The decomposition of dead organic matter is a major determinant of carbon and nutrient cycling in ecosystems, and of carbon fluxes between the biosphere and the atmosphere. Decomposition is driven by a vast diversity of organisms that are structured in complex food webs. Identifying the mechanisms underlying the effects of biodiversity on decomposition is critical given the rapid loss of species worldwide and the effects of this loss on human well-being. Yet despite comprehensive syntheses of studies on how biodiversity affects litter decomposition, key questions remain, including when, where and how biodiversity has a role and whether general patterns and mechanisms occur across ecosystems and different functional types of organism. Here, in field experiments across five terrestrial and aquatic locations, ranging from the subarctic to the tropics, we show that reducing the functional diversity of decomposer organisms and plant litter types slowed the cycling of litter carbon and nitrogen. Moreover, we found evidence of nitrogen transfer from the litter of nitrogen-fixing plants to that of rapidly decomposing plants, but not between other plant functional types, highlighting that specific interactions in litter mixtures control carbon and nitrogen cycling during decomposition. The emergence of this general mechanism and the coherence of patterns across contrasting terrestrial and aquatic ecosystems suggest that biodiversity loss has consistent consequences for litter decomposition and the cycling of major elements on broad spatial scales. Main textBiological diversity that directly influences litter decomposition exists at multiple trophic levels 4 . This diversity includes plants producing litter mixtures of varying quality, microbial decomposers, and invertebrate consumers of varying body size, which selectively exploit the heterogeneous resources provided by litter mixtures 4,13 . Efforts to derive generalities about biodiversity effects on litter decomposition have been elusive, since both pioneering work 14 and recent syntheses have highlighted contrasting effects of litter species richness on 3 decomposition [4][5][6]15,16 . In part, this variation appears to be due to site-specific conditions, including contrasts between aquatic and terrestrial ecosystems as well as geographic settings.Further differences may arise from variation in experimental protocols, selected plant species, and the types of decomposers included in a given experiment. Such methodological discrepancies have complicated syntheses across studies, hindering the emergence of common patterns and mechanisms.Here we report on the results from the first concerted biodiversity experiments on decomposition by manipulating diversity across trophic levels and distinct biomes in both forest floor and stream habitats (Extended Data Table 1). We hypothesised that functional diversity of decomposers (variation in body size) and leaf litter (variation in litter quality) promote C and N cycling across contrasting locations (subarctic to tr...

Running waters are perhaps the most impacted ecosystem on the planet as they have been the focus for human settlement and are heavily exploited for water supplies, irrigation, electricity generation, and waste disposal. Lotic systems also have an intimate contact with their catchments and so land-use alterations affect them directly. Here long-term trends in the factors that currently impact running waters are reviewed with the aim of predicting what the main threats to rivers will be in the year 2025. The main ultimate factors forcing change in running waters (ecosystem destruction, physical habitat and water chemistry alteration, and the direct addition or removal of species) stem from proximate influences from urbanization, industry, land-use change and water-course alterations. Any one river is likely to be subjected to several types of impact, and the management of impacts on lotic systems is complicated by numerous links between different forms of anthropogenic effect. Long-term trends for different impacts vary. Concentrations of chemical pollutants such as toxins and nutrients have increased in rivers in developed countries over the past century, with recent reductions for some pollutants (e.g. metals, organic toxicants, acidification), and continued increases in others (e.g. nutrients); there are no long-term chemical data for developing countries. Dam construction increased rapidly during the twentieth century, peaking in the 1970s, and the number of reservoirs has stabilized since this time, whereas the transfer of exotic species between lotic systems continues to increase. Hence, there have been some success stories in the attempts to reduce the impacts from anthropogenic impacts in developed nations. Improvements in the pH status of running waters should continue with lower sulphurous emissions, although emissions of nitrous oxides are set to continue under current legislation and will continue to contribute to acidification and nutrient loadings. Climate change also will impact running waters through alterations in hydrology and thermal regimes, although precise predictions are problematic; effects are likely to vary between regions and to operate alongside rather than override those from other impacts. Effects from climate change may be more extreme over longer time scales (>50 years). The overriding pressure on running water ecosystems up to 2025 will stem from the predicted increase in the human population, with concomitant increases in urban development, industry, agricultural activities and water abstraction, diversion and damming. Future degradation could be substantial and rapid (c. 10 years) and will be concentrated in those areas of the world where resources for conservation are most limited and knowledge of lotic ecosystems most incomplete; damage will centre on lowland rivers, which are also relatively poorly studied. Changes in management practices and public awareness do appear to be benefiting running water ecosystems in developed countries, and could underpin conservation strategies in developing countries if they were implemented in a relevant way.

