Despite increasing climatic stress, grassland management is intensifying globally to enhance productivity. However, how increased biomass removal and associated shifts in plant composition and plant–microbe feedbacks regulate ecosystem functioning, particularly in nutrient-poor tropical grasslands, remains poorly understood. We hypothesised that increasing biomass removal structures plant communities and their rhizomicrobiome from resource-conservative towards resource-acquisitive dominated systems, and that both low and high management intensities induce unfavourable feedbacks between nutrient cycling and resource allocation, constraining soil microbial activity, plant productivity, and drought resilience. In a native managed tropical grassland, four aboveground biomass removal frequencies (1×, 2×, 3×, and 6× cutting annually) were applied in the field. Intact soil core mesocosms were exposed to drought stress in a split-plot completely randomised design (CRD), and combined with 13CO2 pulse labelling. We assessed plant biomass yield, 13CO2 assimilation and belowground C allocation, arbuscular mycorrhizal fungi (AMF) colonisation, extracellular enzyme activities, and rhizosphere microbial community structure and function. Increased cutting frequency shifted vegetation from resource-conservative, “slow” strategists to more acquisitive, “fast” strategists, accompanied by a transition from oligotrophic, fungi-dominated to copiotrophic, bacteria-dominated rhizosphere communities. Drought reduced total 13C assimilation and biomass production but enhanced allocation of assimilates to symbiotic and stress-alleviating microbes, particularly AMF. Overall, our results demonstrate that biomass removal regulates plant–soil–microbe interactions underpinning ecosystem resilience and productivity under drought, highlighting the importance of optimising management intensity in tropical grasslands under climate change.

ABSTRACT Droughts affect soil microbial abundance and functions—key parameters of plant–soil carbon (C) allocation dynamics. However, the impact of drought may be modified by the mean climatic conditions to which the soil microbiome has previously been exposed. In a future warmer and drier world, effects of drought may therefore differ from those observed in studies that simulate drought under current climatic conditions. To investigate this, we used the field experiment ‘Hohenheim Climate Change,’ an arable field where predicted drier and warmer mean climatic conditions had been simulated for 12 years. In April 2021, we exposed this agroecosystem to 8 weeks of drought with subsequent rewetting. Before drought, at peak drought, and after rewetting, we pulse‐labelled winter wheat in situ with 13 CO 2 to trace recently assimilated C from plants to soil microorganisms and back to the atmosphere. Severe drought decreased soil respiration (−35%) and abundance of gram‐positive bacteria (−15%) but had no effect on gram‐negative bacteria, fungi, and total microbial biomass C. This pattern was not affected by the mean precipitation regime to which the microbes had been pre‐exposed. Reduced mean precipitation had, however, a legacy effect by decreasing the proportion of recently assimilated C allocated to the microbial biomass C pool (−50%). Apart from that, continuous soil warming was an important driver of C fluxes throughout our experiment, increasing plant biomass, root sugar concentration, labile C, and respiration. Warming also shifted microorganisms toward utilizing soil organic matter as a C source instead of recently assimilated compounds. Our study found that moderate shifts in mean precipitation patterns can impose a legacy on how plant‐derived C is allocated in the microbial biomass of a temperate agroecosystem during drought. The overarching effect of soil warming, however, suggests that how temperate agroecosystems respond to drought will mainly be affected by future temperature increases.

ABSTRACT Plant communities and soil microbial communities influence each other directly and indirectly via the resource pools they modify. Despite apparent sensitivities of plants and microbes to climate, little is known concerning how climate change will affect plant–soil relationships. We conducted a downslope translocation of intact soil–plant mesocosms in subalpine grasslands to mid‐ and low‐elevation sites to determine how climate change (warmer and drier conditions) influences plant–soil relationships. While soil nutrient pools and microbial composition were key determinants of plant community characteristics under control and moderate climate change (+1°C, +8 days growing season), these relationships diminished under major climate change (+3°C, +21 days growing season). Positive correlations of fungi and nitrogen‐fixing bacteria for plant growth emerged under moderate climate change and diminished under major climate change. Our findings indicate that climate change effects do not solely impact plant community metrics, soil nutrient pools, and soil microbial community composition, but also a breakdown in the ecological coupling among them. We found evidence of threshold‐like behavior for plant–soil relationships in response to major versus moderate environmental change and that plant community metrics and soil microbial dynamics may become more independent in subalpine grasslands following environmental shifts that accompany climate change.

