{"paper_id":"03e9e5de-be04-4254-8b6f-69f9dc950714","body_text":"The Soil pH and Micronutrients Drive Ageratina Adenophora Invasion in Areas with Acidic and Nutrient-Poor Soils | 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 The Soil pH and Micronutrients Drive Ageratina Adenophora Invasion in Areas with Acidic and Nutrient-Poor Soils Juan Wang, Jingying Lu, Yuehua Zhang, Xianyong Dong, Xiaogang Wu, and 3 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8012262/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 5 You are reading this latest preprint version Abstract Ageratina adenophora poses significant threats to agricultural and forestry production and biodiversity conservation worldwide. However, its expansion mechanism in highly acidic environments has not been studied in depth. To address this issue, we investigated the impacts of A. adenophora invasion on soil nutrient content and enzyme activity levels across 24 samples from Yunnan Province, China. The sampling sites were categorized into four invasion levels, namely, noninvaded (C), lightly invaded (L), moderately invaded (M), and severely invaded (S). Our findings revealed that the soil pH in severely invaded areas was 8.37% greater than that in noninvaded areas, with pH values ranging from 4.8 to 5.3. Notably, severely invaded soils demonstrated relatively high levels of soil organic carbon (SOC), available potassium (AK), aluminum (Al), available iron (Fe), available zinc (Zn), available copper (Cu), available manganese (Mn), exchangeable calcium (Ca), and exchangeable magnesium (Mg). However, the levels of available Al, boron (B), and phosphorus were significantly lower in these areas. Additionally, variations in the total nitrogen (TN), total potassium (TK), sucrase, and nitrate reductase activity levels were observed across the areas with different invasion levels. Correlation analysis underscored the pivotal role of pH in regulating soil nutrient availability and microbial activity levels. The random forest model (RFM) and structural equation modeling (SEM) results indicated that available Mn, AK, and available Zn are the dominant factors in noninvaded areas (p < 0.05), while Mg, B, and available Cu were the main factors in severely invaded areas (p < 0.05). These findings collectively demonstrate that A. adenophora invasion establishes favorable habitat conditions by altering soil pH, nutrient cycling, and enzyme activity levels, thereby suppressing the growth of native plants and ultimately displacing them from their ecological niches. This study provides a sound foundation for the formulation of stage-specific control strategies against A. adenophora invasion. Level of invasion Ageratina adenophora soil pH soil nutrients soil enzyme activity structural equation model Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 1. Introduction Invasive alien plants pose a significant threat to global ecological security, with the potential to reduce regional biodiversity and drastically alter the structure and function of natural ecosystems (Yang et al., 2013 ; Xu et al., 2023 ). Owing to intensified global climate change and increased international exchanges, the spread of invasive plants continues to increase (Osunkoya and Perrett, 2011 ; Ren et al., 2021 ). Such plants compete with native flora for essential resources, including water, light, and soil nutrients (Jones et al., 2019 ; Puissant et al., 2019 ; DeForest and Moorhead, 2020 ). A substantial body of evidence indicates that invasive species can alter microbial community composition and soil enzyme activity through root secretions, litter decomposition, and allelopathy, thus creating nutrient environments favorable for their proliferation (Esch et al., 2017 ; Ahmad Dar et al., 2023 ; Guo et al., 2023 ). Changes in soil environments subsequently enhance the competitive advantage of these species, thereby influencing the dynamics between exotic and native plants (Esch et al., 2017 ; Ahmad Dar et al., 2023 ; Guo et al., 2023 ). The complex feedback relationships between invasive alien plants and soil nutrient cycling continue to attract broad research interest (Gasch et al., 2013 ; Wu et al., 2020 ; Kumar et al., 2021 ). Ageratina adenophora , a perennial herbaceous or semishrubby plant native to Mexico, is recognized globally as a highly pernicious invasive species (Wu et al., 2020 ; Kumar et al., 2021 ). This plant exhibits significant environmental adaptability and has been demonstrated to substantially increase soil fertility after prolonged establishment (Zhao et al., 2019 ; Zhang et al., 2023 ). It has been demonstrated that A. adenophora can mitigate unfavorable soil conditions, thereby increasing its uptake of essential nutrients such as carbon (C), nitrogen (N), and phosphorus (P) (Bojórquez-Quintal et al., 2017 ; Ahmad Dar et al., 2023 ). This, in turn, accelerates its growth and development (Zhao et al., 2019 ; Verma et al., 2023 ). In addition to macronutrients, A. adenophora requires various micronutrients for optimal growth and development of physiological functions. Notably, nutrient deficiency can significantly impair health (Bojórquez-Quintal et al., 2017 ; Wu et al., 2023 ). Compared with native flora, invasive species typically exhibit accelerated growth and increased biomass, necessitating greater intake of micronutrients and consequently intensifying the activation or depletion of soil micronutrient pools (Pizzeghello et al., 2011 ; Wu et al., 2023 ). For example, invasion by Acacia dealbata and Lantana camara L. significantly increases soil calcium (Ca) and magnesium (Mg) levels while altering the concentrations of iron (Fe), copper (Cu), manganese (Mn), and sulfur (S) (Osunkoya and Perrett, 2011 ; Pizzeghello et al., 2011 ; Kaur et al., 2012 ; Souza-Alonso et al., 2014 ). These findings suggest that there may occur positive feedback between these two plants and soil micronutrients, in turn facilitating their invasion (Esch et al., 2017 ; Kim et al., 2018 ; Puissant et al., 2019 ). Unfortunately, research on soil micronutrient dynamics during A. adenophora invasion, particularly in nutrient-poor soils, remains scarce (Bursali et al., 2009 ; Wu et al., 2020 ; Chen et al., 2022 ). The soil pH is closely associated with nutrient biogeochemical processes and availability (Zhao et al., 2019 ; Xiao et al., 2023 ). Under acidic soil conditions, salt-based ions such as Zn, Mg, and Ca are readily displaced by acid ions (H + and Al 3+ ), resulting in their loss through leaching or precipitation events (Merry et al., 2002 ; Bojórquez-Quintal et al., 2017 ). Concurrently, Mo can form iron phosphate precipitates, which results in a reduction in availability (Jones et al., 2019 ; Wu et al., 2020 ). High acidity can even lead to aluminum and manganese toxicity issues, which inhibits plant growth. In response to low soil pH values, plants effectively neutralize or buffer soil acidity during growth by promoting ammonification reactions in the soil, thus inhibiting nitrification reactions and regulating organic components within their own tissues. (Xiao et al., 2017 ; Zhao et al., 2019 ). Previous studies have indicated that A. adenophora may exhibit a similar ability (Bills et al., 2010 ; Kim et al., 2018 ), which becomes increasingly important as invasion deepens or expands over time. For example, Xiao et al. ( 2017 ) revealed that pH promoted the colonization of A. adenophora , thus increasing soil fertility, as indicated by elevated organic carbon (C org ), nitrate (NO 3 − ), ammonium (NH 4 + ), available potassium (AK), and available phosphorus (AP) contents. Additionally, Jones et al. ( 2019 ) indicated that pH is the main factor regulating the carbon utilization efficiency, and Puissant et al. ( 2019 ) reported that pH can affect enzyme activity, which can increase the activities of key soil enzymes such as urease (SUE), phosphatase, and invertase (Zhao et al., 2019 ). Moreover, Wu et al. ( 2020 ) demonstrated that the soil pH and enzyme activity levels were notably greater in areas invaded by A. adenophora than in uninvaded areas (Wu et al., 2020 ). Nevertheless, the precise manner in which increasing the soil pH contributes to changes in soil nutrient contents to match the increased invasion of A. adenophora remains unclear (DeForest and Moorhead, 2020 ; Lhamo et al., 2023 ). Starting during the 1940s, the range of A. adenophora increased from Burma to Yunnan Province in China, and this invasive plant rapidly proliferated throughout Southwest China. Presently, A. adenophora has emerged as one of the most formidable threats to biodiversity and agricultural production in inland China (Pizzeghello et al., 2011 ; Fan et al., 2022 ). Pinus yunnanensis , an endemic tree species in Southwest China, dominates the regional forested landscape. These forests, which mainly encompass homogeneous stands, are characterized by notable soil acidification (with soil pH values typically ranging from 4 to 6) (Merry et al., 2002 ; Kabała and Łabaz, 2018 ) underneath the canopy and limited soil nutrient availability, which significantly restricts plant growth (Esch et al., 2017 ; Darji et al., 2021 ). Interestingly, over the past two decades, field surveys have identified Pinus yunnanensis forests as primary sites for A. adenophora invasion, and invasive populations continue to expand, posing a serious threat to forest regeneration and subcanopy biodiversity. (Wang et al., 2015 ; Sun et al., 2021; Zhang et al., 2022 ). Although a few studies have revealed changes in soil nutrient levels and interspecific associations (Hu et al., 2021 ; Xiao et al., 2023 ; Niu et al., 2007 ; Liu et al., 2023 ) after A. adenophora invasion into Pinus yunnanensis forests, the nutrient dynamics throughout the spread process have not yet been explored in depth. In our study, we considered different invasion levels of A. adenophora in Pinus yunnanensis forests to explore changes in the levels of major soil nutrients, micronutrients, and enzyme activity. Specifically, we aimed to determine (1) whether the soil pH increases with increasing invasion level; (2) how the contents of major soil micronutrients change with increasing invasion level; and (3) the mechanisms through which changes in soil pH and micronutrient levels may facilitate increased invasion. We aimed to enhance the understanding of the expansion mechanisms of A. adenophora in areas with acidic soil conditions and to provide a theoretical basis for developing preventive and control strategies. 2. Materials and Methods 2.1 Study area The research site is located in Jiyi town, Wuding County, Yunnan Province, Southwest China, at coordinates of 26°6′N and 102°14′E. Jiyi town is situated on the banks of the Jinsha River and exhibits an average elevation of 1900 m (Fig. 1 ). The study area experiences a dry-hot valley climate characterized by abundant sunshine, high evaporation rates and an average annual temperature of 15.2°C. Precipitation occurs mainly from May to October, with annual totals ranging from 800 to 1000 mm (Wu et al., 2020 ). Previous studies have identified the soil type at the research site as dry red soil (Chen et al., 2013 ). Specifically, the sampling site is located in the upper-middle part of a Pinus yunnanensis forest with slope gradients ranging from 25° to 40° and faces northeast. The average height of this forest is approximately 10.5 m, and the average diameter at breast height is approximately 13.5 cm. The canopy density levels range from 0.6 to 0.9 units per unit area (m²). Resident surveys indicate that A. adenophora has been invading the understory of this forest for more than two decades. 2.2 Sampling method In August 2023, the extent of A. adenophora invasion was quantified by assessing the relative projected area covered by this species with reference to previous studies (Fig. 1 ). Sample plots were selected randomly and categorized into four invasion levels on the basis of the percentage of the area covered by A. adenophora , namely, noninvaded (C, 0.2%-1.2%), lightly invaded (L, 25%-35%), moderately invaded (M, 50%-70%), and severely invaded (S, 88%-97%) (Table 1 ) (Xiao et al., 2017 ; Wu et al., 2020 ; Zhang et al., 2023 ). For each invasion level, six plots were established, each measuring 1×1 m, characterized by similar topographic features and canopy closure conditions. The plots were spaced at least 20 m apart to avoid spatial overlap. The plant composition varied according to the invasion level (refer to Table 1 for details). Table 1 Distribution of plants in the A. adenophora sample plots under different invasion levels. Invasion level Community overview Coverage of A. adenophora % Noninvaded (C) Pogonatherum crinitum Thunb.; Artemisia lavandulifolia DC.; Grona heterocarpos L.; Scutellaria indica L.; Cynoglossum amabile Stapf & Drummond; Pararuellia delavayana Baill.; Paspalum scrobiculatum Var.; Polygala persicariifolia DC.; Dodonaea viscosa Jacquem.; Origanum vulgare L.; Reinwardtia indica Dumort. 0.68 ± 0.35 Lightly invaded (L) Ageratina adenophora Spreng.; Artemisia lavandulifolia DC.; Ainsliaea spicata Vaniot, Pogonatherum crinitum Thunb.; Clinopodium megalanthum Diels; Oxalis corniculata L.; Dioscorea subcalva Prain & Burkill 29.83 ± 3.39 Moderately invaded (M) Ageratina adenophora Spreng.; Elsholtzia rugulosa Hemsl.; Reinwardtia indica Dumort.; Pogonatherum crinitum Thunb.; Grona heterocarpos L.; Pimpinella candolleana Wight & Arn.; Arisaema erubescens Wall. 60 ± 6.51 Severely invaded (S) Ageratina adenophora Spreng.; Elsholtzia rugulosa Hemsl.; Reinwardtia indica Dumort.; Pogonatherum crinitum Thunb. 92.67 ± 3.04 To assess soil properties, aboveground vegetation and litter were removed in each plot, and five soil samples were collected from the top 20 cm with a soil auger (5-cm diameter) following the five-point sampling method. The samples were thoroughly mixed, transported to the laboratory, and sieved through a 2-mm mesh. The sieved samples were then divided into two portions: one was dried naturally for the analysis of soil physicochemical properties and micronutrient contents, while the other was refrigerated at a temperature of 4°C for two weeks to analyze soil enzyme activity and alkaline dissolved nitrogen levels. 2.3 Soil analysis Soil physicochemical properties encompass a comprehensive range of parameters, including the soil pH, soil organic carbon (SOC), total nitrogen (TN), total phosphorus (TP), total potassium (TK), available nitrogen (AN), available phosphorus (AP), and AK (Xiao et al., 2017 ). In addition, the analysis was extended to soil total microelements, such as Fe, Mn, zinc (Zn), Cu, boron (B), Ca, Mg, and aluminum (Al) (Osunkoya and Perrett, 2011 ; DeForest and Moorhead, 2020 ), as well as soil available microelements, including available Fe, available Mn, available Zn, available Cu, available B, exchangeable Ca, exchangeable Mg, and available Al (Osunkoya and Perrett, 2011 ; Fan et al., 2022 ). Furthermore, the activity of soil enzymes, including SUE, alkaline phosphatase (ALP), sucrase (SSC), nitrate reductase (NR), and catalase (CAT) activities, was evaluated (Kim et al., 2018 ; Zhang et al., 2022 ). In this study, macronutrient indices were determined according to the method of Xiao et al. ( 2017 ), micronutrient indices were determined according to the methods of Osunkoya and Perrett ( 2011 ), DeForest and Moorhead ( 2020 ) and Fan et al. ( 2022 ), and enzyme activity indices were determined according to the methods of Kim et al. ( 2018 ) and Zhang et al. ( 2022 ). These parameters collectively provide a robust framework for assessing soil health and nutrient dynamics in the studied ecosystem. 2.4 Data analysis In this study, the experimental data were systematically analyzed and organized in Microsoft Excel 2021. The normality of the data distribution was evaluated via SPSS 25.0. Graphical representations of the soil pH, SOC, nitrogen (N), phosphorus (P), potassium (K), and enzyme activity data were generated with the ggplot2 package in R version 4.2.2. The soil micronutrient data were analyzed via one-way analysis of variance (ANOVA) using SPSS 25.0. Additionally, correlation analysis, random forest model (RFM) analysis and structural equation modeling (SEM) were performed in R, thereby employing the piecewiseSEM, corrplot, and pacman packages, respectively. In SEM analysis, a P value greater than 0.05 indicated that all path coefficients in the model were less than 1, with lower values of Fisher's exact test statistic, the Akaike information criterion (AIC), and the Bayesian information criterion (BIC) indicating a better model fit (Zhao et al., 2019 ). This comprehensive analytical approach ensured robust data interpretation and provided a solid foundation for understanding the relationships between soil properties and their influence on the studied ecosystem. 3. Results 3.1 Effects of different levels of A. adenophora invasion on soil physicochemical properties As shown in Fig. 2 , significant changes ( P < 0.05) were observed in the soil pH, TN, TK, and AP levels across the areas with varying degrees of A. adenophora invasion compared with those in noninvaded areas. Specifically, the soil pH increased with increasing invasion level (Fig. 2 a), with increases of 1.34%, 3.19%, and 8.37% in lightly, moderately, and severely invaded areas, respectively, relative to those in noninvaded areas. The soil TN content significantly fluctuated ( P < 0.05) with increasing invasion level, with a 35.38% increase in lightly invaded areas, a 10.73% decrease in moderately invaded areas, and a 9.59% increase in severely invaded areas (Fig. 2 b). Conversely, the soil TK content increased with increasing invasion level, peaking (12.04 g/kg) in severely invaded areas. In contrast to the aforementioned parameters, the AP content significantly decreased ( P < 0.05) with increasing invasion level, decreasing by 47.24%, 39.08%, and 40.76% in lightly, moderately, and severely invaded areas, respectively (Fig. 2 g, P < 0.05). Notably, although not statistically significant, the SOC content increased with increasing invasion level. 