Abstract
BACKGROUND AND AIMS: Facilitation is an important ecological process for plant community structure and functional composition. Although direct facilitation has accrued most of the evidence so far, indirect facilitation is ubiquitous in nature and it has an enormous potential to explain community structuring. In this study, we assess the effect of direct and indirect facilitation on community productivity via taxonomic and functional diversity. METHODS: In an alpine community on the Tibetan Plateau, we manipulated the presence of the shrub Dasiphora fruticosa and graminoids in a fenced meadow and a grazed meadow to quantify the effects of direct and indirect facilitation. We measured four plant traits: height, lateral spread, specific leaf area (SLA) and leaf dry matter content (LDMC) of forbs; calculated two metrics of functional diversity [range of trait and community-weighted mean (CWM) of trait]; and assessed the responses of functional diversity to shrub facilitation. We used structural equation modelling to explore how shrubs directly and indirectly drove community productivity via taxonomic diversity and functional diversity. KEY RESULTS: We found stronger effects from herbivore-mediated indirect facilitation than direct facilitation on productivity and taxonomic diversity, regardless of the presence of graminoids. For functional diversity, the range and CWM of height and SLA, rather than lateral spread and LDMC, generally increased due to direct and indirect facilitation. Moreover, we found that the range of traits played a primary role over taxonomic diversity and CWM of traits in terms of shrub effects on community productivity. CONCLUSIONS: Our study reveals that the mechanism of shrub direct and indirect facilitation of community productivity in this alpine community is expanding the realized niche (i.e. expanding range of traits). Our findings indicate that facilitators might increase trait dispersion in the local community, which could alleviate the effect of environmental filters on trait values in harsh environments, thereby contributing to ecosystem functioning.