Designing resilient and sustainable grasslands for a drier future: Adaptive strategies, functional traits and biotic interactions

• Forage mixtures should combine plant strategies for greater drought resilience.
• A conceptual framework to analyze adaptive responses of forage species under drought.
• A trade-off between resistance to moderate drought and survival of intense drought.
• Concepts of community ecology are useful to the design of multi-specific grasslands.
• Defining rules to associate genotypes should enhance functional complementarity.

In many regions of the world, such as Southern Europe and most Mediterranean areas, the frequency and magnitude of droughts and heat waves are expected to increase under global warming and will challenge the sustainability of both native and sown grasslands. To analyze the adaptive strategies of species, genotypes and cultivars, we aim both (i) to understand the composition and functioning of natural grasslands and (ii) to propose ideotypes of cultivars and optimal composition for mixtures of species/genotypes under water deficit and high temperatures. This review presents a conceptual framework to analyze adaptive responses of perennial herbaceous species, starting from resistance to moderate drought with growth maintenance (dehydration avoidance and tolerance of lamina) to growth cessation and survival of plants under severe stress (dehydration avoidance and tolerance of meristems). The most discriminating functional traits vary according to these contrasting strategies because of a trade-off between resistance to moderate moisture deficit and survival of intense drought. Consequently it is crucial to measure the traits of interest in the right organs and as a function of soil water use, in order to avoid misleading interpretations of plant responses. Furthermore, collaboration between ecologists, eco-physiologists, and agronomists is required to study the combination of plant strategies in natural grasslands as only this will provide the necessary rules for species and cultivars or ecotypes assemblage. This ‘agro-ecological’ approach aims to identify and enhance functional complementarity and limit competition within the multi-specific or multi-genotypic material associated in mixtures since using plant biodiversity should contribute to improving grassland resistance and resilience.