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A recent and prominent claim for the value of biodiversity is its importance for sustaining multiple ecosystem functions. The general idea is intuitively appealing: since all species are to some extent unique, each will be important for a different set of functions. Therefore, as more functions are considered, a greater diversity of species is necessary to sustain all functions simultaneously. However, we show here that the relationship between biodiversity and ecosystem functioning does not change with the number of functions considered. Biodiversity affects the level of multifunctionality via non-additive effects on individual functions, and the effect on multifunctionality equals the average effect on single functions. These insights run counter to messages in the literature. In the light of our simulations, we present limitations and pitfalls with current methods used to study biodiversity–multifunctionality, which together provide a perspective for future studies. Biodiversity and ecosystem functioning relationships remain constant no matter how many functions are considered. Biodiversity affects the level of multifunctionality and the effect on multifunctionality equals the average effect on single functions.

Biodiversity is proposed to be important for the rate of ecosystem functions. Most biodiversity—ecosystem function studies, however, consider only one response variable at a time, and even when multiple variables are examined they are analyzed separately. This means that a very important aspect of biodiversity is overlooked: the possibility for different species to carry out different functions at any one time. We propose a conceptual model to explore the effects of species loss on overall ecosystem functioning, where overall functioning is defined as the joint effect of many ecosystem functions. We show that, due to multifunctional complementarity among species, overall functioning is more susceptible to species loss than are single functions. Modeled relationships between species richness and overall ecosystem functioning using five empirical data sets on monocultures reflected the range of effects of species loss on multiple functions predicted by the model. Furthermore, an exploration of the correlations across functions and the degree of redundancy within functions revealed that multifunctional redundancy was generally lower than single-function redundancy in these empirical data sets. We suggest that by shifting the focus to the variety of functions maintained by a diversity of species, the full importance of biodiversity for the functioning of ecosystems can be uncovered. Our results are thus important for conservation and management of biota and ecosystem services.

The importance of biodiversity for the integrated functioning of ecosystems remains unclear because most evidence comes from analyses of biodiversity’s effect on individual functions. Here we show that the effects of biodiversity on ecosystem function become more important as more functions are considered. We present the first systematic investigation of biodiversity’s effect on ecosystem multifunctionality across multiple taxa, trophic levels and habitats using a comprehensive database of 94 manipulations of species richness. We show that species-rich communities maintained multiple functions at higher levels than depauperate ones. These effects were stronger for herbivore biodiversity than for plant biodiversity, and were remarkably consistent across aquatic and terrestrial habitats. Despite observed tradeoffs, the overall effect of biodiversity on multifunctionality grew stronger as more functions were considered. These results indicate that prior research has underestimated the importance of biodiversity for ecosystem functioning by focusing on individual functions and taxonomic groups. The influence of biodiversity on multiple ecosystem processes is not well understood. Analysing 94 biodiversity-ecosystem functioning experiments, Lefcheck et al . find that increased species richness maintains more ecological functions, across multiple taxa, trophic levels and habitats.

Marine ecosystems are experiencing rapid and pervasive changes in biodiversity and species composition. Understanding the ecosystem consequences of these changes is critical to effectively managing these systems. Over the last several years, numerous experimental manipulations of species richness have been performed, yet existing quantitative syntheses have focused on a just a subset of processes measured in experiments and, as such, have not summarized the full data available from marine systems. Here, we present the results of a meta-analysis of 110 marine experiments from 42 studies that manipulated the species richness of organisms across a range of taxa and trophic levels and analysed the consequences for various ecosystem processes (categorised as production, consumption or biogeochemical fluxes). Our results show that, generally, mixtures of species tend to enhance levels of ecosystem function relative to the average component species in monoculture, but have no effect or a negative effect on functioning relative to the ‘highest-performing' species. These results are largely consistent with those from other syntheses, and extend conclusions to ecological functions that are commonly measured in the marine realm (e.g. nutrient release from sediment bioturbation). For experiments that manipulated three or more levels of richness, we attempted to discern the functional form of the biodiversity–ecosystem functioning relationship. We found that, for response variables related to consumption, a powerfunction best described the relationship, which is also consistent with previous findings. However, we identified a linear relationship between richness and production. Combined, our results suggest that changes in the number of species will, on average, tend to alter the functioning of marine ecosystems. We outline several research frontiers that will allow us to more fully understand how, why, and when diversity may drive the functioning of marine ecosystems. Synthesis The oceans host an incredible number and variety of species. However, human activities are driving rapid changes in the marine environment. It is imperative we understand ecosystem consequences of any associated loss of species. We summarized data from 110 experiments that manipulated species diversity and evaluated resulting changes to a range of ecosystem responses. We show that losing species, on average, decreases productivity, growth, and a myriad of other processes related to how marine organisms capture and utilize resources. Finally, we suggest that the loss of species may have stronger consequences for some processes than others.