3.2 Effects of different levels of A. adenophora invasion on soil micronutrient content and availability levels The mean concentrations of soil trace elements under different invasion levels are provided in Table 2 . Although the differences in micronutrient concentrations across invasion levels did not reach statistical significance ( P > 0.05), certain trends were observed. Except boron (B), which exhibited the highest concentration in noninvaded areas, the remaining seven micronutrients demonstrated increasing trends following A. adenophora invasion, with increases ranging from 0.99% to 18.08%. Specifically, the maximum concentrations of Zn, Ca, Mg, and Al occurred in severely invaded areas, whereas Fe and Cu peaked in moderately invaded areas. The highest Mn concentration only occurred in lightly invaded areas among all the sampled areas. Table 2 Effects of micronutrients on A. adenophora under different invasion levels. Different levels of invasion Mean ± SE Fe mg/kg Mn mg/kg Zn mg/kg Cu mg/kg B mg/kg Ca mg/kg Mg mg/kg Al mg/kg C 21.2 ± 0.6a 902.33 ± 119.08a 44.47 ± 0.95 27.88 ± 0.45a 86.55 ± 5.57a 1.82 ± 0.21a 2.6 ± 0.1a 64.37 ± 1.29a L 20.98 ± 1.03a 1065.47 ± 163.62a 44.1 ± 1.01a 27.73 ± 1.04a 73.48 ± 4.99a 1.7 ± 0.22a 2.52 ± 0.08a 66.55 ± 2.29a M 21.41 ± 0.51a 822.4 ± 106.57a 45.32 ± 0.87a 29.85 ± 1.01a 81.32 ± 4.39a 1.54 ± 0.11a 2.56 ± 0.1a 66.05 ± 3.96a S 20.77 ± 1.09a 948.38 ± 128.17a 44.82 ± 0.98a 29 ± 1.31a 78.62 ± 6.18a 2.01 ± 0.16a 2.72 ± 0.11a 70.71 ± 3.5a With respect to the availability of soil micronutrients (Table 3 ), significant differences ( P < 0.05) were observed in available Mn, Zn, and Cu contents across the areas with different invasion levels, with an overall increasing trend. The concentrations of available Mn, Zn, and Cu in severely invaded areas were 45.48%, 98.52%, and 72.87% higher, respectively, than those in noninvaded areas. Moreover, although the differences in available Fe, Ca, and Mg concentrations did not reach statistical significance (P > 0.05), the values generally peaked in severely invaded areas. Conversely, compared with that in noninvaded areas, the available Al concentration decreased by 19.04%, 25.06%, and 18.56% with increasing invasion levels. Table 3 Effects of different levels of A. adenophora invasion on available micronutrients. Different levels of invasion Mean ± SE available Fe mg/kg available Mn mg/kg available Zn mg/kg available Cu mg/kg available B mg/kg Exchangeable Ca mg/kg Exchangeable Mg mg/kg available Al mg/kg C 40.22 ± 5.05a 64.54 ± 5.18b 0.22 ± 0.02b 0.57 ± 0.03c 0.1 ± 0.03a 3.16 ± 0.18a 1.15 ± 0.09a 52.07 ± 5.63a L 45.24 ± 2.89a 76.62 ± 3.99b 0.31 ± 0.08ab 0.7 ± 0.04bc 0.12 ± 0.03a 4.24 ± 0.75a 1.26 ± 0.22a 42.16 ± 4.95a M 42.72 ± 8.41a 68.43 ± 7.91b 0.25 ± 0.05ab 0.78 ± 0.08b 0.13 ± 0.03a 4 ± 0.18a 1.13 ± 0.02a 39.02 ± 7.23a S 44.71 ± 1.97a 93.89 ± 3.25a 0.44 ± 0.09a 0.98 ± 0.08a 0.1 ± 0.03a 4.8 ± 0.69a 1.37 ± 0.19a 42.41 ± 3.72a Note: C, noninvaded; L, lightly invaded; M, moderately invaded; S, severely invaded. 3.3 Effects of different levels of A. adenophora invasion on soil enzymes The invasion of A. adenophora significantly affected soil SSC and NR activity levels ( P < 0.05), whereas the impacts on soil SUE, CAT, and ALP activity levels were relatively minor ( P > 0.05). In severely invaded areas, soil sucrase activity reached its peak value (26.09 mg/kg), whereas moderately invaded areas exhibited the lowest activity (12.90 mg/kg). Compared with those in noninvaded areas, lightly invaded areas exhibited comparable soil sucrase activity levels (Fig. 3 b). Compared with those in noninvaded areas, soil nitrate reductase activity levels increased by 4.85% and 0.76% in lightly and moderately invaded areas, respectively, but decreased by 4.67% in severely invaded areas (Fig. 3 c). Additionally, soil CAT (Fig. 3 d) and ALP (Fig. 3 e) activity levels increased with increasing invasion level. 3.4 Determination of the primary factors driving coverage change under different levels of A. adenophora invasion This study revealed the complex interactions among soil trace elements, enzyme activity, and nutrient dynamics through comprehensive correlation analysis of soil enzymatic activity and physicochemical properties (Fig. 4 ). The results revealed significant correlations ( P < 0.05) among various soil trace elements. Notably, TK was significantly positively correlated with B, Mg, and available Fe ( P < 0.05) but significantly negatively correlated with AK ( P < 0.05). TP was significantly positively correlated with AP and TN ( P < 0.05) but significantly negatively correlated with available Ca and available Mg ( P < 0.05). Furthermore, with respect to activity, significant positive correlations ( P < 0.05) were observed among four soil enzymes (SSC, SUE, NR, and ALP). Importantly, the soil pH was significantly positively correlated with the SOC, SSC, ALP, CAT, AK, available Cu, and available Mn levels ( P < 0.05). The RFM, which is a robust machine learning tool, was employed in this study to systematically identify and predict the key environmental drivers influencing the invasion and spread of A. adenophora . Our model results revealed significant variations in the driving factors influencing the invasion coverage of A. adenophora across the areas with different invasion levels (Fig. 5 ). Specifically, in noninvaded areas (Fig. 5 a), available Mn, AK, available Zn, available B, and Zn were identified as the most significant factors influencing the invasion coverage of A. adenophora ( P < 0.05). In lightly invaded areas (Fig. 5 b), Al, available Al, available Fe, available Mg, and available Ca emerged as the primary drivers ( P < 0.05). In moderately invaded areas (Fig. 5 c), Cu, Mg, Al, B, available Zn, available Cu, available Ca, and available Mn significantly affected the invasion coverage of A. adenophora ( P < 0.05). In severely invaded areas (Fig. 5 d), Mg, B, Zn, Fe, available Cu, and available Al were identified as the key drivers influencing invasion coverage ( P < 0.05). 3.5 What are the pathways or mechanisms that contribute to the increase in the invasion coverage of A. adenophora ? SEM analysis was employed to elucidate the significant and nonsignificant pathways through which soil properties influence the coverage of A. adenophora (A. adenophora) under varying invasion levels, providing critical insights into its invasion mechanisms. In this study, we observed a significant linear increase in the soil pH following A. adenophora invasion ( P < 0.05), indicating that its inclusion was an independent variable in the model (Fig. 2 a). The key soil factors identified through the RFM were selected as observed variables, while the aboveground coverage of A. adenophora served as the dependent variable (Table 1 ). The results revealed distinct response pathways of soil physicochemical properties for A. adenophora across the different invasion levels (Fig. 6 ). Specifically, in noninvaded areas, pH indirectly influenced A. adenophora coverage by regulating the availability of soil nutrients (R² = 0.7), with AK emerging as a critical driver ( P < 0.05). In lightly invaded areas, soil nutrients directly and significantly affected A. adenophora coverage (R² = 0.69), with a path coefficient of 0.64 ( P < 0.05). In moderately invaded areas, both pH and soil nutrient content indirectly promoted A. adenophora coverage by influencing the availability of soil nutrients (R² = 0.52). Notably, the effects of soil nutrients (Al) and available nutrients (available Ca and available Cu) were significantly greater than that of pH, with path coefficients of 0.38 ( P < 0.05) and 0.96 ( P < 0.001), respectively. In severely invaded areas, both available nutrients and soil nutrients significantly influenced A. adenophora coverage (R² = 0.91), with path coefficients of 0.63 ( P < 0.001) and 0.46 ( P < 0.01), respectively. Importantly, B, Fe, and available Al played particularly significant roles in influencing A. adenophora coverage in severely invaded areas, highlighting the critical role of soil properties at advanced invasion stages. 4. Discussion 4.1 A. adenophora invasion increases the soil pH and alters the levels of soil macronutrients The soil water content and pH significantly influence plant growth and development (Puissant et al., 2019; Guo et al., 2022). Invasive plants modify soil pH conditions through changes in soil physical properties, such as increased water uptake and root infiltration (Niu et al., 2007; Xiao et al., 2017; Puissant et al., 2019). For example, Zhang and Suseela (2021) reported that the soil pH was significantly lower (pH = 5.3) in fertilized and invaded areas than in noninvaded areas (pH = 5.8) and other invaded areas (pH = 6.1). Additionally, invasive species have been shown to regulate the soil pH and alter the nutrient status, with Xia et al. (2022) demonstrating that significant releases of soil cations in invaded areas led to an increase in the soil pH due to the consumption of hydrogen ions (Bojórquez-Quintal et al., 2017; Fan et al., 2022; Wang et al., 2019; Chen et al., 2022; Marchante et al., 2023). The habitats in our study, which are located in Pinus yunnanensis forests, are characterized by high soil acidification (Xiao et al., 2017; Wu et al., 2020). We investigated whether A. adenophora can leverage its characteristics to promote invasion by improving habitat conditions. Our findings indicated that invasion significantly altered the soil pH in invaded areas, with a notable difference between severely invaded areas (with pH values ranging from approximately 5.4–6.2) and noninvaded areas (with pH values ranging from approximately 4.8–5.3) (Jones et al., 2019). Furthermore, research from Kashmir Himalaya has indicated that the soil pH in invaded areas is significantly greater than that in noninvaded areas after invasion (Kumar et al., 2021; Ahmad Dar et al., 2023; Dar et al., 2023), confirming that A. adenophora can increase the soil pH. In highly acidic and alkaline soils, invasive plants modify their physicochemical properties, affecting nutrient availability, which is crucial for rapid growth and development (Osunkoya and Perrett, 2011; Ren et al., 2021; Yang et al., 2013; Souza-Alonso et al., 2014). The growth of A. adenophora relies on N- and P-enriched soils, leading to significant reductions in soil TN and AP concentrations in invaded areas (Khatri et al., 2023; Zhang and Suseela, 2021). This trend is supported by the gradual increase in the SOC content observed with increasing invasion level in our study (Sun et al., 2019; McLeod et al., 2021), as shown in Fig. 2e. Additionally, studies have indicated that plant invasion alters soil enzyme activity levels, thus affecting nutrient effectiveness (Zhang et al., 2022). Changes in soil properties and plant diversity due to A. adenophora invasion have also been observed in Punakha, Bhutan, and in evergreen broad-leaved forests in southwestern Yunnan Province, China (Lhamo et al., 2023; Zhao et al., 2019; Zhang et al., 2023). Our findings revealed that the soil TN content is significantly greater in lightly invaded areas than in severely invaded areas, whereas the soil TK content shows the opposite pattern (Zhang and Suseela, 2021). This difference may stem from the distinct nutrient requirements of A. adenophora ; notably, in lightly invaded areas, A. adenophora may demand more TK for rapid growth, whereas in severely invaded areas, it may need more TN to increase biomass and soil organic matter (Chen et al., 2022; Cotrufo et al., 2022). 4.2 Invasion of A. adenophora significantly affects soil available micronutrient contents Soil micronutrients, which are essential for plant growth, are effectively used by invasive species such as A. adenophora for growth and development (Liu et al., 2023). The relationship between soil enzyme activity and micronutrient levels often results in synergistic or antagonistic interactions with plants, influencing micronutrient uptake (Niu et al., 2007; Kaur et al., 2012; Jones et al., 2019). In this study, we investigated the preferences of A. adenophora for specific micronutrients and revealed that all micronutrients except available Al occur at higher concentrations in invaded areas than in noninvaded areas, with Mn, Zn, and Cu exhibiting significant variability across the areas with different invasion levels (Pizzeghello et al., 2011; Wu et al., 2023). Excessive or insufficient soil micronutrient levels can be lethal to plants (Osunkoya and Perrett, 2011; Marchante et al., 2023). Recent research has indicated that A. adenophora can absorb copper ions from aquatic environments, acting as an adsorbent (Fan et al., 2022). Additionally, A. adenophora requires substantial amounts of boron (B) for growth, and B deficiency can be fatal to native plants (Bursali et al., 2009; Chen et al., 2021). Our findings also revealed a decreasing trend in the soil B content with increasing invasion level, whereas the Al content increased (Jones et al., 2019). Previous research has demonstrated that reduced Fe and available Al levels in soil can lead to P loss (Pizzeghello et al., 2011). Consistent with these findings, our study revealed lower AP, Fe, and Al levels in invaded areas than in uninvaded areas (Chapuis-Lardy et al., 2006; Lin et al., 2023), which conforms with findings of Wu et al. (2020). There is consensus that changes in soil micronutrient levels impact soil properties, a phenomenon also observed in areas invaded by Lantana camara L. (Osunkoya and Perrett, 2011; Datta et al., 2017). Our results demonstrated that A. adenophora affects soil micronutrient levels as invasion progresses, particularly those of Mn, Zn, Al, B, Fe, and Cu (Wang et al., 2015; Shen et al., 2019; Souza-Alonso et al., 2014; Jones et al., 2019; Lin et al., 2023). This adaptation enables A. adenophora to alter soil properties and manipulate nutrient cycling, thus suppressing the development of native plants (Yang et al., 2013; Xiao et al., 2017; Fan et al., 2022). 4.3 Correlations between soil enzyme activity and soil nutrient levels altered by A. adenophora invasion Soil enzyme activity is significantly correlated with both the ambient temperature and soil pH (Esch et al., 2017; Kim et al., 2018). Puissant et al. (2019) reported that the optimal pH for enzyme activity did not always coincide with the environmental soil pH. It is widely acknowledged that invasive plants can modify soil habitats, thereby altering soil properties and nutrient cycling via soil enzyme activities (Shen et al., 2019; Hu et al., 2021; Khatri et al., 2023). In our research, a strong correlation was observed between the soil pH and three soil enzymes, excluding nitrate reductase and urease (Fig. 4). Invasions by Amaranthus palmeri and Phragmites australis have been shown to significantly impact soil enzyme activity levels (Zhang et al., 2022; Kim et al., 2018). Similarly, our findings indicated that A. adenophora invasion significantly increased the activity of soil sucrase, catalase, and alkaline phosphatase (Wang et al., 2015; Puissant et al., 2019). However, changes in soil enzyme activity can be attributed to various mechanisms (Kim et al., 2018). Soil enzyme activity, which is influenced by microorganisms, root secretions, and the decomposition of organic material, plays crucial roles in shaping plant invasion dynamics (DeForest and Moorhead, 2020; Lhamo et al., 2023). In the A. adenophora community, variations in enzyme activities across different invasion levels are likely linked to root secretions and apoplastic materials (Wherry, 1920; Yang et al., 2013; Zhao et al., 2019). Our study demonstrated that soil urease activity decreased gradually with increasing invasion level, while alkaline phosphatase activity increased slowly (Fig. 3). These observations conform with the results of other studies, where invasions by Mimosa pudica and Falcataria moluccana led to significant increases in soil urease, catalase, and acid phosphatase activity levels (Wang et al., 2015; Esch et al., 2017; DeForest and Moorhead, 2020; Hu et al., 2021). Furthermore, A. adenophora invasion significantly increased the activities of soil sucrase, catalase, and alkaline phosphatase, contributing to a sharp decline in soil TN and AP contents (Sun et al., 2019; Wang et al., 2019), which agrees with findings for Mimosa pudica invasion (Wang et al., 2015). This suggests that optimizing the nutrient acquisition efficiency by regulating soil enzyme activity is a common strategy for successful invasion (Zhao et al., 2019; Zhang and Suseela, 2021; Zhang et al., 2022). 4.4 Soil pH and micronutrients are important factors for A. adenophora invasion in acid-poor areas A. adenophora invasion poses significant ecological threats, including biodiversity loss (Yang et al., 2013; Nunes et al., 2022; Guo et al., 2023) and adverse impacts on agriculture, forestry, and livestock health (Datta et al., 2017; Cotrufo et al., 2022). In the study area, the low soil pH under the canopy of Pinus yunnanensis has been associated with the increase in soil pH following A. adenophora invasion (Wu et al., 2020; Zhang et al., 2022). In this study, we integrated RFM and SEM analysis methods to systematically analyze the driving mechanisms of the influence of soil physicochemical properties on the aboveground coverage of A. adenophora during invasion (Xiao et al., 2017; Jones et al., 2019; Puissant et al., 2019). The results revealed that the invasion strategy of A. adenophora exhibits distinct stage-specific characteristics, with its success relying on dynamic adaptation and the regulation of soil environmental factors across different invasion stages (Ren et al., 2021; SUN et al., 2021). Previous studies have demonstrated that A. adenophora modulates the soil pH to suit its optimal acclimation range, thereby increasing the soil pH when it is less than 6 and decreasing it when it exceeds 7.5 (Kumar et al., 2021; Wu et al., 2020; Zhang and Suseela, 2021). This raises the question of whether stabilizing the soil pH could effectively control A. adenophora invasion (Niu et al., 2007; Shen et al., 2019; Xu et al., 2023). The RFM results revealed stage-specific differentiation of driving factors during A. adenophora invasion (Fig. 5) (Malik et al., 2018; Guo et al., 2023; Marchante et al., 2023). In noninvaded areas, available Mn, Zn, and B play dominant roles in shaping invasion coverage (p < 0.05), likely because of the rapid nutrient acquisition strategy of A. adenophora during early invasion (Zhang and Suseela, 2021; Guo et al., 2022; Marchante et al., 2023). At the light invasion stage, the increased significance of Al and available Fe (p < 0.05) suggests that A. adenophora mitigates aluminum toxicity by altering soil metal ion speciation while enhancing competitiveness through iron-mediated redox processes (Yang et al., 2013). At the moderate to severe invasion stages, the significant influence of Cu, Mg, and available Ca (p < 0.05) may be linked to the regulation of the soil cation exchange capacity (CEC) via root exudates to optimize the nutrient acquisition efficiency (Xiao et al., 2017). This dynamic shift in driving factors reflects the evolution of A. adenophora strategies from nutrient exploitation to habitat modification, which conforms closely with the positive feedback loop theory of invasive plants (Niu et al., 2007). This feedback mechanism may operate through the following pathways: (1) chelation, which reduces aluminum toxicity and facilitates boron-dependent cell wall synthesis (Chen et al., 2021), and (2) nutrient availability optimization through soil pH modification (Wu et al., 2020). These adaptive strategies enable A. adenophora to more efficiently absorb soil nutrients at severe invasion stages, thereby suppressing native plant growth and ultimately displacing them from their ecological niches (Marchante et al., 2023). Invasive plants disrupt native plant growth by modifying soil enzyme activity levels to favor their own development (Niu et al., 2007). SEM analysis revealed the synergistic effects of the soil pH and nutrient pathways (Fig. 6). In noninvaded areas, pH indirectly influences coverage by regulating available nutrients (R² = 0.7), with the key role of AK (path coefficient = 0.64, p < 0.05) likely related to the efficient use of potassium ion channels by A. adenophora during early invasion (Jones et al., 2019). With increasing invasion level, the direct effect of pH decreases, while the path coefficients of soil nutrients (e.g., Al and available Cu) significantly increase (p < 0.001), indicating that A. adenophora actively modifies the soil environment by altering rhizosphere microbial communities (e.g., acid-producing bacteria abundance) (Puissant et al., 2019; Lin et al., 2023). Particularly at severe invasion stages, the significant influences of available Al and B (path coefficient = 0.63, p < 0.001) may be attributed to their roles in reducing aluminum toxicity through chelation and promoting boron-dependent cell wall synthesis (Chen et al., 2021). This mechanism is particularly advantageous in regions with frequent acid rainfall (pH < 5.5) (Wu et al., 2020). By combining RFM and SEM analysis methods, we quantified, for the first time, the threshold effects of the nutrient competition-habitat modification dual-stage model during A. adenophora invasion (Malik et al., 2018; Puissant et al., 2019; Guo et al., 2023). The light invasion stage (R² = 0.69) is characterized primarily by direct nutrient competition, which can be mitigated through early intervention targeting available Fe and Al (Lin et al., 2023). In contrast, the severe invasion stage (R² = 0.91) requires targeted strategies to disrupt the biogeochemical cycles of trace elements such as Cu and B (Kumar et al., 2021). Notably, the active regulation of pH by A. adenophora (with postinvasion pH increases ranging from 0.3–0.9 units) may exacerbate the vicious cycle of soil acidification and nutrient loss (Dar et al., 2023), providing a critical target for invasion control in acidic soil regions. Although this study identified key driving factors and their pathways, the impacts of soil microbial functional groups and faunal disturbances on nutrient cycling remain to be incorporated into the model (Lhamo et al., 2023). Future research could aim to integrate metagenomics and stable isotope labeling techniques to elucidate the regulatory mechanisms of rhizosphere microbe‒plant interactions for the turnover of specific nutrients (e.g., available Cu) (Guo et al., 2023). Additionally, on the basis of the stage-specific thresholds established in this study, biochar- or chelator-driven targeted remediation technologies could be developed to assess the feasibility of inhibiting A. adenophora expansion by disrupting the Al‒Cu‒B cycles (Fan et al., 2022). 