The increase in atmospheric nitrogen (N) deposition has profound effects on soil respiration (SR). However, the responses of SR to the addition of different N compounds, particularly in saline–alkaline grasslands remain unclear. A 3-year controlled field experiment was conducted to investigate the responses of SR to different N compounds (NH₄NO₃, (NH₄)₂SO₄ and NH₄HCO₃) during the growing seasons in a saline–alkaline grassland located in the agro-pastoral ecotone of northern China. Our results demonstrated that SR showed a bimodal pattern and a significant interannual difference that was regulated by air or soil temperature and precipitation. Nitrogen addition had a significant effect on SR, and the effect of N addition on SR varied yearly, which was related to seasonal precipitation. The mean SR across 3 years (2017–2019) was increased by 19.9%, 13.0% and 16.6% with the addition of NH₄NO₃, (NH₄)₂SO₄ and NH₄HCO₃, respectively. The highest effect of NH₄NO₃ addition on SR across 3 years was ascribed to the highest aboveground net primary production, belowground net primary production (BNPP) and soil NO₃⁻ concentrations. SR (C loss) was significantly increased while plant productivity (C input) did not significantly change under NH₄HCO₃ addition, indicating a decrease in C sequestration. In addition, BNPP was the main direct factor influencing SR in this saline–alkaline grassland, and soil salinization (e.g. soil base cations and pH) indirectly affected SR through soil microorganisms. Notably, NH₄NO₃ addition overestimated the response of SR to N addition, and different N compounds should be considered, especially in saline–alkaline grassland.

Ecological intensification (EI) is the enhancement of ecosystem services to complement or substitute for the role of anthropogenic inputs in maintaining or increasing yields. EI has potential to increase farming’s environmental sustainability, e.g. reducing environmentally harmful management activities while sustaining yields. EI is based upon ecological processes which in turn are influenced by biodiversity. We review how biodiversity, particularly vascular plant diversity, can regulate ecosystem processes relevant to EI at multiple spatial scales. At an individual plant genotype level, complementarity in functional traits has a direct impact on productivity. At in-field, population level, mixtures of crop types confer resilience to minimize the risk of pest and disease incidence and spread. Scaling up to the field level, a diversity of non-crop plants (i.e. weeds) provides resources necessary for in-field functional processes, both below ground (carbon inputs, decomposition) and above ground (resource continuity for pollinators and natural enemies). At the landscape scale, mosaics of semi-natural and managed vegetation provide buffers against extreme events through flood and drought risk mitigation, climate amelioration and pest population regulation. Overall this emphasizes the importance of heterogeneity across scales in maintaining ecosystem functions in farmland. Major research challenges highlighted by our review include the need: to better integrate plant functional diversity (from traits to habitat scales) into cropping system design; to quantify the (likely interactive) contribution of plant diversity for effective EI relative to other management options; and to optimize through targeted management the system function benefits of biodiversity for resilient, efficient and productive agroecosystems.

The concept of nestedness originated from the field of biogeography decades ago and has been widely used in metacommunities and biological interaction networks, but there is still a lack of research within local communities. Moreover, studies on nestedness usually rarely incorporate the functional traits of the species and the environmental characteristics of the sites. In this study, we constructed a species presence–absence matrix of a 50-ha forest plot, used the simulated annealing algorithm to reveal the maximum nested structure and further tested the significance of nestedness patterns by constructing null ensembles. The nested ranks were used to represent the orders of species and quadrats in the maximum nestedness matrix. The regression tree analysis was used to reveal the relationships of nested ranks with environmental factors and functional traits. We found that the co-occurrence pattern of local plant communities was significantly nested. The regression tree results showed that the nested ranks of quadrats were determined by soil available phosphorus, soil water content, soil organic carbon and soil pH. Intraspecific variation of functional traits, including leaf C, leaf pH, leaf dry matter content and maximum photosynthetic rate rather than means of functional traits, provided a better explanation for the formation of species’ nested ranks. Understanding the causes of species and quadrats nested ranks provides novel lens and useful insights into ecological processes underlying nestedness, and further improves our knowledge of how local plant communities are assembled.

Species coculture can increase agro-biodiversity and therefore constitutes an ecological intensification measure for agriculture. Rice-aquatic animal coculture, one type of species coculture, has been practiced and researched widely. Here, we review recent studies and present results of a quantitative analysis of literature on rice-aquatic animal coculture systems. We address three questions: (i) can rice yield and soil fertility be maintained or increased with less chemical input through rice-aquatic animal coculture? (ii) how do aquatic animals benefit the paddy ecosystem? (iii) how can coculture be implemented for ecological intensification? Meta-analysis based on published papers showed that rice-aquatic animal cocultures increased rice yield, soil organic carbon and total nitrogen and decreased insect pests and weeds compared with rice monocultures. Studies also showed that rice-aquatic animal cocultures reduced pesticide and fertilizer application compared with rice monocultures. Rice plants provide a beneficial environment for aquatic animals, leading to high animal activities in the field. Aquatic animals, in turn, help remove rice pests and act as ecological engineers that affect soil conditions, which favor the growth of rice plants. Aquatic animals promote nutrient cycling and the complementary use of nutrients between rice and aquatic animals, which enhances nutrient-use efficiency in the coculture. To generate beneficial outcomes, how to develop compatible partnerships between rice and aquatic animals, and compatible culturing strategies for coculture systems are the key points. Investigating which traits of aquatic animals and rice varieties could best match to create productive and sustainable coculture systems could be one of the future focuses.

Tropical forests are characterized by vast biomass, complex structures and mega-biodiversity. However, the adaptation processes of these forests to seasonal water availability are less understood, especially those located in the monsoonal and mountainous regions of tropical Southeast Asia. This study used four representative tropical forests spanning from 2° N to 22° N in continental Southeast Asia to address dry-condition photosynthesis at the seasonal scale. We first provided novel and reliable estimations of ecosystem photosynthesis (gross primary production; GPP) seasonality at all four sites. As expected, both evergreen and deciduous seasonal forests exhibited higher GPPs during the rainy season than during the dry season. A bimodal pattern corresponding to solar radiation occurred in the GPP of the perhumid forest. The surface conductance (G_s) was consistently lower both in the dry season and during dry spells (DSPs) than during the wet season and non-dry spells. However, this did not prevent GPP from increasing alongside increasing irradiance in the perhumid forest, suggesting that other ecosystem physiological properties, for example, the light-saturated photosynthetic rate, must have increased, thus surpassing the effect of G_s reduction. Thus, perhumid forests could be defined as light-demanding ecosystems with regard to their seasonal dynamics. Seasonal forests are water-stressed ecosystems in the dry season, as shown by the reductions in GPP, G_s and related ecosystem physiological properties. At all four forest sites, we observed a lack of consistent adaptive strategy to fit the water seasonality due to the diversity in leaf phenology, soil nutrient availability, root depth and other potential factors.

