Positive effects of desert shrub in ecosystem multifunctionality primarily depend on increasing soil nutrients

Positive effects of desert shrub in ecosystem multifunctionality primarily depend on increasing soil nutrients | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Positive effects of desert shrub in ecosystem multifunctionality primarily depend on increasing soil nutrients Shuang-Fu Zhou, Xue-Ni Zhang, Xiao-Hui Li, Yu-Fei Chen, Hui-Xia Li This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9474137/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 4 You are reading this latest preprint version Abstract Aims Shrub encroachment in desert ecosystems is projected to intensify worldwide because of rapid climate change and anthropogenic activity, leading to significant alterations of ecosystem functions. The study investigates the effects of shrub on various ecosystem functionality relevant to plant diversity, productivity, and soil nutrients under different water and salt environment in desert ecosystems. Methods Soil environments are classified into high, medium, and low water and salt environments based on soil moisture content and electrical conductivity. Differences in soil and plant traits between the understory and bare within sample plots across different water and salt environments were compared. The effects of shrubs on individual ecosystem functions and ecosystem multifunctionality, and their water and salt responses, are analyzed using averaging and multiple-threshold approach. The drivers of shrub influence on ecosystem multifunctionality are analyzed using generalized linear mixed model (GLMM). Results Our findings suggest that desert shrub increased herbaceous aboveground biomass and Soil organic carbon (SOC), total nitrogen (TN), total phosphorus (TP), the positive effects of desert shrubs on ecosystem multifunctionality varied with water-salinity conditions. Shrubs increased plant productivity and soil nutrients for high (HSW) and medium water and salt environment (MSW) while reducing plant diversity for low environment (LSW). The enhancement of ecosystem multifunctionality by desert shrubs was closely related to the promotion of soil nutrient content and plant productivity. Conclusion Understanding the effect of desert shrubs on ecosystem functions will help us screen native species from a functional perspective for ecological restoration, thereby enhancing the ecological benefits of plant-based remediation. shrub encroachment desert ecosystem individual ecosystem function ecosystem multifunctionality water and salt environment Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 INTRODUCTION Desert ecosystems are an important component of global terrestrial ecosystems, playing a prominent role in maintaining global ecological balance and serving the entire human population (Liang et al., 2019 ). As the dominant plant species in desert ecosystems, shrubs may profoundly influence multiple ecosystem functions (ecosystem multifunctionality) through multidimensional ecological processes (Yue et al., 2025 ). Currently, global desert ecosystems are undergoing widespread scrub encroachment (Deng et al., 2021 ), leading to mutations in the desert vegetation diversity, productivity, and soil nutrients, accompanied by transformations in ecosystem multifunctionality (Karaoud & Noumi, 2025 ; Zhang et al., 2016 ; Zhang et al., 2023 ). However, the mechanisms by which shrubs drive these changes and alter both the singular and multifaceted aspects of ecosystem functionality remain poorly understood (Ding et al., 2024 ; Liu et al., 2019 ). Shrubs play a significant role in soil nutrient dynamics, plant composition, and ecological interactions between vegetation and soil within desert ecosystems (Thomas et al., 2018 ). In desert environments, shrubs are often regarded as nurse plants, providing a stable and temperate growth environment for understory herbaceous plants, thereby enhancing their richness, diversity, and productivity (La Montagna et al., 2025 ). However, in grassland ecosystems, shrub encroachment often leads to decline of herbaceous richness, diversity, and biomass, which ultimately resulting grassland degradation (Valencia et al., 2015 ). Thus, the effect of shrubs on herbaceous plants may differ with ecosystem types. In semi-arid regions, shrubs improve soil moisture conditions and soil nutrients availability beneath the canopy through the “fertility island” effect, enhancing spatial heterogeneity of soil nutrients (Yao et al., 2019 ). During years with lower precipitation, the positive effect of shrubs on soil moisture becomes more pronounced, helping mitigate the negative impacts of drought on plants (Liu et al., 2020a ). These soil and vegetation changes induced by shrubs further influence ecosystem functions. For example, shrub encroachment increases soil activity in tundra ecosystem, enhancing nutrient cycling functions (Aguirre et al., 2021 ); In the Tibetan Plateau region, leguminous shrubs enhance the multifunctionality of grassland ecosystems, closely related to improved soil nutrients (Yang et al., 2024 ). In desert ecosystems, shrubs facilitated the development of their associated herbaceous communities via accumulating more soil resources (Xie et al., 2022 ). In addition, ecosystem multifunctionality was positively related to plant diversity (Wang et al., 2020 ). Shrubs exert positive effects on plant diversity, productivity, and soil nutrients in desert ecosystems, suggesting their influence may also be beneficial on desert ecosystem multifunctionality, though this requires further research and practical validation. Desert ecosystems also exhibit significant environmental heterogeneity, making the role of shrubs in desert ecosystem functions is highly susceptible to soil moisture and salinity conditions. The extent of these effects may vary with soil moisture fluctuations (Butterfield et al., 2015 ; Wang et al., 2024 ). Consequently, the role of soil environment also cannot be overlooked in the process of shrub impacts on desert ecosystem multifunctionality. The positive effects of shrubs on ecosystem functions may diminish with increasing drought stress. In forest ecosystems, reduced soil moisture caused by human activities diminishes the effect of plant diversity on ecosystem multifunctionality (Zhao et al., 2025 ). In alpine meadow and grassland ecosystems, the positive effect of shrubs on multiple ecosystem functions greatly depended on increasing soil nutrient (Yang et al., 2024 ). This influence of soil moisture and nutrient on ecosystem multifunctionality has also been confirmed in alpine ecosystems (Yang et al., 2023 ). Plants and soil, as two primary drivers of ecosystem function, the effects of plant diversity, productivity, and soil nutrients on ecosystem multifunctionality are also synergistic and complementary (Delgado-Baquerizo et al., 2020 ). Changes in the soil environment alter the effect of shrubs on plant diversity, productivity, and soil nutrients, potentially affecting shifts in ecosystem multifunctionality. In arid regions, drought stress weakens the positive influence of plant diversity on ecosystem multifunctionality (Bhadauria et al., 2022 ). The relationship between plant diversity and ecosystem multifunctionality may vary along environmental gradients (Fanin et al., 2018 ). Most studies indicate that the effect of plant diversity on ecosystem multifunctionality decreases with increasing environmental stress, though some reports suggest no change or even an increase (Hu et al., 2021 ). In arid regions, Maestre et al. demonstrated that shrubs enhance ecosystem function by mitigating the negative impacts of drought stress (Maestre et al., 2012 ). Given the multidimensional nature of ecosystem functionality, the effect mechanisms of shrubs plants on the multifunctionality of desert ecosystems may exhibit complex variations along water-salt gradients (Xu et al., 2024 ), though our understanding remains limited. Nowadays, the intensification of extreme drought climate events has become a global reality (Huang et al., 2024 ). However, research on the effects of shrub plants on the multifunctionality of desert ecosystems remains scarce, and the roles of desert shrubs in the single function and multifunctionality of the ecosystem are still unclear. Therefore, this study takes the arid desert ecosystem of the Ebinur Lake Basin as the research subject and explores the effect pathways of different desert shrub plants on various functions and multifunctionality of the ecosystem at drought stress gradients. It aims to address the following questions: (1) How do desert shrubs affect herbaceous plants and soil environments? (2) What are the pathways through which shrubs influence single or multiple desert ecosystem functions? Addressing these questions will enhance the predictability of desert ecosystem functions and services under changing environmental conditions. MATERIAL AND METHODS Study site The Ebinur Lake Wetland National Nature Reserve (44°30′-45°09′N, 82°36′-83°50′E) is located in northwest Jinghe County, Bortala Mongolian Autonomous Prefecture, Xinjiang, China, in the lowest depression in the southwest margin of Junggar Basin. The region has a dry typical continental climate with an annual precipitation of only about 89.80–169.70 mm and strong potential annual evaporation is higher than precipitation, which ranges from 1569–3421mm. The soil parent material is sandy soil, grey-brown desert soil, and grey desert soil, which are severely affected by desertification. The main vegetation types in the study area include Populus euphratica , Haloxylon ammodendron , Halimodendron halodendron , Nitraria tangutorum , Alhagi sparsifolia , Reaumuria songarica , Calligonum mongolicum Turcz , Apocynum Venetum , Phragmites australis , Suaeda glauca . Experimental design In July 2024, on the north side of the Aqikesu River within the Ebinur Lake, the locations approximately 1km, 2.5km and 4.5km from the Aqikesu River were selected as high water and salt environment (HSW), medium water and salt environment (MSW) and low water and salt environment (LSW), respectively (Fig. 1 ). Each water-salt environment included 3 replicated plots (30×30) and separated by approximately 20m, each plot was further divided into 9 smaller sample plots (10m×10m), totaling 81 plots. Shrub environment (SE) and without shrub environment (WE) were set up in each smaller sample plots. The understory environment beneath shrub canopies served as shrub environment (SE), while bare ground areas constituted without shrub environment (WE). Field sampling and measurements Record the quantity, height, coverage, base diameter of shrub species and the species name, quantity, height (CH), base diameter (D) and coverage of herbaceous plants in each square (10×10). Then, we subsequently harvested herbaceous plant materials at ground level