5. Conclusion In this study, we systematically investigated the dynamic changes and driving mechanisms of soil environmental factors in the invasion process of A. adenophora by integrating soil physicochemical properties, enzyme activities, and advanced modeling approaches, including RFM and SEM analysis methods. The results demonstrated that A. adenophora invasion significantly altered soil pH, nutrient content, and enzyme activity levels, with the driving factors exhibiting distinct stage-specific characteristics. Specifically, the soil pH increased with increasing invasion level, with an 8.37% increase in severely invaded areas compared with that in noninvaded regions ( P < 0.05). The soil TN and AP contents significantly fluctuated with invasion level, with the TN content increasing by 35.38% in lightly invaded areas and the AP content decreasing by 40.76% in severely invaded regions ( P < 0.05). Additionally, soil TK and micronutrients (e.g., Zn, Cu, and Mn) reached peak levels in severely invaded areas. Correlation analysis underscored the pivotal role of pH in regulating soil nutrient availability and microbial activity levels. The RFM results revealed significant differences in key driving factors across invasion stages: available Mn, AK, and available Zn were dominant in noninvaded areas ( P < 0.05), whereas Mg, B, and available Cu were the primary drivers in severely invaded regions ( P < 0.05). This highlights a shift in A. adenophora strategies from nutrient competition to habitat modification. SEM analysis was employed to quantify the synergistic effects of the soil pH and nutrient pathways and revealed that the path coefficient of available nutrients for coverage in severely invaded areas reached 0.63 ( P < 0.001), indicating that A. adenophora exacerbates invasion through plant‒soil feedback mechanisms. In summary, A. adenophora invasion establishes favorable habitat conditions by altering soil pH, nutrient cycling, and enzyme activity levels, thereby suppressing the growth of native plants and ultimately displacing them from their ecological niches. This study provides a systematic basis for the formulation of stage-specific control strategies against A. adenophora invasion. Early intervention through the regulation of the contents of available Fe and Al is recommended at the light invasion stage, whereas targeted strategies to disrupt the biogeochemical cycles of Mg, B, and available Cu are essential for managing severe invasion. Declarations Acknowledge This research was funded by the financial support of Sponsored by Funded Projects: Scientific research projects of China Three Gorges (Construction) Group Corporation (JG-EP-030222001 and JG-EP-030222002); Western Light Project of the Chinese Academy of Sciences (2022XBZG_XBQNXZ_A_003), Sichuan Science and Technology Program (2025ZNSFSC1017). CRediT author statement Juan Wang & Jingying Lu & Yuehua Zhang & Xianyong Dong & Xiaogang Wu & Lumei Xiao & Kaiwen Pan & Lin Zhang: Sample-plot Investigation, Conceptualization, Methodology, Software, Data curation, Visualization, Writing- Original draft preparation, Writing- Reviewing and Editing, sample-plot Investigation, Supervision, Revise, Writing- Reviewing and Editing, Funding acquisition. Declaration of interests ☒ 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. ☐ The authors declare the following financial interests/personal relationships which may be considered as potential competing interests: data availability statement Data will be made available upon request References Ahmad Dar, M., Ahmad, M., Singh, R., Kumar Kohli, R., Singh, H.P., Batish, D.R., 2023. Invasive plants alter soil properties and nutrient dynamics: A case study of Anthemis cotula invasion in Kashmir Himalaya. Catena 226. Bills, J.S., Jacinthe, P.-A., Tedesco, L.P., 2010. Soil organic carbon pools and composition in a wetland complex invaded by reed canary grass. Biology and Fertility of Soils 46, 697-706. Bojórquez-Quintal, E., Escalante-Magaña, C., Echevarría-Machado, I., Martínez-Estévez, M., 2017. Aluminum, a friend or foe of higher plants in acid soils. Frontiers in plant science 8, 1767. Bursali, E.A., Cavas, L., Seki, Y., Bozkurt, S.S., Yurdakoc, M., 2009. Sorption of boron by invasive marine seaweed: Caulerpa racemosa var. cylindracea. Chemical Engineering Journal 150, 385-390. Chapuis-Lardy, L., Vanderhoeven, S., Dassonville, N., Koutika, L.-S., Meerts, P., 2006. Effect of the exotic invasive plant Solidago gigantea on soil phosphorus status. Biology and Fertility of Soils 42, 481-489. Chen A Q, Zhang D, Peng H, et al., 2013. Experimental study on the development of colapse of ovemhanging lavers of auly in Yuanmou Valevchinaly. Gatena, 109, 177-185. Chen, L., Xia, F., Wang, M., Mao, P., 2021. Physiological and proteomic analysis reveals the impact of boron deficiency and surplus on alfalfa (Medicago sativa L.) reproductive organs. Ecotoxicology and Environmental Safety 214, 112083. Chen, S., Gao, D., Zhang, J., Müller, C., Li, X., Zheng, Y., Dong, H., Yin, G., Han, P., Liang, X., 2022. Invasive Spartina alterniflora accelerates soil gross nitrogen transformations to optimize its nitrogen acquisition in an estuarine and coastal wetland of China. Soil Biology and Biochemistry 174, 108835. Cotrufo, M.F., Haddix, M.L., Kroeger, M.E., Stewart, C.E., 2022. The role of plant input physical-chemical properties, and microbial and soil chemical diversity on the formation of particulate and mineral-associated organic matter. Soil Biology and Biochemistry 168, 108648. Dar, M.A., Ahmad, M., Singh, R., Kohli, R.K., Singh, H.P., Batish, D.R., 2023. Invasive plants alter soil properties and nutrient dynamics: A case study of Anthemis cotula invasion in Kashmir Himalaya. Catena 226, 107069. Darji, T.B., Adhikari, B., Pathak, S., Neupane, S., Thapa, L.B., Bhatt, T.D., Pant, R.R., Pant, G., Pal, K.B., Bishwakarma, K., 2021. Phytotoxic effects of invasive Ageratina adenophora on two native subtropical shrubs in Nepal. Scientific Reports 11, 13663. Datta, A., Kühn, I., Ahmad, M., Michalski, S., Auge, H., 2017. Processes affecting altitudinal distribution of invasive Ageratina adenophora in western Himalaya: The role of local adaptation and the importance of different life-cycle stages. PloS one 12, e0187708. DeForest, J.L., Moorhead, D.L., 2020. Effects of elevated pH and phosphorus fertilizer on soil C, N and P enzyme stoichiometry in an acidic mixed mesophytic deciduous forest. Soil Biology and Biochemistry 150, 107996. Esch, E.H., Lipson, D., Cleland, E.E., 2017. Direct and indirect effects of shifting rainfall on soil microbial respiration and enzyme activity in a semi-arid system. Plant and soil 411, 333-346. Fan, L., Miao, J., Yang, J., Zhao, X., Shi, W., Xie, M., Wang, X., Chen, W., An, X., Luo, H., 2022. Invasive plant-crofton weed as adsorbent for effective removal of copper from aqueous solution. Environmental Technology & Innovation 26, 102280. Gasch, C.K., Enloe, S.F., Stahl, P.D., Williams, S.E., 2013. An aboveground-belowground assessment of ecosystem properties associated with exotic annual brome invasion. Biology and Fertility of Soils 49, 919-928. Guo, K., Pyšek, P., Chytrý, M., Divíšek, J., Lososová, Z., van Kleunen, M., Pierce, S., Guo, W.Y., 2022. Ruderals naturalize, competitors invade: Varying roles of plant adaptive strategies along the invasion continuum. Functional Ecology 36, 2469-2479. Guo, K., Zheng, M.-M., Liu, R.-L., Wang, Y.-Y., Gao, Y., Shu, L., Wang, X.-R., Zhang, J., Guo, W.-Y., 2023. Intraspecific variations of adaptive strategies of native and invasive plant species along an elevational gradient. Flora 304, 152297. Hu, Z., Li, J., Shi, K., Ren, G., Dai, Z., Sun, J., Zheng, X., Zhou, Y., Zhang, J., Li, G., 2021. Effects of Canada goldenrod invasion on soil extracellular enzyme activities and ecoenzymatic stoichiometry. Sustainability 13, 3768. Jones, D.L., Cooledge, E.C., Hoyle, F.C., Griffiths, R.I., Murphy, D.V., 2019. pH and exchangeable aluminum are major regulators of microbial energy flow and carbon use efficiency in soil microbial communities. Soil Biology and Biochemistry 138, 107584. Kabała, C., Łabaz, B., 2018. Relationships between soil pH and base saturation-conclusions for Polish and international soil classifications. Soil Science Annual 69. Kaur, R., Malhotra, S., Inderjit, 2012. Effects of invasion of Mikania micrantha on germination of rice seedlings, plant richness, chemical properties and respiration of soil. Biology and Fertility of Soils 48, 481-488. Khatri, K., Negi, B., Bargali, K., Bargali, S.S., 2023. Phenotypic variation in morphology and associated functional traits in Ageratina adenophora along an altitudinal gradient in Kumaun Himalaya, India. Biologia 78, 1333-1347. Kim, S., Kang, J., Megonigal, J.P., Kang, H., Seo, J., Ding, W., 2018. Impacts of Phragmites australis invasion on soil enzyme activities and microbial abundance of tidal marshes. Microbial ecology 76, 782-790. Kumar, M., Kumar, S., Verma, A.K., Joshi, R.K., Garkoti, S.C., 2021. Invasion of Lantana camara and Ageratina adenophora alters the soil physico-chemical characteristics and microbial biomass of chir pine forests in the central Himalaya, India. Catena 207, 105624. Lhamo, S., Thinley, U., Dorji, U., 2023. Impacts of Invasion by Ageratina adenophora on Soil Properties and Plant Diversity. Bhutan Journal of Natural Resources and Development 10, 1-9. Li, Q., Wan, F., Zhao, M., 2022. Distinct soil microbial communities under Ageratina adenophora invasions. Plant Biology 24, 430-439. Li, Y.-P., Li, W.-T., Li, J., Feng, Y.-L., 2023. Temporal dynamics of plant− soil feedback and related mechanisms depend on environmental context during invasion processes of a subtropical invader. Plant and soil, 1-16. Lin, M., Chen, Y., Cheng, L., Zheng, Y., Wang, W., Sardans, J., Song, Z., Guggenberger, G., Zou, Y., Ding, X., 2023. Response of topsoil Fe-bound organic carbon pool and microbial community to Spartina alterniflora invasion in coastal wetlands. Catena 232, 107414. Liu, Y.-m., Li, W.-t., Zheng, Y.-l., 2023. The effect of acid rain and fertilization on the performance of invasive Chromolaena odorata and two native plants. Acta Oecologica 120, 103938. Malik, A.A., Puissant, J., Buckeridge, K.M., Goodall, T., Jehmlich, N., Chowdhury, S., Gweon, H.S., Peyton, J.M., Mason, K.E., van Agtmaal, M., 2018. Land use driven change in soil pH affects microbial carbon cycling processes. Nature communications 9, 3591. Marchante, H., Marchante, E., Verbrugge, L., Lommen, S., Shaw, R., 2023. Knowledge and perceptions of invasive plant biocontrol in Europe versus the rest of the world. Journal of Environmental Management 327, 116896. McLeod, M.L., Bullington, L., Cleveland, C.C., Rousk, J., Lekberg, Y., 2021. Invasive plant-derived dissolved organic matter alters microbial communities and carbon cycling in soils. Soil Biology and Biochemistry 156, 108191. Merry, R.H., Spouncer, L.R., Fitzpatrick, R.W., Davies, P.J., Bruce, D.A., McVicar, T., Rui, L., Walker, J., 2002. Regional prediction of soil profile acidity and alkalinity. ACIAR MONOGRAPH SERIES 84, 155-164. Niu, H.-b., Liu, W.-x., Wan, F.-h., Liu, B., 2007. An invasive aster (Ageratina adenophora) invades and dominates forest understories in China: altered soil microbial communities facilitate the invader and inhibit natives. Plant and soil 294, 73-85. Nunes, M., Lemley, D.A., Adams, J.B., 2022. Flow regime and nutrient input control invasive alien aquatic plant distribution and species composition in small closed estuaries. Science of the Total Environment 819, 152038. Osunkoya, O.O., Perrett, C., 2011. Lantana camara L.(Verbenaceae) invasion effects on soil physicochemical properties. Biology and Fertility of Soils 47, 349-355. Pizzeghello, D., Berti, A., Nardi, S., Morari, F., 2011. Phosphorus forms and P-sorption properties in three alkaline soils after long-term mineral and manure applications in north-eastern Italy. Agriculture, ecosystems & environment 141, 58-66. Puissant, J., Jones, B., Goodall, T., Mang, D., Blaud, A., Gweon, H.S., Malik, A., Jones, D.L., Clark, I.M., Hirsch, P.R., 2019. The pH optimum of soil exoenzymes adapt to long term changes in soil pH. Soil Biology and Biochemistry 138, 107601. Ren, Z., Okyere, S.K., Wen, J., Xie, L., Cui, Y., Wang, S., Wang, J., Cao, S., Shen, L., Ma, X., 2021. An overview: the toxicity of Ageratina adenophora on animals and its possible interventions. International Journal of Molecular Sciences 22, 11581. Shen, S., Xu, G., Li, D., Jin, G., Liu, S., Clements, D.R., Yang, Y., Rao, J., Chen, A., Zhang, F., 2019. Ipomoea batatas (sweet potato), a promising replacement control crop for the invasive alien plant Ageratina adenophora (Asteraceae) in China. Management of Biological Invasions 10, 559-572. Souza-Alonso, P., Novoa, A., González, L., 2014. Soil biochemical alterations and microbial community responses under Acacia dealbata Link invasion. Soil Biology and Biochemistry 79, 100-108. Sun, F., Ou, Q., Yu, H., Li, N., Peng, C., 2019. The invasive plant Mikania micrantha affects the soil foodweb and plant-soil nutrient contents in orchards. Soil Biology and Biochemistry 139, 107630. SUN, Y.-y., ZHANG, Q.-x., ZHAO, Y.-p., DIAO, Y.-h., GUI, F.-r., YANG, G.-q., 2021. Beneficial rhizobacterium provides positive plant-soil feedback effects to Ageratina adenophora. Journal of Integrative Agriculture 20, 1327-1335. Verma, A.K., Nayak, R., Manika, N., Bargali, K., Pandey, V.N., Chaudhary, L.B., Behera, S.K., 2023. Monitoring the distribution pattern and invasion status of Ageratina adenophora across elevational gradients in Sikkim Himalaya, India. Environmental Monitoring and Assessment 195, 152. Wang, C., Wu, B., Jiang, K., Zhou, J., Liu, J., Lv, Y., 2019. Canada goldenrod invasion cause significant shifts in the taxonomic diversity and community stability of plant communities in heterogeneous landscapes in urban ecosystems in East China. Ecological Engineering 127, 504-509. Wang, R., Dai, T., Quan, G., Zhang, J., 2015. Changes in soil physico-chemical properties, enzyme activities and soil microbial communities under Mimosa pudica invasion. Allelopathy J 36, 15-24. Wu, X., Duan, C., Fu, D., Peng, P., Zhao, L., Jones, D.L., 2020. Effects of Ageratina adenophora invasion on the understory community and soil phosphorus characteristics of different forest types in southwest China. Forests 11, 806. Wu, X., Xing, H., Wang, X., Yang, J., Chen, J., Liu, X., Dai, D., Zhang, M., Yang, Q., Dong, S., 2023. Changes in soil microbial communities are linked to metal elements in a subtropical forest. Applied Soil Ecology 188, 104919. Xia, H., Riaz, M., Liu, B., Li, Y., El-Desouki, Z., Jiang, C., 2022. Over two years study: Peanut biochar promoted potassium availability by mediating the relationship between bacterial community and soil properties. Applied Soil Ecology 176, 104485. Xiao, H., Schaefer, D.A., Yang, X., 2017. pH drives ammonia oxidizing bacteria rather than archaea thereby stimulate nitrification under Ageratina adenophora colonization. Soil Biology and Biochemistry 114, 12-19. Xiao, L., Min, X., Liu, G., Li, P., Xue, S., 2023. Effect of plant-plant interactions and drought stress on the response of soil nutrient contents, enzyme activities and microbial metabolic limitations. Applied Soil Ecology 181, 104666. Xu, C.-W., Yang, M.-Z., Chen, Y.-J., Chen, L.-M., Zhang, D.-Z., Mei, L., Shi, Y.-T., Zhang, H.