Nutritional changes in trees during drought-induced mortality: A comprehensive meta-analysis and a field study.

Both macronutrients and micronutrients are essential for tree growth and development through participating in various ecophysiological processes. However, the impact of the nutritional status of trees on their ability to withstand drought-induced mortality remains inconclusive. We thus conducted a comprehensive meta-analysis, compiling data on 11 essential nutrients from 44 publications (493 independent observations). Additionally, a field study was conducted on Pinus sylvestris L. trees with varying drought-induced vitality loss in the "Visp" forest in southern Switzerland. No consistent decline in tree nutritional status was observed during tree mortality. The meta-analysis revealed significantly lower leaf potassium (K), iron (Fe), and copper (Cu) concentrations with tree mortality. However, the field study showed no causal relationships between nutritional levels and the vitality status of trees. This discrepancy is mainly attributed to the intrinsic differences in the two types of experimental designs and the ontogenetic stages of target trees. Nutrient reductions preceding tree mortality were predominantly observed in non-field conditions, where the study was conducted on seedlings and saplings with underdeveloped root systems. It limits the nutrient uptake capacity of these young trees during drought. Furthermore, tree nutritional responses are also influenced by many variables. Specifically, (a) leaf nutrients are more susceptible to drought stress than other organs; (b) reduced tree nutrient concentrations are more prevalent in evergreen species during drought-induced mortality; (c) of all biomes, Mediterranean forests are most vulnerable to drought-induced nutrient deficiencies; (d) soil types affect the direction and extent of tree nutritional responses. We identified factors that influence the relationship between tree nutritional status and drought survival, and proposed potential early-warning indicators of impending tree mortality, for example, decreased K concentrations with declining vitality. These findings contribute to our understanding of tree responses to drought and provide practical implications for forest management strategies in the context of global change.

Functional structure mediates the responses of productivity to addition of three nitrogen compounds in a meadow steppe.

Atmospheric nitrogen (N) deposition is altering grassland productivity and community structure worldwide. Deposited N comes in different forms, which can have different consequences for productivity due to differences in their fertilization and acidification effects. We hypothesize that these effects may be mediated by changes in plant functional traits. We investigated the responses of aboveground primary productivity and community functional composition to addition of three nitrogen compounds (NH4NO3, [NH4]2SO4, and CO[NH2]2) at the rates of 0, 5, 10, 20 g N m-2 yr-1. We used structural equation modeling (SEM) to evaluate how functional structure influences the responses of productivity to the three N compounds. Nitrogen addition increased community-level leaf chlorophyll content but decreased leaf dry matter content and phosphorus concentration. These changes were mainly due to intra-specific variation. Functional dispersion of traits was reduced by N addition through changes in species composition. SEM revealed that fertilization effects were more important than soil acidification for the responses of productivity to CO(NH2)2 addition, which enhanced productivity by decreasing functional trait dispersion. In contrast, the effects of (NH4)2SO4 and NH4NO3 were primarily due to soil acidification, influencing productivity via community-weighted means of functional traits. Our results suggest that N forms with different fertilizing and acidifying effects influence productivity via different functional traits pathways. Our study also emphasizes the need for in situ experiments with the relevant N compounds to accurately understand and predict the ecological effects of atmospheric N deposition on ecosystems.

Rhizosphere activity in an old-growth forest reacts rapidly to changes in soil moisture and shapes whole-tree carbon allocation.

Drought alters carbon (C) allocation within trees, thereby impairing tree growth. Recovery of root and leaf functioning and prioritized C supply to sink tissues after drought may compensate for drought-induced reduction of assimilation and growth. It remains unclear if C allocation to sink tissues during and following drought is controlled by altered sink metabolic activities or by the availability of new assimilates. Understanding such mechanisms is required to predict forests' resilience to a changing climate. We investigated the impact of drought and drought release on C allocation in a 100-y-old Scots pine forest. We applied 13CO2 pulse labeling to naturally dry control and long-term irrigated trees and tracked the fate of the label in above- and belowground C pools and fluxes. Allocation of new assimilates belowground was ca. 53% lower under nonirrigated conditions. A short rainfall event, which led to a temporary increase in the soil water content (SWC) in the topsoil, strongly increased the amounts of C transported belowground in the nonirrigated plots to values comparable to those in the irrigated plots. This switch in allocation patterns was congruent with a tipping point at around 15% SWC in the response of the respiratory activity of soil microbes. These results indicate that the metabolic sink activity in the rhizosphere and its modulation by soil moisture can drive C allocation within adult trees and ecosystems. Even a subtle increase in soil moisture can lead to a rapid recovery of belowground functions that in turn affects the direction of C transport in trees.