for aboveground biomass (AGB) determination. Collect soil samples from 0–20 cm depth under canopy and in open areas. Determine soil moisture content using the oven-drying method. Soil samples were screened through a 2mm sieve to remove roots and stones for the measurements of soil physicochemical properties. Soil electrical conductivity (EC) was determined using conductivity meter; soil organic carbon (SOC) content by the potassium dichromate-sulfate oxidation method (Nelson & Sommers, 1982 ); soil total nitrogen (TN) by the Kjeldahl method (Lynch & Barbano, 1999 ); soil total phosphorus (TP) by the molybdenum antimony colorimetric method (Murphy & Riley, 1962 ); soil available phosphorus (AP) by sodium bicarbonate extraction-molybdenum-antimony anti-spectrophotometric method; soil ammonium nitrogen (NH 4 + -N) and nitrate nitrogen (NO 3 − -N) by ultraviolet spectrophotometry. Data analysis The calculation formula for the relative importance value (IV) of shrubs: $$\:IV=(RD+RC+RF)/3$$ Where RD represents the relative density of the shrub, RC represents the relative coverage of the shrub, and RF represents the relative frequency of the shrub. We used the Species richness index (R), Shannon-Wiener diversity index (H), and Pielou evenness index (J) as our plant diversity indicators, which can be calculated as follows: Species richness index: $$\:R=S$$ Shannon-Wiener index: $$\:H=-{\sum\:}_{i=1}^{s}pi{ln}pi$$ Pielou index: $$\:J=H/{ln}S$$ Where S represents the total number of plant species, and Pi represents the importance value (IV) of the ith species. To evaluate the differences in ecosystem functions between shrub and bare land, we assessed the ecosystem multifunctionality (EMF) index based on six specifically chosen individual ecosystem functions (ABG、SOC、SW、D、CH、TP), which were selected following a thorough analysis using the Random Forest method (Fig. 2 ). The EMF was calculated using two methods, averaging approach and multiple-threshold approach. We further classified these individual ecosystem functions into three functional groups: (1) functions relating to diversity (R, H, and J), (2) plant productivity (AGB、CH and D), (3) soil nutrients (SOC、TN、NH 4 + -N、NO 3 − -N、TP、AP). The EMF index assessing using averaging approach that can be calculated as: $$\:EMF=\frac{1}{N}{\sum\:}_{i=1}^{N}Zij$$ Where N is the total number of selected ecosystem plots and Zij represents the standardization of the evaluated ith ecosystem function at the jth plot. In addition, the ecosystem multifunctionality index, calculated using the multiple-threshold approach (Byrnes et al., 2014 ), is determined by counting the number of functions that simultaneously surpass a specified threshold of the maximum observed value for a given function across all study plots. As any choice of a threshold, that determines whether a given ecosystem function contributes to multifunctionality, is likely arbitrary, three different thresholds (25%, 50%, and 75%) were used, thus representing a wide spectrum in the analyses (Delgado-Baquerizo et al., 2020 ). This approach overcomes the weakness of the average multifunctionality method. We used independent samples t-tests were employed to analyze and evaluate significant differences in ecosystem functions under shrub canopies in desert ecosystems. Structural equation modeling (SEM) was employed to construct a pathway diagram of ecosystem multifunctionality drivers. Generalized linear mixed models (GLMM) were applied to analyze the multivariate explained variance of ecosystem functions along the water-salt gradient and the individual effects of its drivers. All statistical analyses were processed using SPSS and R software, with Origin software utilized for graphical representation. RESULTS Characteristics of shrubs in different water-salt environments and their effects on herbaceous plants The composition of shrubs and their importance values vary with different water-salinity environments. HSW is dominated by Halimodendron halodendron , with a relative importance value (IV) of 0.7126. While MSW is primarily composed of Alhagi sparsifolia and Reaumuria songarica , with IV of 0.6113 and 0.6044, respectively. Shrubs of LSW mainly are Alhagi sparsifolia and Haloxylon ammodendron , with IV of 0.6417 and 0.5148, respectively (Fig. 3 a-c). HSW refers to tall shrubs, MSW are short shrubs, and shrubs of LSW are tall but have a smaller crown spread (Fig. 3 d-f). When comparing the shrubs canopy with the bare land, shrubs significantly reduced herbaceous species richness but increased aboveground biomass of herbaceous plants in desert ecosystems. Shrubs decreased the Shannon-Wiener index of LSW herbaceous plants while increasing the Pielou index of HSW herbaceous plants. Shrubs increased plant height but reduced their basal diameter of MSW herbaceous plants (Fig. 4 ). Effects of shrubs on soil properties under different water-salt environments Table 1 Physical and chemical properties of soil under shrub canopy and bare land in different water and salt environments Environment Variables SE WE P value HSW SW 14.16 ± 1.56 10.51 ± 1.21 P < 0.001 SOC 1.66 ± 0.41 1.06 ± 0.32 P < 0.001 TN 1.65 ± 0.60 0.97 ± 0.28 P < 0.001 NH 4 + -N 0.004 ± 0.002 0.004 ± 0.002 P = 0.994 NO 3 − -N 0.006 ± 0.002 0.007 ± 0.008 P = 0.382 TP 0.65 ± 0.07 0.63 ± 0.04 P = 0.236 AP 0.08 ± 0.04 0.06 ± 0.04 P = 0.056 MSW SW 7.61 ± 0.90 5.46 ± 0.59 P < 0.001 SOC 0.36 ± 0.11 0.33 ± 0.15 P = 0.447 TN 0.56 ± 0.26 0.40 ± 0.20 P = 0.015 NH 4 + -N 0.004 ± 0.002 0.003 ± 0.002 P = 0.103 NO 3 − -N 0.008 ± 0.002 0.008 ± 0.002 P = 0.982 TP 0.46 ± 0.05 0.42 ± 0.06 P = 0.003 AP 0.09 ± 0.03 0.06 ± 0.02 P < 0.001 LSW SW 0.83 ± 0.20 0.96 ± 0.37 P = 0.002 SOC 0.12 ± 0.03 0.09 ± 0.34 P < 0.001 TN 0.45 ± 0.13 0.32 ± 0.10 P < 0.001 NH 4 + -N 0.002 ± 0.0008 0.002 ± 0.004 P = 0.967 NO 3 − -N 0.008 ± 0.001 0.007 ± 0.002 P = 0.577 TP 0.34 ± 0.04 0.30 ± 0.02 P < 0.001 AP 0.02 ± 0.01 0.03 ± 0.03 P = 0.064 Note: Data are presented as mean ± standard deviation. The significance level for independent samples t-tests was set at 0.05, with significant values shown in bold. HSW, the high water-salt environment; MSW, the medium water-salt environment; LSW, the low water-salt environment. SE, shrub environment; WE, without shrub environment; SW, soil water content; SOC, soil organic carbon; TN, soil total nitrogen; NH₄⁺-N, soil ammonium nitrogen; NO₃⁻-N, soil nitrate nitrogen; TP, soil total phosphorus; AP, soil available phosphorus. As shown in Table 1 , with shrub canopies showing relatively higher TN than bare land in desert ecosystems. The SW under shrub canopies exceeded bare land (p < 0.001) in HSW and MSW, and the SOC under shrub canopies was significantly higher than bare land in HSW and LSW (p < 0.001). In the MSW, both TP (p = 0.003) and AP (p < 0.001) under shrub canopies were higher than bare land. Relatively higher TP (p < 0.001) but lower SW (p = 0.002) under shrub canopies than bare land in LSW. Along with the water-salinity gradient, the positive effect of shrubs showed an increasing trend for TP, while their positive effect exhibited a decreasing trend for SW. Even in LSW, this effect was reversed. Response of individual and multiple ecosystem functions to desert shrub The EMF indexes varied greatly across water-salt gradient, The mean EMF index decreased along water and salt environment. Based on two calculation methods of multifunctionality, shrubs significantly enhanced the EMF, EMF-productivity, and EMF-nutrient indexes of HSW and MSW, and reduced the EMF-diversity indexes of LSW. However, the EMF indexes and EMF-nutrient indexes calculated by two methods show differences. After evaluating the EMF and the EMF-nutrient through the multiple-threshold method, it is observed that selecting the 25% threshold results in shrubs enhanced the EMF and the EMF-nutrient of LSW (Fig. 5 ). Effect of shrub traits and environmental factors on individual and multiple ecosystem functions Shrub cover (E = 0.03, p = 0.013) and height (E = 0.02, p = 0.035) both promote the multifunctionality of desert ecosystems. Shrub cover positively influenced soil moisture content (E = 0.20, p ≤ 0.001) and soil nutrient function (E = 0.16, p = 0.008); shrub height directly negatively affected plant diversity (E=-0.26, p ≤ 0.001) (Fig. 6 a-d). The effect of individual ecosystem functions on multifunctionality varies significantly across water-salinity gradients. Plant productivity exerted the strongest influence on ecosystem multifunctionality, while the effect of soil nutrient diminished along the water-salinity gradient (Fig. 6 e-g). In desert ecosystems, shrubs primarily enhance multifunctionality by increasing soil nutrient content. DISCUSSION Desert shrubs exert positive effects on plant productivity and soil nutrient content in non-extreme environments The research findings indicate that the composition and growth of desert shrubs vary with the water-salt environment. Shrub species richness and growth performance of MSW was lower than in other water and salt environment. This was primarily due to drought and salt stress in reducing height, basal diameter, and species richness of shrubs (Gatica et al., 2020 ; Shi et al., 2019 ). Shrub cover exhibited heightened sensitivity to moisture fluctuations, with drought stress being the primary factor why significantly higher shrub cover of HSW than MSW and LSW (Chunyuan et al., 2024 ). The relative importance of Alhagi sparsifolia and Haloxylon ammodendron were significantly higher than other shrubs, owing to their stronger drought-tolerant genetic endowment (Gao et al., 2020 ). Consequently, the inherent tolerance of shrub species themselves is also one of the reasons for the differences in shrub composition along the water and salt environment. The presence or absence of shrubs significantly influences changes in soil nutrient content within desert ecosystems (Johnson et al., 2016 ), The research results also confirmed that desert shrubs increase the content of SOC, TN, and TP under their canopy, while having no significant effect on soil nutrient availability. This finding indicates that the increase in soil nutrient content under shrub canopies primarily originates from external sources. Shrubs deposit these nutrients into the understory soil through litterfall and canopy interception (Ding & Eldridge, 2020 ), thereby increasing the nutrient content in the soil. However, this positive effect may change along with the variations in the water and salt environment of desert ecosystems. Soil moisture content under shrub canopies was lower than bare land in LSW, which is consistent with studies from other desert ecosystems (Butterfield et al., 2015 ). Under moderate drought stress, shrubs enhance soil water retention by reducing evaporation and providing canopy shade, thereby increasing SW beneath their canopies (Noumi et al., 2015 ). Additionally, In desert ecosystems, shrubs improve the