-B., 2012. Changes in non-symbiotic nitrogen-fixing bacteria inhabiting rhizosphere soils of an invasive plant Ageratina adenophora. Applied Soil Ecology 54, 32-38. Xu, Z., Xu, J., Chen, P., Zhong, S., Xu, Z., Yu, Y., Wang, C., Du, D., 2023. Heavy metal pollution is more conducive to the independent invasion of Solidago canadensis L. than the co-invasion of two Asteraceae invasive plants. Acta Oecologica 120, 103934. Yang, Q., Carrillo, J., Jin, H., Shang, L., Hovick, S.M., Nijjer, S., Gabler, C.A., Li, B., Siemann, E., 2013. Plant-soil biota interactions of an invasive species in its native and introduced ranges: Implications for invasion success. Soil Biology and Biochemistry 65, 78-85. Zhang, M., Li, X., Qiu, Z., Shi, C., Wang, K., Fukuda, K., Shi, F., 2022. Effects of Amaranthus palmeri invasion on soil extracellular enzyme activities and enzymatic stoichiometry. Journal of Soil Science and Plant Nutrition 22, 5183-5194. Zhang, P., Nie, M., Li, B., Wu, J., 2017. The transfer and allocation of newly fixed C by invasive Spartina alterniflora and native Phragmites australis to soil microbiota. Soil Biology and Biochemistry 113, 231-239. Zhang, X., Wang, G., Peng, P., Zhou, Y., Chen, Z., Feng, Y., Wang, Y., Shi, S., Li, J., 2023. Influences of environment, human activity, and climate on the invasion of Ageratina adenophora (Spreng.) in Southwest China. PeerJ 11, e14902. Zhang, Z., Suseela, V., 2021. Nitrogen availability modulates the impacts of plant invasion on the chemical composition of soil organic matter. Soil Biology and Biochemistry 156, 108195. Zhao, M., Lu, X., Zhao, H., Yang, Y., Hale, L., Gao, Q., Liu, W., Guo, J., Li, Q., Zhou, J., 2019. Ageratina adenophora invasions are associated with microbially mediated differences in biogeochemical cycles. Science of the Total Environment 677, 47-56. Supplementary Files file.rhistory RFmodelresult.xlsx SEMCSresult.xlsx SETC.xlsx SETL.xlsx SETM.xlsx SETS.xlsx ZJZLLW.r e juzhengcor.csv renamedbbb69.xlsx Cite Share Download PDF Status: Under Review Version 1 posted Reviewers agreed at journal 11 Dec, 2025 Reviewers invited by journal 25 Nov, 2025 Editor invited by journal 15 Nov, 2025 Editor assigned by journal 03 Nov, 2025 First submitted to journal 02 Nov, 2025 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. 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Note: C, noninvaded; L, lightly invaded; M, moderately invaded; S, severely invaded.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"1.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-8012262/v1/bb97f8db2db23339c93b0313.png\"},{\"id\":97138923,\"identity\":\"c4ce8ee2-cf97-45fd-8803-3b6bd212a09d\",\"added_by\":\"auto\",\"created_at\":\"2025-12-01 09:59:26\",\"extension\":\"png\",\"order_by\":2,\"title\":\"Figure 2\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":139552,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eEffects of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion on physicochemical properties under different invasion levels. Note: C, noninvaded; L, lightly invaded; M, moderately invaded; S, severely invaded.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"2.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-8012262/v1/82e952a127319104ac595ebf.png\"},{\"id\":97007696,\"identity\":\"678c392c-fa54-416c-b673-996d38185d80\",\"added_by\":\"auto\",\"created_at\":\"2025-11-28 14:57:17\",\"extension\":\"png\",\"order_by\":3,\"title\":\"Figure 3\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":98293,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eEffects of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e on enzyme activities under different invasion levels.\\u003c/p\\u003e\\n\\u003cp\\u003eNote: C, noninvaded; L, lightly invaded; M, moderately invaded; S, severely invaded. SSC, soil sucrase; NR, nitrate reductase; SUE, soil urease; CAT, catalase; ALP, alkaline phosphatase.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"3.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-8012262/v1/ae35268e6aea2861303d6a36.png\"},{\"id\":97139321,\"identity\":\"4141a14f-4ec6-4116-9f0c-dc060100e28a\",\"added_by\":\"auto\",\"created_at\":\"2025-12-01 10:00:01\",\"extension\":\"png\",\"order_by\":4,\"title\":\"Figure 4\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":155815,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eCorrelation analysis of the soil physicochemical properties and enzyme activity.\\u003c/p\\u003e\\n\\u003cp\\u003eNote: SSC, soil sucrase; NR, nitrate reductase; SUE, soil urease; CAT, catalase; ALP, alkaline phosphatase; B, boron; Zn, zinc; Ca, calcium; Mg, magnesium; Al, aluminum; Fe, iron; Cu, copper; Mn, manganese; SOC, soil organic carbon; TN, total nitrogen; TP, total phosphorus; TK, total potassium; AN, available nitrogen; AP, available phosphorus; AK, available potassium.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"4.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-8012262/v1/4c52f0dbf26e7de88d8a7cf9.png\"},{\"id\":97139881,\"identity\":\"03af10af-2931-4857-9399-551792251148\",\"added_by\":\"auto\",\"created_at\":\"2025-12-01 10:02:55\",\"extension\":\"png\",\"order_by\":5,\"title\":\"Figure 5\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":92554,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eAnalysis of the driving factors influencing \\u003cem\\u003eA. adenophora \\u003c/em\\u003ecoverage across areas with different invasion levels. Note: (a), noninvaded; (b), lightly invaded; (c), moderately invaded; (d), severely invaded.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"5.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-8012262/v1/128b3a538dd584c0de72da4f.png\"},{\"id\":97007698,\"identity\":\"8c72dd10-a412-48a8-be32-2bf492be303f\",\"added_by\":\"auto\",\"created_at\":\"2025-11-28 14:57:17\",\"extension\":\"png\",\"order_by\":6,\"title\":\"Figure 6\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":440394,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003ePathway analysis of factors influencing \\u003cem\\u003eA. adenophora\\u003c/em\\u003e coverage under different invasion levels. In SEM analysis, pH was incorporated as an independent variable, the key factors identified with the random forest model were adopted as observed variables, and the aboveground coverage of \\u003cem\\u003eA. adenophora \\u003c/em\\u003ewas employed as the dependent variable. The red and black arrows denote significant and nonsignificant relationships, respectively. 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09:58:40\",\"extension\":\"\",\"order_by\":20,\"title\":\"\",\"display\":\"\",\"copyAsset\":false,\"role\":\"supplement\",\"size\":11964390,\"visible\":true,\"origin\":\"\",\"legend\":\"\",\"description\":\"\",\"filename\":\"e\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-8012262/v1/98e5d78ea3a4ed636dcd158c\"},{\"id\":97007732,\"identity\":\"a0e21b3e-e32f-4766-bf16-1c2b21d41e7a\",\"added_by\":\"auto\",\"created_at\":\"2025-11-28 14:57:18\",\"extension\":\"csv\",\"order_by\":21,\"title\":\"\",\"display\":\"\",\"copyAsset\":false,\"role\":\"supplement\",\"size\":10429,\"visible\":true,\"origin\":\"\",\"legend\":\"\",\"description\":\"\",\"filename\":\"juzhengcor.csv\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-8012262/v1/e808040ce36958bd9121acb6.csv\"},{\"id\":97007709,\"identity\":\"ae3e9881-4794-4f46-ac85-89dfd630857f\",\"added_by\":\"auto\",\"created_at\":\"2025-11-28 14:57:18\",\"extension\":\"xlsx\",\"order_by\":22,\"title\":\"\",\"display\":\"\",\"copyAsset\":false,\"role\":\"supplement\",\"size\":33869,\"visible\":true,\"origin\":\"\",\"legend\":\"\",\"description\":\"\",\"filename\":\"renamedbbb69.xlsx\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-8012262/v1/02cc93fcbb9210eeafa9e27a.xlsx\"}],\"financialInterests\":\"\",\"formattedTitle\":\"The Soil pH and Micronutrients Drive Ageratina Adenophora Invasion in Areas with Acidic and Nutrient-Poor Soils\",\"fulltext\":[{\"header\":\"1. Introduction\",\"content\":\"\\u003cp\\u003eInvasive alien plants pose a significant threat to global ecological security, with the potential to reduce regional biodiversity and drastically alter the structure and function of natural ecosystems (Yang et al., \\u003cspan citationid=\\\"CR55\\\" class=\\\"CitationRef\\\"\\u003e2013\\u003c/span\\u003e; Xu et al., \\u003cspan citationid=\\\"CR54\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e). Owing to intensified global climate change and increased international exchanges, the spread of invasive plants continues to increase (Osunkoya and Perrett, \\u003cspan citationid=\\\"CR37\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e; Ren et al., \\u003cspan citationid=\\\"CR40\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e). Such plants compete with native flora for essential resources, including water, light, and soil nutrients (Jones et al., \\u003cspan citationid=\\\"CR20\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e; Puissant et al., \\u003cspan citationid=\\\"CR39\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e; DeForest and Moorhead, \\u003cspan citationid=\\\"CR13\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). A substantial body of evidence indicates that invasive species can alter microbial community composition and soil enzyme activity through root secretions, litter decomposition, and allelopathy, thus creating nutrient environments favorable for their proliferation (Esch et al., \\u003cspan citationid=\\\"CR14\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e; Ahmad Dar et al., \\u003cspan citationid=\\\"CR1\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e; Guo et al., \\u003cspan citationid=\\\"CR18\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e). Changes in soil environments subsequently enhance the competitive advantage of these species, thereby influencing the dynamics between exotic and native plants (Esch et al., \\u003cspan citationid=\\\"CR14\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e; Ahmad Dar et al., \\u003cspan citationid=\\\"CR1\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e; Guo et al., \\u003cspan citationid=\\\"CR18\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e). The complex feedback relationships between invasive alien plants and soil nutrient cycling continue to attract broad research interest (Gasch et al., \\u003cspan citationid=\\\"CR16\\\" class=\\\"CitationRef\\\"\\u003e2013\\u003c/span\\u003e; Wu et al., \\u003cspan citationid=\\\"CR48\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e; Kumar et al., \\u003cspan citationid=\\\"CR25\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e).\\u003c/p\\u003e\\u003cp\\u003e\\u003cem\\u003eAgeratina adenophora\\u003c/em\\u003e, a perennial herbaceous or semishrubby plant native to Mexico, is recognized globally as a highly pernicious invasive species (Wu et al., \\u003cspan citationid=\\\"CR48\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e; Kumar et al., \\u003cspan citationid=\\\"CR25\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e). This plant exhibits significant environmental adaptability and has been demonstrated to substantially increase soil fertility after prolonged establishment (Zhao et al., \\u003cspan citationid=\\\"CR60\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e; Zhang et al., \\u003cspan citationid=\\\"CR58\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e). It has been demonstrated that \\u003cem\\u003eA. adenophora\\u003c/em\\u003e can mitigate unfavorable soil conditions, thereby increasing its uptake of essential nutrients such as carbon (C), nitrogen (N), and phosphorus (P) (Boj\\u0026oacute;rquez-Quintal et al., \\u003cspan citationid=\\\"CR3\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e; Ahmad Dar et al., \\u003cspan citationid=\\\"CR1\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e). This, in turn, accelerates its growth and development (Zhao et al., \\u003cspan citationid=\\\"CR60\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e; Verma et al., \\u003cspan citationid=\\\"CR45\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e). In addition to macronutrients, \\u003cem\\u003eA. adenophora\\u003c/em\\u003e requires various micronutrients for optimal growth and development of physiological functions. Notably, nutrient deficiency can significantly impair health (Boj\\u0026oacute;rquez-Quintal et al., \\u003cspan citationid=\\\"CR3\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e; Wu et al., \\u003cspan citationid=\\\"CR49\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e). Compared with native flora, invasive species typically exhibit accelerated growth and increased biomass, necessitating greater intake of micronutrients and consequently intensifying the activation or depletion of soil micronutrient pools (Pizzeghello et al., \\u003cspan citationid=\\\"CR38\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e; Wu et al., \\u003cspan citationid=\\\"CR49\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e). For example, invasion by \\u003cem\\u003eAcacia dealbata\\u003c/em\\u003e and \\u003cem\\u003eLantana camara L.\\u003c/em\\u003e significantly increases soil calcium (Ca) and magnesium (Mg) levels while altering the concentrations of iron (Fe), copper (Cu), manganese (Mn), and sulfur (S) (Osunkoya and Perrett, \\u003cspan citationid=\\\"CR37\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e; Pizzeghello et al., \\u003cspan citationid=\\\"CR38\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e; Kaur et al., \\u003cspan citationid=\\\"CR22\\\" class=\\\"CitationRef\\\"\\u003e2012\\u003c/span\\u003e; Souza-Alonso et al., \\u003cspan citationid=\\\"CR42\\\" class=\\\"CitationRef\\\"\\u003e2014\\u003c/span\\u003e). These findings suggest that there may occur positive feedback between these two plants and soil micronutrients, in turn facilitating their invasion (Esch et al., \\u003cspan citationid=\\\"CR14\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e; Kim et al., \\u003cspan citationid=\\\"CR24\\\" class=\\\"CitationRef\\\"\\u003e2018\\u003c/span\\u003e; Puissant et al., \\u003cspan citationid=\\\"CR39\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e). Unfortunately, research on soil micronutrient dynamics during \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion, particularly in nutrient-poor soils, remains scarce (Bursali et al., \\u003cspan citationid=\\\"CR4\\\" class=\\\"CitationRef\\\"\\u003e2009\\u003c/span\\u003e; Wu et al., \\u003cspan citationid=\\\"CR48\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e; Chen et al., \\u003cspan citationid=\\\"CR8\\\" class=\\\"CitationRef\\\"\\u003e2022\\u003c/span\\u003e).\\u003c/p\\u003e\\u003cp\\u003eThe soil pH is closely associated with nutrient biogeochemical processes and availability (Zhao et al., \\u003cspan citationid=\\\"CR60\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e; Xiao et al., \\u003cspan citationid=\\\"CR52\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e). Under acidic soil conditions, salt-based ions such as Zn, Mg, and Ca are readily displaced by acid ions (H\\u003csup\\u003e+\\u003c/sup\\u003e and Al\\u003csup\\u003e3+\\u003c/sup\\u003e), resulting in their loss through leaching or precipitation events (Merry et al., \\u003cspan citationid=\\\"CR34\\\" class=\\\"CitationRef\\\"\\u003e2002\\u003c/span\\u003e; Boj\\u0026oacute;rquez-Quintal et al., \\u003cspan citationid=\\\"CR3\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e). Concurrently, Mo can form iron phosphate precipitates, which results in a reduction in availability (Jones et al., \\u003cspan citationid=\\\"CR20\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e; Wu et al., \\u003cspan citationid=\\\"CR48\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). High acidity can even lead to aluminum and manganese toxicity issues, which inhibits plant growth. In response to low soil pH values, plants effectively neutralize or buffer soil acidity during growth by promoting ammonification reactions in the soil, thus inhibiting nitrification reactions and regulating organic components within their own tissues. (Xiao et al., \\u003cspan citationid=\\\"CR51\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e; Zhao et al., \\u003cspan citationid=\\\"CR60\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e). Previous studies have indicated that \\u003cem\\u003eA. adenophora\\u003c/em\\u003e may exhibit a similar ability (Bills et al., \\u003cspan citationid=\\\"CR2\\\" class=\\\"CitationRef\\\"\\u003e2010\\u003c/span\\u003e; Kim et al., \\u003cspan citationid=\\\"CR24\\\" class=\\\"CitationRef\\\"\\u003e2018\\u003c/span\\u003e), which becomes increasingly important as invasion deepens or expands over time. For example, Xiao et al. (\\u003cspan citationid=\\\"CR51\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e) revealed that pH promoted the colonization of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e, thus increasing soil fertility, as indicated by elevated organic carbon (C\\u003csub\\u003eorg\\u003c/sub\\u003e), nitrate (NO\\u003csub\\u003e3\\u003c/sub\\u003e\\u003csup\\u003e\\u0026minus;\\u003c/sup\\u003e), ammonium (NH\\u003csub\\u003e4\\u003c/sub\\u003e\\u003csup\\u003e+\\u003c/sup\\u003e), available potassium (AK), and available phosphorus (AP) contents. Additionally, Jones et al. (\\u003cspan citationid=\\\"CR20\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e) indicated that pH is the main factor regulating the carbon utilization efficiency, and Puissant et al. (\\u003cspan citationid=\\\"CR39\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e) reported that pH can affect enzyme activity, which can increase the activities of key soil enzymes such as urease (SUE), phosphatase, and invertase (Zhao et al., \\u003cspan citationid=\\\"CR60\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e). Moreover, Wu et al. (\\u003cspan citationid=\\\"CR48\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e) demonstrated that the soil pH and enzyme activity levels were notably greater in areas invaded by \\u003cem\\u003eA. adenophora\\u003c/em\\u003e than in uninvaded areas (Wu et al., \\u003cspan citationid=\\\"CR48\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). Nevertheless, the precise manner in which increasing the soil pH contributes to changes in soil nutrient contents to match the increased invasion of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e remains unclear (DeForest and Moorhead, \\u003cspan citationid=\\\"CR13\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e; Lhamo et al., \\u003cspan citationid=\\\"CR26\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e).