Evidence is mounting that extinctions are altering key processes important to the productivity and sustainability of Earth's ecosystems. Further species loss will accelerate change in ecosystem processes, but it is unclear how these effects compare to the direct effects of other forms of environmental change that are both driving diversity loss and altering ecosystem function. Here we use a suite of meta-analyses of published data to show that the effects of species loss on productivity and decomposition--two processes important in all ecosystems--are of comparable magnitude to the effects of many other global environmental changes. In experiments, intermediate levels of species loss (21-40%) reduced plant production by 5-10%, comparable to previously documented effects of ultraviolet radiation and climate warming. Higher levels of extinction (41-60%) had effects rivalling those of ozone, acidification, elevated CO(2) and nutrient pollution. At intermediate levels, species loss generally had equal or greater effects on decomposition than did elevated CO(2) and nitrogen addition. The identity of species lost also had a large effect on changes in productivity and decomposition, generating a wide range of plausible outcomes for extinction. Despite the need for more studies on interactive effects of diversity loss and environmental changes, our analyses clearly show that the ecosystem consequences of local species loss are as quantitatively significant as the direct effects of several global change stressors that have mobilized major international concern and remediation efforts.

Litter decomposition provides the primary source of mineral nitrogen (N) for biological activity in most terrestrial ecosystems. A 10-year decomposition experiment in 21 sites from seven biomes found that net N release from leaf litter is dominantly driven by the initial tissue N concentration and mass remaining regardless of climate, edaphic conditions, or biota. Arid grasslands exposed to high ultraviolet radiation were an exception, where net N release was insensitive to initial N. Roots released N linearly with decomposition and exhibited little net N immobilization. We suggest that fundamental constraints on decomposer physiologies lead to predictable global-scale patterns in net N release during decomposition.

As atmospheric CO 2 increases, ecosystem carbon sequestration will largely depend on how global changes in climate will alter the balance between net primary production and decomposition. The response of primary production to climatic change has been examined using well-validated mechanistic models, but the same is not true for decomposition, a primary source of atmospheric CO 2 . We used the Long-term Intersite Decomposition Experiment Team (LIDET) dataset and model-selection techniques to choose and parameterize a model that describes global patterns of litter decomposition. Mass loss was best represented by a three-pool negative exponential model, with a rapidly decomposing labile pool, an intermediate pool representing cellulose, and a recalcitrant pool. The initial litter lignin/nitrogen ratio defined the size of labile and intermediate pools. Lignin content determined the size of the recalcitrant pool. The decomposition rate of all pools was modified by climate, but the intermediate pool's decomposition rate was also controlled by relative amounts of litter cellulose and lignin (indicative of ligninencrusted cellulose). The effect of climate on decomposition was best represented by a composite variable that multiplied a water-stress function by the Lloyd and Taylor variable Q 10 temperature function. Although our model explained nearly 70% of the variation in LIDET data, we observed systematic deviations from model predictions. Below-and aboveground material decomposed at notably different rates, depending on the decomposition stage. Decomposition in certain ecosystem-specific environmental conditions was not well represented by our model; this included roots in very wet and cold soils, and aboveground litter in N-rich and arid sites. Despite these limitations, our model may still be extremely useful for global modeling efforts, because it accurately (R 2 5 0.6804) described general patterns of long-term global decomposition for a wide array of litter types, using relatively minimal climatic and litter quality data.

Summary 1.Litter decomposition recycles nutrients and causes large fluxes of carbon dioxide into the atmosphere. It is typically assumed that climate, litter quality and decomposer communities determine litter decay rates, yet few comparative studies have examined their relative contributions in tropical forests. 2. We used a short-term litterbag experiment to quantify the effects of litter quality, placement and mesofaunal exclusion on decomposition in 23 tropical forests in 14 countries. Annual precipitation varied among sites (760-5797 mm). At each site, two standard substrates ( Raphia farinifera and Laurus nobilis ) were decomposed in fine-and coarse-mesh litterbags both above and below ground for approximately 1 year. 3. Decomposition was rapid, with >95% mass loss within a year at most sites. Litter quality, placement and mesofaunal exclusion all independently affected decomposition, but the magnitude depended upon site. Both the average decomposition rate at each site and the ratio of above-to below-ground decay increased linearly with annual precipitation, explaining 60-65% of among-site variation. Excluding mesofauna had the largest impact on decomposition, reducing decomposition rates by half on average, but the magnitude of decrease was largely independent of climate. This suggests that the decomposer community might play an important role in explaining patterns of decomposition among sites. Which litter type decomposed fastest varied by site, but was not related to climate. 4. Synthesis . A key goal of ecology is to identify general patterns across ecological communities, as well as relevant site-specific details to understand local dynamics. Our pan-tropical study shows that certain aspects of decomposition, including average decomposition rates and the ratio of above-to below-ground decomposition are highly correlated with a simple climatic index: mean annual precipitation. However, we found no relationship between precipitation and effects of mesofaunal exclusion or litter type, suggesting that site-specific details may also be required to understand how these factors affect decomposition at local scales.

