Direct and indirect effects of shrub encroachment on alpine grasslands mediated by plant-pollinator interactions

Abstract

1. Mutualistic interactions structure ecological communities and they are strongly influenced by the combined effect of different drivers of global change. Land-use changes and global warming have elicited rapid shrub encroachment in alpine grasslands in recent decades, which may have detrimental outcomes for native alpine forbs. In spite of the importance of this process, we lack knowledge about how shrub encroachment modifies community-wide patterns of plant–pollinator mutualistic interactions. 2. Based on the functional biodiversity hypothesis (FBH), which predicts higher pollinator biodiversity in species-rich plant communities, we asked whether the increase in nutritional resources available for pollinators due to shrub expansion modifies pollinator niche breadth and species richness, and whether these changes affect plant–plant interactions. 3. For this purpose, we compared quantitative plant–flower visitor interaction network assemblages at replicated plots in two habitat types in dry cryophilic grasslands of Sierra de Guadarrama (Spain): (i) encroached pastures (EP) and (ii) pastures dominated by forb species where shrub species are absent (PA). 4. As predicted by FBH, flower visitor richness increased in EP, but their niche breadth did not vary. Furthermore, shrubs had more interactions with flower visitors and received more visits per plant than forbs in EP in agreement with their significantly higher linkage and strength. 5. Overall, results revealed that moderate levels of shrub encroachment affected the flower visitation patterns of forb species in alpine grasslands as flower visitor diversity increased and plant–plant competition for shared flower visitors became greater. These findings highlight the need to use an integrative approach to study the cascading effects of global change drivers on species interactions and their impact on the structure and functioning of threatened ecological communities.