\\u003c/p\\u003e\\u003cp\\u003eStarting during the 1940s, the range of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e increased from Burma to Yunnan Province in China, and this invasive plant rapidly proliferated throughout Southwest China. Presently, \\u003cem\\u003eA. adenophora\\u003c/em\\u003e has emerged as one of the most formidable threats to biodiversity and agricultural production in inland China (Pizzeghello et al., \\u003cspan citationid=\\\"CR38\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e; Fan et al., \\u003cspan citationid=\\\"CR15\\\" class=\\\"CitationRef\\\"\\u003e2022\\u003c/span\\u003e). \\u003cem\\u003ePinus yunnanensis\\u003c/em\\u003e, an endemic tree species in Southwest China, dominates the regional forested landscape. These forests, which mainly encompass homogeneous stands, are characterized by notable soil acidification (with soil pH values typically ranging from 4 to 6) (Merry et al., \\u003cspan citationid=\\\"CR34\\\" class=\\\"CitationRef\\\"\\u003e2002\\u003c/span\\u003e; Kabała and Łabaz, \\u003cspan citationid=\\\"CR21\\\" class=\\\"CitationRef\\\"\\u003e2018\\u003c/span\\u003e) underneath the canopy and limited soil nutrient availability, which significantly restricts plant growth (Esch et al., \\u003cspan citationid=\\\"CR14\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e; Darji et al., \\u003cspan citationid=\\\"CR11\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e). Interestingly, over the past two decades, field surveys have identified \\u003cem\\u003ePinus yunnanensis\\u003c/em\\u003e forests as primary sites for \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion, and invasive populations continue to expand, posing a serious threat to forest regeneration and subcanopy biodiversity. (Wang et al., \\u003cspan citationid=\\\"CR47\\\" class=\\\"CitationRef\\\"\\u003e2015\\u003c/span\\u003e; Sun et al., 2021; Zhang et al., \\u003cspan citationid=\\\"CR56\\\" class=\\\"CitationRef\\\"\\u003e2022\\u003c/span\\u003e). Although a few studies have revealed changes in soil nutrient levels and interspecific associations (Hu et al., \\u003cspan citationid=\\\"CR19\\\" class=\\\"CitationRef\\\"\\u003e2021\\u003c/span\\u003e; Xiao et al., \\u003cspan citationid=\\\"CR52\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e; Niu et al., \\u003cspan citationid=\\\"CR35\\\" class=\\\"CitationRef\\\"\\u003e2007\\u003c/span\\u003e; Liu et al., \\u003cspan citationid=\\\"CR30\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e) after \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion into \\u003cem\\u003ePinus yunnanensis\\u003c/em\\u003e forests, the nutrient dynamics throughout the spread process have not yet been explored in depth. In our study, we considered different invasion levels of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e in \\u003cem\\u003ePinus yunnanensis\\u003c/em\\u003e forests to explore changes in the levels of major soil nutrients, micronutrients, and enzyme activity. Specifically, we aimed to determine (1) whether the soil pH increases with increasing invasion level; (2) how the contents of major soil micronutrients change with increasing invasion level; and (3) the mechanisms through which changes in soil pH and micronutrient levels may facilitate increased invasion. We aimed to enhance the understanding of the expansion mechanisms of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e in areas with acidic soil conditions and to provide a theoretical basis for developing preventive and control strategies.\\u003c/p\\u003e\"},{\"header\":\"2. Materials and Methods\",\"content\":\"\\u003cdiv id=\\\"Sec3\\\" class=\\\"Section2\\\"\\u003e\\u003ch2\\u003e2.1 Study area\\u003c/h2\\u003e\\u003cp\\u003eThe research site is located in Jiyi town, Wuding County, Yunnan Province, Southwest China, at coordinates of 26\\u0026deg;6\\u0026prime;N and 102\\u0026deg;14\\u0026prime;E. Jiyi town is situated on the banks of the Jinsha River and exhibits an average elevation of 1900 m (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e). The study area experiences a dry-hot valley climate characterized by abundant sunshine, high evaporation rates and an average annual temperature of 15.2\\u0026deg;C. Precipitation occurs mainly from May to October, with annual totals ranging from 800 to 1000 mm (Wu et al., \\u003cspan citationid=\\\"CR48\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e). Previous studies have identified the soil type at the research site as dry red soil (Chen et al., \\u003cspan citationid=\\\"CR6\\\" class=\\\"CitationRef\\\"\\u003e2013\\u003c/span\\u003e). Specifically, the sampling site is located in the upper-middle part of a \\u003cem\\u003ePinus yunnanensis\\u003c/em\\u003e forest with slope gradients ranging from 25\\u0026deg; to 40\\u0026deg; and faces northeast. The average height of this forest is approximately 10.5 m, and the average diameter at breast height is approximately 13.5 cm. The canopy density levels range from 0.6 to 0.9 units per unit area (m\\u0026sup2;). Resident surveys indicate that \\u003cem\\u003eA. adenophora\\u003c/em\\u003e has been invading the understory of this forest for more than two decades.\\u003c/p\\u003e\\u003cp\\u003e\\u003c/p\\u003e\\u003c/div\\u003e\\u003cdiv id=\\\"Sec4\\\" class=\\\"Section2\\\"\\u003e\\u003ch2\\u003e2.2 Sampling method\\u003c/h2\\u003e\\u003cp\\u003eIn August 2023, the extent of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion was quantified by assessing the relative projected area covered by this species with reference to previous studies (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e). Sample plots were selected randomly and categorized into four invasion levels on the basis of the percentage of the area covered by \\u003cem\\u003eA. adenophora\\u003c/em\\u003e, namely, noninvaded (C, 0.2%-1.2%), lightly invaded (L, 25%-35%), moderately invaded (M, 50%-70%), and severely invaded (S, 88%-97%) (Table\\u0026nbsp;\\u003cspan refid=\\\"Tab1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e) (Xiao et al., \\u003cspan citationid=\\\"CR51\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e; Wu et al., \\u003cspan citationid=\\\"CR48\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e; Zhang et al., \\u003cspan citationid=\\\"CR58\\\" class=\\\"CitationRef\\\"\\u003e2023\\u003c/span\\u003e). For each invasion level, six plots were established, each measuring 1\\u0026times;1 m, characterized by similar topographic features and canopy closure conditions. The plots were spaced at least 20 m apart to avoid spatial overlap. The plant composition varied according to the invasion level (refer to Table\\u0026nbsp;\\u003cspan refid=\\\"Tab1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e for details).\\u003c/p\\u003e\\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\\u003eDistribution of plants in the \\u003cem\\u003eA. adenophora\\u003c/em\\u003e sample plots under different invasion levels.\\u003c/p\\u003e\\u003c/div\\u003e\\u003c/caption\\u003e\\u003ccolgroup cols=\\\"3\\\"\\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\\u003cthead\\u003e\\u003ctr\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c1\\\"\\u003e\\u003cp\\u003eInvasion level\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c2\\\"\\u003e\\u003cp\\u003eCommunity overview\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c3\\\"\\u003e\\u003cp\\u003eCoverage of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e %\\u003c/p\\u003e\\u003c/th\\u003e\\u003c/tr\\u003e\\u003c/thead\\u003e\\u003ctbody\\u003e\\u003ctr\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e\\u003cp\\u003eNoninvaded (C)\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e\\u003cp\\u003e\\u003cem\\u003ePogonatherum crinitum\\u003c/em\\u003e Thunb.; \\u003cem\\u003eArtemisia lavandulifolia\\u003c/em\\u003e DC.; \\u003cem\\u003eGrona heterocarpos\\u003c/em\\u003e L.; \\u003cem\\u003eScutellaria indica\\u003c/em\\u003e L.; \\u003cem\\u003eCynoglossum amabile\\u003c/em\\u003e Stapf \\u0026amp; Drummond; \\u003cem\\u003ePararuellia delavayana\\u003c/em\\u003e Baill.; \\u003cem\\u003ePaspalum scrobiculatum\\u003c/em\\u003e Var.; \\u003cem\\u003ePolygala persicariifolia\\u003c/em\\u003e DC.; \\u003cem\\u003eDodonaea viscosa\\u003c/em\\u003e Jacquem.; \\u003cem\\u003eOriganum vulgare\\u003c/em\\u003e L.; \\u003cem\\u003eReinwardtia indica\\u003c/em\\u003e Dumort.\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"char\\\" char=\\\"\\u0026plusmn;\\\" colname=\\\"c3\\\"\\u003e\\u003cp\\u003e0.68\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.35\\u003c/p\\u003e\\u003c/td\\u003e\\u003c/tr\\u003e\\u003ctr\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e\\u003cp\\u003eLightly invaded (L)\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e\\u003cp\\u003e\\u003cem\\u003eAgeratina adenophora\\u003c/em\\u003e Spreng.; \\u003cem\\u003eArtemisia lavandulifolia\\u003c/em\\u003e DC.; \\u003cem\\u003eAinsliaea spicata\\u003c/em\\u003e Vaniot, \\u003cem\\u003ePogonatherum crinitum\\u003c/em\\u003e Thunb.; \\u003cem\\u003eClinopodium megalanthum\\u003c/em\\u003e Diels; \\u003cem\\u003eOxalis corniculata\\u003c/em\\u003e L.; \\u003cem\\u003eDioscorea subcalva\\u003c/em\\u003e Prain \\u0026amp; Burkill\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"char\\\" char=\\\"\\u0026plusmn;\\\" colname=\\\"c3\\\"\\u003e\\u003cp\\u003e29.83\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;3.39\\u003c/p\\u003e\\u003c/td\\u003e\\u003c/tr\\u003e\\u003ctr\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e\\u003cp\\u003eModerately invaded (M)\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e\\u003cp\\u003e\\u003cem\\u003eAgeratina adenophora\\u003c/em\\u003e Spreng.; \\u003cem\\u003eElsholtzia rugulosa\\u003c/em\\u003e Hemsl.; \\u003cem\\u003eReinwardtia indica\\u003c/em\\u003e Dumort.; \\u003cem\\u003ePogonatherum crinitum\\u003c/em\\u003e Thunb.; \\u003cem\\u003eGrona heterocarpos\\u003c/em\\u003e L.; \\u003cem\\u003ePimpinella candolleana\\u003c/em\\u003e Wight \\u0026amp; Arn.; \\u003cem\\u003eArisaema erubescens\\u003c/em\\u003e Wall.\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"char\\\" char=\\\"\\u0026plusmn;\\\" colname=\\\"c3\\\"\\u003e\\u003cp\\u003e60\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;6.51\\u003c/p\\u003e\\u003c/td\\u003e\\u003c/tr\\u003e\\u003ctr\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e\\u003cp\\u003eSeverely invaded (S)\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e\\u003cp\\u003e\\u003cem\\u003eAgeratina adenophora\\u003c/em\\u003e Spreng.; \\u003cem\\u003eElsholtzia rugulosa\\u003c/em\\u003e Hemsl.; \\u003cem\\u003eReinwardtia indica\\u003c/em\\u003e Dumort.; \\u003cem\\u003ePogonatherum crinitum\\u003c/em\\u003e Thunb.\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"char\\\" char=\\\"\\u0026plusmn;\\\" colname=\\\"c3\\\"\\u003e\\u003cp\\u003e92.67\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;3.04\\u003c/p\\u003e\\u003c/td\\u003e\\u003c/tr\\u003e\\u003c/tbody\\u003e\\u003c/colgroup\\u003e\\u003c/table\\u003e\\u003c/div\\u003e\\u003c/p\\u003e\\u003cp\\u003eTo assess soil properties, aboveground vegetation and litter were removed in each plot, and five soil samples were collected from the top 20 cm with a soil auger (5-cm diameter) following the five-point sampling method. The samples were thoroughly mixed, transported to the laboratory, and sieved through a 2-mm mesh. The sieved samples were then divided into two portions: one was dried naturally for the analysis of soil physicochemical properties and micronutrient contents, while the other was refrigerated at a temperature of 4\\u0026deg;C for two weeks to analyze soil enzyme activity and alkaline dissolved nitrogen levels.\\u003c/p\\u003e\\u003c/div\\u003e\\u003cdiv id=\\\"Sec5\\\" class=\\\"Section2\\\"\\u003e\\u003ch2\\u003e2.3 Soil analysis\\u003c/h2\\u003e\\u003cp\\u003eSoil physicochemical properties encompass a comprehensive range of parameters, including the soil pH, soil organic carbon (SOC), total nitrogen (TN), total phosphorus (TP), total potassium (TK), available nitrogen (AN), available phosphorus (AP), and AK (Xiao et al., \\u003cspan citationid=\\\"CR51\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e). In addition, the analysis was extended to soil total microelements, such as Fe, Mn, zinc (Zn), Cu, boron (B), Ca, Mg, and aluminum (Al) (Osunkoya and Perrett, \\u003cspan citationid=\\\"CR37\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e; DeForest and Moorhead, \\u003cspan citationid=\\\"CR13\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e), as well as soil available microelements, including available Fe, available Mn, available Zn, available Cu, available B, exchangeable Ca, exchangeable Mg, and available Al (Osunkoya and Perrett, \\u003cspan citationid=\\\"CR37\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e; Fan et al., \\u003cspan citationid=\\\"CR15\\\" class=\\\"CitationRef\\\"\\u003e2022\\u003c/span\\u003e). Furthermore, the activity of soil enzymes, including SUE, alkaline phosphatase (ALP), sucrase (SSC), nitrate reductase (NR), and catalase (CAT) activities, was evaluated (Kim et al., \\u003cspan citationid=\\\"CR24\\\" class=\\\"CitationRef\\\"\\u003e2018\\u003c/span\\u003e; Zhang et al., \\u003cspan citationid=\\\"CR56\\\" class=\\\"CitationRef\\\"\\u003e2022\\u003c/span\\u003e). In this study, macronutrient indices were determined according to the method of Xiao et al. (\\u003cspan citationid=\\\"CR51\\\" class=\\\"CitationRef\\\"\\u003e2017\\u003c/span\\u003e), micronutrient indices were determined according to the methods of Osunkoya and Perrett (\\u003cspan citationid=\\\"CR37\\\" class=\\\"CitationRef\\\"\\u003e2011\\u003c/span\\u003e), DeForest and Moorhead (\\u003cspan citationid=\\\"CR13\\\" class=\\\"CitationRef\\\"\\u003e2020\\u003c/span\\u003e) and Fan et al. (\\u003cspan citationid=\\\"CR15\\\" class=\\\"CitationRef\\\"\\u003e2022\\u003c/span\\u003e), and enzyme activity indices were determined according to the methods of Kim et al. (\\u003cspan citationid=\\\"CR24\\\" class=\\\"CitationRef\\\"\\u003e2018\\u003c/span\\u003e) and Zhang et al. (\\u003cspan citationid=\\\"CR56\\\" class=\\\"CitationRef\\\"\\u003e2022\\u003c/span\\u003e). These parameters collectively provide a robust framework for assessing soil health and nutrient dynamics in the studied ecosystem.\\u003c/p\\u003e\\u003c/div\\u003e\\u003cdiv id=\\\"Sec6\\\" class=\\\"Section2\\\"\\u003e\\u003ch2\\u003e2.4 Data analysis\\u003c/h2\\u003e\\u003cp\\u003eIn this study, the experimental data were systematically analyzed and organized in Microsoft Excel 2021. The normality of the data distribution was evaluated via SPSS 25.0. Graphical representations of the soil pH, SOC, nitrogen (N), phosphorus (P), potassium (K), and enzyme activity data were generated with the ggplot2 package in R version 4.2.2. The soil micronutrient data were analyzed via one-way analysis of variance (ANOVA) using SPSS 25.0. Additionally, correlation analysis, random forest model (RFM) analysis and structural equation modeling (SEM) were performed in R, thereby employing the piecewiseSEM, corrplot, and pacman packages, respectively. In SEM analysis, a P value greater than 0.05 indicated that all path coefficients in the model were less than 1, with lower values of Fisher's exact test statistic, the Akaike information criterion (AIC), and the Bayesian information criterion (BIC) indicating a better model fit (Zhao et al., \\u003cspan citationid=\\\"CR60\\\" class=\\\"CitationRef\\\"\\u003e2019\\u003c/span\\u003e). This comprehensive analytical approach ensured robust data interpretation and provided a solid foundation for understanding the relationships between soil properties and their influence on the studied ecosystem.\\u003c/p\\u003e\\u003c/div\\u003e\"},{\"header\":\"3. Results\",\"content\":\"\\u003cdiv id=\\\"Sec8\\\" class=\\\"Section2\\\"\\u003e\\u003ch2\\u003e3.1 Effects of different levels of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion on soil physicochemical properties\\u003c/h2\\u003e\\u003cp\\u003eAs shown in Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003e, significant changes (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05) were observed in the soil pH, TN, TK, and AP levels across the areas with varying degrees of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion compared with those in noninvaded areas. Specifically, the soil pH increased with increasing invasion level (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003ea), with increases of 1.34%, 3.19%, and 8.37% in lightly, moderately, and severely invaded areas, respectively, relative to those in noninvaded areas. The soil TN content significantly fluctuated (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05) with increasing invasion level, with a 35.38% increase in lightly invaded areas, a 10.73% decrease in moderately invaded areas, and a 9.59% increase in severely invaded areas (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003eb). Conversely, the soil TK content increased with increasing invasion level, peaking (12.04 g/kg) in severely invaded areas. In contrast to the aforementioned parameters, the AP content significantly decreased (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05) with increasing invasion level, decreasing by 47.24%, 39.08%, and 40.76% in lightly, moderately, and severely invaded areas, respectively (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003eg, P\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05). Notably, although not statistically significant, the SOC content increased with increasing invasion level.\\u003c/p\\u003e\\u003cp\\u003e\\u003c/p\\u003e\\u003c/div\\u003e\\u003cdiv id=\\\"Sec9\\\" class=\\\"Section2\\\"\\u003e\\u003ch2\\u003e3.2 Effects of different levels of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion on soil micronutrient content and availability levels\\u003c/h2\\u003e\\u003cp\\u003eThe mean concentrations of soil trace elements under different invasion levels are provided in Table\\u0026nbsp;\\u003cspan refid=\\\"Tab2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003e. Although the differences in micronutrient concentrations across invasion levels did not reach statistical significance (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026gt;\\u0026thinsp;0.05), certain trends were observed. Except boron (B), which exhibited the highest concentration in noninvaded areas, the remaining seven micronutrients demonstrated increasing trends following \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion, with increases ranging from 0.99% to 18.08%. Specifically, the maximum concentrations of Zn, Ca, Mg, and Al occurred in severely invaded areas, whereas Fe and Cu peaked in moderately invaded areas. The highest Mn concentration only occurred in lightly invaded areas among all the sampled areas.\\u003c/p\\u003e\\u003cp\\u003e\\u003cdiv class=\\\"gridtable\\\"\\u003e\\u003ctable float=\\\"Yes\\\" id=\\\"Tab2\\\" border=\\\"1\\\"\\u003e\\u003ccaption language=\\\"En\\\"\\u003e\\u003cdiv class=\\\"CaptionNumber\\\"\\u003eTable 2\\u003c/div\\u003e\\u003cdiv class=\\\"CaptionContent\\\"\\u003e\\u003cp\\u003eEffects of micronutrients on \\u003cem\\u003eA. adenophora\\u003c/em\\u003e under different invasion levels.