The carbon and water cycle, an important biophysical process of terrestrial ecosystems, is changed by anthropogenic revegetation in arid and semiarid areas. However, there is still a lack of understanding of the mechanisms of carbon and water coupling in intrinsic ecosystems in the context of human activities. Based on the CO₂ and H₂O flux measurements of the desert steppe with the planted shrub Caragana liouana, this study explored the carbon and water flux coupling of the ecosystem by analyzing the variations in gross primary productivity (GPP), evapotranspiration (ET) and water use efficiency (WUE) and discussing the driving mechanisms of biological factors. The seasonal variation in climate factors induced a periodic variation pattern of biophysical traits and carbon and water fluxes. The GPP and ET fluctuated in seasons, but the WUE was relatively stable in the growing season. The GPP, ET and WUE were significantly driven by global radiation (R_g), temperature (T_a and T_s), water vapor pressure deficit, leaf area index and plant water stress index (PWSI). However, R_g, temperature and PWSI were the most important factors regulating WUE. R_g and temperature directly affected WUE with a positive effect but indirectly inhibited WUE by rising PWSI. Plant water stress inhibited photosynthesis and transpiration of the planted shrub community in the desert steppe. When the plant water stress exceeded a threshold (PWSI >0.54), the WUE would decrease since the GPP responded more quickly to the plant water stress than ET. Our findings suggest that policies related to large-scale carbon sequestration initiatives under afforestation must first fully consider the status of water consumption and WUE.

Given the current scenario of climate change and anthropogenic impacts, spatial predictions of biodiversity are fundamental to support conservation and restoration actions. Here, we compared different stacked species distribution models (S-SDMs) to forest inventories to assess if S-SDMs capture emerging properties and geographic patterns of species richness and composition of local communities in a biodiversity hotspot. We generated SDMs for 1499 tree species sampled in 151 sites across the Atlantic Forest. We applied four model stacking approaches to reconstruct the plant communities: binary SDMs (bS-SDMs), binary SDMs cropped by minimum convex polygons (bS-SDMs-CROP), stacked SDMs constrained by the observed species richness (cS-SDMs) and minimum convex polygons of species occurrences (MCPs). We compared the stacking methods with local communities in terms of species richness, composition, community prediction metrics and components of beta diversity—nestedness and turnover. S-SDMs captured general patterns, with bS-SDMs-CROP being the most parsimonious approach. Species composition differed between local communities and all stacking methods, in which bS-SDMs, bS-SDMs-CROP and MCPs followed a nested pattern, whereas species turnover was most important in cS-SDMs. S-SDMs varied in terms of performance, omission and commission errors, leading to a misprediction of some vulnerable, endangered and critically endangered species. Despite differing from forest inventory data, S-SDMs captured part of the variation from local communities, representing the potential species pool. Our results support the use of S-SDMs to endorse biodiversity synthesis and conservation planning at coarse scales and warn of potential misprediction at local scales in megadiverse regions.

Plant density and nitrogen (N) availability influence plant survival and nutrient use strategies, but the interaction between these two factors for plant growth and the balance of elements remains poorly addressed. Here, we conducted experimental manipulations using Arabidopsis thaliana, with the combination of four levels of plant density and four levels of N addition, and then examined the corresponding changes in plant biomass production (indicated by total plant biomass and biomass partitioning) and nutrient use strategies (indicated by leaf N and phosphorus (P) stoichiometry). The biomass-density relationship was regulated by N availability, with a negative pattern in low N availability but an asymptotic constant final yield pattern at high N availability. Excessive N addition reduced plant growth at low plant density, but this effect was alleviated by increasing plant density. The root to shoot biomass ratio increased with plant density at low N availability, but decreased at high N availability. N availability was more important than plant density in regulating leaf N and P stoichiometry, with the increasing leaf N concentration and decreasing leaf P concentration under increasing N addition, resulting in a negative scaling relationship between these two elemental concentrations. Our results show that N availability and plant density interactively regulate plant biomass production and leaf stoichiometry of A. thaliana, and highlight that the interactive effects of these two factors should be considered when predicting plant growth behaviour under intraspecific competitive environments in the context of nutrient changes.

Understanding large-scale patterns of biodiversity and their drivers remains central in ecology. Many hypotheses have been proposed, including hydrothermal dynamic hypothesis, tropical niche conservatism hypothesis, Janzen’s hypothesis and a combination model containing energy, water, seasonality and habitat heterogeneity. Yet, their relative contributions to groups with different lifeforms and range sizes remain controversial, which have limited our ability to understand the general mechanisms underlying species richness patterns. Here we evaluated how lifeforms and species range sizes influenced the relative contributions of these three hypotheses to species richness patterns of a tropical family Moraceae. The distribution data of Moraceae species at a spatial resolution of 50 km × 50 km and their lifeforms (i.e. shrubs, small trees and large trees) were compiled. The species richness patterns were estimated for the entire family, different life forms and species with different range sizes separately. The effects of environmental variables on species richness were analyzed, and relative contributions of different hypotheses were evaluated across life forms and species range size groups. The species richness patterns were consistent across different species groups and the species richness was the highest in Sichuan, Guangzhou and Hainan provinces, making these provinces the hotspots of this family. Climate seasonality is the primary factor in determining richness variation of Moraceae. The best combination model gave the largest explanatory power for Moraceae species richness across each group of range size and life forms followed by the hydrothermal dynamic hypothesis, Janzen’s hypothesis and tropical niche conservatism hypothesis. All these models has a large shared effects but a low independent effect (< 5%), except rare species. These findings suggest unique patterns and mechanisms underlying rare species richness and provide a theoretical basis for protection of the Moraceae species in China.

