Trait-dependent modification of facilitation on cobble beaches.

Fundamental gaps remain in our knowledge of how positive species interactions, such as facilitation and mutualism, structure and maintain populations and communities. Foundation species create extensive biogenic habitats, but we know little of how their traits, such as density, age, and patch size, modify their ability to facilitate other species. We tested the role of facilitator traits in cobble beach plant communities in New England, USA. In this system, intertidal beds of the cordgrass Spartina alterniflora facilitate populations of halophytic forbs at higher shore elevations by buffering wave action, stabilizing cobbles, and limiting physical disturbance. Using descriptive and experimental techniques, we tested the hypotheses that (1) the density and height of cordgrass shoots modify the strength of the cordgrass-forb facilitation, and (2) cordgrass peat alone contributes to the facilitation of forbs. Increased shoot density, as well as the combination of cordgrass peat and shoots, positively affected two life history stages (seedling and adult) of the abundant forb Suaeda linearis, demonstrating that cordgrass traits modify the strength of facilitation in this system. Since the expression of traits varies within and among patches of any given foundation species, traits can and should be used to predict the strength of facilitation, to guide the development of conservation strategies, and to develop more accurate models of species interactions.