\\u003c/p\\u003e\\u003c/div\\u003e\\u003c/caption\\u003e\\u003ccolgroup cols=\\\"9\\\"\\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=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c3\\\" colnum=\\\"3\\\"\\u003e\\u003c/div\\u003e\\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c4\\\" colnum=\\\"4\\\"\\u003e\\u003c/div\\u003e\\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c5\\\" colnum=\\\"5\\\"\\u003e\\u003c/div\\u003e\\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c6\\\" colnum=\\\"6\\\"\\u003e\\u003c/div\\u003e\\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c7\\\" colnum=\\\"7\\\"\\u003e\\u003c/div\\u003e\\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c8\\\" colnum=\\\"8\\\"\\u003e\\u003c/div\\u003e\\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c9\\\" colnum=\\\"9\\\"\\u003e\\u003c/div\\u003e\\u003cthead\\u003e\\u003ctr\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c1\\\" morerows=\\\"1\\\" rowspan=\\\"2\\\"\\u003e\\u003cp\\u003eDifferent levels of invasion\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colspan=\\\"8\\\" nameend=\\\"c9\\\" namest=\\\"c2\\\"\\u003e\\u003cp\\u003eMean\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;SE\\u003c/p\\u003e\\u003c/th\\u003e\\u003c/tr\\u003e\\u003ctr\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c2\\\"\\u003e\\u003cp\\u003eFe mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c3\\\"\\u003e\\u003cp\\u003eMn mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u003cp\\u003eZn mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c5\\\"\\u003e\\u003cp\\u003eCu mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c6\\\"\\u003e\\u003cp\\u003eB mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c7\\\"\\u003e\\u003cp\\u003eCa mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c8\\\"\\u003e\\u003cp\\u003eMg mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c9\\\"\\u003e\\u003cp\\u003eAl mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003c/tr\\u003e\\u003c/thead\\u003e\\u003ctbody\\u003e\\u003ctr\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e\\u003cp\\u003eC\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e\\u003cp\\u003e21.2\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.6a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c3\\\"\\u003e\\u003cp\\u003e902.33\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;119.08a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u003cp\\u003e44.47\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.95\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c5\\\"\\u003e\\u003cp\\u003e27.88\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.45a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c6\\\"\\u003e\\u003cp\\u003e86.55\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;5.57a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c7\\\"\\u003e\\u003cp\\u003e1.82\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.21a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c8\\\"\\u003e\\u003cp\\u003e2.6\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.1a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c9\\\"\\u003e\\u003cp\\u003e64.37\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;1.29a\\u003c/p\\u003e\\u003c/td\\u003e\\u003c/tr\\u003e\\u003ctr\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e\\u003cp\\u003eL\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e\\u003cp\\u003e20.98\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;1.03a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c3\\\"\\u003e\\u003cp\\u003e1065.47\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;163.62a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u003cp\\u003e44.1\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;1.01a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c5\\\"\\u003e\\u003cp\\u003e27.73\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;1.04a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c6\\\"\\u003e\\u003cp\\u003e73.48\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;4.99a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c7\\\"\\u003e\\u003cp\\u003e1.7\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.22a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c8\\\"\\u003e\\u003cp\\u003e2.52\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.08a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c9\\\"\\u003e\\u003cp\\u003e66.55\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;2.29a\\u003c/p\\u003e\\u003c/td\\u003e\\u003c/tr\\u003e\\u003ctr\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e\\u003cp\\u003eM\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e\\u003cp\\u003e21.41\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.51a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c3\\\"\\u003e\\u003cp\\u003e822.4\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;106.57a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u003cp\\u003e45.32\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.87a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c5\\\"\\u003e\\u003cp\\u003e29.85\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;1.01a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c6\\\"\\u003e\\u003cp\\u003e81.32\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;4.39a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c7\\\"\\u003e\\u003cp\\u003e1.54\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.11a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c8\\\"\\u003e\\u003cp\\u003e2.56\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.1a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c9\\\"\\u003e\\u003cp\\u003e66.05\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;3.96a\\u003c/p\\u003e\\u003c/td\\u003e\\u003c/tr\\u003e\\u003ctr\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e\\u003cp\\u003eS\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e\\u003cp\\u003e20.77\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;1.09a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c3\\\"\\u003e\\u003cp\\u003e948.38\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;128.17a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u003cp\\u003e44.82\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.98a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c5\\\"\\u003e\\u003cp\\u003e29\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;1.31a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c6\\\"\\u003e\\u003cp\\u003e78.62\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;6.18a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c7\\\"\\u003e\\u003cp\\u003e2.01\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.16a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c8\\\"\\u003e\\u003cp\\u003e2.72\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.11a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c9\\\"\\u003e\\u003cp\\u003e70.71\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;3.5a\\u003c/p\\u003e\\u003c/td\\u003e\\u003c/tr\\u003e\\u003c/tbody\\u003e\\u003c/colgroup\\u003e\\u003c/table\\u003e\\u003c/div\\u003e\\u003c/p\\u003e\\u003cp\\u003eWith respect to the availability of soil micronutrients (Table\\u0026nbsp;\\u003cspan refid=\\\"Tab3\\\" class=\\\"InternalRef\\\"\\u003e3\\u003c/span\\u003e), significant differences (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05) were observed in available Mn, Zn, and Cu contents across the areas with different invasion levels, with an overall increasing trend. The concentrations of available Mn, Zn, and Cu in severely invaded areas were 45.48%, 98.52%, and 72.87% higher, respectively, than those in noninvaded areas. Moreover, although the differences in available Fe, Ca, and Mg concentrations did not reach statistical significance (P\\u0026thinsp;\\u0026gt;\\u0026thinsp;0.05), the values generally peaked in severely invaded areas. Conversely, compared with that in noninvaded areas, the available Al concentration decreased by 19.04%, 25.06%, and 18.56% with increasing invasion levels.\\u003c/p\\u003e\\u003cp\\u003e\\u003cdiv class=\\\"gridtable\\\"\\u003e\\u003ctable float=\\\"Yes\\\" id=\\\"Tab3\\\" border=\\\"1\\\"\\u003e\\u003ccaption language=\\\"En\\\"\\u003e\\u003cdiv class=\\\"CaptionNumber\\\"\\u003eTable 3\\u003c/div\\u003e\\u003cdiv class=\\\"CaptionContent\\\"\\u003e\\u003cp\\u003eEffects of different levels of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion on available micronutrients.\\u003c/p\\u003e\\u003c/div\\u003e\\u003c/caption\\u003e\\u003ccolgroup cols=\\\"9\\\"\\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=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c3\\\" colnum=\\\"3\\\"\\u003e\\u003c/div\\u003e\\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c4\\\" colnum=\\\"4\\\"\\u003e\\u003c/div\\u003e\\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c5\\\" colnum=\\\"5\\\"\\u003e\\u003c/div\\u003e\\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c6\\\" colnum=\\\"6\\\"\\u003e\\u003c/div\\u003e\\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c7\\\" colnum=\\\"7\\\"\\u003e\\u003c/div\\u003e\\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c8\\\" colnum=\\\"8\\\"\\u003e\\u003c/div\\u003e\\u003cdiv align=\\\"left\\\" class=\\\"colspec\\\" colname=\\\"c9\\\" colnum=\\\"9\\\"\\u003e\\u003c/div\\u003e\\u003cthead\\u003e\\u003ctr\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c1\\\" morerows=\\\"1\\\" rowspan=\\\"2\\\"\\u003e\\u003cp\\u003eDifferent levels of invasion\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colspan=\\\"8\\\" nameend=\\\"c9\\\" namest=\\\"c2\\\"\\u003e\\u003cp\\u003eMean\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;SE\\u003c/p\\u003e\\u003c/th\\u003e\\u003c/tr\\u003e\\u003ctr\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c2\\\"\\u003e\\u003cp\\u003eavailable Fe mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c3\\\"\\u003e\\u003cp\\u003eavailable Mn mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u003cp\\u003eavailable Zn mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c5\\\"\\u003e\\u003cp\\u003eavailable Cu mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c6\\\"\\u003e\\u003cp\\u003eavailable B mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c7\\\"\\u003e\\u003cp\\u003eExchangeable Ca mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c8\\\"\\u003e\\u003cp\\u003eExchangeable Mg mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003cth align=\\\"left\\\" colname=\\\"c9\\\"\\u003e\\u003cp\\u003eavailable Al mg/kg\\u003c/p\\u003e\\u003c/th\\u003e\\u003c/tr\\u003e\\u003c/thead\\u003e\\u003ctbody\\u003e\\u003ctr\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e\\u003cp\\u003eC\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e\\u003cp\\u003e40.22\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;5.05a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c3\\\"\\u003e\\u003cp\\u003e64.54\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;5.18b\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u003cp\\u003e0.22\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.02b\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c5\\\"\\u003e\\u003cp\\u003e0.57\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.03c\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c6\\\"\\u003e\\u003cp\\u003e0.1\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.03a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c7\\\"\\u003e\\u003cp\\u003e3.16\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.18a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c8\\\"\\u003e\\u003cp\\u003e1.15\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.09a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c9\\\"\\u003e\\u003cp\\u003e52.07\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;5.63a\\u003c/p\\u003e\\u003c/td\\u003e\\u003c/tr\\u003e\\u003ctr\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e\\u003cp\\u003eL\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e\\u003cp\\u003e45.24\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;2.89a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c3\\\"\\u003e\\u003cp\\u003e76.62\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;3.99b\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u003cp\\u003e0.31\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.08ab\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c5\\\"\\u003e\\u003cp\\u003e0.7\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.04bc\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c6\\\"\\u003e\\u003cp\\u003e0.12\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.03a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c7\\\"\\u003e\\u003cp\\u003e4.24\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.75a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c8\\\"\\u003e\\u003cp\\u003e1.26\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.22a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c9\\\"\\u003e\\u003cp\\u003e42.16\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;4.95a\\u003c/p\\u003e\\u003c/td\\u003e\\u003c/tr\\u003e\\u003ctr\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e\\u003cp\\u003eM\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e\\u003cp\\u003e42.72\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;8.41a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c3\\\"\\u003e\\u003cp\\u003e68.43\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;7.91b\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u003cp\\u003e0.25\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.05ab\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c5\\\"\\u003e\\u003cp\\u003e0.78\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.08b\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c6\\\"\\u003e\\u003cp\\u003e0.13\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.03a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c7\\\"\\u003e\\u003cp\\u003e4\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.18a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c8\\\"\\u003e\\u003cp\\u003e1.13\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.02a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c9\\\"\\u003e\\u003cp\\u003e39.02\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;7.23a\\u003c/p\\u003e\\u003c/td\\u003e\\u003c/tr\\u003e\\u003ctr\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c1\\\"\\u003e\\u003cp\\u003eS\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c2\\\"\\u003e\\u003cp\\u003e44.71\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;1.97a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c3\\\"\\u003e\\u003cp\\u003e93.89\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;3.25a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c4\\\"\\u003e\\u003cp\\u003e0.44\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.09a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c5\\\"\\u003e\\u003cp\\u003e0.98\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.08a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c6\\\"\\u003e\\u003cp\\u003e0.1\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.03a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c7\\\"\\u003e\\u003cp\\u003e4.8\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.69a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c8\\\"\\u003e\\u003cp\\u003e1.37\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.19a\\u003c/p\\u003e\\u003c/td\\u003e\\u003ctd align=\\\"left\\\" colname=\\\"c9\\\"\\u003e\\u003cp\\u003e42.41\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;3.72a\\u003c/p\\u003e\\u003c/td\\u003e\\u003c/tr\\u003e\\u003c/tbody\\u003e\\u003c/colgroup\\u003e\\u003ctfoot\\u003e\\u003ctr\\u003e\\u003ctd colspan=\\\"9\\\"\\u003eNote: C, noninvaded; L, lightly invaded; M, moderately invaded; S, severely invaded.\\u003c/td\\u003e\\u003c/tr\\u003e\\u003c/tfoot\\u003e\\u003c/table\\u003e\\u003c/div\\u003e\\u003c/p\\u003e\\u003c/div\\u003e\\u003cdiv id=\\\"Sec10\\\" class=\\\"Section2\\\"\\u003e\\u003ch2\\u003e3.3 Effects of different levels of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion on soil enzymes\\u003c/h2\\u003e\\u003cp\\u003eThe invasion of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e significantly affected soil SSC and NR activity levels (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05), whereas the impacts on soil SUE, CAT, and ALP activity levels were relatively minor (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026gt;\\u0026thinsp;0.05). In severely invaded areas, soil sucrase activity reached its peak value (26.09 mg/kg), whereas moderately invaded areas exhibited the lowest activity (12.90 mg/kg). Compared with those in noninvaded areas, lightly invaded areas exhibited comparable soil sucrase activity levels (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig3\\\" class=\\\"InternalRef\\\"\\u003e3\\u003c/span\\u003eb). Compared with those in noninvaded areas, soil nitrate reductase activity levels increased by 4.85% and 0.76% in lightly and moderately invaded areas, respectively, but decreased by 4.67% in severely invaded areas (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig3\\\" class=\\\"InternalRef\\\"\\u003e3\\u003c/span\\u003ec). Additionally, soil CAT (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig3\\\" class=\\\"InternalRef\\\"\\u003e3\\u003c/span\\u003ed) and ALP (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig3\\\" class=\\\"InternalRef\\\"\\u003e3\\u003c/span\\u003ee) activity levels increased with increasing invasion level.\\u003c/p\\u003e\\u003cp\\u003e\\u003c/p\\u003e\\u003c/div\\u003e\\u003cdiv id=\\\"Sec11\\\" class=\\\"Section2\\\"\\u003e\\u003ch2\\u003e3.4 Determination of the primary factors driving coverage change under different levels of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion\\u003c/h2\\u003e\\u003cp\\u003eThis study revealed the complex interactions among soil trace elements, enzyme activity, and nutrient dynamics through comprehensive correlation analysis of soil enzymatic activity and physicochemical properties (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig4\\\" class=\\\"InternalRef\\\"\\u003e4\\u003c/span\\u003e). The results revealed significant correlations (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05) among various soil trace elements. Notably, TK was significantly positively correlated with B, Mg, and available Fe (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05) but significantly negatively correlated with AK (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05). TP was significantly positively correlated with AP and TN (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05) but significantly negatively correlated with available Ca and available Mg (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05). Furthermore, with respect to activity, significant positive correlations (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05) were observed among four soil enzymes (SSC, SUE, NR, and ALP). Importantly, the soil pH was significantly positively correlated with the SOC, SSC, ALP, CAT, AK, available Cu, and available Mn levels (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05).\\u003c/p\\u003e\\u003cp\\u003e\\u003c/p\\u003e\\u003cp\\u003eThe RFM, which is a robust machine learning tool, was employed in this study to systematically identify and predict the key environmental drivers influencing the invasion and spread of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e. Our model results revealed significant variations in the driving factors influencing the invasion coverage of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e across the areas with different invasion levels (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig5\\\" class=\\\"InternalRef\\\"\\u003e5\\u003c/span\\u003e). Specifically, in noninvaded areas (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig5\\\" class=\\\"InternalRef\\\"\\u003e5\\u003c/span\\u003ea), available Mn, AK, available Zn, available B, and Zn were identified as the most significant factors influencing the invasion coverage of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05). In lightly invaded areas (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig5\\\" class=\\\"InternalRef\\\"\\u003e5\\u003c/span\\u003eb), Al, available Al, available Fe, available Mg, and available Ca emerged as the primary drivers (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05). In moderately invaded areas (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig5\\\" class=\\\"InternalRef\\\"\\u003e5\\u003c/span\\u003ec), Cu, Mg, Al, B, available Zn, available Cu, available Ca, and available Mn significantly affected the invasion coverage of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05). In severely invaded areas (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig5\\\" class=\\\"InternalRef\\\"\\u003e5\\u003c/span\\u003ed), Mg, B, Zn, Fe, available Cu, and available Al were identified as the key drivers influencing invasion coverage (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05).