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Despite the importance of litter decomposition for ecosystem fertility and carbon balance, key uncertainties remain about how this fundamental process is affected by nitrogen (N) availability. Resolving such uncertainties is critical for predicting the ecosystem consequences of increased anthropogenic N deposition. Toward that end, we decomposed green leaves and senesced litter of northern pin oak (Quercus ellipsoidalis) in three forested stands dominated by northern pin oak or white pine (Pinus strobus) to compare effects of substrate N (as it differed between leaves and litter) and externally supplied N (inorganic or organic forms) on decomposition and decomposer community structure and function over four years. Asymptotic decomposition models fit the data equally well as single exponential models and allowed us to compare effects of N on both the initial decomposition rate (ka) and the level of asymptotic mass remaining (A, proportion of mass remaining at which decomposition approaches zero, i.e., the fraction of slowly decomposing litter). In all sites, both substrate N and externally supplied N (regardless of form) accelerated the initial decomposition rate. Faster initial decomposition rates corresponded to higher activity of polysaccharide‐degrading enzymes associated with externally supplied N and greater relative abundances of Gram‐negative and Gram‐positive bacteria associated with green leaves and externally supplied organic N (assessed using phospholipid fatty acid analysis, PLFA). By contrast, later in decomposition, externally supplied N slowed decomposition, increasing the fraction of slowly decomposing litter (A) and reducing lignin‐degrading enzyme activity and relative abundances of Gram‐negative and Gram‐positive bacteria. Higher‐N green leaves, on the other hand, had lower levels of A (a smaller slow fraction) than lower‐N litter. Contrasting effects of substrate and externally supplied N during later stages of decomposition likely occurred because higher‐N leaves also had considerably lower lignin, causing them to decompose more quickly throughout decomposition. In conclusion, elevated atmospheric N deposition in forest ecosystems may have contrasting effects on the dynamics of different soil carbon pools, decreasing mean residence times of active fractions in fresh litter (which would be further reduced if deposition increased litter N concentrations), while increasing those of more slowly decomposing fractions, including more processed litter.

Through litter decomposition enormous amounts of carbon is emitted to the atmosphere. Numerous large-scale decomposition experiments have been conducted focusing on this fundamental soil process in order to understand the controls on the terrestrial carbon transfer to the atmosphere. However, previous studies were mostly based on site-specific litter and methodologies, adding major uncertainty to syntheses, comparisons and meta-analyses across different experiments and sites. In the TeaComposition initiative, the potential litter decomposition is investigated by using standardized substrates (Rooibos and Green tea) for comparison of litter mass loss at 336 sites (ranging from -9 to +26 °C MAT and from 60 to 3113 mm MAP) across different ecosystems. In this study we tested the effect of climate (temperature and moisture), litter type and land-use on early stage decomposition (3 months) across nine biomes. We show that litter quality was the predominant controlling factor in early stage litter decomposition, which explained about 65% of the variability in litter decomposition at a global scale. The effect of climate, on the other hand, was not litter specific and explained <0.5% of the variation for Green tea and 5% for Rooibos tea, and was of significance only under unfavorable decomposition conditions (i.e. xeric versus mesic environments). When the data were aggregated at the biome scale, climate played a significant role on decomposition of both litter types (explaining 64% of the variation for Green tea and 72% for Rooibos tea). No significant effect of land-use on early stage litter decomposition was noted within the temperate biome. Our results indicate that multiple drivers are affecting early stage litter mass loss with litter quality being dominant. In order to be able to quantify the relative importance of the different drivers over time, long-term studies combined with experimental trials are needed.

Summary 1.Understanding the relationship between species diversity and productivity is central to linking compositional and functional aspects of terrestrial ecosystems, and little is known about such issues in boreal forests. We used structural equation modelling (SEM) to test several hypotheses about direct and indirect influences of productivity, its correlate basal area, and resources on understorey vascular plant diversity on 2025 plots in 81 southern boreal forests in Minnesota, USA. 2. We first examined the hypothesis that increasing basal area reduces plot-scale species richness due to competitive exclusion from the most limiting resource, light. As expected, light pre-emption increased with total basal area, which directly reduced understorey species richness. However, complex relations between basal area, dominant understorey species, and resource supply to the understorey can also influence understorey communities. Hence, we addressed whether plots with low light availability in the understorey were associated with low abundance of dominant understorey species and alleviation of competitive exclusion of other understorey species. SEM results showed that low light decreased total understorey cover, alleviating resource competition from this stratum and thus increasing understorey species richness. Furthermore, the cover of four dominant understorey species was positively correlated with light availability and negatively correlated with plotscale species richness. 3. Aggregating data for the 25 plots at each stand, SEM showed that stand-scale species richness was positively influenced by light heterogeneity, which in turn increased with annual above-ground productivity. 4. Species richness was positively influenced by litter %N, considered an index of nitrogen availability at the plot and stand scale. 5. Synthesis. These results suggest that understorey species richness in boreal forests is regulated by productivity, but is primarily mediated by the indirect effects of productivity of the dominant producers on resource availability and heterogeneity.