Optimality as a concept to understand and model vegetation at different scales

Experimental studies with grassland species found a positive relationship between species richness and community biomass production, however the response of individual species was highly variable. The mechanisms behind these patterns are poorly understood. Here we studied aboveground biomass production and plasticity of growth characteristics of four legumes with similar morphology (Lotus corniculatus, Medicago lupulina, Onobrychis viciifolia, Trifolium hybridum) in experimental grasslands varying in species richness (1, 2, 4, 8, 16 and 60) and composition. We identified O. viciifolia and T. hybridum as species that reached higher biomass production in mixtures than expected from monoculture yields, while L. corniculatus and M. lupulina mostly had lower yields than expected. Variation of morphological traits across the species-richness gradient was lowest in the highly competitive O. viciifolia, but increased in the smaller species. The tall-growing O. viciifolia achieved higher biomass production by both, a higher number of plant individuals and an increase in mean mass per individual. Mean shoot number per individual remained constant, but individuals produced heavier shoots. The medium-sized T. hybridum also increased the number of plant individuals, but mean mass per individual did not respond to community species richness. The average mass per shoot was increased in mixtures, but the species developed less shoots per individual. Shoot length and stem weight ratio of T. hybridum increased with community species richness. Morphological changes in the less successful L. corniculatus and M. lupulina with a smaller growth stature were similarly directed as those of T. hybridum. The observed morphological changes are known as typical shade-avoidance mechanisms in dense vegetation. Our study shows that stress responses to changes in resource availability may be a mechanism to enforce higher aboveground biomass production of individual species in mixtures, but it depends on species identity whether trait plasticity is large enough to exceed stress-induced growth limitations