Previous studies showed that moss stoichiometric characteristics were influenced by moss patch size, shrubs and the environment in the desert. The study of moss stoichiometry in different spatial distribution areas is crucial for an understanding of growth and adaptation strategy of the mosses. In this study, the dominant moss (Syntrichia caninervis) of biological soil crusts and soil under the moss patches in the Gurbantunggut Desert were selected to determine their stoichiometry in different dunes and sites. Moss stoichiometry and soil available nutrients were significantly influenced by different distribution areas except for moss C. The N_aboveground vs. N_belowground' P_aboveground vs. P_belowground and K_aboveground vs. K_belowground scaling exponents of moss were 0.251, 0.389 and 0.442, respectively. The N vs. P scaling exponents were 0.71, 0.84 in above- and below-ground parts of moss. Moss stoichiometry was disproportionately distributed in the above-ground and below-ground parts. Moreover, moss N, P and K elements were influenced by mean annual precipitation (MAP), longitude and soil nutrients. The nutrients of moss were affected by spatial distribution, mean annual temperature (MAT), MAP and soil nutrients. The growth of moss was limited by N element in the temperate desert. This study provides the stoichiometric characteristics of C, N, P and K of moss at different spatial scales and explores their relationships with environmental variables, which can help understand nutrient patterns and utilization strategy of N, P and K, and their potential responses to global climate changes in desert.

Dominant species may strongly influence biotic conditions and interact with other species, and thus are important drivers of community dynamics and ecosystem functioning, particularly in the stressed environment of alpine grasslands. However, the effects of dominant species on the community stability of different ecosystems remain poorly understood. We examined the mechanisms underlying temporal stability (2014-2020 year) of aboveground productivity and community stability in four alpine grasslands (alpine meadow, alpine meadow steppe, alpine steppe and alpine desert steppe) of the northern Tibetan with different species composition and dominance. Our results showed that community stability was significantly higher in the alpine meadow than in the other three types of grasslands. This difference was mainly attributed to the higher compensatory effect and selection effect in the alpine meadows. Furthermore, dominant species strongly affected community stability by increasing dominant species stability and species asynchrony. However, species richness had little effect on community stability. Our findings demonstrate that dominant species, as foundation species, may play leading roles in shaping community stability in the alpine grasslands, highlighting the importance of conserving dominant species for stable ecosystem functioning in these fragile ecosystems under increasing environmental fluctuations.

Leaf size varies conspicuously within and among species under different environments. However, it is unclear how leaf size would change with elevation, whether there is a general elevational pattern, and what determines the altitudinal variation of leaf size. We thus aimed to address these questions by focusing on the broad-leaved herbaceous species at high altitudes on the northeastern Qinghai-Tibetan Plateau. We measured the leaf size, leaf length, leaf width and leaf mass per area for 39 broad-leaved herbaceous species inhabited in the open areas along two mountain slopes from 3200 to 4400 m at the Lenglongling and the Daban Mountain, the northeastern Qinghai-Tibetan Plateau. We analyzed the altitudinal patterns in leaf size in relation to leaf inclination and leaf surface features, and applied a leaf energy balance model to discuss the underlying mechanisms. Leaf size decreased significantly at higher altitudes. The altitudinal reduction of leaf size was mainly attributed to the reduction of leaf length, and differed in different species, and in leaves with different inclination and leaf surface features. A leaf energy balance model with local environmental measurements demonstrates that leaf temperature tracks air temperature more closely in small than in large leaves, and that the leaf-size impact is stronger at higher latitudes. Based on the observational findings, we propose that the distribution upper-limit for broad-leaved herbaceous species would be at an elevation of about 5400 m on the northeastern Qinghai-Tibetan Plateau.

Nutrient resorption is a crucial mechanism for plant nutrient conservation, but most previous studies did not consider the leaf-mass loss during senescence due to lack of measured data. This would lead to an underestimation of nutrient resorption efficiency (NuRE), or calculating NuRE of various species based on the average mass loss at plant-functional-group level in the literature, thus affecting its accuracy. Here we measured the leaf-mass loss to correct NuRE with the species-specific mass loss correction factor (MLCF), so as to foster a more accurate calculation of the nutrient fluxes within and between plants and the soil. Green leaves and senesced leaves were collected from 35 dominant woody plants in northern China. Mass of green and senesced leaves were measured to calculate the MLCF at species level. The MLCF was reported for each of the 35 dominant woody plants in northern China. These species averagely lost 17% of the green-leaf mass during leaf senescence, but varied greatly from 1.3% to 36.8% mass loss across the 35 species, or 11.7% to 19.6% loss across the functional types. Accordingly, the MLCF varied from 0.632 to 0.987 across the 35 species with an average value 0.832. The NuRE corrected with MLCF was remarkably increased on the whole (e.g. both the average nitrogen and phosphorus NuRE became about 9% higher, or more accurate), compared with the uncorrected ones, especially in the case of low resorption efficiencies. Our field data provide reliable references for the MLCF of plants in related regions at both species and functional-type levels, and are expected to promote more accurate calculations of NuRE.

The Tibetan Plateau is generally referred to as the Chinese water tower, and evapotranspiration (ET) affects the water budget and stability of alpine meadows on the Tibetan Plateau. However, its variability and controlling mechanisms have not been well documented under the drier conditions induced by global warming. Therefore, this study aimed to clarify whether meteorological or biological factors primarily affected the variability in ET under contrasting water conditions in the alpine meadow ecosystem on the Tibetan Plateau. Based on 6-year (2013–2018) eddy covariance observations and the corresponding meteorological and biological data, linear perturbation analyses were employed to isolate the contributions of meteorological and biological factors to the variability in evapotranspiration (δET). The results showed that δET was mainly driven by meteorological factors in wet peak seasons (July and August), and was dominated by net radiation (Rn) and air temperature (Ta), indicating that the inadequate available energy is the factor limiting ET. However, the dominant factors affecting δET shifted from meteorological to biological in dry peak seasons when the canopy stomatal conductance (gs) and leaf area index were dominant. At this point, the ecosystem was limited by the water conditions. These results provide empirical insights into how meteorological and biological factors regulate variability in ET under contrasting water conditions. These findings can further improve our understanding of water cycle processes and can help effectively manage water resources in alpine meadow ecosystems under future climate change conditions.