local environment to provide relatively better conditions for herbaceous plants, promoting their biomass (Franco et al., 2019 ; Zhang et al., 2025a). The finding that AGB under shrub canopies was significantly higher than bare land further corroborated the conclusion. As shrubs promote the accumulation of soil organic matter and nutrients beneath their canopies, enabling sustained access to growth resources for understory plants (Weber-Grullon et al., 2022 ). Shrubs may also limit grazing by herbivores through their canopy structure, thereby increasing AGB of herbaceous plants. However, plant richness on bare land is significantly higher than plant community shrub canopies. as shrubs expand, they simultaneously suppress the growth of some plants (Sun et al., 2023 ). Shrub canopy setting up physical barriers to prevent the entry of some plants (Kang et al., 2025 ). and shading amplifies their negative effect on plant richness (Xu et al., 2025 ), leading to a decline in herbaceous plant richness, this aligns with global grassland ecosystem research (Wieczorkowski & Lehmann, 2022 ). The intensification of environmental stress increases competition between shrubs and herbaceous plants for survival resources (Shi et al., 2025 ), thereby the inhibitory effect of shrubs on plant diversity intensifies along water-salt gradients. Together, desert shrubs reduced herbaceous species diversity while promoting biomass growth. This change correlates with effect of shrubs on soil nutrient content. The effect of shrubs on soil and herbaceous plants subsequently influences both the single-function and multifunctional aspects of ecosystems (Chen et al., 2023 ). In desert ecosystems, shrubs increase plant productivity and soil nutrients, which aligns with the view that shrubs, as nurse plants in arid areas, enhance ecosystem functions. within a certain range of shrub canopy cover (5% − 45%) (Liancourt et al., 2020 ), greater canopy cover exhibits stronger positive effects on understory plant productivity and soil nutrients (liu et al., 2020b ). In the study area, shrubs decrease plant diversity, consistent with studies on desert shrub invasions and grassland vegetation succession (Hector et al., 2008 ; Wu et al., 2024 ). Research indicates that desert shrubs have a positive effect on ecosystem multifunctionality, and this effect diminishes along water-salt gradients. Which consistent with recent studies from the stress gradient hypothesis (Zhang et al., 2022 ). This shift occurs primarily due to extreme drought stress and the reduction shrub canopy cover, which cause the relationship between shrubs and understory herbaceous plants to shift toward competition (Sun et al., 2023 ). The effects of shrubs on plant diversity, plant productivity, and soil nutrients drive changes in ecosystem multifunctionality. Desert shrubs enhance ecosystem multifunctionality by increasing soil nutrient In desert ecosystems, we found that plant diversity exerts a positive influence on ecosystem multifunctionality, though its effect is not significant. This is contrary to the prevailing view that plant diversity is generally considered the primary driver of ecosystem multifunctionality (Li et al., 2025b ; Ma et al., 2022 ). However, from another perspective, When measured by plant richness and density, plant diversity effectively predicts trends in ecosystem multifunctionality under precipitation variations in desert grasslands, exhibiting a significant positive correlation with ecosystem multifunctionality (Hu et al., 2022 ). Additionally, the research also revealed a significant negative effect of shrub height on plant diversity. Along with the water-salinity gradient, the influence of plant diversity on ecosystem multifunctionality first decreased and then increased, which aligns with the nonlinear relationship between plant diversity and ecosystem multifunctionality across changing of environmental gradients (Liu et al., 2025 ). Changes in water and salt levels trigger changes in many environmental variables, impacting on the distribution and diversity of plant species and thus largely determine the stability of the ecosystem in the study area (Zou et al., 2024 ). The effect of shrubs on ecosystem multifunctionality is also reflected in the changes of plant productivity. Observations in this study that plant productivity positively influences ecosystem multifunctionality support this assertion. Our results show that aboveground biomass of understory herbaceous is significantly higher than that of bare lands, though shrubs did not significantly enhance plant productivity directly. In fact, the positive effect of shrubs on the productivity of herbaceous plants stems from the fact that the canopy of shrubs provides relatively favorable soil conditions for plant growth (Thakur & Kumar, 2021 ), thereby indirectly promoting positive effect of plant productivity on ecosystem multifunctionality. Furthermore, the positive effect of plant diversity on plant productivity was not significant, contradicting the general view that plant richness correlates positively with plant productivity (Tilman et al., 2014). Shrub cover promoted plant productivity, while shrub height exerted a negative effect on plant diversity. In desert ecosystems, plant productivity and diversity are controlled by different traits of shrub, potentially differentiating their relationship (Sanaei et al., 2018 ). In the study area, the effect of plant productivity on the multifunctionality of the ecosystem is the greatest, but no significant variation across the gradient. Soil nutrients and moisture content, as primary indirect drivers of plant productivity, alleviated the effect of water-salinity fluctuations on plant productivity (La Pierre et al., 2016 ), thereby stabilizing the positive role of plant productivity in ecosystem multifunctionality. The shrub canopy plays a pivotal role in the effect of shrubs on ecosystem functions (soil nutrients) (Lu et al., 2018 ; Maestre et al., 2009 ). Compared to bare lands, shrubs significantly improve soil nutrient status, further validating this point. Soil nutrients directly promote ecosystem multifunctionality while also indirectly influencing it by enhancing plant productivity. This is the same mechanism by which soil nutrient loss leads to the decline of ecosystem multifunctionality during land degradation (Li et al., 2025a). Furthermore, Soil microorganisms play a crucial role in the changing mechanisms of ecosystem multifunctionality in arid regions (Zhang et al., 2025b; Zhou et al., 2025), therefore, necessitating investigation into their role in the process of ecosystem multifunctionality variation in desert ecosystems. The establishment of herbaceous plants significantly increases soil nutrient content in arid areas (Liu et al., 2023 ), providing evidence that plant productivity enhances soil nutrient content. Higher levels of plant biomass are more likely to increase soil nutrients, with the plant litter input and accumulation contributing to nutrient sequestration and soil moisture retention(Liu et al., 2021b ; Ward et al., 2018 ). In summary, the mutual promotion of plant productivity and soil nutrients primarily benefits from a temperate microenvironment created by shrub canopy in desert ecosystems, providing stable conditions for a positive feedback loop between plants and soil. Both plant functional traits and soil nutrients are equally important for the variation in ecosystem multifunctionality on the water-salt gradient. In summary, this research results provide clear evidence that shrub facilitates ecosystem functioning of desert ecosystem, which had a stronger facilitative effect in HSW and MSW than LSW. However, we recognize there are still limitations in the present study. We intentionally focused on the shrub induced changes in the herbaceous community, soil properties, and ecosystem functioning of desert ecosystem. Previous studies have demonstrated that shrubs positively influence soil microbial activity and abundance (Liu et al., 2021a ). Therefore, it is necessary to consider the role of soil microorganisms in the ecological impact of desert shrubs. Elevated temperature or altered precipitation have a significant impact on multiple ecosystem functions in desert regions. This suggests that ongoing climate change and anthropogenic activity may mediate future effectiveness of shrub encroachment (Tu et al., 2024 ; Wang et al., 2016 ). CONCLUSIONS Research indicates that shrub vegetation significantly alters the plant community structure and soil environment of desert ecosystems, enhancing their multifunctionality. Shrubs increase soil nutrient content and herbaceous aboveground biomass while reducing herbaceous species richness. Furthermore, the effect of shrubs on desert ecosystem functions varies along water-salinity gradients. Desert shrubs enhance plant productivity and soil nutrients of HSW and MSW (SW: 5%~12%, EC: 11 ~ 27 ms/cm) while reducing plant diversity of LSW. This research provides crucial insights into the interactions between shrub vegetation characteristics and ecosystem functions in desert ecosystems, offering decision-making guidance for desert ecosystem utilization and management. Given the threat of global climate change, widespread shrub encroachment in desert ecosystems has become a critical issue. Enhancing our ability to predict the impacts of these phenomena on the structure and function of global desert ecosystems is paramount. Addressing these challenges at larger spatio-temporal scales to improve our capacity to understand, mitigate, or plan for the effects of such vegetation changes on desert ecosystem functions requires further empirical research and modeling. Declarations CRediT authorship contribution statement Shuangfu Zhou : Writing – original draft, Methodology, Investigation, Software, Experiment, Data curation. Xiaohui Li : Investigation, Software, Experiment, Data curation. Huixia Li : Investigation, Experiment, Data curation. Yufei Chen : Investigation, Experiment, Data curation. Xueni Zhang (corresponding author) : Writing – review & editing, Methodology, Supervision, Funding acquisition. Declaration of competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Acknowledgments This work was supported by the Natural Science Foundation of China (32360277). Data availability Data will be made available on request. References Aguirre D, Benhumea AE, McLaren JR (2021) Shrub encroachment affects tundra ecosystem properties through their living canopy rather than increased litter inputs. 