This data [1] represents ACi curves of twelve winter wheat varieties, which were grown under elevated and ambient CO2 concentrations within a FACE experiment and the subsequent F1 generation was exposed to ambient and elevated CO2 concentrations in a highly controlled environment using climate chambers. The 12 winter wheat genotypes (Triticum aestivum L.) were selected based on their susceptibilty to leaf rust (Puccinia triticina Eriks.) and Fusarium head blight (Fusarium graminearum Schwabe) according to the descriptive variety list of the German Federal Office of Plant Varietes (Beschreibende Sortenliste, Bundessortenamt 2024). The aim was to obtain a diverse set of varieties with the widest possible range of susceptibilities to leaf rust and fusarium head blight. Photosynthesis was measured using the novel Dynamic Assimilation Technique, thus not with the common steady-state approach. The individual wheat plants were measured twice, once under saturating soil water availability (θFC) and once under reduced soil water availability (θcsoil). θcsoil represents the gravimetric water content when the soil matric potential drops below the root matric potential, thus the onset of plant drought stress (sensu Cai et al. [2]). The photosynthesis data was used to fit ACi curves and extract the maximum Rubisco carboxylation rate [Vcmax], maximum rate of electron transport [Jmax] and dark respiration [Rd]. At both measurements we determined BBCH and plant height to quantify plant morphological development, as well as leaf water potential to quantify plant ecohydrologic status. At the end of the experiment, biomass was harvested and reported. Further, we provide environmental data of the climate chambers in use.Within the data repository, we provide comprehensive experimental data on the investigation of transgenerational memory effects on photosynthetic efficiency. We provide photosynthetic raw data as well as processed (merged) and derived (extracted ACi fit) data. Additionally, we provide the R-code to reproduce the calculation of the derived parameters.Data on transgenerational memory effects (that is, the influence of the parental environment on offspring phenotype and performance) are scarce, i.e. on the adaptive capacity of the photosynthetic apparatus. Thus, the data provided here can contribute to closing this gap. The highly controlled environment allows to closely investigate cause-effect relationships, thereby contributing to a mechanistic understanding of the transgenerational memory effects on photosynthetic efficiency and how this is altered by reduced soil water availability. By using a recently developed methodological approach, the data contributes to further investigate the quality of the method and establish it within the field of plant ecophysiology.

Although variation in effect sizes and predicted values among studies of similar phenomena is inevitable, such variation far exceeds what might be produced by sampling error alone. One possible explanation for variation among results is differences among researchers in the decisions they make regarding statistical analyses. A growing array of studies has explored this analytical variability in different fields and has found substantial variability among results despite analysts having the same data and research question. Many of these studies have been in the social sciences, but one small “many analyst” study found similar variability in ecology. We expanded the scope of this prior work by implementing a large-scale empirical exploration of the variation in effect sizes and model predictions generated by the analytical decisions of different researchers in ecology and evolutionary biology. We used two unpublished datasets, one from evolutionary ecology (blue tit, Cyanistes caeruleus, to compare sibling number and nestling growth) and one from conservation ecology (Eucalyptus, to compare grass cover and tree seedling recruitment). The project leaders recruited 174 analyst teams, comprising 246 analysts, to investigate the answers to prespecified research questions. Analyses conducted by these teams yielded 141 usable effects (compatible with our meta-analyses and with all necessary information provided) for the blue tit dataset, and 85 usable effects for the Eucalyptus dataset. We found substantial heterogeneity among results for both datasets, although the patterns of variation differed between them. For the blue tit analyses, the average effect was convincingly negative, with less growth for nestlings living with more siblings, but there was near continuous variation in effect size from large negative effects to effects near zero, and even effects crossing the traditional threshold of statistical significance in the opposite direction. In contrast, the average relationship between grass cover and Eucalyptus seedling number was only slightly negative and not convincingly different from zero, and most effects ranged from weakly negative to weakly positive, with about a third of effects crossing the traditional threshold of significance in one direction or the other. However, there were also several striking outliers in the Eucalyptus dataset, with effects far from zero. For both datasets, we found substantial variation in the variable selection and random effects structures among analyses, as well as in the ratings of the analytical methods by peer reviewers, but we found no strong relationship between any of these and deviation from the meta-analytic mean. In other words, analyses with results that were far from the mean were no more or less likely to have dissimilar variable sets, use random effects in their models, or receive poor peer reviews than those analyses that found results that were close to the mean. The existence of substantial variability among analysis outcomes raises important questions about how ecologists and evolutionary biologists should interpret published results, and how they should conduct analyses in the future.

