1. Climate warming is shifting the distributions of mountain plant species to higher elevations. Cold-adapted plant species are under increasing pressure from novel competitors that are encroaching from lower elevations. Plant capacity to adjust to these pressures may be measurable as variation in trait values within a species. In particular, the strength and patterns of intraspecific trait variation along abiotic and biotic gradients can inform us whether and how species can adjust their anatomy and morphology to persist in a changing environment.2. Here, we tested whether species specialized to high elevations or with narrow elevational ranges show more conservative (i.e. less variable) trait responses across their elevational distribution, or in response to neighbours, than species from lower elevations or with wider elevational ranges. We did so by studying intraspecific trait variation of 66 species along 40 elevational gradients in four countries in both hemispheres. As an indication of potential neighbour interactions that could drive trait variation, we also analysed plant species' height ratio, its height relative to its nearest neighbour.

The study of plant phenology has frequently been used to link phenological events to various factors, such as temperature or photoperiod. In the high-alpine environment, proper timing of the phenological cycle has always been crucial to overcome harsh conditions and potential extreme events (i.e. spring frosts) but little is known about the response dynamics of the vegetation, which could shape the alpine landscape in a future of changing climate. Alpine tundra vegetation is composed by an array of species belonging to different phytosociological optima and with various survival strategies, and snowbed communities are a relevant expression of such an extreme-climate adapted flora. We set eight permanent plots with each one in a snowbed located on the Cimalegna plateau in Northwestern Italy and then we selected 10 most recurring species among our plots, all typical of the alpine tundra environment and classified in 3 different pools: snowbed specialists, grassland species and rocky debris species. For 3 years we registered the phenophases of each species during the whole growing season using an adaptation of the BBCH scale. We later focused on the three most biologically relevant phenophases, i.e., flower buds visible, full flowering, and beginning of seed dispersion. Three important season-related variables were chosen to investigate their relationship with the phenological cycle of the studied species: (i) the Day Of Year (DOY), the progressive number of days starting from the 1st of January, used as a proxy of photoperiod, (ii) Days From Snow Melt (DFSM), selected to include the relevance of the snow dynamics, and (iii) Growing Degree Days (GDD), computed as a thermal sum. Our analysis highlighted that phenological development correlated better with DFSM and GDD than with DOY. Indeed, models showed that DOY was always a worse predictor since it failed to overcome interannual variations, while DFSM and marginally GDD were better suited to predict the phenological development of most of the species, despite differences in temperature and snowmelt date among the three years. Even if the response pattern to the three variables was mainly consistent for all the species, the timing of their phenological response was different. Indeed, species such as Salix herbacea and Ranunculus glacialis were always earlier in the achievement of the phenophases, while Agrostis rupestris and Euphrasia minima developed later and the remaining species showed an intermediate behavior. However, we did not detect significant differences among the three functional pools of species.

The study investigated plant-soil interactions along a proglacial chronosequence in the Italian Alps, with a specific focus on pioneer and grassland species structure and biogeochemical processes, with the aim to evaluate the biotic patterns in ecosystem development. We recorded vascular plant frequencies and the mean diameter of one pioneer and one grassland target species in 18 permanent plots distributed along six different stages encompassing a 170-years chronosequence in the Lauson Glacier forefield (NW Italy). We evaluated the main soil properties and measured the C:N:P stoichiometry in the biomass of pioneer and grassland target species and in the underlying soil. For comparative purposes, we analyzed also bare soils sampled near the sampled plant individuals. Pioneer species number and cover significantly increased 10 and 40 years after deglaciation respectively, while alpine grassland species cover and number peaked only after 65 and 140 years, respectively. Along the chronosequence, soils beneath vascular plants were enriched in nutrients, especially under individuals of alpine grassland species, with total organic C contents ranging between 1.3 and 8.9 g·kg−1 compared to 0.2 and 3.3 g·kg−1 in bare soils. Nitrogen content in bare soils was nearly undetectable, while it increased in the plant-affected soils, leading to a more balanced C:N:P stoichiometry in the oldest stages. The colonization of alpine grassland species started immediately, although species number and cover increased only when the soil acquired sufficient nutrient supply and functionality. Although the ecosystem remained C and N limited, the soil could provide adequate conditions for more competitive species establishment, as confirmed by the increasing number and cover of alpine grassland species. Thus, soil nutrient dynamics were strongly influenced by plants, with a major influence triggered by late-successional grassland species.

The snowbed habitats represent a relevant component of the alpine tundra biome, developing in areas characterized by a long-lasting snow cover. Such areas are particularly sensitive to climate changes, because small variations in air temperature, rain, and snowfall may considerably affect the pedoclimate and plant phenology, which control the soil C and N cycling. Therefore, it is fundamental to identify the most sensitive abiotic and biotic variables affecting soil nutrient cycling. This work was performed at seven permanent snowbed sites belonging to Salicetum herbaceae vegetation community in the northwestern Italian Alps, at elevations between 2,686 and 2,840 m.a.s.l. During a four-year study, we investigated climate, pedoclimate, floristic composition, phenology, and soil C and N dynamics. We found that lower soil water content and earlier melt-out day decreased soil N-NH 4 + , N-NO 3 − , dissolved organic carbon (DOC), dissolved organic nitrogen (DON), total dissolved nitrogen (TDN), microbial nitrogen (Nmicr), microbial carbon (Cmicr), and C:Nmicr ratio. The progression of the phenological stages of Salix herbacea reduced soil N-NH 4 + and increased DOC. Our results showed that the snow melt-out day, soil temperature, soil water content, and plant phenological stages were the most important factors affecting soil biogeochemical cycles, and they should be taken into account when assessing the effects of climate change in alpine tundra ecosystems, in the framework of long-term ecological research.