Facilitation between higher plant species in a semiarid environment

A major role of facilitation between higher plant species, particularly in stressful environments, has recently been reported in several papers that suggest that beneficial interactions could be more important in the dynamics of plant communities than has been recognized to date. In a semiarid region in southeastern Spain we determined the effect of a leguminous shrub, Retama sphaerocarpa, on its environment, testing the hypothesis that facilitation by the shrub of one species of its understory, Marrubium vulgare, is reciprocal and that the shrub benefits from sheltering herbs beneath its canopy. Marrubium plants under Retama had greater specific leaf area, leaf mass, shoot mass, leaf area, more flowers, a higher nitrogen (N) concentration in leaf tissue, and more N per plant than isolated plants, suggesting a facilitative effect of Retama on Marrubium including increased availability of resources. Biomass of 1-yr-old cladodes, total biomass, total nitrogen content of 3-yr-old branches, and shoot water potential at midday were higher for Retama shrubs with Marrubium beneath them than in shrubs without Marrubium. Retama strongly improved its own environment, facilitated the growth of Marrubium and other species underneath its canopy, and at the same time obtained benefits from sheltering herbs underneath. The interaction between these two species was indirect, associated with differences in soil properties and with improved nutrient availability under shrubs compared with plants growing on their own. We propose that the mutual benefit of the association between Retama and Marrubium is best termed a facultative mutualism. Each partner benefits from greater availability of resources in the ''island of fertility'' that results from their association. If beneficial plant-plant interactions as described here are widespread, positive relationships may have a major role in determining the pattern and structure of plant communities.