The ecosystem apparent quantum yield (α), maximum rate of gross CO₂ assimilation (P_max) and daytime ecosystem respiration rate (R_d), reflecting the physiological functioning of ecosystem, are vital photosynthetic parameters for the estimation of ecosystem carbon budget. Climatic drivers may affect photosynthetic parameters both directly and indirectly by altering the response of vegetation. However, the relative contribution and regulation pathway of environmental and physiological controls remain unclear, especially in semi-arid grasslands. We analyzed seasonal and interannual variations of photosynthetic parameters derived from eddy-covariance observation in a typical semi-arid grassland in Inner Mongolia, Northern China, over 12 years from 2006 to 2017. Regression analyses and a structural equation model (SEM) were adopted to separate the contributions of environmental and physiological effects. The photosynthetic parameters showed unimodal seasonal patterns and significantly interannual variations. Variations of air temperature (T_a) and soil water content (SWC) drove the seasonal patterns of photosynthetic parameters, while SWC predominated their interannual variations. Moreover, contrasting with the predominant roles of T_a on α and R_d, SWC explained more variance of P_max than T_a. Results of SEM revealed that environmental factors impacted photosynthetic parameters both directly and indirectly through regulating physiological responses reflected by stomatal conductance at the canopy level. Moreover, leaf area index (LAI) directly affected α, P_max and R_d and dominated the variation of P_max. On the other hand, SWC influenced photosynthetic parameters indirectly through LAI and canopy surface conductance (gc). Our findings highlight the importance of physiological regulation on the photosynthetic parameters and carbon assimilation capacity, especially in water-limited grassland ecosystems.

It has been suggested that the importance of network architecture to species diversity and stability should be based on preference networks (comprised of niche differentiations), rather than observational networks, because species abundance may significantly affect interaction frequencies. Considering that resource abundance is usually greater for herbivores than parasites, we hypothesize that the abundance effect is stronger for parasitic than herbivory interactions. To test this hypothesis, we collected 80 quantitative observational networks including 34 herbivorous and 46 parasitic networks from the published literature, and derived preference networks by removing the effects of species abundance. We then determined the network nestedness using both weighted NODF and spectral radius. We also determined species degree distribution, interaction evenness, weighted connectance and robustness for both observational and preference networks. The observational networks (including both herbivory and parasitic networks) were more nested judged by weighted NODF than spectral radius. Preference networks were less nested for parasitic than herbivory networks in terms of both weighted NODF and spectral radius, possibly because removing the abundance effect increased interaction evenness. These trends indicate that the abundance effect on network nestedness is stronger for parasitic than herbivory networks. Weighted connectance and robustness were greater in most preference networks than observational networks, indicating that preference networks may have high network stability and community persistence compared with observational ones. The data indicate that future network analyses should not only address the structural difference between mutualistic and antagonistic interactions, but also between herbivory and parasitic interactions.

Alpine forests in the eastern Tibetan Plateau are important ecological barriers in the upper reaches of the Yangtze River. However, due to continuous high-intensity harvesting, a large number of plantings, and the complete harvesting ban measures in recent decades, the forest tree species and age cohorts have become relatively homogenous, and the biodiversity and ecological functions have been reduced. To design effective forest management options to optimize forest structure and increase carbon sequestration capacity, Mao County in Sichuan Province was selected as the study site and six forest management options (harvesting, planting) of different intensities were tested using the LANDIS-II model to simulate and compare the differences in forest aboveground carbon sequestration rate (ACSR) between these options and the current management option over the next 100 years. Our results showed that (i) the different harvesting and planting intensities significantly changed the ACSR compared with the current management options; (ii) different communities responded differently to the management options, with the ACSR differing significantly in cold temperate conifers and temperate conifers but not in broad-leaved trees (P < 0.05); and (iii) a comprehensive consideration of forest management options at the species, community and landscape levels was necessary. Our results suggest that implementing a longer harvesting and planting interval (20 years) at the study site can maximize forest ACSR. This study provides an important reference for evaluating the ability of forest management options to restore forest ecological functions and increase carbon sequestration capacity and for selecting effective forest management programs in the eastern Tibetan Plateau.

Rodents attack oak (Quercus wutaishanica) seeds based on their sizes and frequencies before germination. However, the predation of oak seeds post-germination (seedling cotyledons) is not well studied. Here, we not only tested the preference of rodents for Q. wutaishanica seedling cotyledons based on the frequency of large- versus small-seeded (FLS), but also evaluated the effects of predation on seedlings growth and survival in different habitats. We transplanted seedlings with the FLS set as 9:1, 7:3, 5:5, 3:7 and 1:9, respectively, in the forest gap and under the canopy in the Liupan Mountains National Nature Reserve in Ningxia Hui Autonomous Region, Northwest China. The results showed that: (i) in 1-7 days after transplanting seedlings, rodents prefer the cotyledon of large-seeded seedings while small-seeded seedlings were preferred in 8-60 days, and the positive frequency-dependent predation was observed. (ii) The cotyledons were preyed on, the apical buds were bitted off, and the whole seedlings were uprooted, which mostly occurred under the forest canopy. At the end of a growing season, the survival rate of seedlings in the forest gaps was more than twice that under forest canopies. (iii) If cotyledons were preyed on, the growth of Q. wutaishanica seedlings would not be affected, but the seedlings growth was severely inhibited when the apical bud was bitten off. These results not only provide new insights into the coexistence between rodents and seedlings of different phenotypes, but also reveal the ecological characteristics of deciduous Quercus regeneration.

In Mongolia, overgrazing and the resulting degradation of rangelands are recognized as serious issues. To address rangeland degradation, we sought to develop a broad-scale vegetation classification of Mongolian rangeland communities focusing on regional characteristics. Moreover, we sought to clarify the spatial distributions of communities and the environmental drivers of the distributions. Between 2012 and 2016, we surveyed vegetation in 278 plots (each 10 m × 10 m) in different regions of Mongolia (43-50° N, 87-119° E) in plots where grazing pressure is low relative to adjacent areas. The data were grouped into vegetation units using a modified two-way indicator species analysis (TWINSPAN). We then explored the regional characteristics of species compositions and community distributions, as well as relationships between distributions and climatic variables. The modified TWINSPAN classified the vegetation data into three cluster groups, each of which corresponds to a particular type of zonal vegetation (i.e. forest steppe, steppe and desert steppe). The aridity index was identified as an important driver of the distributions of all cluster groups, whereas longitude and elevation were important determinants of the distribution of clusters within cluster groups. Western regions, which are characterized by higher elevation and continentality compared with eastern regions, have lower mean temperature and precipitation during the wettest quarter, leading to differences in species composition within cluster groups. Regional differences in species composition reflect differences in phytogeographic origin. Thus, the framework of species composition and distributional patterns in Mongolian rangeland communities was demonstrated in relation to climatic and geographical factors.