Ethylene-regulated leaf lifespan explains divergent responses of plant productivity to warming among three hydrologically different growing seasons.

Leaf senescence is known to be regulated by the plant hormone ethylene, but how leaf lifespan responds to global environmental change and links to ecosystem-level responses remains largely unexplored. Here we investigated the effects of climate warming and nitrogen addition on plant functional traits, plant hormone ethylene and net primary production in a 13-year field experiment in a desert steppe. Across the last 3 years of the experiment (2016-2018), plant productivity increased under warming only in 2016, when there was above normal precipitation, but consistently increased with nitrogen addition. Warming enhanced net photosynthesis, leaf nitrogen and ethylene production and reduced leaf lifespan in 2016 (a wet year), but not in 2017 (a drought year); the effect of warming in 2018 (a year with normal precipitation) was opposite to 2016, likely due to the below-normal precipitation in the mid-growing season in 2018. Nitrogen addition led to increases in leaf nitrogen, ethylene production and net photosynthesis, and declines in leaf lifespan in 2016 and 2018, but not in 2017. The ethylene-regulated lifespan was further evidenced by the addition of CoCl2 (an ethylene biosynthesis inhibitor) that reduced ethylene production and prolonged lifespan. Structural equation modeling showed that leaf lifespan had a negative effect on plant productivity, both directly and indirectly via its negative effect on net photosynthesis, across all 3 years. Our results demonstrate the divergent responses of leaf lifespan and, in turn, plant productivity to warming under inter-annual and intra-annual precipitation variation, thus linking plant hormone production, functional traits and ecosystem functioning in the face of global environmental change.

Homeostatic levels of nonstructural carbohydrates after 13 yr of drought and irrigation in Pinus sylvestris.

Nonstructural carbohydrates (NSCs) are important for the growth and survival of trees. Drought may lead to a decrease in tree growth and to NSC depletion, whereas increased soil moisture in otherwise dry ecosystems may increase growth and NSC concentrations. A long-term (13 yr) irrigation experiment was conducted in a Pinus sylvestris-dominated forest located at the dry margin of the species in southern Switzerland. We measured the relative leaf area, growth, NSCs, needle δ13 C, [N] and [P] in trees on control and irrigated plots. Irrigation resulted in higher growth rates and carbon isotope discrimination, but did not alter NSC levels. Growth and NSC decreased with decreasing leaf area in both treatments, but NSC did not correlate with leaf-level gas exchange indices, such as foliar δ13 C, [N] or [P]. A legacy effect was shown, as trees with initially low leaf area had limited ability to respond to prolonged irrigation. The NSC constancy across treatments provides evidence that carbohydrate storage may stay constant when climate changes are sufficiently slow to allow acclimation. Moreover, we speculate that total leaf area, rather than leaf gas exchange per unit leaf area, drives the variation in whole-tree carbohydrate dynamics in this system.

Differential responses of canopy nutrients to experimental drought along a natural aridity gradient.

