Plant height is a key functional trait related to aboveground biomass, leaf photosynthesis and plant fitness. However, large-scale geographical patterns in community-average plant height (CAPH) of woody species and drivers of these patterns across different life forms remain hotly debated. Moreover, whether CAPH could be used as a predictor of ecosystem primary productivity is unknown.

We compiled mature height and distributions of 11 422 woody species in eastern Eurasia, and estimated geographic patterns in CAPH for different taxonomic groups and life forms. Then we evaluated the effects of environmental (including current climate and historical climate change since the Last Glacial Maximum (LGM)) and evolutionary factors on CAPH. Lastly, we compared the predictive power of CAPH on primary productivity with that of LiDAR-derived canopy-height data from a global survey.

Geographic patterns of CAPH and their drivers differed among taxonomic groups and life forms. The strongest predictor for CAPH of all woody species combined, angiosperms, all dicots and deciduous dicots was actual evapotranspiration, while temperature was the strongest predictor for CAPH of monocots and tree, shrub and evergreen dicots, and water availability for gymnosperms. Historical climate change since the LGM had only weak effects on CAPH. No phylogenetic signal was detected in family-wise average height, which was also unrelated to the tested environmental factors. Finally, we found a strong correlation between CAPH and ecosystem primary productivity. Primary productivity showed a weaker relationship with CAPH of the tallest species within a grid cell and no relationship with LiDAR-derived canopy height reported in the global survey. Our findings suggest that current climate rather than historical climate change and evolutionary history determine the geographical patterns in CAPH. However, the relative effects of climatic factors representing environmental energy and water availability on spatial variations of CAPH vary among plant life forms. Moreover, our results also suggest that CAPH can be used as a good predictor of ecosystem primary productivity.

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.

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 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.

Plants typically experience great losses from their reproductive potential represented by ovule production to the post-dispersal crop of viable seed. We examined seed density and viability in a founder population of mountain birch (Betula pubescens ssp. tortuosa), aiming to quantify losses at different stages and examine potential selection forces on the reproduction success of the founder generation of an isolated population. At the time of the study (2017–2020), the population had recently reached reproductive maturity, following its colonization around 1990 through long-distance dispersal onto an early successional outwash plain in southeast Iceland. Seed densities were high, but 89% of apparently intact seeds did not contain an embryo, despite being visually indistinguishable from flled seeds. Externally evident losses amounted to about 45% of the total seed crop and were mostly due to predation by the gall midge Semudobia betulae. When all losses were accounted for, 2.7% of the seed crop remained viable and germinated. Pollen limitation may partially explain the high incidence of empty seeds. Excessive fower production is compatible with the predator satiation hypothesis but cannot explain pre-dispersal losses. Another adaptation to predation, masting, appears poorly developed in Iceland. Our results suggest the presence of constraints on the reproduction potential of the new island population, that are more limiting than in neighbouring populations, and we discuss their developmental, ecological, and environmental correlates.

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.

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.

Phylosymbiosis, the congruence of microbiome composition with host phylogeny, is a valuable framework for investigating plant–microbe associations and their evolutionary ecology. This review assesses the prevalence of phylosymbiosis across the plant kingdom, elucidates the fundamental ecological and evolutionary processes contributing to its occurrence based on previous research and explores commonly used methods for identifying phylosymbiosis. We find that the presence of phylosymbiosis may be influenced by both phylogenetic distance and the taxonomic level at which host plants are examined, with the strength of associations potentially decreasing as the taxonomic scale becomes finer. Notably, the endophytic microbiome exhibits a stronger phylosymbiosis signal compared with the epiphytic or rhizosphere-associated microbiomes. Microorganisms such as fungi and bacteria can yield highly variable evidence for phylosymbiosis due to differences in colonization, transmission or functional characteristics. We also outline how the four community assembly processes (dispersal, selection, diversification and drift) contribute to the establishment and maintenance of host–microbe phylosymbiosis. Furthermore, we highlight the diversity of methods employed to detect phylosymbiosis, which involves three key processes: constructing host phylogenies, assessing microbial data and statistically evaluating the correlation between host phylogeny and microbial composition. Remarkably different methodologies across studies make comparisons between findings challenging. To advance our understanding, future research is expected to explore phylosymbiosis at lower taxonomic levels and investigate different microbial communities coexisting synergistically within the same host. Understanding the relative importance of community assembly processes in driving phylosymbiosis will be critical for gaining deeper insights into the ecology and evolution of host–microbe interactions.