Plants are frequently exposed to adverse environments during their life span. Among them drought stress is one of the major threats to agricultural productivity. In order to survive in such unstable environment, plants have developed mechanisms through which they recognize the severity of the stress based on the incoming environmental stimuli. To combat the detrimental effects of drought, the plants have evolved various strategies to modulate their physio-hormonal attributes. These strategies that can be modulated by shade and microbes contribute to enhancing tolerance to drought and reducing yield loss. Plant hormones, such as abscisic acid, auxin and ethylene have a major role in the shade- and microbe-associated improvement of drought tolerance through their effects on various metabolic pathways. In this process, the CLAVATA3/EMBRYOSURROUNDING REGION-RELATED 25 peptide has a major role due to its effect on ABA synthesis as shown in our regulatory model.

Wetlands store large amounts of carbon stocks and are essential in both global carbon cycling and regional ecosystem services. Understanding the dynamics of wetland carbon exchange is crucial for assessing carbon budgets and predicting their future evolution. Although many studies have been conducted on the effects of climate change on the ecosystem carbon cycle, little is known regarding carbon emissions from the alpine wetlands in arid northwest China. In this study, we used an automatic chamber system (LI-8100A) to measure ecosystem respiration (ER) in the Bayinbuluk alpine wetland in northwest China. The ER showed a significant bimodal diurnal variation, with peak values appearing at 16:30 and 23:30 (Beijing time, UTC + 8). A clear seasonal pattern in ER was observed, with the highest value (19.38 µmol m⁻² s⁻¹) occurring in August and the lowest value (0.11 µmol m⁻² s⁻¹) occurring in late December. The annual ER in 2018 was 678 g C m⁻² and respiration during the non-growing season accounted for 13% of the annual sum. Nonlinear regression revealed that soil temperature at 5 cm depth and soil water content (SWC) were the main factors controlling the seasonal variation in ER. The diurnal variation in ER was mainly controlled by air temperature and solar radiation. Higher temperature sensitivity (Q₁₀) occurred under conditions of lower soil temperatures and medium SWC (25% ≤ SWC ≤ 40%). The present study deepens our understanding of CO₂ emissions in alpine wetland ecosystems and helps evaluate the carbon budget in alpine wetlands in arid regions.

Soil microorganisms in tropical forests can adapt to phosphorus (P)-poor conditions by changing the activity ratios of different types of phosphatases. We tested whether microorganisms in P-poor tropical forest soils increased the phosphomonoesterase (PME) to phosphodiesterase (PDE) activity ratio, because a one-step enzymatic reaction of monoester P degradation might be more adaptive for microbial P acquisition than a two-step reaction of diester P degradation. A continuous 10-year P addition experiment was performed in three tropical forests. The activities of PME and PDE, and their ratio in soil, were determined under the hypothesis that the P-fertilized plots where P shortage is relieved would have lower PME:PDE ratios than the unfertilized controls. We demonstrated that long-term P addition in tropical forest soil did not alter the PME:PDE ratio in primary and secondary forests, whereas P fertilization elevated the PME:PDE ratio in planted forest. These results were in contrast to previous results. The long-term, large-scale P fertilization in our study may have reduced litter- and/or throughfall-derived PDE, which negated the lowered PME:PDE ratio via exogenous P inputs.

Previous studies have shown that plant-pollinator mutualistic interactions experience highly interannual variation. Given that pollinators often move across multiple plant species, the plant-plant interactions that take place via heterospecific pollen (HP) transfer may also vary temporally, which could have important implications for floral evolution and community assembly. Here, we evaluated the interannual variation in plant-pollinator networks and plant-plant heterospecific pollen transfer (HPT) networks of a subalpine meadow community in Southwest China for three consecutive years. The interactions largely varied among years for both network types. The composition of donor-species HP deposited on the plants varied less than did the visit composition of the pollinators, suggesting that HP could be transferred from identical donor species to recipient species through different shared pollinators among years. The plant species were at more similar positions in the HPT network than they were in the plant-pollinator network across years. Moreover, the more generalized plant species in the plant-pollinator network tended to export their pollen grains and more strongly influence HPT. We evaluated the relatively stable structure of the HPT network compared with the plant-pollinator network, which represents an important step in the integration of plant-pollinator and plant-plant interactions.

Land use and nutrient enrichment can substantially affect biodiversity and ecosystem functioning. However, whether and how the responses of community temporal stability to land use and nutrient enrichment change with time remain poorly understood. As part of a 15-year (2005-2019) field experiment, this study was conducted to explore the effects of mowing, nitrogen (N) and phosphorus (P) additions on community temporal stability in a temperate steppe on the Mongolian Plateau. Over the 15 years, N and P additions decreased community temporal stability by reducing the population stability, especially the shrub and semi-shrub stability. However, mowing increased community temporal stability in the early stage (2005-2009) only. Nitrogen addition suppressed community temporal stability in the early and late (2015-2019) stages, whereas enhanced it in the intermediate stage (2010-2014). Phosphorus addition decreased community temporal stability marginally in the early stage and significantly in the late stage. The fluctuations of N-induced changes in community temporal stability are mainly explained by its diverse effects on species asynchrony and population stability over time. Our findings highlight the important role of plant functional groups and species asynchrony in regulating community temporal stability, suggesting that more long-term studies are needed to accurately forecast ecosystem response patterns in the context of global change.

All organisms need elements in fixed proportions for carrying out normal metabolic processes and how flexible they are depends on how effective they are utilizing these resources from external sources. It is important to understand the interactions among plant, soil and microbial biomass carbon (C), nitrogen (N) and phosphorus (P) stoichiometry under different conditions of resource supply. We conducted a pot experiment on 1-year-old Robinia pseudoacacia seedlings for nearly 5 months under different water, nitrogen and phosphorus supplies, and we determined plant, soil and microbial biomass C, N and P stoichiometry. We found that plant, soil and microbial nutrients and stoichiometry exhibited a certain degree of plasticity in response to the changes in water and nutrient conditions in their environments. Variation partitioning analysis showed that root stoichiometry accounted for a large part of the variance in microbial stoichiometry. Structural equation modeling further revealed that root stoichiometry and leaf stoichiometry were two direct factors affecting microbial biomass C:N and C:P, and that root stoichiometry had the greatest direct effect. In addition, the degree of homeostasis for microbial biomass C and C:P was more sensitive to changes in soil nutrients than changes in other factors, and other elements and elemental ratios displayed strict homeostasis. These results highlight the importance of studying microbial stoichiometry in improving our understanding of nutrient cycling of the plant–soil system under different water and nutrient supply.