\\u003c/p\\u003e\\u003cp\\u003e\\u003c/p\\u003e\\u003cp\\u003e3.5 What are the pathways or mechanisms that contribute to the increase in the invasion coverage of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e?\\u003c/p\\u003e\\u003cp\\u003eSEM analysis was employed to elucidate the significant and nonsignificant pathways through which soil properties influence the coverage of \\u003cem\\u003eA. adenophora (A. adenophora)\\u003c/em\\u003e under varying invasion levels, providing critical insights into its invasion mechanisms. In this study, we observed a significant linear increase in the soil pH following \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05), indicating that its inclusion was an independent variable in the model (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig2\\\" class=\\\"InternalRef\\\"\\u003e2\\u003c/span\\u003ea). The key soil factors identified through the RFM were selected as observed variables, while the aboveground coverage of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e served as the dependent variable (Table\\u0026nbsp;\\u003cspan refid=\\\"Tab1\\\" class=\\\"InternalRef\\\"\\u003e1\\u003c/span\\u003e). The results revealed distinct response pathways of soil physicochemical properties for \\u003cem\\u003eA. adenophora\\u003c/em\\u003e across the different invasion levels (Fig.\\u0026nbsp;\\u003cspan refid=\\\"Fig6\\\" class=\\\"InternalRef\\\"\\u003e6\\u003c/span\\u003e). Specifically, in noninvaded areas, pH indirectly influenced \\u003cem\\u003eA. adenophora\\u003c/em\\u003e coverage by regulating the availability of soil nutrients (R\\u0026sup2; = 0.7), with AK emerging as a critical driver (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05). In lightly invaded areas, soil nutrients directly and significantly affected \\u003cem\\u003eA. adenophora\\u003c/em\\u003e coverage (R\\u0026sup2; = 0.69), with a path coefficient of 0.64 (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05). In moderately invaded areas, both pH and soil nutrient content indirectly promoted \\u003cem\\u003eA. adenophora\\u003c/em\\u003e coverage by influencing the availability of soil nutrients (R\\u0026sup2; = 0.52). Notably, the effects of soil nutrients (Al) and available nutrients (available Ca and available Cu) were significantly greater than that of pH, with path coefficients of 0.38 (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05) and 0.96 (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.001), respectively. In severely invaded areas, both available nutrients and soil nutrients significantly influenced \\u003cem\\u003eA. adenophora\\u003c/em\\u003e coverage (R\\u0026sup2; = 0.91), with path coefficients of 0.63 (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.001) and 0.46 (\\u003cem\\u003eP\\u003c/em\\u003e\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.01), respectively. Importantly, B, Fe, and available Al played particularly significant roles in influencing \\u003cem\\u003eA. adenophora\\u003c/em\\u003e coverage in severely invaded areas, highlighting the critical role of soil properties at advanced invasion stages.\\u003c/p\\u003e\\u003cp\\u003e\\u003c/p\\u003e\\u003c/div\\u003e\"},{\"header\":\"4. Discussion\",\"content\":\"\\u003cdiv id=\\\"Sec13\\\"\\u003e\\n \\u003ch2\\u003e4.1 \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion increases the soil pH and alters the levels of soil macronutrients\\u003c/h2\\u003e\\n \\u003cp\\u003eThe soil water content and pH significantly influence plant growth and development (Puissant et al., 2019; Guo et al., 2022). Invasive plants modify soil pH conditions through changes in soil physical properties, such as increased water uptake and root infiltration (Niu et al., 2007; Xiao et al., 2017; Puissant et al., 2019). For example, Zhang and Suseela (2021) reported that the soil pH was significantly lower (pH\\u0026thinsp;=\\u0026thinsp;5.3) in fertilized and invaded areas than in noninvaded areas (pH\\u0026thinsp;=\\u0026thinsp;5.8) and other invaded areas (pH\\u0026thinsp;=\\u0026thinsp;6.1). Additionally, invasive species have been shown to regulate the soil pH and alter the nutrient status, with Xia et al. (2022) demonstrating that significant releases of soil cations in invaded areas led to an increase in the soil pH due to the consumption of hydrogen ions (Boj\\u0026oacute;rquez-Quintal et al., 2017; Fan et al., 2022; Wang et al., 2019; Chen et al., 2022; Marchante et al., 2023). The habitats in our study, which are located in \\u003cem\\u003ePinus yunnanensis\\u003c/em\\u003e forests, are characterized by high soil acidification (Xiao et al., 2017; Wu et al., 2020). We investigated whether \\u003cem\\u003eA. adenophora\\u003c/em\\u003e can leverage its characteristics to promote invasion by improving habitat conditions. Our findings indicated that invasion significantly altered the soil pH in invaded areas, with a notable difference between severely invaded areas (with pH values ranging from approximately 5.4\\u0026ndash;6.2) and noninvaded areas (with pH values ranging from approximately 4.8\\u0026ndash;5.3) (Jones et al., 2019). Furthermore, research from Kashmir Himalaya has indicated that the soil pH in invaded areas is significantly greater than that in noninvaded areas after invasion (Kumar et al., 2021; Ahmad Dar et al., 2023; Dar et al., 2023), confirming that \\u003cem\\u003eA. adenophora\\u003c/em\\u003e can increase the soil pH.\\u003c/p\\u003e\\n \\u003cp\\u003eIn highly acidic and alkaline soils, invasive plants modify their physicochemical properties, affecting nutrient availability, which is crucial for rapid growth and development (Osunkoya and Perrett, 2011; Ren et al., 2021; Yang et al., 2013; Souza-Alonso et al., 2014). The growth of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e relies on N- and P-enriched soils, leading to significant reductions in soil TN and AP concentrations in invaded areas (Khatri et al., 2023; Zhang and Suseela, 2021). This trend is supported by the gradual increase in the SOC content observed with increasing invasion level in our study (Sun et al., 2019; McLeod et al., 2021), as shown in Fig.\\u0026nbsp;2e. Additionally, studies have indicated that plant invasion alters soil enzyme activity levels, thus affecting nutrient effectiveness (Zhang et al., 2022). Changes in soil properties and plant diversity due to \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion have also been observed in Punakha, Bhutan, and in evergreen broad-leaved forests in southwestern Yunnan Province, China (Lhamo et al., 2023; Zhao et al., 2019; Zhang et al., 2023). Our findings revealed that the soil TN content is significantly greater in lightly invaded areas than in severely invaded areas, whereas the soil TK content shows the opposite pattern (Zhang and Suseela, 2021). This difference may stem from the distinct nutrient requirements of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e; notably, in lightly invaded areas, \\u003cem\\u003eA. adenophora\\u003c/em\\u003e may demand more TK for rapid growth, whereas in severely invaded areas, it may need more TN to increase biomass and soil organic matter (Chen et al., 2022; Cotrufo et al., 2022).\\u003c/p\\u003e\\n\\u003c/div\\u003e\\n\\u003cdiv id=\\\"Sec14\\\"\\u003e\\n \\u003ch2\\u003e4.2 Invasion of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e significantly affects soil available micronutrient contents\\u003c/h2\\u003e\\n \\u003cp\\u003eSoil micronutrients, which are essential for plant growth, are effectively used by invasive species such as \\u003cem\\u003eA. adenophora\\u003c/em\\u003e for growth and development (Liu et al., 2023). The relationship between soil enzyme activity and micronutrient levels often results in synergistic or antagonistic interactions with plants, influencing micronutrient uptake (Niu et al., 2007; Kaur et al., 2012; Jones et al., 2019). In this study, we investigated the preferences of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e for specific micronutrients and revealed that all micronutrients except available Al occur at higher concentrations in invaded areas than in noninvaded areas, with Mn, Zn, and Cu exhibiting significant variability across the areas with different invasion levels (Pizzeghello et al., 2011; Wu et al., 2023).\\u003c/p\\u003e\\n \\u003cp\\u003eExcessive or insufficient soil micronutrient levels can be lethal to plants (Osunkoya and Perrett, 2011; Marchante et al., 2023). Recent research has indicated that \\u003cem\\u003eA. adenophora\\u003c/em\\u003e can absorb copper ions from aquatic environments, acting as an adsorbent (Fan et al., 2022). Additionally, \\u003cem\\u003eA. adenophora\\u003c/em\\u003e requires substantial amounts of boron (B) for growth, and B deficiency can be fatal to native plants (Bursali et al., 2009; Chen et al., 2021). Our findings also revealed a decreasing trend in the soil B content with increasing invasion level, whereas the Al content increased (Jones et al., 2019).\\u003c/p\\u003e\\n \\u003cp\\u003ePrevious research has demonstrated that reduced Fe and available Al levels in soil can lead to P loss (Pizzeghello et al., 2011). Consistent with these findings, our study revealed lower AP, Fe, and Al levels in invaded areas than in uninvaded areas (Chapuis-Lardy et al., 2006; Lin et al., 2023), which conforms with findings of Wu et al. (2020). There is consensus that changes in soil micronutrient levels impact soil properties, a phenomenon also observed in areas invaded by \\u003cem\\u003eLantana camara\\u003c/em\\u003e L. (Osunkoya and Perrett, 2011; Datta et al., 2017). Our results demonstrated that \\u003cem\\u003eA. adenophora\\u003c/em\\u003e affects soil micronutrient levels as invasion progresses, particularly those of Mn, Zn, Al, B, Fe, and Cu (Wang et al., 2015; Shen et al., 2019; Souza-Alonso et al., 2014; Jones et al., 2019; Lin et al., 2023). This adaptation enables \\u003cem\\u003eA. adenophora\\u003c/em\\u003e to alter soil properties and manipulate nutrient cycling, thus suppressing the development of native plants (Yang et al., 2013; Xiao et al., 2017; Fan et al., 2022).\\u003c/p\\u003e\\n\\u003c/div\\u003e\\n\\u003cdiv id=\\\"Sec15\\\"\\u003e\\n \\u003ch2\\u003e4.3 Correlations between soil enzyme activity and soil nutrient levels altered by \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion\\u003c/h2\\u003e\\n \\u003cp\\u003eSoil enzyme activity is significantly correlated with both the ambient temperature and soil pH (Esch et al., 2017; Kim et al., 2018). Puissant et al. (2019) reported that the optimal pH for enzyme activity did not always coincide with the environmental soil pH. It is widely acknowledged that invasive plants can modify soil habitats, thereby altering soil properties and nutrient cycling via soil enzyme activities (Shen et al., 2019; Hu et al., 2021; Khatri et al., 2023). In our research, a strong correlation was observed between the soil pH and three soil enzymes, excluding nitrate reductase and urease (Fig.\\u0026nbsp;4). Invasions by \\u003cem\\u003eAmaranthus palmeri\\u003c/em\\u003e and \\u003cem\\u003ePhragmites australis\\u003c/em\\u003e have been shown to significantly impact soil enzyme activity levels (Zhang et al., 2022; Kim et al., 2018). Similarly, our findings indicated that \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion significantly increased the activity of soil sucrase, catalase, and alkaline phosphatase (Wang et al., 2015; Puissant et al., 2019). However, changes in soil enzyme activity can be attributed to various mechanisms (Kim et al., 2018).\\u003c/p\\u003e\\n \\u003cp\\u003eSoil enzyme activity, which is influenced by microorganisms, root secretions, and the decomposition of organic material, plays crucial roles in shaping plant invasion dynamics (DeForest and Moorhead, 2020; Lhamo et al., 2023). In the \\u003cem\\u003eA. adenophora\\u003c/em\\u003e community, variations in enzyme activities across different invasion levels are likely linked to root secretions and apoplastic materials (Wherry, 1920; Yang et al., 2013; Zhao et al., 2019). Our study demonstrated that soil urease activity decreased gradually with increasing invasion level, while alkaline phosphatase activity increased slowly (Fig. 3). These observations conform with the results of other studies, where invasions by \\u003cem\\u003eMimosa pudica\\u003c/em\\u003e and \\u003cem\\u003eFalcataria moluccana\\u003c/em\\u003e led to significant increases in soil urease, catalase, and acid phosphatase activity levels (Wang et al., 2015; Esch et al., 2017; DeForest and Moorhead, 2020; Hu et al., 2021). Furthermore, \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion significantly increased the activities of soil sucrase, catalase, and alkaline phosphatase, contributing to a sharp decline in soil TN and AP contents (Sun et al., 2019; Wang et al., 2019), which agrees with findings for \\u003cem\\u003eMimosa pudica\\u003c/em\\u003e invasion (Wang et al., 2015). This suggests that optimizing the nutrient acquisition efficiency by regulating soil enzyme activity is a common strategy for successful invasion (Zhao et al., 2019; Zhang and Suseela, 2021; Zhang et al., 2022).\\u003c/p\\u003e\\n \\u003cp\\u003e4.4 Soil pH and micronutrients are important factors for \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion in acid-poor areas\\u003c/p\\u003e\\n \\u003cp\\u003e\\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion poses significant ecological threats, including biodiversity loss (Yang et al., 2013; Nunes et al., 2022; Guo et al., 2023) and adverse impacts on agriculture, forestry, and livestock health (Datta et al., 2017; Cotrufo et al., 2022). In the study area, the low soil pH under the canopy of \\u003cem\\u003ePinus yunnanensis\\u003c/em\\u003e has been associated with the increase in soil pH following \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion (Wu et al., 2020; Zhang et al., 2022). In this study, we integrated RFM and SEM analysis methods to systematically analyze the driving mechanisms of the influence of soil physicochemical properties on the aboveground coverage of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e during invasion (Xiao et al., 2017; Jones et al., 2019; Puissant et al., 2019). The results revealed that the invasion strategy of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e exhibits distinct stage-specific characteristics, with its success relying on dynamic adaptation and the regulation of soil environmental factors across different invasion stages (Ren et al., 2021; SUN et al., 2021).\\u003c/p\\u003e\\n \\u003cp\\u003ePrevious studies have demonstrated that \\u003cem\\u003eA. adenophora\\u003c/em\\u003e modulates the soil pH to suit its optimal acclimation range, thereby increasing the soil pH when it is less than 6 and decreasing it when it exceeds 7.5 (Kumar et al., 2021; Wu et al., 2020; Zhang and Suseela, 2021). This raises the question of whether stabilizing the soil pH could effectively control \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion (Niu et al., 2007; Shen et al., 2019; Xu et al., 2023). The RFM results revealed stage-specific differentiation of driving factors during \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion (Fig.\\u0026nbsp;5) (Malik et al., 2018; Guo et al., 2023; Marchante et al., 2023). In noninvaded areas, available Mn, Zn, and B play dominant roles in shaping invasion coverage (p\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05), likely because of the rapid nutrient acquisition strategy of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e during early invasion (Zhang and Suseela, 2021; Guo et al., 2022; Marchante et al., 2023). At the light invasion stage, the increased significance of Al and available Fe (p\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05) suggests that \\u003cem\\u003eA. adenophora\\u003c/em\\u003e mitigates aluminum toxicity by altering soil metal ion speciation while enhancing competitiveness through iron-mediated redox processes (Yang et al., 2013). At the moderate to severe invasion stages, the significant influence of Cu, Mg, and available Ca (p\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05) may be linked to the regulation of the soil cation exchange capacity (CEC) via root exudates to optimize the nutrient acquisition efficiency (Xiao et al., 2017). This dynamic shift in driving factors reflects the evolution of \\u003cem\\u003eA. adenophora\\u003c/em\\u003e strategies from nutrient exploitation to habitat modification, which conforms closely with the positive feedback loop theory of invasive plants (Niu et al., 2007). This feedback mechanism may operate through the following pathways: (1) chelation, which reduces aluminum toxicity and facilitates boron-dependent cell wall synthesis (Chen et al., 2021), and (2) nutrient availability optimization through soil pH modification (Wu et al., 2020). These adaptive strategies enable \\u003cem\\u003eA. adenophora\\u003c/em\\u003e to more efficiently absorb soil nutrients at severe invasion stages, thereby suppressing native plant growth and ultimately displacing them from their ecological niches (Marchante et al., 2023).\\u003c/p\\u003e\\n \\u003cp\\u003eInvasive plants disrupt native plant growth by modifying soil enzyme activity levels to favor their own development (Niu et al., 2007). SEM analysis revealed the synergistic effects of the soil pH and nutrient pathways (Fig.\\u0026nbsp;6). In noninvaded areas, pH indirectly influences coverage by regulating available nutrients (R\\u0026sup2; = 0.7), with the key role of AK (path coefficient\\u0026thinsp;=\\u0026thinsp;0.64, p\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.05) likely related to the efficient use of potassium ion channels by \\u003cem\\u003eA. adenophora\\u003c/em\\u003e during early invasion (Jones et al., 2019). With increasing invasion level, the direct effect of pH decreases, while the path coefficients of soil nutrients (e.g., Al and available Cu) significantly increase (p\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.001), indicating that \\u003cem\\u003eA. adenophora\\u003c/em\\u003e actively modifies the soil environment by altering rhizosphere microbial communities (e.g., acid-producing bacteria abundance) (Puissant et al., 2019; Lin et al., 2023). Particularly at severe invasion stages, the significant influences of available Al and B (path coefficient\\u0026thinsp;=\\u0026thinsp;0.63, p\\u0026thinsp;\\u0026lt;\\u0026thinsp;0.001) may be attributed to their roles in reducing aluminum toxicity through chelation and promoting boron-dependent cell wall synthesis (Chen et al., 2021). This mechanism is particularly advantageous in regions with frequent acid rainfall (pH\\u0026thinsp;\\u0026lt;\\u0026thinsp;5.5) (Wu et al., 2020).