Excessive nutrient loading is a major threat to aquatic ecosystems worldwide that leads to profound changes in aquatic biodiversity and biogeochemical processes. Systematic quantitative assessment of functional ecosystem measures for river networks is, however, lacking, especially at continental scales. Here, we narrow this gap by means of a pan-European field experiment on a fundamental ecosystem process--leaf-litter breakdown--in 100 streams across a greater than 1000-fold nutrient gradient. Dramatically slowed breakdown at both extremes of the gradient indicated strong nutrient limitation in unaffected systems, potential for strong stimulation in moderately altered systems, and inhibition in highly polluted streams. This large-scale response pattern emphasizes the need to complement established structural approaches (such as water chemistry, hydrogeomorphology, and biological diversity metrics) with functional measures (such as litter-breakdown rate, whole-system metabolism, and nutrient spiraling) for assessing ecosystem health.

Recent experiments, mainly in terrestrial environments, have provided evidence of the functional importance of biodiversity to ecosystem processes and properties. Compared to terrestrial systems, aquatic ecosystems are characterised by greater propagule and material exchange, often steeper physical and chemical gradients, more rapid biological processes and, in marine systems, higher metazoan phylogenetic diversity. These characteristics limit the potential to transfer conclusions derived from terrestrial experiments to aquatic ecosystems whilst at the same time provide opportunities for testing the general validity of hypotheses about effects of biodiversity on ecosystem functioning. Here, we focus on a number of unique features of aquatic experimental systems, propose an expansion to the scope of diversity facets to be considered when assessing the functional consequences of changes in biodiversity and outline a hierarchical classification scheme of ecosystem functions and their corresponding response variables. We then briefly highlight some recent controversial and newly emerging issues relating to biodiversity‐ecosystem functioning relationships. Based on lessons learnt from previous experimental and theoretical work, we finally present four novel experimental designs to address largely unresolved questions about biodiversity‐ecosystem functioning relationships. These include (1) investigating the effects of non‐random species loss through the manipulation of the order and magnitude of such loss using dilution experiments; (2) combining factorial manipulation of diversity in interconnected habitat patches to test the additivity of ecosystem functioning between habitats; (3) disentangling the impact of local processes from the effect of ecosystem openness via factorial manipulation of the rate of recruitment and biodiversity within patches and within an available propagule pool; and (4) addressing how non‐random species extinction following sequential exposure to different stressors may affect ecosystem functioning. Implementing these kinds of experimental designs in a variety of systems will, we believe, shift the focus of investigations from a species richness‐centred approach to a broader consideration of the multifarious aspects of biodiversity that may well be critical to understanding effects of biodiversity changes on overall ecosystem functioning and to identifying some of the potential underlying mechanisms involved.

Restoration schemes often rely on the assumption that enhancing habitat complexity through addition of in‐stream structures such as boulders and woody debris leads to increased biodiversity, but evidence for this assumption is scarce. We compared structural heterogeneity and fish and invertebrate diversity at restored, unrestored, and reference sites on tributaries of the Ume River, northern Sweden, where several kilometers of streams have been restored from channelization through placement of boulders into the channel. Structural heterogeneity at the study sites was assessed using a contour tracer at two spatial resolutions likely to be affected by restoration. These are the patch scale (0.7 m), reflecting substratum characteristics, and the reach scale (50 m), reflecting general channel topography. Fish and invertebrate samples collected via electroshocking were used to assess taxonomic richness, taxonomic density, evenness, and assemblage composition at the study sites. Measures of structural heterogeneity were substantially higher at restored relative to channelized sites; however, components of fish and invertebrate diversity were similar between these treatments. At restored sites, measures of structural heterogeneity and fish and invertebrate diversity were consistent with, or slightly exceeded reference levels. This implies that local (patch to reach) heterogeneity did not structure fish and invertebrate assemblages in the study streams. Our results suggest that restoration might have little beneficial effect on biodiversity if the restoration schemes (and the original impact under amelioration) do not affect structural heterogeneity relevant to the target organisms.