Although increases in precipitation variability in arid ecosystems are projected due to climate change, the response of desert shrub communities to precipitation change has not been fully elucidated. Such knowledge is important since drought-adapted plants exhibit varied mechanisms of survival that may contribute to species coexistence.

We tested the responses of eight drought-adapted plants, a mix of graminoids, shrubs and forbs to three summer precipitation scenarios (1.3, 2.6 and 3.9 cm per month) in a common garden experiment in the Great Basin (Owens Valley, California). Changes in mineral nutrient uptake (carbon, nitrogen, phosphorus, potassium, calcium, magnesium, manganese, copper, boron, zinc, iron and sodium) and gas exchange parameters (photosynthetic rate and stomatal conductance) were investigated in the studied species.

Two graminoids (Sporobolus airoides and Leymus triticoides) and one salt tolerant shrub species (Atriplex confertifolia) responded to increased water availability with increases in photosynthetic rate and/or stomatal conductance. There was a significant correlation between water availability and uptake of nutrients for five out of eight species. Artemisia tridentata, with higher rates of photosynthesis, contained greater amounts of potassium, copper and boron, while Juncus arcticus, with higher rates of photosynthesis, contained greater amounts of magnesium and iron, and less sodium. Juncus arcticus and three salt-adapted species (A. confertifolia, Distichlis spicata and S. airoides) exhibited correlations with stomatal conductance and concentrations of nutrients. Results indicate that differential physiological response mechanisms to increased moisture and associated nutrient uptake strategies in drought-adapted species may mediate coexistence under increased summer precipitation.

The alpine meadow ecosystem in Tibet is fragile and sensitive, and its carbon sink function with respect to climate change has become a matter of widespread concern. Therefore, this study aims to clarify the inter-annual variations (IAVs) in the carbon fluxes in an alpine meadow and to further quantify the contributions of the driving factors to the IAVs. Based on 7 years of flux data (2012–2018) and the corresponding climatic and biotic data, a set of look-up tables was used to separate and quantify the IAV sources. Furthermore, linear perturbation analyses were employed to quantify the contributions of each key factor. During 2012–2018, the net ecosystem productivity (NEP), gross primary productivity (GPP) and ecosystem respiration (Re) of this alpine meadow were 3.31 ± 26.90, 210.18 ± 48.35 and 206.88 ± 28.45 g C m⁻² y⁻¹, respectively, which indicated relatively large IAVs. When the contributions of climatic and biotic effects were distinguished and quantified, the dominant effects of biotic factors emerged. Additionally, negative interactions between climatic and biotic effects were detected. Among the climatic factors, only soil water content contributed relatively more to the IAVs and played a role in regulating the interactions between climatic and biotic effects. These results suggest that biotic effects must be carefully considered to reduce the uncertainties associated with future carbon flux estimates.

Plant litter decomposition is critical for the carbon (C) balance and nutrient turnover in terrestrial ecosystems, and is sensitive to the ongoing anthropogenic biologically nitrogen (N) input. Previous studies evaluating the N effect on litter decomposition relied mostly on short-term experiments (<2 years), which may mask the real N effect on litter decomposition. Therefore, long-lasting experiments are imperative for the overall evaluation of the litter decomposition dynamics under N enrichment. We conducted a relative long-term (4-year) N-addition experiment with N levels ranging from 0 to 50 g N m^-2 yr^-1 to identify the potential abiotic and biotic factors in regulating the decomposition process of litterfall from the dominant species Leymus chinensis. The results showed a consistent decrease of decomposition rate with increasing N-addition rates, providing strong evidence in support of the inhibitory effect of N addition on decomposition. The N-induced alterations in soil environment (acidification and nutrient stoichiometry), microbial activity (microbial biomass and enzyme activity), changes of litter quality (residual lignin and nutrient content) and plant community (aboveground productivity and species richness) jointly contributed to the lowered decomposition. During the whole decomposition process, the changes of litter quality, including accumulation of lignin and the concentrations of nutrient, were mainly driven by the soil and microbial activity in this N-enriched environment. The findings help clarify how increasing N input rates affect long-term litter decomposition, and advance the mechanistic understanding of the linkages between ecosystem N enrichment and terrestrial C cycling.

The reproductive strategies of alpine plants are often altered by environmental changes caused by changes in the spatial distribution of the gradient. However, few studies have investigated whether reproductive patterns of the same species vary with elevation. Three natural populations of Primula atrodentata, which are distributed in the eastern Himalayas and have a long flowering period, were selected along the elevation gradients in Shergyla Mountain, Tibet, China. Morph ratio investigation, floral trait measurement, pollinator observation and manipulated pollination experiments were conducted to explore the changes in self-compatibility and floral traits associated with the selfing syndrome along elevation gradients. We found that the breeding system of the S-morph is facultative outcrossing, and that of the L-morph is obligatory outcrossing. We further found that with increasing elevation, the number of pollen and ovules, anther-stigma distance, and inbreeding depression index first increased and then decreased, whereas the seeds per fruit and seed-setting rate under hand self-pollination, pollen limitation and self-incompatibility index tended to decrease first, but then increased. In addition, pollinator diversity and visiting frequency were the highest at the middle elevation (4050 population), which can better explain the nonlinear change in self-fertility with elevation. Our findings provide insights into the evolutionary pattern of self-compatibility in alpine plants along elevational gradients.