We examine how levels of multiple ecosystem services (ESs) change with succession in forests with different tree species composition. More specifically we ask how ecosystem age interacts with environmental conditions to regulate ES delivery. Using the nationwide Swedish forest inventory, comprising boreal and temperate regions, we investigated how levels of six provisioning, regulating, recreational, and/or cultural forest ESs changed with forest age (10-185 years) in stands of different tree species composition. We also tested whether the number of ESs delivered (i.e. multifunctionality) changed substantially with stand age, using different threshold levels for ES delivery. Accounting for environmental conditions and stand properties, we found that levels of single ESs changed with stand age. Tree biomass production usually peaked in young to medium aged stands. In contrast, production of berries and game, and services related to biodiversity, were typically highest in old stands (120-185 years). Consistent with this strong temporal tradeoff, multifunctionality at lower threshold levels increased with stand age in most monocultures and mixtures, with the highest multifunctionality being reached somewhere between 100 and 185 years, depending on tree species composition. This was not evident for the highest threshold ES level (the top-20%), however. Moreover, multifunctionality usually decreased with warmer climatic conditions, with the exception of spruce-pine-birch mixtures. Taken together, our results show that a reduced forest age, e.g. due to forestry targeting early harvest of stands, most likely would limit the delivery of several ESs valued by society and result in less multifunctional forests. To maintain the capacity of forests to deliver high levels of multiple ESs, the role of stand age and tree species composition should be considered in decisions on how to manage future forests.

Over the past several decades, a rapidly expanding field of research known as biodiversity and ecosystem functioning has begun to quantify how the world's biological diversity can, as an independent variable, control ecological processes that are both essential for, and fundamental to, the functioning of ecosystems. Research in this area has often been justified on grounds that (1) loss of biological diversity ranks among the most pronounced changes to the global environment and that (2) reductions in diversity, and corresponding changes in species composition, could alter important services that ecosystems provide to humanity (e.g., food production, pest/disease control, water purification). Here we review over two decades of experiments that have examined how species richness of primary producers influences the suite of ecological processes that are controlled by plants and algae in terrestrial, marine, and freshwater ecosystems. Using formal meta-analyses, we assess the balance of evidence for eight fundamental questions and corresponding hypotheses about the functional role of producer diversity in ecosystems. These include questions about how primary producer diversity influences the efficiency of resource use and biomass production in ecosystems, how primary producer diversity influences the transfer and recycling of biomass to other trophic groups in a food web, and the number of species and spatial /temporal scales at which diversity effects are most apparent. After summarizing the balance of evidence and stating our own confidence in the conclusions, we outline several new questions that must now be addressed if this field is going to evolve into a predictive science that can help conserve and manage ecological processes in ecosystems.

It is well known that ocean acidification can have profound impacts on marine organisms. However, we know little about the direct and indirect effects of ocean acidification and also how these effects interact with other features of environmental change such as warming and declining consumer pressure. In this study, we tested whether the presence of consumers (invertebrate mesograzers) influenced the interactive effects of ocean acidification and warming on benthic microalgae in a seagrass community mesocosm experiment. Net effects of acidification and warming on benthic microalgal biomass and production, as assessed by analysis of variance, were relatively weak regardless of grazer presence. However, partitioning these net effects into direct and indirect effects using structural equation modeling revealed several strong relationships. In the absence of grazers, benthic microalgae were negatively and indirectly affected by sediment-associated microalgal grazers and macroalgal shading, but directly and positively affected by acidification and warming. Combining indirect and direct effects yielded no or weak net effects. In the presence of grazers, almost all direct and indirect climate effects were nonsignificant. Our analyses highlight that (i) indirect effects of climate change may be at least as strong as direct effects, (ii) grazers are crucial in mediating these effects, and (iii) effects of ocean acidification may be apparent only through indirect effects and in combination with other variables (e.g., warming). These findings highlight the importance of experimental designs and statistical analyses that allow us to separate and quantify the direct and indirect effects of multiple climate variables on natural communities.

