Question: Positive interactions are predicted to be common in communities developing under high physical stress or high herbivory pressure due to neighbour amelioration of limiting physical and consumer stresses, respectively. However, when both stress sources meet in the same community, the relative importance of the two facilitation mechanisms is poorly understood. We ask: What is the relative importance of abiotic vs. biotic mechanisms of facilitation of tree saplings by shrubs in Mediterranean mountain forests? Location: Sierra Nevada, SE Spain (1800-1850 m a.s.l.) Methods: Saplings of four tree taxa (Acer opalus ssp. grana tense, Quercus ilex, Pinus nigra ssp. salzmanii and P. sylvestris var. nevadensis) were planted following a 2 × 2 factorial design: two levels of herbivory (control and ungulate exclusion) and two microhabitats (under shrubs and in open areas). Sapling survival and growth were monitored for five years. Results: Shrubs had positive effects on sapling survival both in control and ungulate excluded plots. This effect was speciesspecific, with shrubs increasing the survival of Acer opalus and Quercus ilex three and twofold, respectively, but having a minor effect on the Pinus species. Herbivory damage was also species-specific, being much higher for Acer opalus than for any other species. Shrubs did not protect saplings of any species against ungulates. Thus, all Acer saplings (the most damaged species) suffered herbivory outside the exclosures, which largely reduced sapling height. Conclusions: Protection from abiotic stress (summer drought and winter frost) was much more relevant than protection from biotic stress (herbivory). However, we propose that the final balance between the two mechanisms can be expected to vary strongly between sites, depending on the relative magnitude of the different sources of stress and the intrinsic traits (e.g. palatability) of the species interacting.Keywords: Abiotic stress; Facilitation mechanism; Herbivory pressure; Mediterranean mountain; Nurse shrub; Sapling survival; Species-specific effect.Nomenclature: Castroviejo et al. (1986Castroviejo et al. ( -2001 for tree species, and Molero-Mesa et al. (1992) for shrub species.Abbreviations: PPFD = Photosynthetic photon flux density; RII = Relative interaction index; VPD = Vapour pressure deficit.
IntroductionThe abundance, performance and spatial distribution of plant species are highly linked to the strength and sign of the interactions involving them in their communities (Roughgarden & Diamond 1986;Brown et al. 2001). In 1994, Bertness & Callaway proposed a conceptual model suggesting that strong positive interactions should be particularly common in communities developing under high abiotic stress and in communities with high consumer pressure, whereas competition should be the dominant interaction under mild physical conditions and low consumer pressure. Since then, a large number of empirical studies have supported this hypothesis by reporting that positive interactions are the rule under severe ...