In experimental plant communities, relationships between biodiversity and ecosystem functioning have been found to strengthen over time, a fact often attributed to increased resource complementarity between species in mixtures and negative plant-soil feedbacks in monocultures. Here we show that selection for niche differentiation between species can drive this increasing biodiversity effect. Growing 12 grassland species in test monocultures and mixtures, we found character displacement between species and increased biodiversity effects when plants had been selected over 8 years in species mixtures rather than in monocultures. When grown in mixtures, relative differences in height and specific leaf area between plant species selected in mixtures (mixture types) were greater than between species selected in monocultures (monoculture types). Furthermore, net biodiversity and complementarity effects were greater in mixtures of mixture types than in mixtures of monoculture types. Our study demonstrates a novel mechanism for the increase in biodiversity effects: selection for increased niche differentiation through character displacement. Selection in diverse mixtures may therefore increase species coexistence and ecosystem functioning in natural communities and may also allow increased mixture yields in agriculture or forestry. However, loss of biodiversity and prolonged selection of crops in monoculture may compromise this potential for selection in the longer term.

Species extinctions from local communities negatively affect ecosystem functioning. Ecological mechanisms underlying these impacts are well studied, but the role of evolutionary processes is rarely assessed. Using a long-term field experiment, we tested whether natural selection in plant communities increased biodiversity effects on productivity. We re-assembled communities with 8-year co-selection history adjacent to communities with identical species composition but no history of co-selection ('na€ ıve communities'). Monocultures, and in particular mixtures of two to four co-selected species, were more productive than their corresponding na€ ıve communities over 4 years in soils with or without co-selected microbial communities. At the highest diversity level of eight plant species, no such differences were observed. Our findings suggest that plant community evolution can lead to rapid increases in ecosystem functioning at low diversity but may take longer at high diversity. This effect was not modified by treatments simulating co-evolutionary processes between plants and soil organisms.

Diverse experimental plant communities are more productive than monocultures. The increase of this biodiversity effect over time has been attributed to evolutionary selection for complementarity in mixtures. Here we show that evolutionary selection for enhanced net facilitative plant interactions occurred only in mixtures, while evolutionary selection for reduced net competition occurred in mixtures with mixture coexistence history and monocultures with monoculture coexistence history. Widespread declines in natural and agricultural biodiversity could therefore compromise potential evolution of facilitative interactions, that is, cornerstone processes in nature conservation and the development of sustainable agriculture.

In grassland biodiversity experiments the positive biodiversity-ecosystem functioning relationship generally increases over time. However, we know little about the underlying short-term evolutionary processes. Using five plant species selected for twelve years in a biodiversity experiment in mixture or monoculture and plants without such a selection history, we assessed whether differential selection altered productivity, biodiversity effects, and functional trait differences within newly assembled monocultures and 2-species mixtures. Plants without past community selection history produced the lowest assemblage biomass and showed the weakest biodiversity effects. In newly assembled mixtures, plants with a selection history in mixtures produced more biomass than plants with a monoculture selection history. Biodiversity effects were generally positive and differed significantly between selection histories. However, contrary to our expectations, biodiversity effects were not stronger for mixture-type plants. Biodiversity effects were influenced by both trait differences between plants and community-weighted means, but these relationships were mostly independent of selection history. Our findings suggest that twelve years of selection history in monocultures or species mixtures differentiated plants of each species into monoculture-and mixture-types. Such rapid evolution of different community-types within grassland species and its effect on ecosystem services and functioning are likely to be important for species conservation practice..

Plant-plant and plant-soil interactions can help maintain plant diversity and ecosystem functions. Changes in these interactions may underlie experimentally-observed increases in biodiversity effects over time via the selection of genotypes adapted to low or high plant diversity. However, little is known about such community-history effects and particularly the role of plant-soil interactions in this process. Soil-legacy effects may occur if co-evolved interactions with soil communities either positively or negatively modify plant biodiversity effects. We tested how plant selection and soil legacy influence biodiversity effects on productivity, and whether such effects increase the resistance of the communities to invasion by weeds. We used two plant selection treatments: parental plants growing in monoculture or in mixture over 8 years in a grassland biodiversity experiment in the field, which we term monoculture types and mixture types. The two soil-legacy treatments used in this study were neutral soil inoculated with live or sterilized soil inocula collected from the same plots in the biodiversity experiment. For each of the four factorial combinations, seedlings of eight species were grown in monocultures or 4-species mixtures in pots in an experimental garden over fifteen weeks. Soil legacy (live inoculum) strongly increased biodiversity complementarity effects for communities of mixture types, and to a significantly weaker extent for communities of monoculture types. This may be attributed to negative plant-soil feedbacks suffered by mixture types in monocultures whereas monoculture types had positive plant-soil feedbacks in both monocultures and mixtures. Monocultures of mixture types were most strongly invaded by weeds, presumably due to increased pathogen susceptibility of mixture types and thereby reduced biomass and altered plant-soil interactions. These results show that biodiversity effects in experimental grassland communities can be modified by the evolution of positive vs. negative plant-soil feedbacks of plant monoculture vs. mixture types.