Human disturbances both decrease the number of species in ecosystems and change their relative abundances. Here we present field evidence demonstrating that shifts in species abundances can have effects on ecosystem functioning that are as great as those from shifts in species richness. We investigated spatial and temporal variability of leaf decomposition rates and community metrics of leaf-eating invertebrates (shredders) in streams. The shredder community composition dramatically influenced the diversityfunction relationship; decomposition was much higher for a given species richness at sites with high species dominance than at sites where dominance was low. Decomposition rates also markedly depended on the identity of the dominant species. Further, dominance effects on decomposition varied seasonally and the number of species required for maintaining decomposition increased with increasing evenness. These findings reveal important but less obvious aspects of the biodiversity-ecosystem functioning relationship.

2000. Ecosystem process rate increases with animal species richness: evidence from leaf-eating, aquatic insects. -Oikos 89: 519 -523.Effects of species number and identity on the breakdown rate of leaf litter were estimated in a laboratory experiment using leaf-eating insects, three species of Plecoptera, as detritivores. We found significant differences between the different species on this process in single-species experiment, but not when animal biomass was accounted for. When species were combined the effect of species identity was strongly reduced and rendered insignificant, whereas the number of species had a significant effect. This shows that rates of ecosystem processes may benefit from species richness even when all species belong to the same guild, which is in contrast to hypotheses predicting redundancy within guilds. Facilitation between species and negative interactions, where intraspecific interactions are greater than interspecific interactions, are two potential mechanisms which could explain increasing decomposition rates with species richness.

1. Riverine systems consist of a mosaic of patches and habitats linked by diverse processes and supporting highly complex communities. Invertebrates show a high taxonomic and functional diversity in riverine systems and are in several ways important components of these systems. Their distribution patterns, movements and effects on ecological flows, testify to their importance in various landscape ecological processes. This paper reviews the invertebrate literature with respect to patterns and processes in the riverine landscape. 2. The distribution of invertebrates in riverine habitats is governed by a number of factors that typically act at different scales. Hence, the local community structure can be seen as the result of a continuous sorting process through environmental filters ranging from regional or catchment‐wide processes, involving speciation, geological history and climate, to the small‐scale characteristics of individual patches, such as local predation risk, substratum porosity and current velocity. 3. Dispersal is an important process driving invertebrate distribution, linking different ecological systems across boundaries. Dispersal occurs within the aquatic habitat as well as into the terrestrial surrounding, and also over land to other waterbodies. New genetic techniques have contributed significantly to the understanding of aquatic invertebrate dispersal and revealed the importance of factors such as physical barriers, synchrony of emergence and taxonomic affiliation. 4. Invertebrates affect the cycling of nutrients and carbon by being a crucial intermediate link between primary producers, detritus pools or primary consumers, and predators higher up in the trophic hierarchy. Suspension feeders increase the retention of carbon. The subsidies of aquatic invertebrates to the terrestrial ecosystem have been shown to be important, as are reciprocal processes such as the supply of terrestrial invertebrates that fall into the water. 5. Future studies are needed both to advance theoretical aspects of landscape ecology pertaining to the invertebrates in riverine systems and to intensify the experimental testing of hypotheses, for example with respect to the scaling of processes and to linkages between the terrestrial and aquatic systems. Another promising avenue is to take advantage of naturally steep environmental gradients, and of systems disturbed by humans, such as regulated rivers. By comparison with unimpaired reference sites, the mechanisms involved might be identified. The use of `natural' experiments, especially where environmental gradients are steep, is another technique with great potential.