover rate of aboveground forest biomass, which averaged 0.014 yr(-1) among sites. Consistent with these findings, forest biomass increased with site MAT at a rate of 5-13 Mg C x ha(-1) x degrees C(-1). Despite greater productivity in warmer forests, soil organic matter accumulations decreased with site MAT, with a slope of -8 Mg C x ha(-1) x degrees C(-1), indicating that decomposition rates of soil organic matter increased with MAT faster than did rates of NPP. Turnover rates of surface litter also increased with temperature among forests. We found no detectable effect of temperature on total carbon storage among moist-tropical evergreen forests, but rather a shift in ecosystem structure, from low-biomass forests with relatively large accumulations of detritus in cooler sites, to large-biomass forests with relatively smaller detrital stocks in warmer locations. These results imply that, in a warmer climate, conservation of forest biomass will be critical to the maintenance of carbon stocks in moist tropical forests.

We use a mechanistically based ecosystem simulation model to describe and analyze the spatial and temporal patterns of terrestrial net primary productivity (NPP) in South America. The Terrestrial Ecosystem Model (TEM) is designed to predict major carbon and nitrogen fluxes and pool sizes in terrestrial ecosystems at continental to global scales. Information from intensively studies field sites is used in combination with continental-scale information on climate, soils, and vegetation to estimate NPP in each of 5888 non-wetland, 0.5° latitude °0.5° longitude grid cells in South America, at monthly time steps. Preliminary analyses are presented for the scenario of natural vegetation throughout the continent, as a prelude to evaluating human impacts on terrestrial NPP. The potential annual NPP of South America is estimated to be 12.5 Pg/yr of carbon (26.3 Pg/yr of organic matter) in a non-wetland area of 17.0 ° 10 6 km 2 . More than 50% of this production occurs in the tropical and subtropical evergreen forest region. Six independent model runs, each based on an independently derived set of model parameters, generated mean annual NPP estimates for the tropical evergreen forest region ranging from 900 to 1510 g°m - 2 °yr - 1 of carbon, with an overall mean of 1170 g°m - 2 °yr - 1 . Coefficients of variation in estimated annual NPP averaged 20% for any specific location in the evergreen forests, which is probably within the confidence limits of extant NPP measurements. Predicted rates of mean annual NPP in other types of vegetation ranged from 95 g°m - 2 °yr - 1 in arid shrublands to 930 g°m @ ?yr - 1 in savannas, and were within the ranges measured in empirical studies. The spatial distribution of predicted NPP was directly compared with estimates made using the Miami mode of Lieth (1975). Overall, TEM predictions were °10% lower than those of the Miami model, but the two models agreed closely on the spatial patterns of NPP in south America. Unlike previous models, however, TEM estimates NPP monthly, allowing for the evaluation of seasonal phenomena. This is an important step toward integration of ecosystem models with remotely sensed information, global climate models, and atmospheric transport models, all of which are evaluated at comparable spatial and temporal scales. Seasonal patterns of NPP in South America are correlated with moisture availability in most vegetation types, but are strongly influenced by seasonal differences in cloudiness in the tropical evergreen forests. On an annual basis, moisture availability was the factor that was correlated most strongly with annual NPP in South America, but differences were again observed among vegetation types. These results allow for the investigation and analysis of climatic controls over NPP at continental scales, within and among vegetation types, and within years. Further model validation is needed. Nevertheless, the ability to investigate NPP-environment interactions with a high spatial and temporal resolution at continental scales should prove useful if not essential for rigorous analysis of the potential effects of global climate changes on terrestrial ecosystems., (© 1991 by the Ecological Society of America.)