Summary1 Ecological and agronomic research suggests that increased crop diversity in speciespoor intensive systems may improve their provision of ecosystem services. Such general predictions can have critical importance for worldwide food production and agricultural practice but are largely untested at higher levels of diversity. 2 We propose new methodology for the design and analysis of experiments to quantify diversity-function relationships. Our methodology can quantify the relative strength of inter-specific interactions that contribute to a functional response, and can disentangle the separate contributions of species richness and relative abundance. 3 Applying our methodology to data from a common experiment at 28 European sites, we show that the above-ground biomass of four-species mixtures (two legumes and two grasses) in intensive grassland systems was consistently greater than that expected from monoculture performance, even at high productivity levels. The magnitude of this effect generally resulted in transgressive overyielding. 4 A combined analysis of first-year results across sites showed that the additional performance of mixtures was driven by the number and strength of pairwise inter-specific interactions and the evenness of the community. In general, all pairwise interactions contributed equally to the additional performance of mixtures; the grass-grass and legume-legume interactions were as strong as those between grasses and legumes. 5 The combined analysis across geographical and temporal scales in our study provides a generality of interpretation of our results that would not have been possible from individual site analyses or experimentation at a single site. 6 Our four-species agricultural grassland communities have proved a simple yet relevant model system for experimentation and development of methodology in diversity-function research. Our study establishes that principles derived from biodiversity research in extensive, semi-natural grassland systems are applicable in intensively managed grasslands with agricultural plant species.

Summary 1.A coordinated continental-scale field experiment across 31 sites was used to compare the biomass yield of monocultures and four species mixtures associated with intensively managed agricultural grassland systems. To increase complementarity in resource use, each of the four species in the experimental design represented a distinct functional type derived from two levels of each of two functional traits, nitrogen acquisition (N 2 -fixing legume or nonfixing grass) crossed with temporal development (fast-establishing or temporally persistent). Relative abundances of the four functional types in mixtures were systematically varied at sowing to vary the evenness of the same four species in mixture communities at each site and sown at two levels of seed density. 2. Across multiple years, the total yield (including weed biomass) of the mixtures exceeded that of the average monoculture in >97% of comparisons. It also exceeded that of the best monoculture (transgressive overyielding) in about 60% of sites, with a mean yield ratio of mixture to best-performing monoculture of 1Á07 across all sites. Analyses based on yield of sown species only (excluding weed biomass) demonstrated considerably greater transgressive overyielding (significant at about 70% of sites, ratio of mixture to best-performing monoculture = 1Á18).3. Mixtures maintained a resistance to weed invasion over at least 3 years. In mixtures, median values indicate <4% of weed biomass in total yield, whereas the median percentage of weeds in monocultures increased from 15% in year 1 to 32% in year 3. 4. Within each year, there was a highly significant relationship (P < 0Á0001) between sward evenness and the diversity effect (excess of mixture performance over that predicted from the monoculture performances of component species). At lower evenness values, increases in community evenness resulted in an increased diversity effect, but the diversity effect was not significantly different from the maximum diversity effect across a wide range of higher evenness values. The latter indicates the robustness of the diversity effect to changes in species' relative abundances. 5. Across sites with three complete years of data (24 of the 31 sites), the effect of interactions between the fast-establishing and temporal persistent trait levels of temporal development was highly significant and comparable in magnitude to effects of interactions between N 2 -fixing and nonfixing trait levels of nitrogen acquisition. 6. Synthesis and applications. The design of grassland mixtures is relevant to farm-level strategies to achieve sustainable intensification. Experimental evidence indicated significant yield benefits of four species agronomic mixtures which yielded more than the highest-yielding monoculture at most sites. The results are relevant for agricultural practice and show how grassland mixtures can be designed to improve resource complementarity, increase yields and reduce weed invasion. The yield benefits were robust to considerable changes in the relative propo...

Increased biodiversity may improve ecosystem services, including herbage yield. A mixture experiment was carried out at five sites in Northern Europe and one in Canada to investigate whether mixtures of grasses and legumes would give higher herbage yield than monocultures. Resistance of the mixtures to weed invasion and nutritive value of the herbage were also investigated. The experimental layout followed a simplex design, where four species differing in specific functional traits, timothy (Phleum pratense L.), smooth meadow grass (Poa pratensis L.), red clover (Trifolium pratense L.) and white clover (Trifolium repens L.), were grown in monocultures and eleven different mixtures with systematically varying proportions of the four species. Positive diversity effects (DE) were observed, leading to greater herbage dry‐matter (DM) yield in mixtures than expected from species sown in monocultures. For centroid mixtures, the DE generated on average an additional 32, 25 and 21% of the DM yield than would be expected from the monocultures in the first, second and third year respectively. On average, the mixtures were 9, 15 and 7% more productive than the most productive monoculture (transgressive overyielding) in the first, second and third year respectively. These benefits persisted over the three harvest years of the experiment and were consistent among most sites. This positive effect was not accompanied by a reduction in herbage digestibility and crude protein concentration that is usually observed with increased DM yield. Mixtures also reduced the invasion of weeds to <5% of herbage yield compared to monocultures (10–60% of herbage yield).

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Climate change and its effects on grassland productivity vary across Europe. The Mediterranean and Nordic regions represent the opposite ends of a gradient of changes in temperature and precipitation patterns, with increasingly warmer and wetter winters in the north and increasingly warmer and drier summers in the south. Warming and elevated concentration of atmospheric CO2 may boost forage production in the Nordic region. Production in many Mediterranean areas is likely to become even more challenged by drought in the future, but elevated CO2 can to some extent alleviate drought limitation on photosynthesis and growth. In both regions, climate change will affect forage quality and lead to modifications of the annual productivity cycles, with an extended growing season in the Nordic region and a shift towards winter in the Mediterranean region. This will require adaptations in defoliation and fertilization strategies. The identity of species and mixtures with optimal performance is likelyto shift somewhat inresponse to altered climate and management systems. Itis argued that breeding of grassland species should aimto (i) improve plantstrategies to cope with relevant abiotic stresses and (ii) optimize growth and phenology to new seasonal variation, and that plant diversity at all levels is a good adaptation strategy