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Forest Ecosystems , 11. http://10.1016/j.fecs.2024.100187 Cite Share Download PDF Status: Under Review Version 1 posted Reviewers invited by journal 03 May, 2026 Editor invited by journal 27 Apr, 2026 Editor assigned by journal 27 Apr, 2026 First submitted to journal 26 Apr, 2026 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-9474137","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":633696134,"identity":"bef5b2f2-25e0-4e0f-a206-1f982ca23e04","order_by":0,"name":"Shuang-Fu Zhou","email":"data:image/png;base64,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","orcid":"","institution":"Xinjiang University College of Ecology and Environment","correspondingAuthor":true,"prefix":"","firstName":"Shuang-Fu","middleName":"","lastName":"Zhou","suffix":""},{"id":633696135,"identity":"22ca10e9-7ecf-4acb-93d5-bc877a534fa9","order_by":1,"name":"Xue-Ni Zhang","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Xue-Ni","middleName":"","lastName":"Zhang","suffix":""},{"id":633696136,"identity":"677af9b7-31ed-43aa-a55b-96053741b982","order_by":2,"name":"Xiao-Hui Li","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Xiao-Hui","middleName":"","lastName":"Li","suffix":""},{"id":633696137,"identity":"1565eb3a-0a0c-49da-8086-9956b8de3a4a","order_by":3,"name":"Yu-Fei Chen","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Yu-Fei","middleName":"","lastName":"Chen","suffix":""},{"id":633696138,"identity":"2922c393-e1b4-440f-a5fa-5e67beb40589","order_by":4,"name":"Hui-Xia Li","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Hui-Xia","middleName":"","lastName":"Li","suffix":""}],"badges":[],"createdAt":"2026-04-20 15:17:48","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-9474137/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9474137/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":108981772,"identity":"1247c7e0-19a3-4492-8b9a-4953acaee896","added_by":"auto","created_at":"2026-05-11 12:19:50","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1780817,"visible":true,"origin":"","legend":"\u003cp\u003eOverview map of the study area (a); experimental plot distribution map (b); high water-salinity environment (HSW) (c); medium water-salinity environment (MSW) (d); low water-salinity environment (LSW) (e).\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-9474137/v1/5c5dbacf0aa6d0c8c8f07d62.png"},{"id":108981764,"identity":"e0af327d-499a-46ba-9db8-a4c4d6d6318f","added_by":"auto","created_at":"2026-05-11 12:19:40","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":48708,"visible":true,"origin":"","legend":"\u003cp\u003eThe potential driving factors for overall ecosystem multifunctionality (calculated using 13 relevant indicators) were explored. Random Forest (RF) mean predictor importance, represented by the percentage increase of mean square error, was assessed for the EMF index as a driver for the 13 indicators (R, herbaceous species richness; H, Shannon-Wiener index; J, Pielou index; AGB, herbaceous aboveground biomass; CH, herbaceous plant height; D, herbaceous plant basal diameter; SW, soil water content; SOC, soil organic carbon; TN, soil total nitrogen; NH₄⁺-N, soil ammonium nitrogen; NO₃⁻-N, soil nitrate nitrogen; TP, soil total phosphorus; AP, soil available phosphorus).\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-9474137/v1/d682a53614c2ed460ee02c64.png"},{"id":108981885,"identity":"f9007e39-ede2-454a-98a9-ab2ac98cdd5b","added_by":"auto","created_at":"2026-05-11 12:20:38","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":340850,"visible":true,"origin":"","legend":"\u003cp\u003eDesert shrub species (a-c) and their characteristics (d-f) across different water-salt gradients. Mean values ± standard errors are indicated by error bars. Different letters denote significant differences in shrub traits across water-salt gradients (p \u0026lt; 0.05).\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-9474137/v1/cccb3717c119c2bb03062ce0.png"},{"id":108982840,"identity":"47caa04d-9022-4dc9-9b48-697224654d26","added_by":"auto","created_at":"2026-05-11 12:28:11","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":232758,"visible":true,"origin":"","legend":"\u003cp\u003eDifferences in herbaceous plant diversity and morphological traits between shrub canopy and bare land. HSW, the high water-salt environment; MSW, the medium water-salt environment; LSW, the low water-salt environment; SE, shrub environment; WE, without shrub environment; R, herbaceous species richness; H, Shannon-Wiener index; J, Pielou index; AGB, herbaceous aboveground biomass; CH, herbaceous plant height; D, herbaceous plant basal diameter. Means ± standard deviation is indicated by error bars. Asterisks represent the significant differences between shrub canopy and bare land. *, p ≤ 0.05; **, p ≤ 0.01; ***, p ≤ 0.001.\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-9474137/v1/ac20a7e73b772569984d5e37.png"},{"id":108983070,"identity":"a1d6deae-9d84-4bf9-82e5-aab964699e9a","added_by":"auto","created_at":"2026-05-11 12:30:03","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":450543,"visible":true,"origin":"","legend":"\u003cp\u003eDifferences in the ecosystem multifunctionality (EMF), EMF-diversity, EMF-productivity, EMF-nutrient indexes between the shrub canopy and bare land. Calculate using the averaging approach (a; e; i; m) and the multiple-threshold approach (b-d; f-h; j-l; n-p), choosing 25 %, 50 %, and 75 % thresholds. HSW, the high water-salt environment; MSW, the medium water-salt environment; LSW, the low water-salt environment; SE, shrub environment; WE, without shrub environment. Asterisks indicate significant differences between shrub plots and open plots. *, P ≤ 0.05; **, P ≤ 0.01; ***, P ≤ 0.001.\u003c/p\u003e","description":"","filename":"floatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-9474137/v1/ff5826c35e87e88e62305520.png"},{"id":108981888,"identity":"17d1ae84-28e1-4559-ad6c-cd2a2d438c6d","added_by":"auto","created_at":"2026-05-11 12:20:41","extension":"jpeg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":565017,"visible":true,"origin":"","legend":"\u003cp\u003eStructural equation model of ecosystem multifunctionality drivers (a–d) and individual effects of ecosystem functional drivers along the water-salt gradient (e–g). Arrows indicate significant effects, with red denoting positive effects and black denoting negative effects, p ≤ 0.05.\u003c/p\u003e","description":"","filename":"floatimage6.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-9474137/v1/452e4da98b956823cbbdc3b7.jpeg"},{"id":108985196,"identity":"4078a00f-9493-4ae8-9735-69aed722dc65","added_by":"auto","created_at":"2026-05-11 12:44:32","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":4564483,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9474137/v1/b380b1d3-9d91-4652-89e4-2cc34d062b39.pdf"}],"financialInterests":"","formattedTitle":"Positive effects of desert shrub in ecosystem multifunctionality primarily depend on increasing soil nutrients","fulltext":[{"header":"INTRODUCTION","content":"\u003cp\u003eDesert ecosystems are an important component of global terrestrial ecosystems, playing a prominent role in maintaining global ecological balance and serving the entire human population (Liang et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). As the dominant plant species in desert ecosystems, shrubs may profoundly influence multiple ecosystem functions (ecosystem multifunctionality) through multidimensional ecological processes (Yue et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Currently, global desert ecosystems are undergoing widespread scrub encroachment (Deng et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), leading to mutations in the desert vegetation diversity, productivity, and soil nutrients, accompanied by transformations in ecosystem multifunctionality (Karaoud \u0026amp; Noumi, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Zhang et al., \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Zhang et al., \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). However, the mechanisms by which shrubs drive these changes and alter both the singular and multifaceted aspects of ecosystem functionality remain poorly understood (Ding et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Liu et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eShrubs play a significant role in soil nutrient dynamics, plant composition, and ecological interactions between vegetation and soil within desert ecosystems (Thomas et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). In desert environments, shrubs are often regarded as nurse plants, providing a stable and temperate growth environment for understory herbaceous plants, thereby enhancing their richness, diversity, and productivity (La Montagna et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). However, in grassland ecosystems, shrub encroachment often leads to decline of herbaceous richness, diversity, and biomass, which ultimately resulting grassland degradation (Valencia et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Thus, the effect of shrubs on herbaceous plants may differ with ecosystem types. In semi-arid regions, shrubs improve soil moisture conditions and soil nutrients availability beneath the canopy through the \u0026ldquo;fertility island\u0026rdquo; effect, enhancing spatial heterogeneity of soil nutrients (Yao et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). During years with lower precipitation, the positive effect of shrubs on soil moisture becomes more pronounced, helping mitigate the negative impacts of drought on plants (Liu et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2020a\u003c/span\u003e). These soil and vegetation changes induced by shrubs further influence ecosystem functions. For example, shrub encroachment increases soil activity in tundra ecosystem, enhancing nutrient cycling functions (Aguirre et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2021\u003c/span\u003e); In the Tibetan Plateau region, leguminous shrubs enhance the multifunctionality of grassland ecosystems, closely related to improved soil nutrients (Yang et al., \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In desert ecosystems, shrubs facilitated the development of their associated herbaceous communities via accumulating more soil resources (Xie et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In addition, ecosystem multifunctionality was positively related to plant diversity (Wang et al., \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Shrubs exert positive effects on plant diversity, productivity, and soil nutrients in desert ecosystems, suggesting their influence may also be beneficial on desert ecosystem multifunctionality, though this requires further research and practical validation. Desert ecosystems also exhibit significant environmental heterogeneity, making the role of shrubs in desert ecosystem functions is highly susceptible to soil moisture and salinity conditions. The extent of these effects may vary with soil moisture fluctuations (Butterfield et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Wang et al., \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Consequently, the role of soil environment also cannot be overlooked in the process of shrub impacts on desert ecosystem multifunctionality.