Non-native trees are of major importance in forestry and urban greening but are controversially discussed with regard to their effects on native biodiversity. Existing literature reports positive as well as negative biodiversity effects of non-native trees. Geographic, climatic and phylogenetic distance between the non-native and native tree species are named as important factors shaping these biodiversity effects. However, studies with a large geographic and taxonomic coverage are missing. Here we tested the effects of geographic, climatic and phylogenetic distance on resident insect abundance and diversity for 77 native and non-native tree species in a common garden trial. We identified phylogenetic distance of tree species to be the main driver of insect abundance and diversity with a negative effect of increasing phylogenetic distance and stronger effects for phytophagous than for non-phytophagous insects. This highlights the necessity to account for phylogenetic history shared between native and non-native congeners when managing non-native trees.

Drought threatens plant growth and related ecosystem services. The emergence of plant drought stress under edaphic drought is well studied, whilst the importance of atmospheric drought only recently gained momentum. Yet, little is known about the interaction and relative contribution of edaphic and atmospheric drought on the emergence of plant drought stress. We conducted a gradient experiment, fully crossing gravimetric water content (GWC: maximum water holding capacity-permanent wilting point) and vapour pressure deficit (VPD: 1-2.25 kPa) using five wheat varieties from three species (Triticum monococcum, T. durum & T. aestivum). We quantified the occurrence of plant drought stress on molecular (abscisic acid), cellular (stomatal conductance), organ (leaf water potential) and stand level (evapotranspiration). Plant drought stress increased with decreasing GWC across all organizational levels. This effect was magnified nonlinearly by VPD after passing a critical threshold of soil water availability. At around 20%GWC (soil matric potential 0.012 MPa), plants lost their ability to regulate leaf water potential via stomata regulation, followed by the emergence of hydraulic dysfunction. The emergence of plant drought stress is characterized by changing relative contributions of soil versus atmosphere and their non-linear interaction. This highly non-linear response is likely to abruptly alter plant-related ecosystem services in a drying world.

Soil microorganisms are the major players controlling the soil carbon (C) and nutrient cycling, however the extent to which the functional properties of soil microorganisms will alter with climate change in mountain grasslands is still unclear. To simulate future climatic conditions (higher temperature and less precipitation), intact plant-soil mesocosms of three mountain grassland sites, Esterberg (1260 m a.s.l.), Graswang (860 m a.s.l.), and Fendt 550 m a.s.l.), were translocated to the lowest elevation Bayreuth (350 m a.s.l.). After two years of translocation to warmer and drier environments, specific growth rates of soil microorganisms were 25 % slower than those at higher elevations. This indicated a shift towards to slow-growing K-strategists with increasing temperature and decreasing soil moisture, presumably attributed to the depletion of available C for microorganisms. Simultaneously, higher activity and faster substrate turnover time of C-acquiring enzymes in the warmer and drier soils implied a stronger C rather nutrient limitation. This was supported by the increased vector length, which was induced by decreased dissolved organic compounds at lower elevation. In contrast, microorganisms were limited by nitrogen (N) at higher elevation because of the stronger competition for N between plants and microorganisms. In short, translocated microorganisms increased their active fraction, enzymatic activity and altered enzyme systems but decreased specific growth rates. This indicated that soil microbial community adjusted to the changing climate resulting in faster substrate turnover, and in turn caused 15 % reduction in soil C content from Esterberg to Bayreuth. We therefore conclude that climate warming in combination with decreased precipitation increases microbial activity, leads to shifts in microbial life strategies and induces alterations in microbial nutrient limitation, i.e. shift from N- to C and P co-limitation with translocation to lower elevation sites. This in turn, is likely to result in C losses of mountain grassland soils under future climate change, which will amplify negative impacts on global warming.