The allocation and stoichiometry of plant nutrients in leaves reflect fundamental ecosystem processes, biotic interactions, and environmental drivers such as water availability. Climate change will lead to increases in drought severity and frequency, but how canopy nutrients will respond to drought, and how these responses may vary with community composition along aridity gradients is poorly understood. We experimentally addressed this issue by reducing precipitation amounts by 66% during two consecutive growing seasons at three sites located along a natural aridity gradient. This allowed us to assess drought effects on canopy nitrogen (N) and phosphorus (P) concentrations in arid and semiarid grasslands of northern China. Along the aridity gradient, canopy nutrient concentrations were positively related to aridity, with this pattern was driven primarily by species turnover (i.e., an increase in the relative biomass of N- and P-rich species with increasing aridity). In contrast, drought imposed experimentally increased N but decreased P concentrations in plant canopies. These changes were driven by the combined effects of species turnover and intraspecific variation in leaf nutrient concentrations. In addition, the sensitivity of canopy N and P concentrations to drought varied across the three sites. Canopy nutrient concentrations were less affected by drought at drier than wetter sites, because of the opposing effects of species turnover and intraspecific variation, as well as greater drought tolerance for nutrient-rich species. These contrasting effects of long-term aridity vs. short-term drought on canopy nutrient concentrations, as well as differing sensitivities among sites in the same grassland biome, highlight the challenge of predicting ecosystem responses to future climate change.

Warming-induced upward migration of the alpine treeline in the Changbai Mountains, northeast China.

Treeline responses to environmental changes describe an important phenomenon in global change research. Often conflicting results and generally too short observations are, however, still challenging our understanding of climate-induced treeline dynamics. Here, we use a state-of-the-art dendroecological approach to reconstruct long-term changes in the position of the alpine treeline in relation to air temperature at two sides in the Changbai Mountains in northeast China. Over the past 160 years, the treeline increased by around 80 m, a process that can be divided into three phases of different rates and drives. The first phase was mainly influenced by vegetation recovery after an eruption of the Tianchi volcano in 1702. The slowly upward shift in the second phase was consistent with the slowly increasing temperature. The last phase coincided with rapid warming since 1985, and shows with 33 m per 1°C, the most intense upward shift. The spatial distribution and age structure of trees beyond the current treeline confirm the latest, warming-induced upward shift. Our results suggest that the alpine treeline will continue to rise, and that the alpine tundra may disappear if temperatures will increase further. This study not only enhances mechanistic understanding of long-term treeline dynamics, but also highlights the effects of rising temperatures on high-elevation vegetation dynamics.

Effects of experimentally-enhanced precipitation and nitrogen on resistance, recovery and resilience of a semi-arid grassland after drought.

Resistance, recovery and resilience are three important properties of ecological stability, but they have rarely been studied in semi-arid grasslands under global change. We analyzed data from a field experiment conducted in a native grassland in northern China to explore the effects of experimentally enhanced precipitation and N deposition on both absolute and relative measures of community resistance, recovery and resilience--calculated in terms of community cover--after a natural drought. For both absolute and relative measures, communities with precipitation enhancement showed higher resistance and lower recovery, but no change in resilience compared to communities with ambient precipitation in the semi-arid grassland. The manipulated increase in N deposition had little effect on these community stability metrics except for decreased community resistance. The response patterns of these stability metrics to alterations in precipitation and N are generally consistent at community, functional group and species levels. Contrary to our expectations, structural equation modeling revealed that water-driven community resistance and recovery result mainly from changes in community species asynchrony rather than species diversity in the semi-arid grassland. These findings suggest that changes in precipitation regimes may have significant impacts on the response of water-limited ecosystems to drought stress under global change scenarios.

Microplastics stimulated soil bacterial alpha diversity and nitrogen cycle: A global hierarchical meta-analysis.

Microplastics (MPs) pollution is recognized as a global emerging threat with serious potential impacts on ecosystems. Our meta-analysis was conducted based on 117 carefully selected publications, from which 2160 datasets were extracted. These publications described experiments in which MPs were added to soil (in laboratory or greenhouse experiments or in the field) after which the soil microbial community was analyzed and compared to a control group. From these publications, we extracted 1315 observations on soil bacterial alpha diversity and richness indices and 845 datasets on gene abundance of bacterial genes related to the soil nitrogen cycle. These data were analyzed using a multiple hierarchical mixed effects meta-analysis. The mean effect of microplastic exposure was a significant decrease of soil bacterial community diversity and richness. We explored these responses for different regulators, namely MPs addition rates, particle size and plastic type, soil texture and land use, and study type. Of the bacterial processes involved in the soil nitrogen cycle, MPs addition significantly promoted assimilation of ammonium, nitrogen fixation and urea decomposition, but significantly inhibited nitrification. These results suggest that MPs contamination may have considerable impacts on soil bacterial community structure and function as well as on the soil nitrogen cycle.

