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Alpine plant communities are predicted to face range shifts and possibly extinctions with climate change. Fine-scale environmental variation such as nutrient availability or snowmelt timing may contribute to the ability of plant species to persist locally; however, variation in nutrient availability in alpine landscapes is largely unmeasured. On three mountains around Davos, Switzerland, we deployed Plant Root Simulator probes around 58 Salix herbacea plants along an elevational and microhabitat gradient to measure nutrient availability during the first 5 weeks of the summer growing season, and used in situ temperature loggers and observational data to determine date of spring snowmelt. We also visited the plants weekly to assess performance, as measured by stem number, fruiting, and herbivory damage. We found a wide snowmelt gradient which determined growing season length, as well as variations of an order of magnitude or more in the accumulation of 12 nutrients between different microhabitats. Higher nutrient availability had negative effects on most shrub performance metrics, for instance decreasing stem number and the proportion of stems producing fruits. High nutrient availability was associated with increased herbivory damage in early-melting microhabitats, but among late-emerging plants this pattern was reversed. We demonstrate that nutrient availability is highly variable in alpine settings, and that it strongly influences performance in an alpine dwarf shrub, sometimes modifying the response of shrubs to snowmelt timing. As the climate warms and human-induced nitrogen deposition continues in the Alps, these factors may contribute to patterns of local plants persistence.

Alpine dwarf shrub communities are phenologically linked with snowmelt timing, so early spring exposure may increase risk of freezing damage during early development, and consequently reduce seasonal growth. We examined whether environmental factors (duration of snow cover, elevation) influenced size and the vulnerability of shrubs to spring freezing along elevational gradients and snow microhabitats by modelling the past frequency of spring freezing events. We sampled biomass and measured the size of Salix herbacea, Vaccinium myrtillus, Vaccinium uliginosum and Loiseleuria procumbens in late spring. Leaves were exposed to freezing temperatures to determine the temperature at which 50% of specimens are killed for each species and sampling site. By linking site snowmelt and temperatures to long-term climate measurements, we extrapolated the frequency of spring freezing events at each elevation, snow microhabitat and per species over 37 years. Snowmelt timing was significantly driven by microhabitat effects, but was independent of elevation. Shrub growth was neither enhanced nor reduced by earlier snowmelt, but decreased with elevation. Freezing resistance was strongly species dependent, and did not differ along the elevation or snowmelt gradient. Microclimate extrapolation suggested that potentially lethal freezing events (in May and June) occurred for three of the four species examined. Freezing events never occurred on late snow beds, and increased in frequency with earlier snowmelt and higher elevation. Extrapolated freezing events showed a slight, non-significant increase over the 37-year record. We suggest that earlier snowmelt does not enhance growth in four dominant alpine shrubs, but increases the risk of lethal spring freezing exposure for less freezing-resistant species.

We evaluated the impacts of elevated CO2 in a treeline ecosystem in the Swiss Alps in a 9-year free-air CO2 enrichment (FACE) study. We present new data and synthesize plant and soil results from the entire experimental period. Light-saturated photosynthesis (A max) of ca. 35-year-old Larix decidua and Pinus uncinata was stimulated by elevated CO2 throughout the experiment. Slight down-regulation of photosynthesis in Pinus was consistent with starch accumulation in needle tissue. Above-ground growth responses differed between tree species, with a 33 % mean annual stimulation in Larix but no response in Pinus. Species-specific CO2 responses also occurred for abundant dwarf shrub species in the understorey, where Vaccinium myrtillus showed a sustained shoot growth enhancement (+11 %) that was not apparent for Vaccinium gaultherioides or Empetrum hermaphroditum. Below ground, CO2 enrichment did not stimulate fine root or mycorrhizal mycelium growth, but increased CO2 effluxes from the soil (+24 %) indicated that enhanced C assimilation was partially offset by greater respiratory losses. The dissolved organic C (DOC) concentration in soil solutions was consistently higher under elevated CO2 (+14 %), suggesting accelerated soil organic matter turnover. CO2 enrichment hardly affected the C-N balance in plants and soil, with unaltered soil total or mineral N concentrations and little impact on plant leaf N concentration or the stable N isotope ratio. Sustained differences in plant species growth responses suggest future shifts in species composition with atmospheric change. Consistently increased C fixation, soil respiration and DOC production over 9 years of CO2 enrichment provide clear evidence for accelerated C cycling with no apparent consequences on the N cycle in this treeline ecosystem.

Neighbor interactions are likely to play an important role in subarctic plant communities. We conducted experiments in Interior Alaska in which we crossed species removal with greenhouse warming manipulations. We examined changes in community biomass, and in plant survival and growth of individual species in response to experimental warming and to: (1) removal of whole species versus an equivalent amount of biomass across many species, and (2) removal of subdominant (locally common) versus minor (locally uncommon) plants. Community biomass indicated compensation in growth after removal of minor species and after biomass removal without elimination of entire species, but under-compensation after removal of subdominants. Growth and survival of individual species showed facilitation between some species. Warming increased growth of dominant vascular plants, but at the same time reduced survival, and these impacts were greater for larger, more mesic species than for the smaller species associated with drier habitats. Growth of mosses was reduced by the warming. Removal effects did not differ between warming and ambient conditions. The results indicate that common species are able to reduce resources for others (competitive effect) and increase their growth after neighbor removal, whereas locally uncommon species are not able to respond rapidly to increased resources made available by neighbor removal. Therefore, the impact of the presence of common species on locally uncommon species was facilitative overall, but not vice versa. The balance between disturbances such as changes in temperature and species losses from the community will likely be crucial in determining shifts in subsequent community composition.

A positive relationship between plant species richness and ecosystem functioning has been found in a number of experimental studies. Positive species interactions at high species numbers have been suggested as a cause, but mechanisms driving positive interactions have not often been tested. In this experiment we asked three questions: (1) What is the relationship between species richness and productivity in experimentally constructed moss communities? (2) Is this relationship affected by plant density? and (3) Can changes in moisture absorption and retention explain observed relationships? To answer these questions we exposed arctic tundra moss communities of different species richness levels (1-11 species) and two different densities in the greenhouse to two levels of drought (short and long). Biomass (by the community and individual species), height and community moisture absorption and retention were measured as response variables. High species diversity increased productivity (more so in low-density plots than in high-density plots), but only when plots were watered regularly. Plot moisture retention was improved at high species richness as well, and plant height and variation in height was increased compared to plants in monoculture. Under high-density and short-drought conditions 10 out of 12 species grew better in mixture than in monoculture, but under the long drought treatment only six species did. A positive feedback loop between biomass and improved humidity under high diversity was supported by path analysis. We conclude that in this community the relationship between species richness and productivity depends on moisture availability and density, with improved water absorption and retention likely to be the mechanism for increased plant growth when drought periods are short. Furthermore, since this is the opposite of what has been found for temperate moss communities, conclusions from one system cannot automatically be extrapolated to other systems.