Mountainous grassland soils are considered one of the most unique biological hotspots, rich in organic carbon (OC). At the same time, they are exposed to great threats, as climate warming is more pronounced in mountainous regions than in lowland areas. In this study, we assessed the effect of simulated warming (+1K, +2K, and + 3 K) on OC stocks and soil structure in grassland soils of the Northern Limestone Alps in Germany by translocating plant-soil mesocosms from high- (1260 m a.s.l., Rendzic Phaeozem) and mid- (860 m a. s. l., Haplic Cambisol) to low-elevation (600 m a.s.l). Plant-soil mesocosms were exposed to both extensive and intensive grassland management practices. Four years after translocation, we observed a rapid decrease of topsoil SOC stocks under intensive (−1.0 t C ha yr−1) and extensive management (-2.2 t C ha yr−1), under the highest temperature increase. Intensive management with about 1 t C ha−1 yr−1 higher manure C return than extensive management (1.6 vs. 0.8 t C ha−1 yr−1 intensive and extensive, respectively) may explain the difference in SOC losses between different management treatments. Under both management practices, the loss of SOC was mainly associated with a decrease of large macroaggregates, at both management practices. In addition, different aggregate specific OC loss rates resulted in an altered distribution of OC among the aggregate size classes. Our study provides evidence that simulated climate change induced a rapid and substantial decline of SOC in mountainous, OC-rich grassland soils, which may be attributed to decreased physical OC protection within large macroaggregates. Optimized grassland management in form of increased application of organic fertilizers could only partially offset the SOC loss by improved formation of small macroaggregates.

Warming in mountain regions is projected to be three times faster than the global average. Pronounced climate change will likely lead to species reshuffling in mountain plant communities and consequently change ecosystem resilience and functioning. Yet, little is known about the role of inter‐ versus intraspecific changes of plant traits and their consequences for functional richness and evenness of mountain plant communities under climate change. We performed a downslope translocation experiment of intact plant‐soil mesocosms from an alpine pasture and a subalpine grassland in the Swiss and Austrian Alps to simulate an abrupt shift in climate and removal of dispersal barriers. Translocated plant communities experienced warmer and dryer climatic conditions. We found a considerable shift from resource conservative to resource acquisitive leaf‐economy in the two climate change scenarios. However, shifts in leaf‐economy were mainly attributable to species turnover, namely colonization by novel lowland species with trait expressions for a wider range of resource use. We also found an increase in vegetative height of the warmed and drought‐affected alpine plant community, while trait plasticity to warming and drought was limited to few graminoid species of the subalpine plant community. Our results highlight the contrast between the strong competitive potential of novel lowland species in quickly occupying available niche space and native species' lack of both the intraspecific trait variability and the plant functional trait expressions needed to increase functional richness under warming and drought. This is particularly important for the trailing range of many mountain species (i.e. subalpine zone) where upward moving lowland species are becoming more abundant and abiotic climate stressors are likely to become more frequent in the near future. Our study emphasizes mountain plant communities' vulnerability to novel climates and biotic interactions under climate change and highlights graminoid species as potential winners of a warmer and dryer future. Keywords: alpine grassland, functional diversity, invasion, species turnover, traitspace, translocation

With ongoing climate change and the increase in extreme weather events, especially droughts, the challenge of maintaining food security is becoming ever greater. Locally adapted landraces of crops represent a valuable source of adaptation to stressful environments. In the light of future droughts-both by altered soil water supply and increasing atmospheric water demand (vapor pressure deficit [VPD])-plants need to improve their water efficiency. To do so, plants can enhance their access to soil water by improving rhizosphere hydraulic conductivity via the exudation of mucilage. Furthermore, plants can reduce transpirational water loss via stomatal regulation. Although the role of mucilage and stomata regulation on plant water management have been extensively studied, little is known about a possible coordination between root mucilage properties and stomatal sensitivity as well as abiotic drivers shaping the development of drought resistant trait suits within landraces. Mucilage properties and stomatal sensitivity of eight Mexican landraces of Zea mays in contrast with one inbred line were first quantified under controlled conditions and second related to water demand and supply at their respective site of origin. Mucilage physical properties-namely, viscosity, contact angle, and surface tension-differed between the investigated maize varieties. We found strong influences of precipitation seasonality, thus plant water availability, on mucilage production (R 2 = .88, p < .01) and mucilage viscosity (R 2 = .93, p < .01). Further, stomatal sensitivity to increased atmospheric water demand was related to mucilage viscosity and contact angle, both of which are crucial in determining mucilage's water repellent, thus maladaptive, behavior upon soil drying. The identification of landraces with pre-adapted suitable trait sets with regard to drought resistance is of utmost importance, for example, trait combinations such as exhibited in one of the here investigated landraces. Our results suggest a strong environmental selective force of seasonality in plant water availability on mucilage properties as well as regulatory stomatal effects to avoid mucilage's maladaptive potential upon drying and likely delay critical levels of hydraulic dysfunction. By this, landraces from highly seasonal climates may exhibit beneficial mucilage and stomatal traits to prolong plant functioning under edaphic drought. These findings may help breeders to efficiently screen for local landraces with pre-adaptations to drought to ultimately increase crop yield resistance under future climatic variability.

