Why are many anthropogenic agroecosystems particularly species-rich?

Species-rich meadow and pasture habitats are recognised by the European Union Habitats Directive as targets for biodiversity conservation. High species richness is hypothesised to be associated with diversity in plant functional traits and life-history strategies, which are potentially restricted in situations of extremely high and low biomass production. However, variability in functional traits has yet to be investigated across a broad biomass range in nature. We measured variability in a range of functional traits and Grime's competitor, stress-tolerator, ruderal (CSR) strategies for species comprising lowland meadows, subalpine pastures, abandoned grassland and field margins at sites in northern Italy, alongside peak above-ground biomass. The factor most highly and positively correlated with species richness was strategy richness (the number of CSR strategies; Pearson's r = 0.864, P < 0.0001, n = 39), followed by variance in traits involved in leaf resource economics and the timing of flowering. Species richness, trait variance and strategy richness were greatest at intermediate biomass. Thus whilst extremes of biomass production were associated with relatively few taxa exhibiting similar trait values and specialised strategies, greater species richness was apparent in meadows and pastures in which species exhibited divergence in resource economics trait values, reproductive timing and strategy richness.