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.

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.

Knowledge concerning the processes involved in defining the boundaries between rainforests (fire-sensitive) and open formations (fire-tolerant) is essential to safeguarding biodiversity and ecosystem services, especially under climate change and increased anthropogenic pressure. Here, we investigated the main environmental factors involved in the co-occurrence of forest islands and humid grasslands located in a protected area in the Espinhaço Biosphere Reserve, southeastern Brazil. We used permanent plots to collect the soil variables (moisture and chemical properties) in the forest islands. For sampling in wet grasslands, we installed four lines of 30 m from the edge of the islands in different directions. Subsequently, we delimited three points on each line10 m apart, totaling 12 points per area. We also surveyed the vegetation cover before and after prescribed burns. The environmental variables were subjected to tests of means and principal component analysis. We observed higher values of potassium, sum of bases, cation exchange capacity and organic matter in soils from forest islands than in wet grasslands. Therefore, the boundaries’ definition between the two vegetation types appeared to be primarily related to soil fertility and moisture gradients. After prescribed burning of the areas, no regeneration of arboreal individuals was detected near the edges of the islands. Therefore, our results suggest that forest islands are unable to expand due to well-defined edapho-climatic conditions. Thus, these environments should be a target focus for designing public conservation policies because they increase the complexity of the landscape of Campos Rupestres vegetation (mountain rocky grasslands).

Crop variety mixtures can provide many benefits, including pathogen suppression and increased yield and yield stability. However, these benefits do not necessarily occur in all mixtures, and the benefits of diversity may be compromised by disadvantages due to increased crop heterogeneity. In-field development of mixtures by assembling many combinations of crop genotypes without prior expectation about which genotypes need to be combined to produce well-performing mixtures results in prohibitively large designs. Therefore, effective tools are required to narrow down the number of promising variety mixtures, and to then identify in experiments which of these deliver the highest benefits. Here, we first review current knowledge about the mechanisms underlying effects in ecological diversity experiments and in current agricultural applications. We then discuss some of the principal difficulties arising in the application of this knowledge to develop good variety mixtures. We also discuss non-conventional approaches to solve some of these issues. In particular, we highlight the potential and limitations of trait-based methods to determine good variety mixing partners, and argue that nontraditional traits and trait-derived metrics may be needed for the trait-based approach to deliver its full potential. Specifically, we argue that good mixing partners can be identified using modern genetic and genomic approaches. Alternatively, good mixtures may be obtained by combining varieties that respond differently to environmental variation; such varieties could easily be identified in standard variety testing trials. Preliminary analyses show that niche differences underlying the different environmental responses can indicate functional complementarity and promote mixture yield and yield stability.

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.

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.

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.

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.

Non-structural carbohydrates (NSCs) are plant storage compounds used for metabolism, transport, osmoregulation and regrowth following the loss of plant tissue. Even in conditions suitable for optimal growth, plants continue to store NSCs. This storage may be due to passive accumulation from sink-inhibited growth or active reserves that come at the expense of growth. The former pathway implies that NSCs may be a by-product of sink limitation, while the latter suggests a functional role of NSCs for use during poor conditions.

Using 13C pulse labelling, we traced the source of soluble sugars in stem and root organs during drought and everwet conditions for seedlings of two tropical tree species that differ in drought tolerance to estimate the relative allocation of NSCs stored prior to drought versus NSCs assimilated during drought. We monitored growth, stomatal conductance, stem water potential and NSC storage to assess a broad carbon response to drought.