\u003c/p\u003e \u003cp\u003eThe positive effects of shrubs on ecosystem functions may diminish with increasing drought stress. In forest ecosystems, reduced soil moisture caused by human activities diminishes the effect of plant diversity on ecosystem multifunctionality (Zhao et al., \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). In alpine meadow and grassland ecosystems, the positive effect of shrubs on multiple ecosystem functions greatly depended on increasing soil nutrient (Yang et al., \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). This influence of soil moisture and nutrient on ecosystem multifunctionality has also been confirmed in alpine ecosystems (Yang et al., \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Plants and soil, as two primary drivers of ecosystem function, the effects of plant diversity, productivity, and soil nutrients on ecosystem multifunctionality are also synergistic and complementary (Delgado-Baquerizo et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Changes in the soil environment alter the effect of shrubs on plant diversity, productivity, and soil nutrients, potentially affecting shifts in ecosystem multifunctionality. In arid regions, drought stress weakens the positive influence of plant diversity on ecosystem multifunctionality (Bhadauria et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The relationship between plant diversity and ecosystem multifunctionality may vary along environmental gradients (Fanin et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Most studies indicate that the effect of plant diversity on ecosystem multifunctionality decreases with increasing environmental stress, though some reports suggest no change or even an increase (Hu et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In arid regions, Maestre et al. demonstrated that shrubs enhance ecosystem function by mitigating the negative impacts of drought stress (Maestre et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Given the multidimensional nature of ecosystem functionality, the effect mechanisms of shrubs plants on the multifunctionality of desert ecosystems may exhibit complex variations along water-salt gradients (Xu et al., \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), though our understanding remains limited.\u003c/p\u003e \u003cp\u003eNowadays, the intensification of extreme drought climate events has become a global reality (Huang et al., \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). However, research on the effects of shrub plants on the multifunctionality of desert ecosystems remains scarce, and the roles of desert shrubs in the single function and multifunctionality of the ecosystem are still unclear. Therefore, this study takes the arid desert ecosystem of the Ebinur Lake Basin as the research subject and explores the effect pathways of different desert shrub plants on various functions and multifunctionality of the ecosystem at drought stress gradients. It aims to address the following questions: (1) How do desert shrubs affect herbaceous plants and soil environments? (2) What are the pathways through which shrubs influence single or multiple desert ecosystem functions? Addressing these questions will enhance the predictability of desert ecosystem functions and services under changing environmental conditions.\u003c/p\u003e"},{"header":"MATERIAL AND METHODS","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy site\u003c/h2\u003e \u003cp\u003eThe Ebinur Lake Wetland National Nature Reserve (44\u0026deg;30\u0026prime;-45\u0026deg;09\u0026prime;N, 82\u0026deg;36\u0026prime;-83\u0026deg;50\u0026prime;E) is located in northwest Jinghe County, Bortala Mongolian Autonomous Prefecture, Xinjiang, China, in the lowest depression in the southwest margin of Junggar Basin. The region has a dry typical continental climate with an annual precipitation of only about 89.80\u0026ndash;169.70 mm and strong potential annual evaporation is higher than precipitation, which ranges from 1569\u0026ndash;3421mm. The soil parent material is sandy soil, grey-brown desert soil, and grey desert soil, which are severely affected by desertification. The main vegetation types in the study area include \u003cem\u003ePopulus euphratica\u003c/em\u003e, \u003cem\u003eHaloxylon ammodendron\u003c/em\u003e, \u003cem\u003eHalimodendron halodendron\u003c/em\u003e, \u003cem\u003eNitraria tangutorum\u003c/em\u003e, \u003cem\u003eAlhagi sparsifolia\u003c/em\u003e, \u003cem\u003eReaumuria songarica\u003c/em\u003e, \u003cem\u003eCalligonum mongolicum Turcz\u003c/em\u003e, \u003cem\u003eApocynum Venetum\u003c/em\u003e, \u003cem\u003ePhragmites australis\u003c/em\u003e, \u003cem\u003eSuaeda glauca\u003c/em\u003e.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eExperimental design\u003c/h3\u003e\n\u003cp\u003eIn July 2024, on the north side of the Aqikesu River within the Ebinur Lake, the locations approximately 1km, 2.5km and 4.5km from the Aqikesu River were selected as high water and salt environment (HSW), medium water and salt environment (MSW) and low water and salt environment (LSW), respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Each water-salt environment included 3 replicated plots (30\u0026times;30) and separated by approximately 20m, each plot was further divided into 9 smaller sample plots (10m\u0026times;10m), totaling 81 plots. Shrub environment (SE) and without shrub environment (WE) were set up in each smaller sample plots. The understory environment beneath shrub canopies served as shrub environment (SE), while bare ground areas constituted without shrub environment (WE).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e\n\u003ch3\u003eField sampling and measurements\u003c/h3\u003e\n\u003cp\u003eRecord the quantity, height, coverage, base diameter of shrub species and the species name, quantity, height (CH), base diameter (D) and coverage of herbaceous plants in each square (10\u0026times;10). Then, we subsequently harvested herbaceous plant materials at ground level for aboveground biomass (AGB) determination. Collect soil samples from 0\u0026ndash;20 cm depth under canopy and in open areas. Determine soil moisture content using the oven-drying method. Soil samples were screened through a 2mm sieve to remove roots and stones for the measurements of soil physicochemical properties. Soil electrical conductivity (EC) was determined using conductivity meter; soil organic carbon (SOC) content by the potassium dichromate-sulfate oxidation method (Nelson \u0026amp; Sommers, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1982\u003c/span\u003e); soil total nitrogen (TN) by the Kjeldahl method (Lynch \u0026amp; Barbano, \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e1999\u003c/span\u003e); soil total phosphorus (TP) by the molybdenum antimony colorimetric method (Murphy \u0026amp; Riley, \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e1962\u003c/span\u003e); soil available phosphorus (AP) by sodium bicarbonate extraction-molybdenum-antimony anti-spectrophotometric method; soil ammonium nitrogen (NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e-N) and nitrate nitrogen (NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e-N) by ultraviolet spectrophotometry.\u003c/p\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eData analysis\u003c/h2\u003e \u003cp\u003eThe calculation formula for the relative importance value (IV) of shrubs:\u003cdiv id=\"Equa\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equa\" name=\"EquationSource\"\u003e\n$$\\:IV=(RD+RC+RF)/3$$\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003eWhere RD represents the relative density of the shrub, RC represents the relative coverage of the shrub, and RF represents the relative frequency of the shrub.\u003c/p\u003e \u003cp\u003eWe used the Species richness index (R), Shannon-Wiener diversity index (H), and Pielou evenness index (J) as our plant diversity indicators, which can be calculated as follows:\u003c/p\u003e \u003cp\u003eSpecies richness index:\u003cdiv id=\"Equb\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equb\" name=\"EquationSource\"\u003e\n$$\\:R=S$$\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003eShannon-Wiener index:\u003cdiv id=\"Equc\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equc\" name=\"EquationSource\"\u003e\n$$\\:H=-{\\sum\\:}_{i=1}^{s}pi{ln}pi$$\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003ePielou index:\u003cdiv id=\"Equd\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equd\" name=\"EquationSource\"\u003e\n$$\\:J=H/{ln}S$$\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003eWhere S represents the total number of plant species, and Pi represents the importance value (IV) of the ith species.\u003c/p\u003e \u003cp\u003eTo evaluate the differences in ecosystem functions between shrub and bare land, we assessed the ecosystem multifunctionality (EMF) index based on six specifically chosen individual ecosystem functions (ABG、SOC、SW、D、CH、TP), which were selected following a thorough analysis using the Random Forest method (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The EMF was calculated using two methods, averaging approach and multiple-threshold approach. We further classified these individual ecosystem functions into three functional groups: (1) functions relating to diversity (R, H, and J), (2) plant productivity (AGB、CH and D), (3) soil nutrients (SOC、TN、NH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e-N、NO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e-N、TP、AP). The EMF index assessing using averaging approach that can be calculated as:\u003cdiv id=\"Eque\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Eque\" name=\"EquationSource\"\u003e\n$$\\:EMF=\\frac{1}{N}{\\sum\\:}_{i=1}^{N}Zij$$\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003eWhere N is the total number of selected ecosystem plots and Zij represents the standardization of the evaluated ith ecosystem function at the jth plot.\u003c/p\u003e \u003cp\u003eIn addition, the ecosystem multifunctionality index, calculated using the multiple-threshold approach (Byrnes et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), is determined by counting the number of functions that simultaneously surpass a specified threshold of the maximum observed value for a given function across all study plots. As any choice of a threshold, that determines whether a given ecosystem function contributes to multifunctionality, is likely arbitrary, three different thresholds (25%, 50%, and 75%) were used, thus representing a wide spectrum in the analyses (Delgado-Baquerizo et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). This approach overcomes the weakness of the average multifunctionality method.