Assembly of plant communities has long been scrutinized through the lens of trait-based ecology. Studies generally analyze functional traits related to the vegetative growth, survival and resource acquisition and thus ignore how ecological processes may affect plants at other stages of their lifecycle, particularly when seeds disperse, persist in soil and germinate. Here, we analyzed an extensive data set of 16 traits for 167 species measured in-situ in 36 grasslands located along an elevational gradient and compared the impact of abiotic filtering, biotic interactions and dispersal on traits reflecting different trait categories: plant vegetative growth, germination, dispersal, and seed morphology. For each community, we quantified community weighted mean (CWM) and functional diversity (FD) for all traits and established their relationships to mean annual temperature. The seed traits were weakly correlated to vegetative traits and thus constituted independent axes of plant phenotypical variation that were affected differently by the ecological processes considered. Abiotic filtering impacted mostly the vegetative traits and to a lesser extent on seed germination and morphological traits. Increasing low-temperature stress towards colder sites selected for short-stature, slow-growing and frost-tolerant species that produce small quantity of smaller seeds with higher degree of dormancy, high temperature requirements for germination and comparatively low germination speed. Biotic interactions, specifically competition in the lowlands and facilitation in uplands, also filtered certain functional traits in the study communities. The benign climate in lowlands promoted plant with competitive strategies including fast growth and resource acquisition (vegetative growth traits) and early and fast germination (germination traits), whereas the effects of facilitation on the vegetative and germination traits were cancelled out by the strong abiotic filtering. The changes in the main dispersal vector from zoochory to anemochory along the gradient strongly affected the dispersal and the seed morphological trait structure of the communities. Specifically, stronger vertical turbulence and moderate warm-upwinds combined with low grazing intensity selected for light and non-round shaped seeds with lower terminal velocity and endozoochorous potential. Synthesis: We clearly demonstrate that, in addition to vegetation traits, seed traits can substantially contribute to functional structuring of plant communities along environmental gradients. Thus, the hard seed traits related to germination and dispersal are critical to detect multiple, complex community assembly rules. Consequently, such traits should be included in core lists of plant traits and, when applicable, be incorporated into analysis of community assembly.
Integrating succession and community assembly perspectives
