Elevation, moisture and shade drive the functional and phylogenetic meadow communities' assembly in the northeastern Tibetan Plateau

Despite a long history of alpine meadows studies, uncertainty remains about the importance of environmental factors in structuring their assembly. We examined the functional and phylogenetic structure of 170 alpine Tibetan meadow communities in relation to elevation, soil moisture and shade. Functional community structure was estimated with both community-weighted mean (CWM) trait values for specific leaf area (SLA), plant height and seed mass and functional diversity (Rao's quadratic index) for their traits individually and in combination (multivariate functional diversity). We found that shade induced by woody plants significantly increased the phylogenetic diversity and functional diversity of SLA of co-occurring species, suggesting that woody plants behave as "ecosystem engineers" creating a different environment that allows the existence of shade tolerant species and thereby facilitates the coexistence of plant species with different light resource acquisition strategies. We also found evidence for a clear decrease in phylogenetic diversity, CWM and functional diversity related to plant height in the two extreme, both the dry and wet, soil moisture conditions. This indicates that both drought and excess moisture may act as environmental filters selecting species with close phylogenetic relationships and similar height. Moreover, we detected significant decreases in both CWM and functional diversity for seed mass along elevational gradients, suggesting that low net primary productivity (NPP) limits seed size. Finally, because of different individual trait responses to environmental factors, the multivariate functional diversity did not change across environmental gradients. This lack of multivariate response supports the hypothesis that multiple processes, such as environmental filtering, competition and facilitation, may operate simultaneously and exert opposing effects on community assembly along different niche (e.g., water use, light acquisition) axes, resulting in no overall functional community structure change. This contrast between individual and multivariate trait patterns highlights the importance of examining individual traits linked with different ecological processes to better understand the mechanisms of community assembly.