\u003c/p\u003e \u003cp\u003eWe used independent samples t-tests were employed to analyze and evaluate significant differences in ecosystem functions under shrub canopies in desert ecosystems. Structural equation modeling (SEM) was employed to construct a pathway diagram of ecosystem multifunctionality drivers. Generalized linear mixed models (GLMM) were applied to analyze the multivariate explained variance of ecosystem functions along the water-salt gradient and the individual effects of its drivers. All statistical analyses were processed using SPSS and R software, with Origin software utilized for graphical representation.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"RESULTS","content":"\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eCharacteristics of shrubs in different water-salt environments and their effects on herbaceous plants\u003c/h2\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe composition of shrubs and their importance values vary with different water-salinity environments. HSW is dominated by \u003cem\u003eHalimodendron halodendron\u003c/em\u003e, with a relative importance value (IV) of 0.7126. While MSW is primarily composed of \u003cem\u003eAlhagi sparsifolia\u003c/em\u003e and \u003cem\u003eReaumuria songarica\u003c/em\u003e, with IV of 0.6113 and 0.6044, respectively. Shrubs of LSW mainly are \u003cem\u003eAlhagi sparsifolia\u003c/em\u003e and \u003cem\u003eHaloxylon ammodendron\u003c/em\u003e, with IV of 0.6417 and 0.5148, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea-c). HSW refers to tall shrubs, MSW are short shrubs, and shrubs of LSW are tall but have a smaller crown spread (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ed-f).\u003c/p\u003e \u003cp\u003eWhen comparing the shrubs canopy with the bare land, shrubs significantly reduced herbaceous species richness but increased aboveground biomass of herbaceous plants in desert ecosystems. Shrubs decreased the Shannon-Wiener index of LSW herbaceous plants while increasing the Pielou index of HSW herbaceous plants. Shrubs increased plant height but reduced their basal diameter of MSW herbaceous plants (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eEffects of shrubs on soil properties under different water-salt environments\u003c/h3\u003e\n\u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003ePhysical and chemical properties of soil under shrub canopy and bare land in different water and salt environments\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEnvironment\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eVariables\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSE\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eWE\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eP value\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"6\" rowspan=\"7\"\u003e \u003cp\u003eHSW\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSW\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e14.16\u0026thinsp;\u0026plusmn;\u0026thinsp;1.56\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e10.51\u0026thinsp;\u0026plusmn;\u0026thinsp;1.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003eP\u0026thinsp;\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSOC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e1.66\u0026thinsp;\u0026plusmn;\u0026thinsp;0.41\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e1.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003eP\u0026thinsp;\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e1.65\u0026thinsp;\u0026plusmn;\u0026thinsp;0.60\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.97\u0026thinsp;\u0026plusmn;\u0026thinsp;0.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003eP\u0026thinsp;\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e-N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.004\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.004\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eP\u0026thinsp;=\u0026thinsp;0.994\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e-N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.006\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.007\u0026thinsp;\u0026plusmn;\u0026thinsp;0.008\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eP\u0026thinsp;=\u0026thinsp;0.382\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.65\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.63\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eP\u0026thinsp;=\u0026thinsp;0.236\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eP\u0026thinsp;=\u0026thinsp;0.056\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"6\" rowspan=\"7\"\u003e \u003cp\u003eMSW\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSW\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e7.61\u0026thinsp;\u0026plusmn;\u0026thinsp;0.90\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e5.46\u0026thinsp;\u0026plusmn;\u0026thinsp;0.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003eP\u0026thinsp;\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSOC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.36\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.33\u0026thinsp;\u0026plusmn;\u0026thinsp;0.15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eP\u0026thinsp;=\u0026thinsp;0.447\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.56\u0026thinsp;\u0026plusmn;\u0026thinsp;0.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.40\u0026thinsp;\u0026plusmn;\u0026thinsp;0.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003eP\u0026thinsp;=\u0026thinsp;0.015\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e-N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.004\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.003\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eP\u0026thinsp;=\u0026thinsp;0.103\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e-N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.008\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.008\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eP\u0026thinsp;=\u0026thinsp;0.982\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.46\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.42\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003eP\u0026thinsp;=\u0026thinsp;0.003\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.09\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003eP\u0026thinsp;\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"6\" rowspan=\"7\"\u003e \u003cp\u003eLSW\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSW\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.83\u0026thinsp;\u0026plusmn;\u0026thinsp;0.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.96\u0026thinsp;\u0026plusmn;\u0026thinsp;0.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003eP\u0026thinsp;=\u0026thinsp;0.002\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSOC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.09\u0026thinsp;\u0026plusmn;\u0026thinsp;0.34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003eP\u0026thinsp;\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.45\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.32\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003eP\u0026thinsp;\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNH\u003csub\u003e4\u003c/sub\u003e\u003csup\u003e+\u003c/sup\u003e-N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.002\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0008\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.002\u0026thinsp;\u0026plusmn;\u0026thinsp;0.004\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eP\u0026thinsp;=\u0026thinsp;0.967\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNO\u003csub\u003e3\u003c/sub\u003e\u003csup\u003e\u0026minus;\u003c/sup\u003e-N\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.008\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.007\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eP\u0026thinsp;=\u0026thinsp;0.577\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.34\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.30\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003eP\u0026thinsp;\u0026lt;\u0026thinsp;0.001\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAP\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e0.02\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.03\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eP\u0026thinsp;=\u0026thinsp;0.064\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"5\"\u003eNote: Data are presented as mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard deviation. The significance level for independent samples t-tests was set at 0.05, with significant values shown in bold. HSW, the high water-salt environment; MSW, the medium water-salt environment; LSW, the low water-salt environment. SE, shrub environment; WE, without shrub environment; SW, soil water content; SOC, soil organic carbon; TN, soil total nitrogen; NH₄⁺-N, soil ammonium nitrogen; NO₃⁻-N, soil nitrate nitrogen; TP, soil total phosphorus; AP, soil available phosphorus.\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eAs shown in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, with shrub canopies showing relatively higher TN than bare land in desert ecosystems. The SW under shrub canopies exceeded bare land (p\u0026thinsp;\u0026lt;\u0026thinsp;0.001) in HSW and MSW, and the SOC under shrub canopies was significantly higher than bare land in HSW and LSW (p\u0026thinsp;\u0026lt;\u0026thinsp;0.001). In the MSW, both TP (p\u0026thinsp;=\u0026thinsp;0.003) and AP (p\u0026thinsp;\u0026lt;\u0026thinsp;0.001) under shrub canopies were higher than bare land. Relatively higher TP (p\u0026thinsp;\u0026lt;\u0026thinsp;0.001) but lower SW (p\u0026thinsp;=\u0026thinsp;0.002) under shrub canopies than bare land in LSW. Along with the water-salinity gradient, the positive effect of shrubs showed an increasing trend for TP, while their positive effect exhibited a decreasing trend for SW. Even in LSW, this effect was reversed.\u003c/p\u003e\n\u003ch3\u003eResponse of individual and multiple ecosystem functions to desert shrub\u003c/h3\u003e\n\u003cp\u003e \u003c/p\u003e \u003cp\u003eThe EMF indexes varied greatly across water-salt gradient, The mean EMF index decreased along water and salt environment. Based on two calculation methods of multifunctionality, shrubs significantly enhanced the EMF, EMF-productivity, and EMF-nutrient indexes of HSW and MSW, and reduced the EMF-diversity indexes of LSW. However, the EMF indexes and EMF-nutrient indexes calculated by two methods show differences. After evaluating the EMF and the EMF-nutrient through the multiple-threshold method, it is observed that selecting the 25% threshold results in shrubs enhanced the EMF and the EMF-nutrient of LSW (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e).