Facilitation mediates species presence beyond their environmental optimum

Species distributions are driven by abiotic conditions that filter species with specific traits and physiological tolerances and match them with their suitable environment. Plant–plant interactions can constrict (through competition) or loosen (through facilitation) the strength of these environmental filters, which in turn inhibit or enhance establishment and recruitment of plant species at a finer spatial scale. Although competition is often the focus of community assembly processes that further impede the entry of plant species into a site, facilitation is also important for potentially loosening environmental filters (especially climatic filters such as temperature and precipitation), ultimately enhancing species occurrence beyond their physiological optimum. We used multiple data sets from the arid site of Rambla del Saltador Valley to test the hypothesis that facilitation by a nurse-plant promotes the presence of herbaceous, beneficiary species beyond their environmental optimum relative to open sites. Furthermore, we propose that the median elevation and community composition of herbaceous species expands with the age of the nurse-plant, and we tested this hypothesis by examining 105 beneficiary species under 50 nurse-plant shrubs varying in age from 6 to 48 years old. We found nurse-plants both facilitate herbaceous species occurrence beyond their median elevation and support more diverse and a distinctly different composition of species in contrast to open sites. Specifically, herbaceous species that originate from a median elevation more than 600 to 700 m above the site only existed beneath nurse-plants, and below this median elevation, half the species only occurred below the nurse-plant. Moreover, the richness and elevation provenance of the herbaceous species increased with increasing nurse-plant age. Our results highlight the importance of facilitation for alleviating physiological strain (in support of the strain hypothesis) and mediating regional spe