Functional traits shape tree species distribution in the Himalayas

1. Plant functional traits determine plant performance and have therefore the potential to shape and predict species distributions along environmental gradients. This study analyses how traits affect tree species distribution along an elevational gradient in the Himalayas, Nepal. We addressed three questions: (a) what plant strategies can be distinguished among tree species? (b) how are plant traits and strategies associated with elevation? and (c) what plant traits are the best predictors of species positions along elevational gradient? 2. We quantified for 31 tree species a set of 39 plant traits related to resource uptake, use and conservation. We analysed how traits cluster into separate functions using a cluster analysis, and how traits and clusters associate into distinct plant strategies using a principal component analysis. 3. The cluster analysis showed five clusters of traits, reflecting (a) efficiency in vertical expansion, (b) efficiency in horizontal expansion, (c) efficiency in metabolism, (d) physical defence and (e) tree functional groups (conifers vs. broadleaf species). The first PCA axis reflects trade-offs in traits and clusters linked to elevation: highland species had trait values that increase safety against freezing induced cavitation, high solar radiation and strong wind, and that increase resource conservation. In contrast, lowland species had trait values that increase efficiency of resource acquisition, metabolism and expansion. Additionally, a bivariate analysis showed that the first PCA axis is more strongly related to elevation than the individual traits, indicating that the combination of traits is more important than the individual traits. An all subsets regression analysis showed that a small set of traits best explain species distribution: highland species had smaller size (low basal area), safer hydraulics (low conduit diameter) and lower leaf and branch display efficiency (low leaf area per xylem area, specific branch length) that increase persistence under harsh conditions. Remarkably, leaf traits were poor predictors of species' elevational positions. 4. Synthesis. Multiple trade-offs in plant size, hydraulics and light competitiveness shape species distribution along the elevational gradient. Along this extreme environmental gradient, stem and branch traits that integrate multiple plant organs and functions are better predictors of species' elevational distributions than leaf traits.