\u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eEffect of shrub traits and environmental factors on individual and multiple ecosystem functions\u003c/h2\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eShrub cover (E\u0026thinsp;=\u0026thinsp;0.03, p\u0026thinsp;=\u0026thinsp;0.013) and height (E\u0026thinsp;=\u0026thinsp;0.02, p\u0026thinsp;=\u0026thinsp;0.035) both promote the multifunctionality of desert ecosystems. Shrub cover positively influenced soil moisture content (E\u0026thinsp;=\u0026thinsp;0.20, p\u0026thinsp;\u0026le;\u0026thinsp;0.001) and soil nutrient function (E\u0026thinsp;=\u0026thinsp;0.16, p\u0026thinsp;=\u0026thinsp;0.008); shrub height directly negatively affected plant diversity (E=-0.26, p\u0026thinsp;\u0026le;\u0026thinsp;0.001) (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ea-d). The effect of individual ecosystem functions on multifunctionality varies significantly across water-salinity gradients. Plant productivity exerted the strongest influence on ecosystem multifunctionality, while the effect of soil nutrient diminished along the water-salinity gradient (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ee-g). In desert ecosystems, shrubs primarily enhance multifunctionality by increasing soil nutrient content.\u003c/p\u003e \u003c/div\u003e"},{"header":"DISCUSSION","content":"\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eDesert shrubs exert positive effects on plant productivity and soil nutrient content in non-extreme environments\u003c/h2\u003e \u003cp\u003eThe research findings indicate that the composition and growth of desert shrubs vary with the water-salt environment. Shrub species richness and growth performance of MSW was lower than in other water and salt environment. This was primarily due to drought and salt stress in reducing height, basal diameter, and species richness of shrubs (Gatica et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Shi et al., \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Shrub cover exhibited heightened sensitivity to moisture fluctuations, with drought stress being the primary factor why significantly higher shrub cover of HSW than MSW and LSW (Chunyuan et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The relative importance of \u003cem\u003eAlhagi sparsifolia\u003c/em\u003e and \u003cem\u003eHaloxylon ammodendron\u003c/em\u003e were significantly higher than other shrubs, owing to their stronger drought-tolerant genetic endowment (Gao et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Consequently, the inherent tolerance of shrub species themselves is also one of the reasons for the differences in shrub composition along the water and salt environment.\u003c/p\u003e \u003cp\u003eThe presence or absence of shrubs significantly influences changes in soil nutrient content within desert ecosystems (Johnson et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), The research results also confirmed that desert shrubs increase the content of SOC, TN, and TP under their canopy, while having no significant effect on soil nutrient availability. This finding indicates that the increase in soil nutrient content under shrub canopies primarily originates from external sources. Shrubs deposit these nutrients into the understory soil through litterfall and canopy interception (Ding \u0026amp; Eldridge, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), thereby increasing the nutrient content in the soil. However, this positive effect may change along with the variations in the water and salt environment of desert ecosystems. Soil moisture content under shrub canopies was lower than bare land in LSW, which is consistent with studies from other desert ecosystems (Butterfield et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Under moderate drought stress, shrubs enhance soil water retention by reducing evaporation and providing canopy shade, thereby increasing SW beneath their canopies (Noumi et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Additionally, In desert ecosystems, shrubs improve the local environment to provide relatively better conditions for herbaceous plants, promoting their biomass (Franco et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Zhang et al., 2025a). The finding that AGB under shrub canopies was significantly higher than bare land further corroborated the conclusion. As shrubs promote the accumulation of soil organic matter and nutrients beneath their canopies, enabling sustained access to growth resources for understory plants (Weber-Grullon et al., \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Shrubs may also limit grazing by herbivores through their canopy structure, thereby increasing AGB of herbaceous plants. However, plant richness on bare land is significantly higher than plant community shrub canopies. as shrubs expand, they simultaneously suppress the growth of some plants (Sun et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Shrub canopy setting up physical barriers to prevent the entry of some plants (Kang et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). and shading amplifies their negative effect on plant richness (Xu et al., \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), leading to a decline in herbaceous plant richness, this aligns with global grassland ecosystem research (Wieczorkowski \u0026amp; Lehmann, \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The intensification of environmental stress increases competition between shrubs and herbaceous plants for survival resources (Shi et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), thereby the inhibitory effect of shrubs on plant diversity intensifies along water-salt gradients. Together, desert shrubs reduced herbaceous species diversity while promoting biomass growth. This change correlates with effect of shrubs on soil nutrient content.\u003c/p\u003e \u003cp\u003eThe effect of shrubs on soil and herbaceous plants subsequently influences both the single-function and multifunctional aspects of ecosystems (Chen et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). In desert ecosystems, shrubs increase plant productivity and soil nutrients, which aligns with the view that shrubs, as nurse plants in arid areas, enhance ecosystem functions. within a certain range of shrub canopy cover (5% \u0026minus;\u0026thinsp;45%) (Liancourt et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), greater canopy cover exhibits stronger positive effects on understory plant productivity and soil nutrients (liu et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2020b\u003c/span\u003e). In the study area, shrubs decrease plant diversity, consistent with studies on desert shrub invasions and grassland vegetation succession (Hector et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Wu et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Research indicates that desert shrubs have a positive effect on ecosystem multifunctionality, and this effect diminishes along water-salt gradients. Which consistent with recent studies from the stress gradient hypothesis (Zhang et al., \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). This shift occurs primarily due to extreme drought stress and the reduction shrub canopy cover, which cause the relationship between shrubs and understory herbaceous plants to shift toward competition (Sun et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The effects of shrubs on plant diversity, plant productivity, and soil nutrients drive changes in ecosystem multifunctionality.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eDesert shrubs enhance ecosystem multifunctionality by increasing soil nutrient\u003c/h2\u003e \u003cp\u003eIn desert ecosystems, we found that plant diversity exerts a positive influence on ecosystem multifunctionality, though its effect is not significant. This is contrary to the prevailing view that plant diversity is generally considered the primary driver of ecosystem multifunctionality (Li et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2025b\u003c/span\u003e; Ma et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). However, from another perspective, When measured by plant richness and density, plant diversity effectively predicts trends in ecosystem multifunctionality under precipitation variations in desert grasslands, exhibiting a significant positive correlation with ecosystem multifunctionality (Hu et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Additionally, the research also revealed a significant negative effect of shrub height on plant diversity. Along with the water-salinity gradient, the influence of plant diversity on ecosystem multifunctionality first decreased and then increased, which aligns with the nonlinear relationship between plant diversity and ecosystem multifunctionality across changing of environmental gradients (Liu et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Changes in water and salt levels trigger changes in many environmental variables, impacting on the distribution and diversity of plant species and thus largely determine the stability of the ecosystem in the study area (Zou et al., \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe effect of shrubs on ecosystem multifunctionality is also reflected in the changes of plant productivity. Observations in this study that plant productivity positively influences ecosystem multifunctionality support this assertion. Our results show that aboveground biomass of understory herbaceous is significantly higher than that of bare lands, though shrubs did not significantly enhance plant productivity directly. In fact, the positive effect of shrubs on the productivity of herbaceous plants stems from the fact that the canopy of shrubs provides relatively favorable soil conditions for plant growth (Thakur \u0026amp; Kumar, \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), thereby indirectly promoting positive effect of plant productivity on ecosystem multifunctionality. Furthermore, the positive effect of plant diversity on plant productivity was not significant, contradicting the general view that plant richness correlates positively with plant productivity (Tilman et al., 2014). Shrub cover promoted plant productivity, while shrub height exerted a negative effect on plant diversity. In desert ecosystems, plant productivity and diversity are controlled by different traits of shrub, potentially differentiating their relationship (Sanaei et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). In the study area, the effect of plant productivity on the multifunctionality of the ecosystem is the greatest, but no significant variation across the gradient. Soil nutrients and moisture content, as primary indirect drivers of plant productivity, alleviated the effect of water-salinity fluctuations on plant productivity (La Pierre et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), thereby stabilizing the positive role of plant productivity in ecosystem multifunctionality.