Resilience is a key concept in ecology and describes the capacity of an ecosystem to maintain its state and recover from disturbances. Numerous metrics have been applied to quantify resilience over a range of ecosystems. However, the way resilience is quantified affects the degree to which different trajectories of ecosystem recovery from disturbance are represented as 'resilient', precluding a comparison of disturbance responses across ecosystems and their properties and functions. To approach a broadly comparable assessment of resilience we suggest using a bivariate framework that jointly considers the disturbance impact and the recovery rate, both normalized to the undisturbed state of a system. We demonstrate the potential of the framework for attribution and integration across the various components underlying resilience.

Mountain grasslands have recently been exposed to substantial changes in land use and climate and in the near future will likely face an increased frequency of extreme droughts. To date, how the drought responses of carbon (C) allocation, a key process in the C cycle, are affected by land‐use changes in mountain grassland is not known.We performed an experimental summer drought on an abandoned grassland and a traditionally managed hay meadow and traced the fate of recent assimilates through the plant–soil continuum. We applied two 13 CO 2 pulses, at peak drought and in the recovery phase shortly after rewetting.Drought decreased total C uptake in both grassland types and led to a loss of above‐ground carbohydrate storage pools. The below‐ground C allocation to root sucrose was enhanced by drought, especially in the meadow, which also held larger root carbohydrate storage pools.The microbial community of the abandoned grassland comprised more saprotrophic fungal and Gram(+) bacterial markers compared to the meadow. Drought increased the newly introduced AM and saprotrophic (A+S) fungi:bacteria ratio in both grassland types. At peak drought, the 13C transfer into AM and saprotrophic fungi, and Gram(−) bacteria was more strongly reduced in the meadow than in the abandoned grassland, which contrasted the patterns of the root carbohydrate pools.In both grassland types, the C allocation largely recovered after rewetting. Slowest recovery was found for AM fungi and their 13C uptake. In contrast, all bacterial markers quickly recovered C uptake. In the meadow, where plant nitrate uptake was enhanced after drought, C uptake was even higher than in control plots. Synthesis. Our results suggest that resistance and resilience (i.e. recovery) of plant C dynamics and plant‐microbial interactions are negatively related, that is, high resistance is followed by slow recovery and vice versa. The abandoned grassland was more resistant to drought than the meadow and possibly had a stronger link to AM fungi that could have provided better access to water through the hyphal network. In contrast, meadow communities strongly reduced C allocation to storage and C transfer to the microbial community in the drought phase, but in the recovery phase invested C resources in the bacterial communities to gain more nutrients for regrowth. We conclude that the management of mountain grasslands increases their resilience to drought.

With 450,000 kmKobresia (syn. Carex) pygmaea dominated pastures in the eastern Tibetan highlands are the world's largest pastoral alpine ecosystem forming a durable turf cover at 3000-6000 m a.s.l. Kobresia's resilience and competitiveness is based on dwarf habit, predominantly below-ground allocation of photo assimilates, mixture of seed production and clonal growth, and high genetic diversity. Kobresia growth is co-limited by livestock-mediated nutrient withdrawal and, in the drier parts of the plateau, low rainfall during the short and cold growing season. Overstocking has caused pasture degradation and soil deterioration over most parts of the Tibetan highlands and is the basis for this man-made ecosystem. Natural autocyclic processes of turf destruction and soil erosion are initiated through polygonal turf cover cracking, and accelerated by soil-dwelling endemic small mammals in the absence of predators. The major consequences of vegetation cover deterioration include the release of large amounts of C, earlier diurnal formation of clouds, and decreased surface temperatures. These effects decrease the recovery potential of Kobresia pastures and make them more vulnerable to anthropogenic pressure and climate change. Traditional migratory rangeland management was sustainable over millennia, and possibly still offers the best strategy to conserve and possibly increase C stocks in the Kobresia turf.

An experimental forest ecosystem drought Drought is affecting many of the world’ s forested ecosystems, but it has proved challenging to develop an ecosystem-level mechanistic understanding of the ways that drought affects carbon and water fluxes through forest ecosystems. Werner et al . used an experimental approach by imposing an artificial drought on an entire enclosed ecosystem: the Biosphere 2 Tropical Rainforest in Arizona (see the Perspective by Eisenhauer and Weigelt). The authors show that ecosystem-scale plant responses to drought depend on distinct plant functional groups, differing in their water-use strategies and their position in the forest canopy. The balance of these plant functional groups drives changes in carbon and water fluxes, as well as the release of volatile organic compounds into the atmosphere. —AMS

Climate extremes and land-use changes can have major impacts on the carbon cycle of ecosystems. Their combined effects have rarely been tested. We studied whether and how the abandonment of traditionally managed mountain grassland changes the resilience of carbon dynamics to drought. In an in situ common garden experiment located in a subalpine meadow in the Austrian Central Alps, we exposed intact ecosystem monoliths from a managed and an abandoned mountain grassland to an experimental early-summer drought and measured the responses of gross primary productivity, ecosystem respiration, phytomass and its components, and of leaf area index during the drought and the subsequent recovery period. Across all these parameters, the managed grassland was more strongly affected by drought and recovered faster than the abandoned grassland. A bivariate representation of resilience confirmed an inverse relationship of resistance and recovery; thus, low resistance was related to high recovery from drought and vice versa. In consequence, the overall perturbation of the carbon cycle caused by drought was larger in the managed than the abandoned grassland. The faster recovery of carbon dynamics from drought in the managed grassland was associated with a significantly higher uptake of nitrogen from soil. Furthermore, in both grasslands leaf nitrogen concentrations were enhanced after drought and likely reflected drought-induced increases in nitrogen availability. Our study shows that ongoing and future land-use changes have the potential to profoundly alter the impacts of climate extremes on grassland carbon dynamics.Electronic supplementary materialThe online version of this article (doi:10.1007/s10021-017-0178-0) contains supplementary material, which is available to authorized users.