Invasive plants are a major threat to biodiversity and may adversely affect food security. Clonal integration enables the sharing of resources between connected ramets and can enhance plant performance in many invasive species. However, few studies have examined the role of clonal integration when weeds are exposed to plant growth regulators (PGRs). PGRs are used extensively in agriculture and may affect nearby weeds through soil leaching, erosion and runoff. Our aim was to investigate the effects of clonal integration on growth in a noxious weed, Alternanthera philoxeroides (alligator weed), in response to two PGRs frequently used in agriculture, gibberellins (GAs) and paclobutrazol (PAC). Ramets of A. philoxeroides were propagated in the greenhouse, and treated with PGRs. PGRs were applied to the older ramets (i.e. ‘basal’ part), with half of the plants having the stems between the apical (younger) and basal parts left connected, while the remaining plants had the stems between the two parts severed. Following the growing period, plants were measured for growth traits. We found that GA and PAC had contrasting effects on plant growth. GA significantly promoted above-ground growth of the apical ramets via clonal integration. Alternatively, PAC inhibited above-ground growth in the basal and apical parts, and enhanced below-ground growth of the basal and apical ramets through clonal integration. Our results highlight how clonal integration can promote growth in A. philoxeroides following the application of PGRs, which is likely an important mechanism for this species to invade new environments.

Evergreen and deciduous species coexist in the subtropical forests in southeastern China. It has been suggested that phosphorus (P) is the main limiting nutrient in subtropical forests, and that evergreen and deciduous species adopt different carbon capture strategies to deal with this limitation. However, these hypotheses have not been examined empirically to a sufficient degree. In order to fill this knowledge gap, we measured leaf photosynthetic and respiration rates, and nutrient traits related to P-, nitrogen (N)- and carbon (C)-use efficiencies and resorption using 75 woody species (44 evergreen and 31 deciduous species) sampled in a subtropical forest. The photosynthetic N-use efficiency (PNUE), respiration rate per unit N and P (R_d,N and R_d,P, respectively) of the deciduous species were all significantly higher than those of evergreen species, but not in the case of photosynthetic P-use efficiency. These results indicate that, for any given leaf P, evergreen species manifest higher carbon-use efficiency (CUE) than deciduous species, a speculation that is empirically confirmed. In addition, no significant differences were observed between deciduous and evergreen species for nitrogen resorption efficiency, phosphorus resorption efficiency or N:P ratios. These results indicate that evergreen species coexist with deciduous species and maintain dominance in P-limited subtropical forests by maintaining CUE. Our results also indicate that it is important to compare the PNUE of deciduous species with evergreen species in other biomes. These observations provide insights into modeling community dynamics in subtropical forests, particularly in light of future climate change.

Variations in plant traits are indicative of plant adaptations to forest environments, and studying their relationships with tree growth provides valuable insights into forest regeneration. The spatial arrangement of plant seeds within the forest litter or soil critically influences the variations of root-leaf traits, thereby affecting the adaptive strategies of emerging seedlings. However, our current understanding of the impacts of individual root-leaf traits on seedling growth in different relative position, and whether these traits together affect growth, remains limited. This study focuses on the dominant tree species, Castanopsis kawakamii, within the Sanming C. kawakamii Nature Reserve of China. The present experiment aimed to examine the variations in root-leaf traits of seedling, focus on the relative positions of seeds within different layers: beneath or above the litter layer, or within the bare soil layer (without litter). Our findings provided evidence supporting a coordinated relationship between root and leaf traits, wherein leaf traits varied in conjunction with root traits in the relative positions of seeds. Specifically, we observed that seedlings exhibited higher values for specific leaf area and average root diameter, while displaying lower root tissue density. The mixed model explained 86.1% of the variation in root-leaf traits, surpassing the variation explained by the relative positions. Furthermore, soil nitrogen acted as a mediator, regulating the relationship between seedling growth and root-leaf traits, specifically leaf dry matter content and root tissue density. Therefore, future studies should consider artificially manipulating tree species diversity based on root-leaf traits characteristics to promote forest recovery.

Widely accepted universal models and hypotheses such as ‘high vein density-faster growth and higher productivity' hold that high leaf vein density may promote higher coupling efficiency of carbon and water, indicating that rapid individual growth and high stand productivity, have attracted huge interest. However, these models and hypotheses do not include enough gymnosperm samples, especially conifers cultivated in subtropics. We here examined the values and scaling relationships between leaf vein density and leaf functional traits sampled from center region of the distribution range of Cunninghamia lanceolate, which has been well known for rapid growth. We also retrieved an empirical dataset that included photosynthetic, biochemical, anatomical and hydraulic traits of Cunninghamia lanceolata. The leaf vein density (ranging from 0.34 to 1.09 mm mm-2) is extremely low compared to the reported global range (1 to 25 mm mm-2), whereas C. lanceolata is famous for both fast-growing and high-yielding in China for a long time. We further verified that higher vein densities were associated with smaller leaves (r = -0.71, P < 0.001), which is consistent with that found in angiosperms. However, we found that vein density-thickness correlations and leaf lifespan plasticity showed opposite trends for C. lanceolate (negative) when compared with global species (positive), and such relationships may indicate the tradeoffs between functional efficiency and productivities. Our results provide an effective complementary assessment of general growth rules, including evaluation of the influence of regional plant trait characterization, configuration of plant species, and traits efficiency for hydraulic potential.

Multiple elements are critical for plant growth and survival, community structure and vegetation function. Chemical diversity, defined as the ranges in element concentrations of plant species within communities, could provide essential insights into plant nutrient strategies and community assembly rules. However, little is known about the chemical diversity of multi-elements besides N and P, and current understanding of chemical diversity is largely based on aboveground plant traits. We investigated understory plant communities in forest swamps along a local soil chemical gradient and determined 11 major and trace elements in leaves and roots of dominant and subordinate plants. Using n-dimensional hypervolume, we examined the changes in leaf and root chemical diversity and their linkages with soil properties. Plant chemical diversity decreased significantly with soil Al, Mn, Mg and Zn concentrations, but showed no relationships with soil N, P, K, Na, and Fe concentrations, soil pH and C:N. These patterns also held after controlling for species richness and soil moisture. Furthermore, leaf and root chemical diversity was positively correlated and showed similar relationships with soil factors. Root chemical diversity was not significantly higher than leaf chemical diversity. Our results emphasized the important role of soil trace elements for plant chemical diversity along the local soil chemical gradient. Similar patterns and extent of leaf and root chemical diversity may indicate similar local-scale environmental constraint on aboveand belowground plant chemical diversity. These findings have important implications for plant community assembly and ecosystem functioning influenced by soil nutrient changes.