Global hierarchical meta-analysis to identify the factors for controlling effects of antibiotics on soil microbiota.

It is widely known that antibiotics can affect the structure and function of soil microbial communities, but the specific degree of impact and controlled factors on different indicators remain inconclusive. We conducted a multiple hierarchical mixed effects meta-analysis on 2564 observations that were extracted from 60 publications, to comprehensively assess the impact of antibiotics on soil microbiota. The results showed that antibiotics had significant negative effects on soil microbial biomass, α-diversity and soil enzyme activity. Under neutral initial soil, when soil was derived from agricultural land or had a fine-textured, the negative impacts of antibiotics on soil microbial community were exacerbated. Both single and mixed additions of antibiotics had significant inhibitory effects on soil microbial enzyme activities. The Random Forest model predicted the following key moderators involved in the effects of antibiotics on the soil microbiome, and antibiotics type, soil texture were key moderators on the severity of soil microbial biomass changes. Soil texture, temperature and single or combined application constitute of antibiotics were the main drivers of effects on soil enzyme activities. The reported results can be helpful to assess the ecological risk of antibiotics in a soil environment and provides a scientific basis for the rational of antibiotics use in the soil environment.

Effects of thinning and understory removal on soil phosphorus fractions in subtropical pine plantations.

Forest management changes the physical environments and nutrient dynamics and then regulates the forest productivity. Soil phosphorus (P) availability is critical for productivity in tropical and subtropical forests. However, it was still poorly understood how soil P content and fraction respond to various forest management practices in these regions. Here, we measured the soil total P, available P, and Hedley's P fractions, including inorganic and organic P (Pi and Po), in subtropical pine plantations treated with understory removal (UR), non-dominant species thinning (NDST) and dominant species thinning (DST) after nine years. Compared to plantations without management (CK), treatments such as UR, NDST, and DST decreased soil total P at 0-10 cm and soil available P at 0-10 cm and 10-20 cm. Increases in resin-Pi, NaOH-Pi, and C.HCl-Pi resulted in a higher total Pi in 0-10 cm (p < 0.05) in treated plots (UR, NDST, and DST) than in CK plots. UR, NDST, and DST treatments increased NaHCO3-Po and NaOH-Po (p < 0.05) but decreased C.HCl-Po at a depth of 0-10 cm. Regardless of management treatments, soil total P, available P, and P fractions in 0-10 cm showed higher contents than those in 10-20 cm. There were positive relationships between total P and total Po (p < 0.01) and between available P and total Pi. There were also positive relationships between total P, available P, NaHCO3-Pi, and NaOH-Pi (p < 0.05). In conclusion, forest management such as UR, NDST, and DST decreased soil total P and available P, and transforming soil P fractions to available P will meet the P demand following management in the pine plantations of subtropical China.

Shrub encroachment leads to accumulation of C, N, and P in grassland soils and alters C:N:P stoichiometry: A meta-analysis.

Soil stoichiometry of carbon (C), nitrogen (N), and phosphorus (P) are indicators for nutrient balance. Shrub encroachment into grasslands could change nutrient concentrations and stoichiometry in soils, but the general patterns remain unclear. With a meta-analysis of a global dataset covering 344 observations from 68 studies, we examined the responses of grassland soil C:N:P stoichiometry to shrub encroachment under various environmental conditions. Our results show that: 1) Shrub encroachment significantly increased the concentrations of soil C (+29 %), N (+25 %), P (+20 %), C:N (+5 %), C:P (+12 %), and N:P (+6 %). The magnitude of such effects varied with climate, soil texture, and soil layer. 2) Increases in SOC and TN concentrations mainly occurred in Mediterranean and very humid climate zones. Soil C:P and N:P decreased in semi-humid climate zone after shrub encroachment. 3) The increases in SOC and TN concentrations and in the C:N, C:P, and N:P ratios after shrub encroachment were greater in the topsoil than in deeper soil layers. 4) Both finest-textured soil (clay) and coarsest-textured soil (sand) are beneficial for increase of soil nutrient concentrations following shrub encroachment. 5) The magnitude of the change in soil C:N was negatively correlated with the duration of shrub encroachment, due to greater increases in soil TN than in SOC concentrations with longer durations of encroachment. Our results indicate that soil stoichiometric shifts in shrub-encroached grasslands are relatively sensitive to environmental factors, including soil texture, soil pH, and climate. These findings help us to better understand the effects of shrub encroachment on biogeochemical cycling, functioning, and services in grasslands across a broad range of spatio-temporal scales.