Warming in mountain regions is projected to occur three times faster than the global average. Recently, a small number of observational studies have reported species loss in mountain plant communities and have explored mechanisms facilitating the colonization by novel species. We monitored translocated mountain plant communities and their novel competitor interactions following five years of exposure to downslope climates. We found increasing species turnover under two future climate scenarios with time. Local loss of native species was followed by the colonization by novel species after a severe drought year in the third year after translocation. Here, the assumed mechanism facilitating novel species colonization in mountain communities is direct environmental filtering. We find a time lag between the local loss of native species and the colonization by novel species, which can be explained by interacting climate stressors—that is, warming and drought—pushing communities across a threshold of resistance to colonization. Interestingly, despite significant changes in species identity and strong colonization by novel species, the proportional contribution of plant functional groups to aboveground biomass stayed stable across both sites of origin and over time. Our study provides experimental evidence of local species loss in mountain plant communities prone to severe climatic change, revealing abrupt threshold dynamics.

Carbon-water trade-offs in plants are adjusted through stomatal regulation. Stomatal opening enables carbon uptake and plant growth, whereas plants circumvent drought by closing stomata. The specific effects of leaf position and age on stomatal behavior remain largely unknown, especially under edaphic and atmospheric drought. Here, we compared stomatal conductance (gs ) across the canopy of tomato during soil drying. We measured gas exchange, foliage ABA level and soil-plant hydraulics under increasing vapor pressure deficit (VPD). Our results indicate a strong effect of canopy position on stomatal behavior, especially under hydrated soil conditions and relatively low VPD. In wet soil (soil water potential > -50 kPa), upper canopy leaves had the highest gs (0.727 ± 0.154 mol m-2 s-1) and assimilation rate (A; 23.4 ± 3.9 µmol m-2 s-1) compared to the leaves at a medium height of the canopy (gs : 0.159 ± 0.060 mol m2 s-1; A: 15.9 ± 3.8 µmol m-2 s-1). Under increasing VPD (from 1.8 to 2.6 kPa), gs , A and transpiration were initially impacted by leaf position rather than leaf age. However, under high VPD (2.6 kPa), age effect outweighed position effect. The soil-leaf hydraulic conductance was similar in all leaves. Foliage ABA levels increased with rising VPD in mature leaves at medium height (217.56 ± 85 ng g-1 FW) compared to upper canopy leaves (85.36 ± 34 ng g-1 FW). Under soil drought (< -50 kPa), stomata closed in all leaves resulting in no differences in gs across the canopy. We conclude that constant hydraulic supply and ABA dynamics facilitate preferential stomatal behavior and carbon-water trade-offs across the canopy. These findings are fundamental in understanding variations within the canopy, which helps in engineering future crops, especially in the face of climate change.

Lacking comparability among rainfall manipulation studies is still a major limiting factor for generalizations in ecological climate change impact research. A common framework for studying ecological drought effects is urgently needed to foster advances in ecological understanding the effects of drought. In this study, we argue, that the soil–plant–atmosphere‐continuum (SPAC), describing the flow of water from the soil through the plant to the atmosphere, can serve as a holistic concept of drought in rainfall manipulation experiments which allows for the reconciliation experimental drought ecology. Using experimental data, we show that investigations of leaf water potential in combination with edaphic and atmospheric drought – as the three main components of the SPAC – are key to understand the effect of drought on plants. Based on a systematic literature survey, we show that especially plant and atmospheric based drought quantifications are strongly underrepresented and integrative assessments of all three components are almost absent in current experimental literature. Based on our observations we argue, that studying dynamics of plant water status in the framework of the SPAC can foster comparability of different studies conducted in different ecosystems and with different plant species and can facilitate extrapolation to other systems, species or future climates.