Species diversity affects the functioning of ecosystems, including the efficiency by which communities capture limited resources, produce biomass, recycle and retain biologically essential nutrients. These ecological functions ultimately support the ecosystem services upon which humanity depends. Despite hundreds of experimental tests of the effect of biodiversity on ecosystem function (BEF), it remains unclear whether diversity effects are sufficiently general that we can use a single relationship to quantitatively predict how changes in species richness alter an ecosystem function across trophic levels, ecosystems and ecological conditions. Our objective here is to determine whether a general relationship exists between biodiversity and standing biomass. We used hierarchical mixed effects models, based on a power function between species richness and biomass production (Y = a × Sb), and a database of 374 published experiments to estimate the BEF relationship (the change in biomass with the addition of species), and its associated uncertainty, in the context of environmental factors. We found that the mean relationship (b = 0.26, 95% CI: 0.16, 0.37) characterized the vast majority of observations, was robust to differences in experimental design, and was independent of the range of species richness levels considered. However, the richness–biomass relationship varied by trophic level and among ecosystems; in aquatic systems b was nearly twice as large for consumers (herbivores and detritivores) compared to primary producers; in terrestrial ecosystems, b for detritivores was negative but depended on few studies. We estimated changes in biomass expected for a range of changes in species richness, highlighting that species loss has greater implications than species gains, skewing a distribution of biomass change relative to observed species richness change. When biomass provides a good proxy for processes that underpin ecosystem services, this relationship could be used as a step in modeling the production of ecosystem services and their dependence on biodiversity.

The study of the interrelationship between productivity and biodiversity is a major research field in ecology. Theory predicts that if essential resources are heterogeneously distributed across a metacommunity, single species may dominate productivity in individual metacommunity patches, but a mixture of species will maximize productivity across the whole metacommunity. It also predicts that a balanced supply of resources within local patches should favor species coexistence, whereas resource imbalance would favor the dominance of one species. We performed an experiment with five freshwater algal species to study the effects of total supply of resources, their ratios, and species richness on biovolume production and evenness at the scale of both local patches and metacommunities. Generally, algal biovolume increased, whereas algal resource use efficiency (RUE) and evenness decreased with increasing total supply of resources in mixed communities containing all five species. In contrast to predictions for biovolume production, the species mixtures did not outperform all monocultures at the scale of metacommunities. In other words, we observed no general transgressive overyielding. However, RUE was always higher in mixtures than predicted from monocultures, and analyses indicate that resource partitioning or facilitation in mixtures resulted in higher-than-expected productivity at high resource supply. Contrasting our predictions for the local scale, balanced supply of resources did not generally favor higher local evenness, however lowest evenness was confined to patches with the most imbalanced supply. Thus, our study provides mixed support for recent theoretical advancements to understand biodiversity-productivity relationships.

Species diversity is important for a range of ecosystem processes and properties, including the resistance to single and multiple stressors. It has been suggested that genetic diversity may play a similar role, but empirical evidence is still relatively scarce. Here, we report the results of a microcosm experiment where four strains of the marine diatom Skeletonema marinoi were grown in monoculture and in mixture under a factorial combination of temperature and salinity stress. The strains differed in their susceptibility to the two stressors and no strain was able to survive both stressors simultaneously. Strong competition between the genotypes resulted in the dominance of one strain under both control and salinity stress conditions. The overall productivity of the mixture, however, was not related to the dominance of this strain, but was instead dependent on the treatment; under control conditions we observed a positive effect of genetic richness, whereas a negative effect was observed in the stress treatments. This suggests that interactions among the strains can be both positive and negative, depending on the abiotic environment. Our results provide additional evidence that the biodiversity-ecosystem functioning relationship is also relevant at the level of genetic diversity.