We found that the drought-sensitive species had reduced growth, conserved NSC concentrations in leaf, stem and root organs and had a larger proportion of soluble sugars in stem and root organs that originated from pre-drought storage relative to seedlings in control conditions. In contrast, the drought-tolerant species maintained growth and stem and root NSC concentrations but had reduced leaf NSCs concentrations with a larger proportion of stem and root soluble sugars originated from freshly assimilated photosynthates relative to control seedlings. These results suggest the drought-sensitive species passively accumulated NSCs during water deficit due to growth inhibition, while the drought-tolerant species actively responded to water deficit by allocating NSCs to stem and root organs. These strategies seem correlated with baseline maximum growth rates, which supports previous research suggesting a trade-off between growth and drought tolerance while providing new evidence for the importance of plasticity in NSC allocation during drought.

Two formerly broadly distributed woody Atriplex species now occur as fragmented populations across a range of microhabitats in the San Joaquin Valley Desert, southern California. We hypothesized that A. lentiformis and A. polycarpa exhibit inter- and intra-specific differences in their leaf and stem structural and functional traits corresponding with differences in soil salinity and aridity. Water potential, xylem structure and function and leaf traits were compared between three populations of A. lentiformis and four populations of A. polycarpa. The two species significantly differed in their xylem traits, with A. lentiformis displaying lower xylem density, wider mean and maximum vessel diameters and higher hydraulic conductivity (Ks). They also differed in their leaf traits, such that A. lentiformis had larger leaves with higher specific leaf area (SLA) than A. polycarpa. Significant intra-specific differences occurred among leaf traits (leaf area, SLA) in A. lentiformis populations. In contrast, populations varied in their stem xylem structural traits (mean vessel wall thickness, mean vessel diameter, maximum vessel length) among A. polycarpa populations. Many of these differences were associated with soil salinity in A. lentiformis, and with minimum seasonal water potential in A. polycarpa. Overall, both saltbush species showed high intra- and inter-specific trait variation. This could be a critical consideration in understanding the evolution of these native species and has important implications for their conservation and restoration.

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.

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.

The mating systems and floral traits often among relatives of hermaphroditic plants can exhibit considerable diversity. This diversity can be influenced by the evolution of selfing and associated floral traits as a form of reproductive assurance (RA) when pollen limitation (PL) results from insufficient pollinator availability. To explore whether the degree of PL primarily drives differences in mating systems and floral traits, we conducted a comprehensive study involving two closely related species, Halenia elliptica and Halenia grandiflora, in three sympatric sites. We investigated floral characteristics, pollinator visitation, PL, autonomous selfing ability, RA and mating system in studied populations. Our findings show that H. elliptica produces smaller flowers and less nectar production than H. grandiflora, making it less attractive to pollinators. Compared with H. grandiflora, H. elliptica experienced more severe outcross pollen limitation (OPL), but compensates with a higher capacity for autonomous selfing, ensuring seed production under natural conditions. Moreover, significant differences in mating systems were detected between them, with H. elliptica exhibiting a higher selfing rate than H. grandiflora across all studied sympatric populations. These differences are also reflected in variations in herkogamy and dichogamy. Our study suggested that the degree of OPL impacts the divergence in mating systems and floral traits between sympatric closely related species, offering valuable insights into the evolution of plant mating systems and floral traits.

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.

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.

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.

Patterns in functional trait variation associated with the ecological strategies of lianas and trees in subtropical montane forests remain poorly understood due to lack of trait comparisons. Here, we filled this gap by investigating trait divergence between lianas and trees with different leaf habits for 13 traits of 33 species (7 deciduous and 4 evergreen liana species, and 10 deciduous and 12 evergreen tree species) in a subtropical montane forest in southwestern China. We found that lianas had significantly larger stem xylem vessel diameter, higher theoretical hydraulic conductivity, specific leaf area, leaf nitrogen, and phosphorus concentrations, but lower leaf dry-matter content and N/P ratio than sympatric trees, indicating that lianas employ an acquisitive ecological strategy, with a more efficient stem hydraulic system and more productive leaves. In contrast to trees, lianas exhibited a larger variation in vessel diameter, with a few wide and many narrow vessels in the xylem, indicating a greater vessel dimorphism. Growth form explained 48.7% of the total trait variation, while leaf habit only explained 3.8% of trait variation, without significant interaction between growth form and leaf habit. In addition, significant stem-leaf trait relationships were only found in trees, but not in lianas, indicating decoupling of stem hydraulic function and leaf traits in subtropical lianas. These results suggest that subtropical montane lianas and trees differ strikingly in stem and leaf functional traits. Further studies are warranted to strengthen our understanding of the mechanisms underlying the strong divergence in ecological strategies between lianas and trees in subtropical forest ecosystems.