Research in global change ecology relies heavily on global climatic grids derived from estimates of air temperature in open areas at around 2 m above the ground. These climatic grids do not reflect conditions below vegetation canopies and near the ground surface, where critical ecosystem functions occur and most terrestrial species reside. Here, we provide global maps of soil temperature and bioclimatic variables at a 1‐km 2 resolution for 0–5 and 5–15 cm soil depth. These maps were created by calculating the difference (i.e. offset) between in situ soil temperature measurements, based on time series from over 1200 1‐km 2 pixels (summarized from 8519 unique temperature sensors) across all the world's major terrestrial biomes, and coarse‐grained air temperature estimates from ERA5‐Land (an atmospheric reanalysis by the European Centre for Medium‐Range Weather Forecasts). We show that mean annual soil temperature differs markedly from the corresponding gridded air temperature, by up to 10°C (mean = 3.0 ± 2.1°C), with substantial variation across biomes and seasons. Over the year, soils in cold and/or dry biomes are substantially warmer (+3.6 ± 2.3°C) than gridded air temperature, whereas soils in warm and humid environments are on average slightly cooler (−0.7 ± 2.3°C). The observed substantial and biome‐specific offsets emphasize that the projected impacts of climate and climate change on near‐surface biodiversity and ecosystem functioning are inaccurately assessed when air rather than soil temperature is used, especially in cold environments. The global soil‐related bioclimatic variables provided here are an important step forward for any application in ecology and related disciplines. Nevertheless, we highlight the need to fill remaining geographic gaps by collecting more in situ measurements of microclimate conditions to further enhance the spatiotemporal resolution of global soil temperature products for ecological applications.

Abstract Warming due to climate change is generally expected to lengthen the growing season in areas of seasonal climate and to advance plant phenology, particularly the onset of leafing and flowering. However, a reduction in aboveground biomass production and reproductive output may occur when warming is accompanied by drought that crosses critical water deficit thresholds. Tracking warmer temperatures has been shown to be species‐specific with unknown impacts on community composition and productivity. The variability in species’ ability to leverage earlier leaf unfolding and flowering into increased aboveground net primary production (ANPP) or increased investments into reproductive organs has heretofore been poorly explored. We tested whether phenological sensitivity to temperature, as a result of experimental warming, directly translated into increased plant performance, as measured by ANPP and flower abundance. In order to experimentally simulate climate warming, we translocated a total of 45 intact soil–plant communities downslope along an elevational gradient of 900 m within the European Alps from 1260 to 350 m asl and weekly recorded flower abundance and total green cover as well as cumulative biomass production at peak growing season. We found that advanced phenology at lower elevations was related to increased reproductive performance and conditional on whether they experienced drought stress. While a temperature increase of +1K had positive effects on the amount of reproductive organs for species with accelerated phenology, temperature increase going along with drier conditions resulted in plants being unable to sustain early investment in reproduction as measured by flower abundance. This finding highlights that the interaction of two climate change drivers, warming and drought, can push communities’ past resistance thresholds. Moreover, we detected biotic competition mechanisms and shifts toward forb‐depressed states with graminoids best taking advantage of experimentally altered increased temperature and reduced precipitation. Our results suggest that while species may track warmer future climates, concurrent drought events post a high risk for failure of temperature‐driven improvement of reproductive performance and biomass production in the European Alps.

Plant community biomass production is co-dependent on climatic and edaphic factors that are often covarying and non-independent. Disentangling how these factors act in isolation is challenging, especially along large climatic gradients that can mask soil effects. As anthropogenic pressure increasingly alters local climate and soil resource supply unevenly across landscapes, our ability to predict concurrent changes in plant community processes requires clearer understandings of independent and interactive effects of climate and soil. To address this, we developed a multispecies phytometer (i.e., standardized plant community) for separating key drivers underlying plant productivity across gradients. Phytometers were composed of three globally cosmopolitan herbaceous perennials, Dactylis glomerata, Plantago lanceolata, and Trifolium pratense. In 2017, we grew phytometer communities in 18 sites across a pan-European aridity gradient in local site soils and a standardized substrate and compared biomass production. Standard substrate phytometers succeeded in providing a standardized climate biomass response independent of local soil effects. This allowed us to factor out climate effects in local soil phytometers, establishing that nitrogen availability did not predict biomass production, while phosphorus availability exerted a strong, positive effect independent of climate. Additionally, we identified a negative relationship between biomass production and potassium and magnesium availability. Species-specific biomass responses to the environment in the climate-corrected biomass were asynchronous, demonstrating the importance of species interactions in vegetation responses to global change. Biomass production was co-limited by climatic and soil drivers, with each species experiencing its own unique set of co-limitations. Our study demonstrates the potential of phytometers for disentangling effects of climate and soil on plant biomass production and suggests an increasing role of P limitation in the temperate regions of Europe.