The four alien farmland weeds of genus Veronica (i.e. V. arvensis, V. didyma, V. hederifolia and V. persica) have successfully colonized in China, but caused different ecological consequences in the colonized habitats. However, the key biological traits conferring bioinvasion differences under different light conditions among the four alien species of Veronica remain unknown. A comprehensive contrastive analysis experiment was conducted to assess the contribution of the intensity of photosynthetic and sexual and asexual reproductive traits of the four alien Veronica weeds to their invasion level in both field trial and laboratory. The field survey showed that V. persica had the highest invasion level, followed by V. didyma, V. hederifolia and V. arvensis. Their invasiveness was mainly attributed to photosynthetic-related parameters (LMA) and asexual reproduction traits (the ratio of adventitious roots) out of all the 22 tested indexes. The photosynthetic-related and some asexual reproduction indexes from separate determinations under both sun and shade conditions showed that V. persica was able to adapt to strong illumination but was more tolerant of shade than the other species. This adaptive differentiation to illumination conferred different competitiveness over crops on the four alien Veronica weeds by allocating resources to the biomass of each organ in farmland. It may be concluded that the adaptability to illumination conditions and the asexual reproduction traits may endow their successful invasion and become different important farmland weeds.

One central challenge for humanity is to mitigate and adapt to an ongoing climate and biodiversity crisis while providing resources to a growing human population. Ecological intensification (EI) aims to maximize crop productivity while minimizing impacts on the environment, especially by using biodiversity to improve ecosystem functions and services. Many EI measures are based on trophic interactions between organisms (e.g. pollination, biocontrol). Here, we investigate how research on multitrophic effects of biodiversity on ecosystem functioning could advance the application of EI measures in agriculture and forestry. We review previous studies and use qualitative analyses of the literature to test how important variables such as landuse parameters or habitat complexity affect multitrophic diversity, ecosystem functions and multitrophic biodiversity-ecosystem functioning relationships. We found that positive effects of biodiversity on ecosystem functions are prevalent in production systems, largely across ecosystem function dimensions, trophic levels, study methodologies and different ecosystem functions, however, with certain context dependencies. We also found strong impacts of land use and management on multitrophic biodiversity and ecosystem functions. We detected knowledge gaps in terms of data from underrepresented geographical areas, production systems, organism groups and functional diversity measurements. Additionally, we identified several aspects that require more attention in the future, such as trade-offs between multiple functions, temporal dynamics, effects of climate change, the spatial scale of the measures and their implementation. This information will be vital to ensure that agricultural and forest landscapes produce resources for humanity sustainably within the environmental limits of the planet.

Biomass allocation to different organs is a fundamental plant ecophysiological process to better respond to changing environments; yet, it remains poorly understood how patterns of biomass allocation respond to nitrogen (N) additions across terrestrial ecosystems worldwide.

We conducted a meta-analysis using 5474 pairwise observations from 333 articles to assess how N addition affected plant biomass and biomass allocation among different organs. We also tested the ‘ratio-based optimal partitioning’ vs. the ‘isometric allocation’ hypotheses to explain potential N addition effects on biomass allocation.

We found that (i) N addition significantly increased whole plant biomass and the biomass of different organs, but decreased root:shoot ratio (RS) and root mass fraction (RMF) while no effects of N addition on leaf mass fraction and stem mass fraction at the global scale; (ii) the effects of N addition on ratio-based biomass allocation were mediated by individual or interactive effects of moderator variables such as experimental conditions, plant functional types, latitudes and rates of N addition and (iii) N addition did not affect allometric relationships among different organs, suggesting that decreases in RS and RMF may result from isometric allocation patterns following increases in whole plant biomass. Despite alteration of ratio-based biomass allocation between root and shoot by N addition, the unaffected allometric scaling relationships among different organs (including root vs. shoot) suggest that plant biomass allocation patterns are more appropriately explained by the isometric allocation hypothesis rather than the optimal partitioning hypothesis. Our findings contribute to better understand N-induced effects on allometric relationships of terrestrial plants, and suggest that these ecophysiological responses should be incorporated into models that aim to predict how terrestrial ecosystems may respond to enhanced N deposition under future global change scenarios.

Fisher discriminant analysis can comprehensively take multiple factors into consideration and effectively conduct separations between two classes. If it can be used to detect the occurrences of drought, drought can be detected more effectively and accurately. Based on 9-year carbon flux and corresponding meteorological data, soil water content (SWC) and vapor pressure deficit (VPD) were selected as the discriminant factors. Drought occurrences were detected by applying the Fisher discriminant analysis method in an alpine ecosystem in Tibet. Fisher discriminant analysis was successfully applied to detect drought occurrence in an alpine meadow ecosystem. The soil water deficit and atmospheric water deficit were comprehensively taken into consideration. Consequently, this method could detect the onset and end date of droughts more accurately and reasonably. Based on the characteristics of drought and non-drought samples, the discriminant equation was constructed as y = 24.46SWC − 4.60VPD. When y > 1, the days were distributed above the critical line. In addition, when y was greater than one for more than 10 days, it was labeled as one drought event. If the interval between two drought processes was less than 2 days, it was considered one drought event. With increasing the study period and continued accumulation of observation data, the discriminant equation could be further optimized in the future, resulting in more accurate drought detection.

Understanding how maximum canopy height is related to forest community assembly is essential yet largely unexplored. Maximum canopy height is affected by competition and abiotic environmental factors through different ecological processes (e.g. niche differentiation and environmental filtering), as well as historical or stochastic factors. However, little has been done to empirically examine the ecological processes that influence maximum canopy height. We set out to examine the relationship between maximum canopy height and community phylogenic structure. We surveyed maximum canopy heights from a regional dataset of forest plots (466 sites of 50 m × 50 m) from the boreal forest of northeastern Alberta, Canada. We then explored the relationships between maximum canopy height as measured by airborne LiDAR (Light Detection and Ranging) and the phylogenetic structure of seed plants, represented by net relatedness index and nearest taxa index. We found stronger phylogenetic clustering among major evolutionary clades for communities with higher maximum canopy height, which implied that environmental filtering by abiotic factors is a driving factor for boreal forests. However, we also found stronger phylogenetic overdispersion within each clade for communities with higher maximum height, indicating more intense niche differentiation. Our results suggest that communities with higher maximum canopy height may have experienced more intense historical abiotic environmental filtering and recent niche differentiation in boreal forests. These findings will contribute to the monitoring and management of forest biodiversity.