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1. Assembly of plant communities has long been scrutinized through the lens of trait-based ecology. Studies generally analyse functional traits related to the vegetative growth, survival and resource acquisition and thus ignore how assembly rules may affect plants at other stages of their life cycle, particularly when seeds disperse, persist in soil and germinate. 2. Here, we analysed an extensive dataset of 16 traits for 167 species measured in-situ in 36 grasslands located along an elevation gradient and studied 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 means (CWMs) and functional diversity (FD) for all traits and established their relationships to mean annual temperature. 3. The seed traits were weakly correlated with vegetative traits. Therefore, these traits constituted independent axes of plant phenotypical variation that could be affected differently by community assembly rules. Abiotic filtering impacted mostly vegetative traits and to a lesser extent seed germination and morphological traits. Increasing low-temperature stress in upland sites selected for shortstature, slow-growing and frost-tolerant species that produce small quantities of small seeds with high degree of dormancy, high temperature requirements for germination and low germination speed. 4. Biotic interactions, specifically competition in the lowlands and facilitation in uplands, also filtered some functional traits in the studied communities. The benign climate in lowlands seems to promote plant with competitive strategies that include 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. 5. The changes in the main dispersal vector from zoochory to anemochory along the elevation gradient strongly affected the dispersal and the seed morphological trait structure of the communities. This may be explained by stronger vertical turbulence and moderate warm upwinds and low grazing intensity in the uplands that select for light and non-round shaped seeds with lower terminal velocity and endozoochorous potential. 6. Synthesis. We demonstrate that, in addition to vegetative 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 the analysis of community assembly. [ABSTRACT FROM AUTHOR]

Assembly of grassland communities has long been scrutinized through the lens of functional diversity. Studies generally point to an overwhelming influence of climate on observed patterns of functional diversity, despite experimental evidence demonstrating the importance of biotic interactions. We postulate that this is because most observational studies neglect both scale dependencies of assembly processes and phenotypic variation between individuals. Here, we test for changes in the importance of abiotic filtering and biotic interactions along a stress gradient by explicitly accounting for different scales. In addition to quantifying intraspecific trait variability (ITV), we also vary the two components of spatial scale, including grain (i.e. community size) and extent (i.e. the geographical area that defines the species pool)., We sampled 20 grassland communities in ten sites distributed along a 975-m elevation gradient. At each site, we measured seven functional traits for a total of 2020 individuals at different spatial grains. We related community functional diversity metrics to the main environmental gradient of our study area, growing season length (GSL), and assessed the dependence of these relationships on spatial grain, spatial extent and ITV., At large spatial grain and extent, the imprint of environmental filtering on functional diversity became more important with increasing stress (i.e. functional diversity decreased with shorter GSL). At small spatial grain and extent, we found a convex relationship between functional diversity and GSL congruent with the hypothesis that competition is dominant at low-stress levels while facilitative interactions are dominant at high-stress levels (i.e. high functional diversity at both extremes of the stress gradient). Importantly, the effect of intraspecific variability on assembly rules was noticeable only at small spatial grain and extent., Synthesis. Our study reveals how the combination of abiotic stress and biotic interactions shapes the functional diversity of alpine grasslands at different spatial scales, and highlights the importance of phenotype variation between individuals for community assembly processes at fine spatial scale. Our results suggest that studies analysing trait-based assembly rules but ignoring ITV and focusing on a single spatial scale are likely to miss essential features of community diversity patterns. [ABSTRACT FROM AUTHOR]