The decline in tree growth has become a global issue. It is critically important to explore the factors affecting tree growth under the background of global climate change to understand tree growth models. A database was established based on Robinia pseudoacacia growth and its driving factors on China’s Loess Plateau. Linear regression and three machine learning methods, including support vector machine, random forest (RF) and gradient boosting machine were used to develop R. pseudoacacia growth models considering forest age, density, climate factors and topographic factors. The root mean square deviation method was adopted to quantitatively assess the relationship between tree growth and soil properties. The average tree height of R. pseudoacacia on the Loess Plateau was 8.8 ± 0.1 m, the average diameter at breast height (DBH) was 10.4 ± 0.1 cm and the average crown diameter was 3.2 ± 0.1 m. The RF model was a fast and effective machine learning method for predicting R. pseudoacacia growth, which showed the best simulation capability and could account for 67% of tree height variability and 55% of DBH variability. Model importance indicated that forest age and stand density were the main factors predicting R. pseudoacacia growth, followed by climate factors. The trade-off between R. pseudoacacia growth and soil properties revealed that soil texture and soil pH were the primary determinants of R. pseudoacacia growth in this region. Our synthesis provides a good framework for sustainable forest management in vulnerable ecological areas under future climate change.

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.

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.

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.

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.

Managing invasions in the context of globalization is a challenge in part because of the difficulty of inferring invader impacts from their invasiveness (i.e. ability to invade ecosystems). Specifically, the relationship between invasiveness and impact may be context-dependent and it has not been explored in such a unique ecosystem as the Tibetan Plateau. Here, we investigated 32 invasive plant species on the Tibetan Plateau in terms of their distribution, abundance, per capita effects on natives and traits across a large geographic transect to test the relationship between invasiveness and impacts on native communities. We decomposed the components (range, R; local abundance, A; per capita effect, E) that drive the impacts, and investigated the relative contributions of plant traits to these components. The results showed that there was no correlation between invasiveness (R × A) and impacts (R × A × E) of invasive species on the Tibetan Plateau. Specifically, plant invasiveness per se did not indicate a serious threat of harmful impact. In this ecosystem, R and A together drove invasiveness, while R alone drove impacts. Fruit type significantly influenced E, and species bearing berry fruits had the most negative per capita effects. However, plant traits did not drive invasiveness or impact through R, A or E. Our results suggest that the mismatch between components driving invasiveness vs. impact prevent the prediction of impacts of invasive species from their invasiveness. Therefore, management actions directed against invasive plants should prioritize broadly distributed species or those with demonstrated high impacts on native species.

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.

When the shoot apical meristem of plants is damaged or removed, fecundity and/or plant growth may suffer (under-compensation), remain unaffected (compensation) or increase (overcompensation). The latter signifies a potential ‘cost’ of apical dominance. Using natural populations of 19 herbaceous angiosperm species with a conspicuously vertical, apically dominant growth form, we removed (clipped) the shoot apical meristem for replicate plants early in the growing season to test for a potential cost of apical dominance. Clipped and unclipped (control) plants had their near neighbours removed, and were harvested after flowering production had finished but before seed dispersal. Dry mass was measured separately for aboveground body size (shoots), leaves, seeds and fruits; and number of leaves, fruits and seeds per plant were counted. We predicted that: (i) our study species (because of their strong apically dominant growth form) would respond to shoot apical meristem removal with greater branching intensity, and thus overcompensation in terms of fecundity and/or biomass; and (ii) overcompensation is particularly enabled for species that produce smaller but more leaves, and hence with a larger bud bank of axillary meristems available for deployment in branching and/or fruit production. Widely variable compensatory capacities were recorded, and with no significant between-species relationship with leaf size or leafing intensity—thus indicating no generalized potential cost of apical dominance. Overall, the results point to species-specific treatment effects on meristem allocation patterns, and suggest importance for effects involving local variation in resource availability, and between-species variation in phenology, life history traits and susceptibility to herbivory.