Contrasting water-use strategies to climate warming in white birch and larch in a boreal permafrost region.

The effects of rising atmospheric CO2 concentrations (Ca) with climate warming on intrinsic water-use efficiency and radial growth in boreal forests are still poorly understood. We measured tree-ring cellulose δ13C, δ18O, and tree-ring width in Larix dahurica (larch) and Betula platyphylla (white birch), and analyzed their relationships with climate variables in a boreal permafrost region of northeast China over past 68 years covering a pre-warming period (1951-1984; base period) and a warm period (1985-2018; warm period). We found that white birch but not larch significantly increased their radial growth over the warm period. The increased intrinsic water-use efficiency in both species was mainly driven by elevated Ca but not climate warming. White birch but not larch showed significantly positive correlations between tree-ring δ13C, δ18O and summer maximum temperature as well as vapor pressure deficit in the warm period, suggesting a strong stomatal response in the broad-leaved birch to temperature changes. The climate warming-induced radial growth enhancement in white birch is primarily associated with a conservative water-use strategy. In contrast, larch exhibits a profligate water-use strategy. It implies an advantage for white birch over larch in the warming permafrost regions.

The Diversity and Composition of Soil Microbial Communities Differ in Three Land Use Types of the Sanjiang Plain, Northeastern China.

In recent years, the Sanjiang Plain has experienced drastic human activities, which have dramatically changed its ecological environment. Soil microorganisms can sensitively respond to changes in soil quality as well as ecosystem function. In this study, we investigated the changes in soil microbial community diversity and composition of three typical land use types (forest, wetland and cropland) in the Sanjiang Plain using phospholipid fatty acid analysis (PLFA) technology, and 114 different PLFA compounds were identified. The results showed that the soil physicochemical properties changed significantly (p < 0.05) among the different land use types; the microbial diversity and abundance in cropland soil were lower than those of the other two land use types. Soil pH, soil water content, total organic carbon and available nitrogen were the main soil physico-chemical properties driving the composition of the soil microbial community. Our results indicate that the soil microbial community response to the three different habitats is complex, and provide ideas for the mechanism by which land use changes in the Sanjiang Plain affect the structure of soil microbial communities, as well as a theoretical basis for the future management and sustainable use of the Sanjiang plain, in the northeast of China.

Soil fungal community characteristics vary with bamboo varieties and soil compartments.

Soil fungi play an important role in nutrient cycling, mycorrhizal symbiosis, antagonism against pathogens, and organic matter decomposition. However, our knowledge about the community characteristics of soil fungi in relation to bamboo varieties is still limited. Here, we compared the fungal communities in different soil compartments (rhizosphere vs. bulk soil) of moso bamboo (Phyllostachys edulis) and its four varieties using ITS high-throughput sequencing technology. The fungal α diversity (Shannon index) in bulk soil was significantly higher than that in rhizosphere soil, but it was not affected by bamboo variety or interactions between the soil compartment and bamboo variety. Soil compartment and bamboo variety together explained 31.74% of the variation in fungal community diversity. Soil compartment and bamboo variety were the key factors affecting the relative abundance of the major fungal taxa at the phylum and genus levels. Soil compartment mainly affected the relative abundance of the dominant fungal phylum, while bamboo variety primarily influenced the dominant fungal genus. Network analysis showed that the fungal network in rhizosphere soil was more complex, stable, and connected than that in bulk soil. A FUNGuild database analysis indicated that both soil compartment and bamboo variety affect fungal functions. Our findings provide new insights into the roles of both soil compartments and plant species (including variety) in shaping soil fungal communities.