Semi-natural, agriculturally used grasslands provide important ecologic and economic services, such as feed supply. In mountain regions, pastures are the dominant agricultural system and face more severe climate change impacts than lowlands. Climate change threatens ecosystem functions, such as aboveground net primary production [ANPP] and its nutrient content. It is necessary to understand the impacts of climate change and land-management on such ecosystems to develop management practices to sustainably maintain provision of ecosystem services under future climatic conditions. We studied the effect of climate change and different land-use intensities on plant-soil communities by the downslope translocation of plant-soil mesocosms along an elevation gradient in 2016, and the subsequent application of two management types (extensive vs. intensive). Communities’ response to ANPP and leaf carbon (C), nitrogen (N), and phosphorus (P) content was quantified over the subsequent two years after translocation. ANPP increased with warming in 2017 under both management intensities, but this effect was amplified by intensive land-use management. In 2018, ANPP of intensively managed communities decreased, in comparison to 2017, from 35% to 42%, while extensively managed communities maintained their production levels. The changes in ANPP are coupled with an exceptionally dry year in 2018, with up to 100 more days of drought conditions. The C:N of extensively managed communities was higher than those of intensively managed ones, and further increased in 2018, potentially indicating shifts in resource allocation strategies that may explain production stability. Our results revealed a low resistance of intensively managed communities’ ANPP under especially dry conditions. The ability to alter resource allocation likely enables a constant level of production under extensive management, but this ability is lost under intensive management. Thus, future drought events may leave intensive management as a non-sustainable farming practice, and ultimately threaten ecosystem services of montane pastures.

Alpine and prealpine grasslands provide various ecosystem services and are hotspots for the storage of soil organic C (SOC) in Central Europe. Yet, information about aggregate-related SOC storage and its controlling factors in alpine and prealpine grassland soils is limited. In this study, the SOC distribution according to the aggregate size classes large macroaggregates (> 2000 μm), small macroaggregates (250–2000 μm), microaggregates (63–250 μm), and silt-/clay-sized particles (< 63 μm) was studied in grassland soils along an elevation gradient in the Northern Limestone Alps of Germany. This was accompanied by an analysis of earthworm abundance and biomass according to different ecological niches. The SOC and N stocks increased with elevation and were associated with relatively high proportions of water-stable macroaggregates due to high contents of exchangeable Ca 2+ and Mg 2+ . At lower elevations, earthworms appeared to act as catalyzers for a higher microaggregate formation. Thus, SOC stabilization by aggregate formation in the studied soils is a result of a joined interaction of organic matter and Ca 2+ as binding agents for soil aggregates (higher elevations), and the earthworms that act as promoters of aggregate formation through the secretion of biogenic carbonates (low elevation). Our study highlights the importance of aggregate-related factors as potential indices to evaluate the SOC storage potential in other mountainous grassland soils. Graphical abstract

Higher biodiversity can stabilize the productivity and functioning of grassland communities when subjected to extreme climatic events. The positive biodiversity-stability relationship emerges via increased resistance and/or recovery to these events. However, invader presence might disrupt this diversity-stability relationship by altering biotic interactions. Investigating such disruptions is important given that invasion by non-native species and extreme climatic events are expected to increase in the future due to anthropogenic pressure. Here we present one of the first multisite invader x biodiversity x drought manipulation experiment to examine combined effects of biodiversity and invasion on drought resistance and recovery at three semi-natural grassland sites across Europe. The stability of biomass production to an extreme drought manipulation (100% rainfall reduction; BE: 88 days, BG: 85 days, DE: 76 days) was quantified in field mesocosms with a richness gradient of 1, 3, and 6 species and three invasion treatments (no invader, Lupinus polyphyllus, Senecio inaequidens). Our results suggest that biodiversity stabilized community productivity by increasing the ability of native species to recover from extreme drought events. However, invader presence turned the positive and stabilizing effects of diversity on native species recovery into a neutral relationship. This effect was independent of the two invader's own capacity to recover from an extreme drought event. In summary, we found that invader presence may disrupt how native community interactions lead to stability of ecosystems in response to extreme climatic events. Consequently, the interaction of three global change drivers, climate extremes, diversity decline, and invasive species, may exacerbate their effects on ecosystem functioning.