Drought and grazing combined: Contrasting shifts in plant interactions at species pair and community level

The combined effects of drought stress and grazing pressure on shaping plant–plant interactions are still poorly understood, while this combination is common in arid ecosystems. In this study we assessed the relative effect of grazing pressure and slope aspect (drought stress) on vegetation cover and soil functioning in semi-arid Mediterranean grassland–shrublands in southeastern Spain. Moreover, we linked these two stress factors to plant co-occurrence patterns at species-pair and community levels, by performing C-score analyses. Vegetation cover and soil functioning decreased with higher grazing pressure and more south-facing (drier) slopes. At the community level, plants at south-facing slopes were negatively associated at no grazing but positively associated at low grazing pressure and randomly associated at high grazing pressure. At north-facing slopes, grazing did not result in a shift in the direction of the association. In contrast, analysis of pairwise species co-occurrence patterns showed that the dominant species Stipa tenacissima and Anthyllis cytisoides shifted from excluding each other to co-occurring with increasing grazing pressure at north-facing slopes. Our findings highlight that for improved understanding of plant interactions along stress gradients, interactions between species pairs and interactions at the community level should be assessed, as these may reveal contrasting results.