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).

Grassland diversity can support sustainable intensification of grassland production through increased yields, reduced inputs and limited weed invasion. We report the effects of diversity on weed suppression from 3 years of a 31‐site continental‐scale field experiment.At each site, 15 grassland communities comprising four monocultures and 11 four‐species mixtures based on a wide range of species' proportions were sown at two densities and managed by cutting. Forage species were selected according to two crossed functional traits, “method of nitrogen acquisition” and “pattern of temporal development”.Across sites, years and sown densities, annual weed biomass in mixtures and monocultures was 0.5 and 2.0 t  DM ha−1 (7% and 33% of total biomass respectively). Over 95% of mixtures had weed biomass lower than the average of monocultures, and in two‐thirds of cases, lower than in the most suppressive monoculture (transgressive suppression). Suppression was significantly transgressive for 58% of site‐years. Transgressive suppression by mixtures was maintained across years, independent of site productivity.Based on models, average weed biomass in mixture over the whole experiment was 52% less (95% confidence interval: 30%–75%) than in the most suppressive monoculture. Transgressive suppression of weed biomass was significant at each year across all mixtures and for each mixture.Weed biomass was consistently low across all mixtures and years and was in some cases significantly but not largely different from that in the equiproportional mixture. The average variability (standard deviation) of annual weed biomass within a site was much lower for mixtures (0.42) than for monocultures (1.77). Synthesis and applications. Weed invasion can be diminished through a combination of forage species selected for complementarity and persistence traits in systems designed to reduce reliance on fertiliser nitrogen. In this study, effects of diversity on weed suppression were consistently strong across mixtures varying widely in species' proportions and over time. The level of weed biomass did not vary greatly across mixtures varying widely in proportions of sown species. These diversity benefits in intensively managed grasslands are relevant for the sustainable intensification of agriculture and, importantly, are achievable through practical farm‐scale actions.

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