Facilitation associated with three contrasting shrub species in heavily grazed pastures on the eastern Tibetan Plateau

Small-scale vegetation patterns are frequently the results of plant-plant interactions such as facilitation and competition. Facilitation should be particularly pronounced when both abiotic and biotic stresses are high, but few studies were conducted in such habitats. In heavily-grazed pastures on the eastern Tibetan Plateau, an area with both high abiotic stress and strong biotic disturbance, we made relevés of herb species both beneath and outside canopies of three shrub species (Spiraea alpina, Sibiraea angustata and Potentilla fruticosa) differing in palatability and canopy structure. Herb species richness (S), pooled cover (PC) of all species, number of flowering species (FS) and number of inflorescences of all species (IN) were greater outside than beneath the shrub canopies. Evenness (J), in contrast, was smaller outside, while Shannon’s diversity index (H) was the same. Differences in S and J between plots beneath and outside the shrub canopies were greater in the case of P. fruticosa than in the cases of S. angustata and S. alpina, but differences in PC, FS or IN did not depend on the shrub species. Among the common species (frequency ≥6), 47–85% were equally frequent beneath and outside the shrubs, 13–39% were more frequent outside and 3–13% were more frequent beneath the shrubs. For the rarest species (frequency < 6), however, more species occurred beneath than outside the shrubs. The ordination diagram showed a clear separation between the relevés outside and beneath the shrubs and a gradient from P. fruticosa via S. alpina to S. angustata, accompanied by a distinct decrease in the extent of the difference between the vegetation beneath and outside the shrub canopies. In conclusion, the three shrub species facilitated some species in the herb layer and each shrub species had a specific impact, related to its canopy structure and palatability but also to the grazing pressure, which was greater around the P. fruticosa shrubs than around S. alpina and S. angustata.