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.

Architectural plasticity in traits should be useful for understanding morphological and allocation plasticity at the whole-plant level and associated growth strategies of plants in dealing with variation in emergence time. To investigate how plants respond to emergence timing via dynamic architectural plasticity, we conducted a field experiment by sowing plants of Abutilon theophrasti on different dates to make them emerge in spring, late spring, summer and late summer, as four treatments of emergence time (ET1-ET4), and measured a series of mass and morphological traits in different vertical layers (with an inter-distance of 10 or 20 cm per layer) of plants at three stages. The ‘vertical arch' and ‘inverted-cone' shapes displayed at reproductive stage by plants emerged between spring and summer (ET1-ET3) and in late summer (ET4), respectively, revealed that they had contrasting timing schemes for vegetative and reproductive growth. Plants emerged earlier had relatively isolated vegetative and reproductive stages, while late-summer emerged plants had overlapped vegetative and reproductive stages due to accelerated leaf and reproductive growth simultaneously, as a result of shortened lifetime. In spite of this, plants emerged earlier were still able to adopt different optimal strategies in dealing with their corresponding circumstances. Architectural analysis should be a useful approach to better understanding diverse plant strategies in the context of varying environmental conditions.

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.

Warming and grazing, and litter quality jointly determine litter decomposition and nutrient releases in grazing ecosystems. However, their effects have previously been studied in isolation. We conducted a two factorial experiment with asymmetric warming using infrared heaters and moderate grazing in an alpine meadow. Litter samples were collected from all plots in each treatment, among which some subsamples were placed in their original plots and other samples were translocated to other treatment plots to test the relative effects of each treatment on litter decomposition and nutrient releases. We found that warming rather than grazing alone significantly increased total losses of litter mass, total organic carbon, total nitrogen (TN) and total phosphorus (TP) per unit area due to increases in both mass loss rates and litter biomass. However, grazing with warming did not affect their total mass losses because increased mass loss was offset by decreased litter biomass compared with the control. Seasonal mean soil temperature better predicted litter decomposition than litter lignin content or carbon to nitrogen ratio. There were interactions between warming and grazing, but there were no interactions between them and litter quality on litter decomposition. The temperature sensitivity of TN loss was higher than that of TP loss per unit area. Our results suggest that increased temperature has a greater effect on litter decomposition and nutrient release than change in litter quality, and that more N release from litter could result in greater P deficiency in the alpine meadow.

A better understanding the mechanisms driving plant biomass allocation in different ecosystems is an important theoretical basis for illustrating the adaptive strategies of plants. To date, the effects of habitat conditions on plant biomass allocation have been widely studied. However, it is less known how plant community traits and functions (PCTF) affect biomass allocation, particularly in alpine grassland ecosystems. In this study, community-weighted means (CWM) were calculated at the community level using five leaf functional traits, and the relationships between PCTF and biomass trade-offs were explored using correlation analysis, variation partitioning analysis and structural equation modeling. We found that the trade-off values were greater than zero in both alpine meadow (AM) and alpine steppe (AS) across the Tibetan Plateau, with different values of 0.203 and 0.088 for AM and AS, respectively. Moreover, the critical factors determining biomass allocation in AS were species richness (SR; scored at 0.69) and leaf dry matter content of CWM (CWM_LDMC’, scored at 0.42), while in AM, the key factors were leaf dry matter content (CWM_LDMC’, scored at 0.48) and leaf carbon content of CWM (CWM_LC’, scored at -0.45). In particular, both CWMLDMC and SR in AS, as well as CWM_LDMC and CWM_LC in AM were primarily regulated by precipitation. In summary, precipitation tends to drive biomass allocation in alpine grasslands through its effects on PCTF, hence highlighting the importance of PCTF in regulating plant biomass allocation strategies along precipitation gradients.