\u003c/p\u003e \u003cp\u003eThe shrub canopy plays a pivotal role in the effect of shrubs on ecosystem functions (soil nutrients) (Lu et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Maestre et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Compared to bare lands, shrubs significantly improve soil nutrient status, further validating this point. Soil nutrients directly promote ecosystem multifunctionality while also indirectly influencing it by enhancing plant productivity. This is the same mechanism by which soil nutrient loss leads to the decline of ecosystem multifunctionality during land degradation (Li et al., 2025a). Furthermore, Soil microorganisms play a crucial role in the changing mechanisms of ecosystem multifunctionality in arid regions (Zhang et al., 2025b; Zhou et al., 2025), therefore, necessitating investigation into their role in the process of ecosystem multifunctionality variation in desert ecosystems. The establishment of herbaceous plants significantly increases soil nutrient content in arid areas (Liu et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), providing evidence that plant productivity enhances soil nutrient content. Higher levels of plant biomass are more likely to increase soil nutrients, with the plant litter input and accumulation contributing to nutrient sequestration and soil moisture retention(Liu et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2021b\u003c/span\u003e; Ward et al., \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). In summary, the mutual promotion of plant productivity and soil nutrients primarily benefits from a temperate microenvironment created by shrub canopy in desert ecosystems, providing stable conditions for a positive feedback loop between plants and soil. Both plant functional traits and soil nutrients are equally important for the variation in ecosystem multifunctionality on the water-salt gradient.\u003c/p\u003e \u003cp\u003eIn summary, this research results provide clear evidence that shrub facilitates ecosystem functioning of desert ecosystem, which had a stronger facilitative effect in HSW and MSW than LSW. However, we recognize there are still limitations in the present study. We intentionally focused on the shrub induced changes in the herbaceous community, soil properties, and ecosystem functioning of desert ecosystem. Previous studies have demonstrated that shrubs positively influence soil microbial activity and abundance (Liu et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2021a\u003c/span\u003e). Therefore, it is necessary to consider the role of soil microorganisms in the ecological impact of desert shrubs. Elevated temperature or altered precipitation have a significant impact on multiple ecosystem functions in desert regions. This suggests that ongoing climate change and anthropogenic activity may mediate future effectiveness of shrub encroachment (Tu et al., \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Wang et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e"},{"header":"CONCLUSIONS","content":"\u003cp\u003eResearch indicates that shrub vegetation significantly alters the plant community structure and soil environment of desert ecosystems, enhancing their multifunctionality. Shrubs increase soil nutrient content and herbaceous aboveground biomass while reducing herbaceous species richness. Furthermore, the effect of shrubs on desert ecosystem functions varies along water-salinity gradients. Desert shrubs enhance plant productivity and soil nutrients of HSW and MSW (SW: 5%~12%, EC: 11\u0026thinsp;~\u0026thinsp;27 ms/cm) while reducing plant diversity of LSW. This research provides crucial insights into the interactions between shrub vegetation characteristics and ecosystem functions in desert ecosystems, offering decision-making guidance for desert ecosystem utilization and management. Given the threat of global climate change, widespread shrub encroachment in desert ecosystems has become a critical issue. Enhancing our ability to predict the impacts of these phenomena on the structure and function of global desert ecosystems is paramount. Addressing these challenges at larger spatio-temporal scales to improve our capacity to understand, mitigate, or plan for the effects of such vegetation changes on desert ecosystem functions requires further empirical research and modeling.\u003c/p\u003e "},{"header":"Declarations","content":"\u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003eCRediT authorship contribution statement\u003c/h2\u003e \u003cp\u003e \u003cb\u003eShuangfu Zhou\u003c/b\u003e: Writing \u0026ndash; original draft, Methodology, Investigation, Software, Experiment, Data curation. \u003cb\u003eXiaohui Li\u003c/b\u003e: Investigation, Software, Experiment, Data curation. \u003cb\u003eHuixia Li\u003c/b\u003e: Investigation, Experiment, Data curation. \u003cb\u003eYufei Chen\u003c/b\u003e: Investigation, Experiment, Data curation. \u003cb\u003eXueni Zhang (corresponding author)\u003c/b\u003e: Writing \u0026ndash; review \u0026amp; editing, Methodology, Supervision, Funding acquisition.\u003c/p\u003e \u003c/div\u003e\u003cp\u003e \u003ch2\u003eDeclaration of competing Interest\u003c/h2\u003e \u003cp\u003eThe authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eAcknowledgments\u003c/h2\u003e \u003cp\u003eThis work was supported by the Natural Science Foundation of China (32360277).\u003c/p\u003e\u003ch2\u003eData availability\u003c/h2\u003e \u003cp\u003eData will be made available on request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAguirre D, Benhumea AE, McLaren JR (2021) Shrub encroachment affects tundra ecosystem properties through their living canopy rather than increased litter inputs. 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Proc Natl Acad Sci USA, 122(41): e2511071122\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZou J, Luo Y, Seidl R, Thom D, Liu J, Geres L, Richter T, Ye L, Zheng W, Ma L, Song J, Xu K, Li D, Gao L, Seibold S (2024) No generality in biodiversity-productivity relationships along elevation in temperate and subtropical forest landscapes. \u003cem\u003eForest Ecosystems\u003c/em\u003e, 11.\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://10.1016/j.fecs.2024.100187\u003c/span\u003e\u003cspan address=\"http://10.1016/j.fecs.2024.100187\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"plant-and-soil","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"plso","sideBox":"Learn more about [Plant and Soil](https://www.springer.com/journal/11104)","snPcode":"11104","submissionUrl":"https://submission.nature.com/new-submission/11104/3","title":"Plant and Soil","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"shrub encroachment, desert ecosystem, individual ecosystem function, ecosystem multifunctionality, water and salt environment","lastPublishedDoi":"10.21203/rs.3.rs-9474137/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9474137/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e\u003cb\u003eAims\u003c/b\u003e\u003c/p\u003e \u003cp\u003eShrub encroachment in desert ecosystems is projected to intensify worldwide because of rapid climate change and anthropogenic activity, leading to significant alterations of ecosystem functions. The study investigates the effects of shrub on various ecosystem functionality relevant to plant diversity, productivity, and soil nutrients under different water and salt environment in desert ecosystems.\u003c/p\u003e\u003cp\u003e\u003cb\u003eMethods\u003c/b\u003e\u003c/p\u003e \u003cp\u003eSoil environments are classified into high, medium, and low water and salt environments based on soil moisture content and electrical conductivity. Differences in soil and plant traits between the understory and bare within sample plots across different water and salt environments were compared. The effects of shrubs on individual ecosystem functions and ecosystem multifunctionality, and their water and salt responses, are analyzed using averaging and multiple-threshold approach. The drivers of shrub influence on ecosystem multifunctionality are analyzed using generalized linear mixed model (GLMM).\u003c/p\u003e\u003cp\u003e\u003cb\u003eResults\u003c/b\u003e\u003c/p\u003e \u003cp\u003eOur findings suggest that desert shrub increased herbaceous aboveground biomass and Soil organic carbon (SOC), total nitrogen (TN), total phosphorus (TP), the positive effects of desert shrubs on ecosystem multifunctionality varied with water-salinity conditions. Shrubs increased plant productivity and soil nutrients for high (HSW) and medium water and salt environment (MSW) while reducing plant diversity for low environment (LSW). The enhancement of ecosystem multifunctionality by desert shrubs was closely related to the promotion of soil nutrient content and plant productivity.\u003c/p\u003e\u003cp\u003e\u003cb\u003eConclusion\u003c/b\u003e\u003c/p\u003e \u003cp\u003eUnderstanding the effect of desert shrubs on ecosystem functions will help us screen native species from a functional perspective for ecological restoration, thereby enhancing the ecological benefits of plant-based remediation.\u003c/p\u003e","manuscriptTitle":"Positive effects of desert shrub in ecosystem multifunctionality primarily depend on increasing soil nutrients","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-05-11 11:42:12","doi":"10.21203/rs.3.rs-9474137/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewersInvited","content":"","date":"2026-05-03T18:31:18+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Plant and Soil","date":"2026-04-27T12:10:33+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-04-27T10:44:01+00:00","index":"","fulltext":""},{"type":"submitted","content":"Plant and Soil","date":"2026-04-26T23:31:49+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"plant-and-soil","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"plso","sideBox":"Learn more about [Plant and Soil](https://www.springer.com/journal/11104)","snPcode":"11104","submissionUrl":"https://submission.nature.com/new-submission/11104/3","title":"Plant and Soil","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"e1808b61-e53a-4ff5-8d7f-33effbf8b4ef","owner":[],"postedDate":"May 11th, 2026","published":true,"recentEditorialEvents":[{"type":"reviewersInvited","content":"","date":"2026-05-03T18:31:18+00:00","index":"","fulltext":""}],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-05-11T11:42:13+00:00","versionOfRecord":[],"versionCreatedAt":"2026-05-11 11:42:12","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9474137","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9474137","identity":"rs-9474137","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}