Abstract. Field measurements of aboveground net primary productivity (ANPP) in temperate grasslands suggest that both positive and negative asymmetric responses to changes in precipitation (P) may occur. Under normal range of precipitation variability, wet years typically result in ANPP gains being larger than ANPP declines in dry years (positive asymmetry), whereas increases in ANPP are lower in magnitude in extreme wet years compared to reductions during extreme drought (negative asymmetry). Whether the current generation of ecosystem models with a coupled carbon–water system in grasslands are capable of simulating these asymmetric ANPP responses is an unresolved question. In this study, we evaluated the simulated responses of temperate grassland primary productivity to scenarios of altered precipitation with 14 ecosystem models at three sites: Shortgrass steppe (SGS), Konza Prairie (KNZ) and Stubai Valley meadow (STU), spanning a rainfall gradient from dry to moist. We found that (1) the spatial slopes derived from modeled primary productivity and precipitation across sites were steeper than the temporal slopes obtained from inter-annual variations, which was consistent with empirical data; (2) the asymmetry of the responses of modeled primary productivity under normal inter-annual precipitation variability differed among models, and the mean of the model ensemble suggested a negative asymmetry across the three sites, which was contrary to empirical evidence based on filed observations; (3) the mean sensitivity of modeled productivity to rainfall suggested greater negative response with reduced precipitation than positive response to an increased precipitation under extreme conditions at the three sites; and (4) gross primary productivity (GPP), net primary productivity (NPP), aboveground NPP (ANPP) and belowground NPP (BNPP) all showed concave-down nonlinear responses to altered precipitation in all the models, but with different curvatures and mean values. Our results indicated that most models overestimate the negative drought effects and/or underestimate the positive effects of increased precipitation on primary productivity under normal climate conditions, highlighting the need for improving eco-hydrological processes in those models in the future.

Droughts strongly affect carbon and nitrogen cycling in grasslands, with consequences for ecosystem productivity. Therefore, we investigated how experimental grassland communities interact with groups of soil microorganisms. In particular, we explored the mechanisms of the drought-induced decoupling of plant photosynthesis and microbial carbon cycling and its recovery after rewetting. Our aim was to better understand how root exudation during drought is linked to pulses of soil microbial activity and changes in plant nitrogen uptake after rewetting. We set up a mesocosm experiment on a meadow site and used shelters to simulate drought. We performed two 13C-CO2 pulse labelings, the first at peak drought and the second in the recovery phase, and traced the flow of assimilates into the carbohydrates of plants and the water extractable organic carbon and microorganisms from the soil. Total microbial tracer uptake in the main metabolism was estimated by chloroform fumigation extraction, whereas the lipid biomarkers were used to assess differences between the microbial groups. Drought led to a reduction of aboveground versus belowground plant growth and to an increase of 13C tracer contents in the carbohydrates, particularly in the roots. Newly assimilated 13C tracer unexpectedly accumulated in the water-extractable soil organic carbon, indicating that root exudation continued during the drought. In contrast, drought strongly reduced the amount of 13C tracer assimilated into the soil microorganisms. This reduction was more severe in the growth-related lipid biomarkers than in the metabolic compounds, suggesting a slowdown of microbial processes at peak drought. Shortly after rewetting, the tracer accumulation in the belowground plant carbohydrates and in the water-extractable soil organic carbon disappeared. Interestingly, this disappearance was paralleled by a quick recovery of the carbon uptake into metabolic and growth-related compounds from the rhizospheric microorganisms, which was probably related to the higher nitrogen supply to the plant shoots. We conclude that the decoupling of plant photosynthesis and soil microbial carbon cycling during drought is due to reduced carbon uptake and metabolic turnover of rhizospheric soil microorganisms. Moreover, our study suggests that the maintenance of root exudation during drought is connected to a fast reinitiation of soil microbial activity after rewetting, supporting plant recovery through increased nitrogen availability.

Nitrous oxide is a powerful greenhouse gas whose atmospheric growth rate has accelerated over the past decade. Most anthropogenic N2O emissions result from soil N fertilization, which is converted to N2O via oxic nitrification and anoxic denitrification pathways. Drought-affected soils are expected to be well oxygenated; however, using high-resolution isotopic measurements, we found that denitrifying pathways dominated N2O emissions during a severe drought applied to managed grassland. This was due to a reversible, drought-induced enrichment in nitrogen-bearing organic matter on soil microaggregates and suggested a strong role for chemo- or codenitrification. Throughout rewetting, denitrification dominated emissions, despite high variability in fluxes. Total N2O flux and denitrification contribution were significantly higher during rewetting than for control plots at the same soil moisture range. The observed feedbacks between precipitation changes induced by climate change and N2O emission pathways are sufficient to account for the accelerating N2O growth rate observed over the past decade.