Leaf stomatal traits rather than anatomical traits regulate gross primary productivity of moso bamboo (Phyllostachys edulis) stands.

Leaf stomatal and anatomical traits strongly influence plant productivity. Understanding the environmental adaptation mechanisms of leaf stomatal and anatomical traits and their relationship with ecosystem productivity is essential to better understand and predict the long-term adaptation strategies to climate change of moso bamboo forests. Here, we selected 6 sites within the moso bamboo distribution area, measured 3 leaf stomatal traits and 10 leaf anatomical traits of unmanaged moso bamboo stands. We explored the spatial variation characteristics of these traits and their response to environmental changes, assessed the relationships among these traits at regional scales through network analysis, and tested the direct and indirect effects of environmental, leaf stomatal and anatomical traits on gross primary productivity (GPP) of bamboo stands using structural equation modeling (SEM). The results showed that both climate and soil factors significantly affected leaf stomatal and anatomical traits of moso bamboo. Solar radiation (SR) and mean annual precipitation (MAP) out of the climatic factors were the key drivers of variation in leaf stomatal and anatomical traits, respectively. Soil moisture and nutrients out of the soil properties significantly affected both leaf stomatal and anatomical traits of moso bamboo. Network analysis further indicated that there was a significant correlation between leaf stomata and anatomical traits. Stomatal size (SS) showed the highest centrality value at the regional scale, indicating that it plays a key role in adjusting the adaptation of plants to external environmental conditions. SEM analysis showed that environment did not directly but indirectly affect GPP via stomatal performance. The environment explained 53.3% and 39.2% of the variation in leaf stomatal and anatomical traits, respectively, and leaf stomatal traits explained 20.8% of the regional variation in GPP. Our results demonstrate a direct effect of leaf stomatal traits rather than leaf anatomical traits on bamboo ecosystem productivity, which provides new insights into model predictions of bamboo forests under global climate change.

Effect of elevation on composition and diversity of fungi in the rhizosphere of a population of Deyeuxia angustifolia on Changbai Mountain, northeastern China.

Soil fungi are a key component of terrestrial ecosystems and play a major role in soil biogeochemical cycling. Although the diversity and composition of fungal communities are regulated by many abiotic and biotic factors, the effect of elevation on soil fungal community diversity and composition remains largely unknown. In this study, the soil fungal composition and diversity in Deyeuxia angustifolia populations along an elevational gradient (1,690 m to 2020 m a.s.l.) were assessed, using Illumina MiSeq sequencing, on the north-facing slope of the Changbai Mountain, northeastern China. Our results showed that soil physicochemical parameters changed significantly along with the elevational gradients. The Ascomycota and Basidiomycota were the most dominant phyla along with the gradient. Alpha diversity of soil fungi decreased significantly with elevation. Soil nitrate nitrogen (NO3--N) was positively correlated with fungal richness and phylogenetic diversity (PD), indicating that soil nitrate nitrogen (NO3--N) is a key soil property determining fungal community diversity. In addition to soil nitrate content, soil pH and soil moisture were the most important environmental properties determining the soil fungal diversity. Our results suggest that the elevational changes in soil physicochemical properties play a key role in shaping the community composition and diversity of soil fungi. This study will allow us to better understand the biodiversity distribution patterns of soil microorganisms in mountain ecosystems.

Resource availability and parasitism intensity influence the response of soybean to the parasitic plant Cuscuta australis.

