Legacy Effects of Climate Extremes in Alpine Grassland.

Climate change is particularly apparent in many mountainous regions, with warming rates of more than twice the global average being reported for the European Alps. As a result, the probability of climate extremes has increased and is expected to rise further. In an earlier study, we looked into immediate impacts of experimentally imposed heat waves in alpine grassland, and found that these systems were able to cope with heat as long as enough water was available. However, concomitant drought led to increased stress, and reduced aboveground biomass production and green plant cover. Here, we studied the legacy effects (lag-effects) of the imposed climate extreme to see whether delayed responses occurred and how fast the alpine grassland could rebound from the initial changes. Green cover continued to be suppressed the two following years in communities that had been exposed to the most intense hot drought, while aboveground biomass production had returned to control levels by year 2. The initial lower resistance of the forb fraction in the communities was not compensated by faster recovery later on. This resulted in alpine communities that became (and remained) relatively enriched with graminoids, which resisted the original extreme better. The responses of alpine grassland to heat extremes with or without drought observed in this study resemble those typically found in lowland grassland in the short term. However, alpine grassland exhibited longer legacy effects from an annual perspective, with delayed recovery of aboveground production and persistent changes in community composition. This suggests that once initial resistance thresholds are exceeded, impacts may be longer-lasting in alpine grassland, where recovery is constrained by both the short growing season and difficult seedling establishment.

High land-use intensity exacerbates shifts in grassland vegetation composition after severe experimental drought.

Climate change projections anticipate increased frequency and intensity of drought stress, but grassland responses to severe droughts and their potential to recover are poorly understood. In many grasslands, high land-use intensity has enhanced productivity and promoted resource-acquisitive species at the expense of resource-conservative ones. Such changes in plant functional composition could affect the resistance to drought and the recovery after drought of grassland ecosystems with consequences for feed productivity resilience and environmental stewardship. In a 12-site precipitation exclusion experiment in upland grassland ecosystems across Switzerland, we imposed severe edaphic drought in plots under rainout shelters and compared them with plots under ambient conditions. We used soil water potentials to scale drought stress across sites. Impacts of precipitation exclusion and drought legacy effects were examined along a gradient of land-use intensity to determine how grasslands resisted to, and recovered after drought. In the year of precipitation exclusion, aboveground net primary productivity (ANPP) in plots under rainout shelters was -15% to -56% lower than in control plots. Drought effects on ANPP increased with drought severity, specified as duration of topsoil water potential ψ < -100 kPa, irrespective of land-use intensity. In the year after drought, ANPP had completely recovered, but total species diversity had declined by -10%. Perennial species showed elevated mortality, but species richness of annuals showed a small increase due to enhanced recruitment. In general, the more resource-acquisitive grasses increased at the expense of the deeper-rooted forbs after drought, suggesting that community reorganization was driven by competition rather than plant mortality. The negative effects of precipitation exclusion on forbs increased with land-use intensity. Our study suggests a synergistic impact of land-use intensification and climate change on grassland vegetation composition, and implies that biomass recovery after drought may occur at the expense of biodiversity maintenance.

Species richness effects on grassland recovery from drought depend on community productivity in a multisite experiment.

Biodiversity can buffer ecosystem functioning against extreme climatic events, but few experiments have explicitly tested this. Here, we present the first multisite biodiversity × drought manipulation experiment to examine drought resistance and recovery at five temperate and Mediterranean grassland sites. Aboveground biomass production declined by 30% due to experimental drought (standardised local extremity by rainfall exclusion for 72-98 consecutive days). Species richness did not affect resistance but promoted recovery. Recovery was only positively affected by species richness in low-productive communities, with most diverse communities even showing overcompensation. This positive diversity effect could be linked to asynchrony of species responses. Our results suggest that a more context-dependent view considering the nature of the climatic disturbance as well as the productivity of the studied system will help identify under which circumstances biodiversity promotes drought resistance or recovery. Stability of biomass production can generally be expected to decrease with biodiversity loss and climate change.

Simulated heat waves affected alpine grassland only in combination with drought.

The Alpine region is warming fast, and concurrently, the frequency and intensity of climate extremes are increasing. It is currently unclear whether alpine ecosystems are sensitive or resistant to such extremes. We subjected Swiss alpine grassland communities to heat waves with varying intensity by transplanting monoliths to four different elevations (2440-660 m above sea level) for 17 d. Half of these were regularly irrigated while the other half were deprived of irrigation to additionally induce a drought at each site. Heat waves had no significant impacts on fluorescence (Fv /Fm , a stress indicator), senescence and aboveground productivity if irrigation was provided. However, when heat waves coincided with drought, the plants showed clear signs of stress, resulting in vegetation browning and reduced phytomass production. This likely resulted from direct drought effects, but also, as measurements of stomatal conductance and canopy temperatures suggest, from increased high-temperature stress as water scarcity decreased heat mitigation through transpiration. The immediate responses to heat waves (with or without droughts) recorded in these alpine grasslands were similar to those observed in the more extensively studied grasslands from temperate climates. Responses following climate extremes may differ in alpine environments, however, because the short growing season likely constrains recovery.

Positive diversity-invasibility relationship in species-rich semi-natural grassland at the neighbourhood scale.

BACKGROUND AND AIMS: Attempts to answer the old question of whether high diversity causes high invasion resistance have resulted in an invasion paradox: while large-scale studies often find a positive relationship between diversity and invasibility, small-scale experimental studies often find a negative relationship. Many of the small-scale studies are conducted in artificial communities of even-aged plants. Species in natural communities, however, do not represent one simultaneous cohort and occur at various levels of spatial aggregation at different scales. This study used natural patterns of diversity to assess the relationship between diversity and invasibility within a uniformly managed, semi-natural community. METHODS: In species-rich grassland, one seed of each of ten species was added to each of 50 contiguous 16 cm(2) quadrats within seven plots (8 × 100 cm). The emergence of these species was recorded in seven control plots, and establishment success was measured in relation to the species diversity of the resident vegetation at two spatial scales, quadrat (64 cm(2)) within plots (800 cm(2)) and between plots within the site (approx. 400 m(2)) over 46 months. KEY RESULTS: Invader success was positively related to resident species diversity and richness over a range of 28-37 species per plot. This relationship emerged 7 months after seed addition and remained over time despite continuous mortality of invaders. CONCLUSIONS: Biotic resistance to plant invasion may play only a sub-ordinate role in species-rich, semi-natural grassland. As possible alternative explanations for the positive diversity-invasibility relationship are not clear, it is recommended that future studies elaborate fine-scale environmental heterogeneity in resource supplies or potential resource flows from resident species to seedlings by means of soil biological networks established by arbuscular mycorrhizal fungi.