Global change, especially land‐use intensification, affects human well‐being by impacting the delivery of multiple ecosystem services (multifunctionality). However, whether biodiversity loss is a major component of global change effects on multifunctionality in real‐world ecosystems, as in experimental ones, remains unclear. Therefore, we assessed biodiversity, functional composition and 14 ecosystem services on 150 agricultural grasslands differing in land‐use intensity. We also introduce five multifunctionality measures in which ecosystem services were weighted according to realistic land‐use objectives. We found that indirect land‐use effects, i.e. those mediated by biodiversity loss and by changes to functional composition, were as strong as direct effects on average. Their strength varied with land‐use objectives and regional context. Biodiversity loss explained indirect effects in a region of intermediate productivity and was most damaging when land‐use objectives favoured supporting and cultural services. In contrast, functional composition shifts, towards fast‐growing plant species, strongly increased provisioning services in more inherently unproductive grasslands.

Many experiments have shown that loss of biodiversity reduces the capacity of ecosystems to provide the multiple services on which humans depend. However, experiments necessarily simplify the complexity of natural ecosystems and will normally control for other important drivers of ecosystem functioning, such as the environment or land use. In addition, existing studies typically focus on the diversity of single trophic groups, neglecting the fact that biodiversity loss occurs across many taxa and that the functional effects of any trophic group may depend on the abundance and diversity of others. Here we report analysis of the relationships between the species richness and abundance of nine trophic groups, including 4,600 above- and below-ground taxa, and 14 ecosystem services and functions and with their simultaneous provision (or multifunctionality) in 150 grasslands. We show that high species richness in multiple trophic groups (multitrophic richness) had stronger positive effects on ecosystem services than richness in any individual trophic group; this includes plant species richness, the most widely used measure of biodiversity. On average, three trophic groups influenced each ecosystem service, with each trophic group influencing at least one service. Multitrophic richness was particularly beneficial for 'regulating' and 'cultural' services, and for multifunctionality, whereas a change in the total abundance of species or biomass in multiple trophic groups (the multitrophic abundance) positively affected supporting services. Multitrophic richness and abundance drove ecosystem functioning as strongly as abiotic conditions and land-use intensity, extending previous experimental results to real-world ecosystems. Primary producers, herbivorous insects and microbial decomposers seem to be particularly important drivers of ecosystem functioning, as shown by the strong and frequent positive associations of their richness or abundance with multiple ecosystem services. Our results show that multitrophic richness and abundance support ecosystem functioning, and demonstrate that a focus on single groups has led to researchers to greatly underestimate the functional importance of biodiversity.

Summary1. Biotic interactions do not occur in isolation but are imbedded in a network of species interactions. Network analysis facilitates the compilation and understanding of the complexity found in natural ecosystems and is a powerful tool to reveal information on the degree of specialization of the interacting partners and their niches. The indices measuring these properties are based on qualitative or quantitative observations of interactions between partners from different trophic levels, which informs about the structure of network patterns, but not about the underlying mechanisms. Functional traits may control the interaction strength between partners and also the (micro-) structure of networks. Here, we ask whether flower visitors specialize on certain plant traits and how this trait specialization contributes to niche partitioning and interaction partner diversity. 2. We introduce two novel statistical approaches suited to evaluate the dimension of the realized niche and to analyse which traits determine niches. As basis for our analysis, we measured 10 quantitative flower traits and evaluated whether 31 arthropod taxa i visited flowers displaying only subsets of the available trait characteristics, indicating a specialization on these traits by narrow trait-widths 〈S i 〉. The product of 10 trait-and species-specific trait-widths 〈S i 〉 was defined as trait-volume V i (expansion of a n-dimensional hypervolume) occupied by each taxon i. These indices are applicable beyond flower-visitor interactions to quantify realized niches based on various biotic and abiotic factors. 3. Each flower visitor species showed some degree of specialization to a unique set of flower traits (realized niche). Overall, our data suggested a hierarchical sequence of flower traits influencing the flower visitors' behaviour and thus network structure: flowering phenology was found to have the strongest effect, followed by flower height, nectar-tube depth and floral reflectance. Less important were pollen-mass/flower, sugar/flower, anther position, phylogeny, display size and abundance. 4. The species-specific specialization on traits suggests that plant communities with more diverse floral niches may sustain a larger number of flower visitors with non-redundant fundamental niches. Our study and statistical approach provide a basis for a better understanding of how plant traits shape interactions between flowers and their visitors and thus network structure.

One contribution of 17 to a theme issue 'Biodiversity and ecosystem functioning in dynamic landscapes'. Species diversity promotes the delivery of multiple ecosystem functions (multifunctionality). However, the relative functional importance of rare and common species in driving the biodiversity-multifunctionality relationship remains unknown. We studied the relationship between the diversity of rare and common species (according to their local abundances and across nine different trophic groups), and multifunctionality indices derived from 14 ecosystem functions on 150 grasslands across a landuse intensity (LUI) gradient. The diversity of above-and below-ground rare species had opposite effects, with rare above-ground species being associated with high levels of multifunctionality, probably because their effects on different functions did not trade off against each other. Conversely, common species were only related to average, not high, levels of multifunctionality, and their functional effects declined with LUI. Apart from the communitylevel effects of diversity, we found significant positive associations between the abundance of individual species and multifunctionality in 6% of the species tested. Speciesspecific functional effects were best predicted by their response to LUI: species that declined in abundance with land use intensification were those associated with higher levels of multifunctionality. Our results highlight the importance of rare species for ecosystem multifunctionality and help guiding future conservation priorities.

1. Rapid growth of the world's human population has increased pressure on landscapes to deliver high levels of multiple ecosystem services, including food and fibre production, carbon storage, biodiversity conservation, and recreation.However, we currently lack general principles describing how to achieve this landscape multifunctionality.2. We combine theoretical simulations and empirical data on 14 ecosystem services measured across 150 grasslands in three German regions. In doing so, we investigate the circumstances under which spatial heterogeneity in a driver of ecosystem functioning (an "ecosystem-driver," e.g., the presence of keystone species, landuse intensification, or habitat types) increases landscape-level ecosystem multifunctionality.