Introduction: Parasitic plants can damage crop plants and consequently cause yield losses and thus threaten food security. Resource availability (e.g., phosphorus, water) has an important role in the response of crop plants to biotic attacks. However, how the growth of crop plants under parasitism are affected by environmental resource fluctuation is poorly understood. Methods: We conducted a pot experiment to test the effects of the intensity of Cuscuta australis parasitism and the availability of water and phosphorus (P) on soybean shoot and root biomass. Results and discussion: We found that low-intensity parasitism caused ~6% biomass reduction, while high-intensity parasitism caused ~26% biomass reduction in soybean. Under 5-15% water holding capacity (WHC), the deleterious effect of parasitism on soybean hosts was ~60% and ~115% higher than that under 45-55% WHC and 85-95% WHC, respectively. When the P supply was 0 µM, the deleterious effect of parasitism on soybean was 67% lower than that when the P supply was 20 µM. Besides, the biomass of C. australis was highest when both the water and the P availability were lowest. Cuscuta australis caused the highest damage to soybean hosts under 5 µM P supply, 5-15% WHC, and high-intensity parasitism. Additionally, C. australis biomass was significantly and negatively related to the deleterious effect of parasitism on soybean hosts and to the total biomass of soybean hosts under high-intensity parasitism, but not under low-intensity parasitism. Although high resource availability can promote soybean growth, the two resources have different impacts on the response of hosts to parasitism. Higher P availability decreased host tolerance to parasites, while higher water availability increased host tolerance. These results indicate that crop management, specifically water and phosphorus supply, can efficiently control C. australis in soybean. To our best knowledge, this appears to be the first study to test the interactive effect of different resources on the growth and response of host plants under parasitism.

Nitrogen addition and mowing alter drought resistance and recovery of grassland communities.

Nitrogen enrichment and land use are known to influence various ecosystems, but how these anthropogenic changes influence community and ecosystem responses to disturbance remains poorly understood. Here we investigated the effects of increased nitrogen input and mowing on the resistance and recovery of temperate semiarid grassland experiencing a three-year drought. Nitrogen addition increased grassland biomass recovery but decreased structural recovery after drought, whereas annual mowing increased grassland biomass recovery and structural recovery but reduced structural resistance to drought. The treatment effects on community biomass/structural resistance and recovery were largely modulated by the stability of the dominant species and asynchronous dynamics among species, and the community biomass resistance and recovery were also greatly driven by the stability of grasses. Community biomass resistance/recovery in response to drought was positively associated with its corresponding structural stability. Our study provides important experimental evidence that both nitrogen addition and mowing could substantially change grassland stability in both functional and structural aspects. Our findings emphasize the need to study changes across levels of ecological organization for a more complete understanding of ecosystem responses to disturbances under widespread environmental changes.

Using leaf traits to explain species co-existence and its consequences for primary productivity across a forest-steppe ecotone.

Trait-based approaches have been widely applied to uncover the mechanisms determining community assembly and biodiversity-ecosystem functioning relationships. However, they have rarely been used in forest-steppe ecotones. These ecosystems are extremely sensitive to disturbances due to their relatively complex ecosystem structures, functionings and processes. In this study, we selected seven sites along a transect from closed canopy forests (CF) to forest-steppe ecotones (FSE) and meadow steppes (MS) in northeast China. Six leaf functional traits (i.e. leaf nitrogen and phosphorus contents, leaf length and thickness, single leaf area and leaf mass per unit area, LMA) as well as the community composition and aboveground biomass at each site were measured. Both functional trait diversity indices (richness, evenness and divergence) and community-weighted mean trait values (CWMs) were calculated to quantify community trait distributions. We found that dominant species in the FSE communities showed acquisitive strategies with highest leaf nitrogen (Mean ± SE: 19.6 ± 0.5 mg g-1) and single leaf area (19.2 ± 1.3 cm2), but the lowest LMA (59.6 ± 1.3 g cm-2) values compared to adjacent CF and MS communities. The ecotone communities also exhibited the largest functional trait richness (TOP), evenness (TED) and divergence (FDis) values (0.46, 0.92 and 0.67, respectively). Overall, niche differentiation emerges as the main mechanism influencing the coexistence of plant species in ecotone ecosystems. In addition, CWMs of leaf traits were the most important predictors for estimating variations in aboveground productivity across the transect, suggesting a major influence of dominant species. Our findings suggest that vegetation management practices in forest-steppe ecotones should increasingly focus on community functional trait diversity, and support the establishment and regeneration of plant species with rapid resource acquisition strategies.