\\u003c/p\\u003e\\n \\u003cp\\u003eBy combining RFM and SEM analysis methods, we quantified, for the first time, the threshold effects of the nutrient competition-habitat modification dual-stage model during \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion (Malik et al., 2018; Puissant et al., 2019; Guo et al., 2023). The light invasion stage (R\\u0026sup2; = 0.69) is characterized primarily by direct nutrient competition, which can be mitigated through early intervention targeting available Fe and Al (Lin et al., 2023). In contrast, the severe invasion stage (R\\u0026sup2; = 0.91) requires targeted strategies to disrupt the biogeochemical cycles of trace elements such as Cu and B (Kumar et al., 2021). Notably, the active regulation of pH by \\u003cem\\u003eA. adenophora\\u003c/em\\u003e (with postinvasion pH increases ranging from 0.3\\u0026ndash;0.9 units) may exacerbate the vicious cycle of soil acidification and nutrient loss (Dar et al., 2023), providing a critical target for invasion control in acidic soil regions. Although this study identified key driving factors and their pathways, the impacts of soil microbial functional groups and faunal disturbances on nutrient cycling remain to be incorporated into the model (Lhamo et al., 2023). Future research could aim to integrate metagenomics and stable isotope labeling techniques to elucidate the regulatory mechanisms of rhizosphere microbe‒plant interactions for the turnover of specific nutrients (e.g., available Cu) (Guo et al., 2023). Additionally, on the basis of the stage-specific thresholds established in this study, biochar- or chelator-driven targeted remediation technologies could be developed to assess the feasibility of inhibiting \\u003cem\\u003eA. adenophora\\u003c/em\\u003e expansion by disrupting the Al‒Cu‒B cycles (Fan et al., 2022).\\u003c/p\\u003e\\n\\u003c/div\\u003e\"},{\"header\":\"5. Conclusion\",\"content\":\"\\u003cp\\u003eIn this study, we systematically investigated the dynamic changes and driving mechanisms of soil environmental factors in the invasion process of \\u003cem\\u003eA.\\u003c/em\\u003e \\u003cem\\u003eadenophora\\u0026nbsp;\\u003c/em\\u003eby integrating soil physicochemical properties, enzyme activities, and advanced modeling approaches, including RFM and SEM analysis methods. The results demonstrated that \\u003cem\\u003eA. adenophora\\u0026nbsp;\\u003c/em\\u003einvasion significantly altered soil pH, nutrient content, and enzyme activity levels, with the driving factors exhibiting distinct stage-specific characteristics. Specifically, the soil pH increased with increasing invasion level, with an 8.37% increase in severely invaded areas compared with that in noninvaded regions (\\u003cem\\u003eP\\u003c/em\\u003e \\u0026lt; 0.05). The soil TN and AP contents significantly fluctuated with invasion level, with the TN content increasing by 35.38% in lightly invaded areas and the AP content decreasing by 40.76% in severely invaded regions (\\u003cem\\u003eP\\u003c/em\\u003e \\u0026lt; 0.05). Additionally, soil TK and micronutrients (e.g., Zn, Cu, and Mn) reached peak levels in severely invaded areas. Correlation analysis underscored the pivotal role of pH in regulating soil nutrient availability and microbial activity levels. The RFM results revealed significant differences in key driving factors across invasion stages: available Mn, AK, and available Zn were dominant in noninvaded areas (\\u003cem\\u003eP\\u003c/em\\u003e \\u0026lt; 0.05), whereas Mg, B, and available Cu were the primary drivers in severely invaded regions (\\u003cem\\u003eP\\u003c/em\\u003e \\u0026lt; 0.05). This highlights a shift in \\u003cem\\u003eA. adenophora\\u0026nbsp;\\u003c/em\\u003estrategies from nutrient competition to habitat modification. SEM analysis was employed to quantify the synergistic effects of the soil pH and nutrient pathways and revealed that the path coefficient of available nutrients for coverage in severely invaded areas reached 0.63 (\\u003cem\\u003eP\\u003c/em\\u003e \\u0026lt; 0.001), indicating that \\u003cem\\u003eA. adenophora\\u0026nbsp;\\u003c/em\\u003eexacerbates invasion through plant‒soil feedback mechanisms.\\u003c/p\\u003e\\n\\u003cp\\u003eIn summary, \\u003cem\\u003eA. adenophora\\u0026nbsp;\\u003c/em\\u003einvasion establishes favorable habitat conditions by altering soil pH, nutrient cycling, and enzyme activity levels, thereby suppressing the growth of native plants and ultimately displacing them from their ecological niches. This study provides a systematic basis for the formulation of stage-specific control strategies against \\u003cem\\u003eA. adenophora\\u0026nbsp;\\u003c/em\\u003einvasion. Early intervention through the regulation of the contents of available Fe and Al is recommended at the light invasion stage, whereas targeted strategies to disrupt the biogeochemical cycles of Mg, B, and available Cu are essential for managing severe invasion.\\u003c/p\\u003e\"},{\"header\":\"Declarations\",\"content\":\"\\u003ch2\\u003eAcknowledge\\u003c/h2\\u003e\\n\\u003cp\\u003eThis research was funded by the financial support of Sponsored by Funded Projects: Scientific research projects of China Three Gorges (Construction) Group Corporation (JG-EP-030222001 and JG-EP-030222002); Western Light Project of the Chinese Academy of Sciences (2022XBZG_XBQNXZ_A_003), Sichuan Science and Technology Program (2025ZNSFSC1017).\\u003c/p\\u003e\\n\\u003ch2\\u003eCRediT author statement\\u003c/h2\\u003e\\n\\u003cp\\u003eJuan Wang \\u0026amp; Jingying Lu \\u0026amp; Yuehua Zhang \\u0026amp; Xianyong Dong \\u0026amp; Xiaogang Wu \\u0026amp; Lumei Xiao \\u0026amp; Kaiwen Pan \\u0026amp; Lin Zhang: Sample-plot Investigation, Conceptualization, Methodology, Software, Data curation, Visualization, Writing- Original draft preparation, Writing- Reviewing and Editing, sample-plot Investigation, Supervision, Revise, Writing- Reviewing and Editing, Funding acquisition.\\u003c/p\\u003e\\n\\u003ch2\\u003eDeclaration of interests\\u003c/h2\\u003e\\n\\u003cp\\u003e☒\\u0026nbsp;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.\\u003c/p\\u003e\\n\\u003cp\\u003e☐\\u0026nbsp;The authors declare the following financial interests/personal relationships which may be considered as potential competing interests:\\u003c/p\\u003e\\n\\u003ch2\\u003edata availability statement\\u003c/h2\\u003e\\n\\u003cp\\u003eData will be made available upon request\\u003c/p\\u003e\"},{\"header\":\"References\",\"content\":\"\\u003col\\u003e\\n\\u003cli\\u003eAhmad Dar, M., Ahmad, M., Singh, R., Kumar Kohli, R., Singh, H.P., Batish, D.R., 2023. Invasive plants alter soil properties and nutrient dynamics: A case study of Anthemis cotula invasion in Kashmir Himalaya. Catena 226.\\u003c/li\\u003e\\n\\u003cli\\u003eBills, J.S., Jacinthe, P.-A., Tedesco, L.P., 2010. Soil organic carbon pools and composition in a wetland complex invaded by reed canary grass. Biology and Fertility of Soils 46, 697-706.\\u003c/li\\u003e\\n\\u003cli\\u003eBoj\\u0026oacute;rquez-Quintal, E., Escalante-Maga\\u0026ntilde;a, C., Echevarr\\u0026iacute;a-Machado, I., Mart\\u0026iacute;nez-Est\\u0026eacute;vez, M., 2017. Aluminum, a friend or foe of higher plants in acid soils. Frontiers in plant science 8, 1767.\\u003c/li\\u003e\\n\\u003cli\\u003eBursali, E.A., Cavas, L., Seki, Y., Bozkurt, S.S., Yurdakoc, M., 2009. Sorption of boron by invasive marine seaweed: Caulerpa racemosa var. cylindracea. Chemical Engineering Journal 150, 385-390.\\u003c/li\\u003e\\n\\u003cli\\u003eChapuis-Lardy, L., Vanderhoeven, S., Dassonville, N., Koutika, L.-S., Meerts, P., 2006. Effect of the exotic invasive plant Solidago gigantea on soil phosphorus status. Biology and Fertility of Soils 42, 481-489.\\u003c/li\\u003e\\n\\u003cli\\u003eChen A Q, Zhang D, Peng H, et al., 2013. Experimental study on the development of colapse of ovemhanging lavers of auly in Yuanmou Valevchinaly. Gatena, 109, 177-185.\\u003c/li\\u003e\\n\\u003cli\\u003eChen, L., Xia, F., Wang, M., Mao, P., 2021. Physiological and proteomic analysis reveals the impact of boron deficiency and surplus on alfalfa (Medicago sativa L.) reproductive organs. Ecotoxicology and Environmental Safety 214, 112083.\\u003c/li\\u003e\\n\\u003cli\\u003eChen, S., Gao, D., Zhang, J., M\\u0026uuml;ller, C., Li, X., Zheng, Y., Dong, H., Yin, G., Han, P., Liang, X., 2022. Invasive Spartina alterniflora accelerates soil gross nitrogen transformations to optimize its nitrogen acquisition in an estuarine and coastal wetland of China. Soil Biology and Biochemistry 174, 108835.\\u003c/li\\u003e\\n\\u003cli\\u003eCotrufo, M.F., Haddix, M.L., Kroeger, M.E., Stewart, C.E., 2022. The role of plant input physical-chemical properties, and microbial and soil chemical diversity on the formation of particulate and mineral-associated organic matter. Soil Biology and Biochemistry 168, 108648.\\u003c/li\\u003e\\n\\u003cli\\u003eDar, M.A., Ahmad, M., Singh, R., Kohli, R.K., Singh, H.P., Batish, D.R., 2023. Invasive plants alter soil properties and nutrient dynamics: A case study of Anthemis cotula invasion in Kashmir Himalaya. Catena 226, 107069.\\u003c/li\\u003e\\n\\u003cli\\u003eDarji, T.B., Adhikari, B., Pathak, S., Neupane, S., Thapa, L.B., Bhatt, T.D., Pant, R.R., Pant, G., Pal, K.B., Bishwakarma, K., 2021. Phytotoxic effects of invasive Ageratina adenophora on two native subtropical shrubs in Nepal. Scientific Reports 11, 13663.\\u003c/li\\u003e\\n\\u003cli\\u003eDatta, A., K\\u0026uuml;hn, I., Ahmad, M., Michalski, S., Auge, H., 2017. Processes affecting altitudinal distribution of invasive Ageratina adenophora in western Himalaya: The role of local adaptation and the importance of different life-cycle stages. PloS one 12, e0187708.\\u003c/li\\u003e\\n\\u003cli\\u003eDeForest, J.L., Moorhead, D.L., 2020. Effects of elevated pH and phosphorus fertilizer on soil C, N and P enzyme stoichiometry in an acidic mixed mesophytic deciduous forest. Soil Biology and Biochemistry 150, 107996.\\u003c/li\\u003e\\n\\u003cli\\u003eEsch, E.H., Lipson, D., Cleland, E.E., 2017. Direct and indirect effects of shifting rainfall on soil microbial respiration and enzyme activity in a semi-arid system. Plant and soil 411, 333-346.\\u003c/li\\u003e\\n\\u003cli\\u003eFan, L., Miao, J., Yang, J., Zhao, X., Shi, W., Xie, M., Wang, X., Chen, W., An, X., Luo, H., 2022. Invasive plant-crofton weed as adsorbent for effective removal of copper from aqueous solution. Environmental Technology \\u0026amp; Innovation 26, 102280.\\u003c/li\\u003e\\n\\u003cli\\u003eGasch, C.K., Enloe, S.F., Stahl, P.D., Williams, S.E., 2013. An aboveground-belowground assessment of ecosystem properties associated with exotic annual brome invasion. Biology and Fertility of Soils 49, 919-928.\\u003c/li\\u003e\\n\\u003cli\\u003eGuo, K., Py\\u0026scaron;ek, P., Chytr\\u0026yacute;, M., Div\\u0026iacute;\\u0026scaron;ek, J., Lososov\\u0026aacute;, Z., van Kleunen, M., Pierce, S., Guo, W.Y., 2022. Ruderals naturalize, competitors invade: Varying roles of plant adaptive strategies along the invasion continuum. Functional Ecology 36, 2469-2479.\\u003c/li\\u003e\\n\\u003cli\\u003eGuo, K., Zheng, M.-M., Liu, R.-L., Wang, Y.-Y., Gao, Y., Shu, L., Wang, X.-R., Zhang, J., Guo, W.-Y., 2023. Intraspecific variations of adaptive strategies of native and invasive plant species along an elevational gradient. Flora 304, 152297.\\u003c/li\\u003e\\n\\u003cli\\u003eHu, Z., Li, J., Shi, K., Ren, G., Dai, Z., Sun, J., Zheng, X., Zhou, Y., Zhang, J., Li, G., 2021. Effects of Canada goldenrod invasion on soil extracellular enzyme activities and ecoenzymatic stoichiometry. Sustainability 13, 3768.\\u003c/li\\u003e\\n\\u003cli\\u003eJones, D.L., Cooledge, E.C., Hoyle, F.C., Griffiths, R.I., Murphy, D.V., 2019. pH and exchangeable aluminum are major regulators of microbial energy flow and carbon use efficiency in soil microbial communities. Soil Biology and Biochemistry 138, 107584.\\u003c/li\\u003e\\n\\u003cli\\u003eKabała, C., Łabaz, B., 2018. Relationships between soil pH and base saturation-conclusions for Polish and international soil classifications. Soil Science Annual 69.\\u003c/li\\u003e\\n\\u003cli\\u003eKaur, R., Malhotra, S., Inderjit, 2012. Effects of invasion of Mikania micrantha on germination of rice seedlings, plant richness, chemical properties and respiration of soil. Biology and Fertility of Soils 48, 481-488.\\u003c/li\\u003e\\n\\u003cli\\u003eKhatri, K., Negi, B., Bargali, K., Bargali, S.S., 2023. Phenotypic variation in morphology and associated functional traits in Ageratina adenophora along an altitudinal gradient in Kumaun Himalaya, India. Biologia 78, 1333-1347.\\u003c/li\\u003e\\n\\u003cli\\u003eKim, S., Kang, J., Megonigal, J.P., Kang, H., Seo, J., Ding, W., 2018. Impacts of Phragmites australis invasion on soil enzyme activities and microbial abundance of tidal marshes. Microbial ecology 76, 782-790.\\u003c/li\\u003e\\n\\u003cli\\u003eKumar, M., Kumar, S., Verma, A.K., Joshi, R.K., Garkoti, S.C., 2021. Invasion of Lantana camara and Ageratina adenophora alters the soil physico-chemical characteristics and microbial biomass of chir pine forests in the central Himalaya, India. Catena 207, 105624.\\u003c/li\\u003e\\n\\u003cli\\u003eLhamo, S., Thinley, U., Dorji, U., 2023. Impacts of Invasion by Ageratina adenophora on Soil Properties and Plant Diversity. Bhutan Journal of Natural Resources and Development 10, 1-9.\\u003c/li\\u003e\\n\\u003cli\\u003eLi, Q., Wan, F., Zhao, M., 2022. Distinct soil microbial communities under Ageratina adenophora invasions. Plant Biology 24, 430-439.\\u003c/li\\u003e\\n\\u003cli\\u003eLi, Y.-P., Li, W.-T., Li, J., Feng, Y.-L., 2023. Temporal dynamics of plant\\u0026minus; soil feedback and related mechanisms depend on environmental context during invasion processes of a subtropical invader. Plant and soil, 1-16.\\u003c/li\\u003e\\n\\u003cli\\u003eLin, M., Chen, Y., Cheng, L., Zheng, Y., Wang, W., Sardans, J., Song, Z., Guggenberger, G., Zou, Y., Ding, X., 2023. Response of topsoil Fe-bound organic carbon pool and microbial community to Spartina alterniflora invasion in coastal wetlands. Catena 232, 107414.\\u003c/li\\u003e\\n\\u003cli\\u003eLiu, Y.-m., Li, W.-t., Zheng, Y.-l., 2023. The effect of acid rain and fertilization on the performance of invasive Chromolaena odorata and two native plants. Acta Oecologica 120, 103938.\\u003c/li\\u003e\\n\\u003cli\\u003eMalik, A.A., Puissant, J., Buckeridge, K.M., Goodall, T., Jehmlich, N., Chowdhury, S., Gweon, H.S., Peyton, J.M., Mason, K.E., van Agtmaal, M., 2018. Land use driven change in soil pH affects microbial carbon cycling processes. Nature communications 9, 3591.\\u003c/li\\u003e\\n\\u003cli\\u003eMarchante, H., Marchante, E., Verbrugge, L., Lommen, S., Shaw, R., 2023. Knowledge and perceptions of invasive plant biocontrol in Europe versus the rest of the world. Journal of Environmental Management 327, 116896.\\u003c/li\\u003e\\n\\u003cli\\u003eMcLeod, M.L., Bullington, L., Cleveland, C.C., Rousk, J., Lekberg, Y., 2021. Invasive plant-derived dissolved organic matter alters microbial communities and carbon cycling in soils. Soil Biology and Biochemistry 156, 108191.\\u003c/li\\u003e\\n\\u003cli\\u003eMerry, R.H., Spouncer, L.R., Fitzpatrick, R.W., Davies, P.J., Bruce, D.A., McVicar, T., Rui, L., Walker, J., 2002. Regional prediction of soil profile acidity and alkalinity. ACIAR MONOGRAPH SERIES 84, 155-164.\\u003c/li\\u003e\\n\\u003cli\\u003eNiu, H.-b., Liu, W.-x., Wan, F.-h., Liu, B., 2007. An invasive aster (Ageratina adenophora) invades and dominates forest understories in China: altered soil microbial communities facilitate the invader and inhibit natives. Plant and soil 294, 73-85.\\u003c/li\\u003e\\n\\u003cli\\u003eNunes, M., Lemley, D.A., Adams, J.B., 2022. Flow regime and nutrient input control invasive alien aquatic plant distribution and species composition in small closed estuaries. Science of the Total Environment 819, 152038.\\u003c/li\\u003e\\n\\u003cli\\u003eOsunkoya, O.O., Perrett, C., 2011. Lantana camara L.(Verbenaceae) invasion effects on soil physicochemical properties. Biology and Fertility of Soils 47, 349-355.\\u003c/li\\u003e\\n\\u003cli\\u003ePizzeghello, D., Berti, A., Nardi, S., Morari, F., 2011. Phosphorus forms and P-sorption properties in three alkaline soils after long-term mineral and manure applications in north-eastern Italy. Agriculture, ecosystems \\u0026amp; environment 141, 58-66.\\u003c/li\\u003e\\n\\u003cli\\u003ePuissant, J., Jones, B., Goodall, T., Mang, D., Blaud, A., Gweon, H.S., Malik, A., Jones, D.L., Clark, I.M., Hirsch, P.R., 2019. The pH optimum of soil exoenzymes adapt to long term changes in soil pH. Soil Biology and Biochemistry 138, 107601.\\u003c/li\\u003e\\n\\u003cli\\u003eRen, Z., Okyere, S.K., Wen, J., Xie, L., Cui, Y., Wang, S., Wang, J., Cao, S., Shen, L., Ma, X., 2021. An overview: the toxicity of Ageratina adenophora on animals and its possible interventions. International Journal of Molecular Sciences 22, 11581.\\u003c/li\\u003e\\n\\u003cli\\u003eShen, S., Xu, G., Li, D., Jin, G., Liu, S., Clements, D.R., Yang, Y., Rao, J., Chen, A., Zhang, F., 2019. Ipomoea batatas (sweet potato), a promising replacement control crop for the invasive alien plant Ageratina adenophora (Asteraceae) in China. Management of Biological Invasions 10, 559-572.\\u003c/li\\u003e\\n\\u003cli\\u003eSouza-Alonso, P., Novoa, A., Gonz\\u0026aacute;lez, L., 2014. Soil biochemical alterations and microbial community responses under Acacia dealbata Link invasion. Soil Biology and Biochemistry 79, 100-108.\\u003c/li\\u003e\\n\\u003cli\\u003eSun, F., Ou, Q., Yu, H., Li, N., Peng, C., 2019. The invasive plant Mikania micrantha affects the soil foodweb and plant-soil nutrient contents in orchards. Soil Biology and Biochemistry 139, 107630.\\u003c/li\\u003e\\n\\u003cli\\u003eSUN, Y.-y., ZHANG, Q.-x., ZHAO, Y.-p., DIAO, Y.-h., GUI, F.-r., YANG, G.-q., 2021. Beneficial rhizobacterium provides positive plant-soil feedback effects to Ageratina adenophora. Journal of Integrative Agriculture 20, 1327-1335.\\u003c/li\\u003e\\n\\u003cli\\u003eVerma, A.K., Nayak, R., Manika, N., Bargali, K., Pandey, V.N., Chaudhary, L.B., Behera, S.K., 2023. Monitoring the distribution pattern and invasion status of Ageratina adenophora across elevational gradients in Sikkim Himalaya, India. Environmental Monitoring and Assessment 195, 152.\\u003c/li\\u003e\\n\\u003cli\\u003eWang, C., Wu, B., Jiang, K., Zhou, J., Liu, J., Lv, Y., 2019. Canada goldenrod invasion cause significant shifts in the taxonomic diversity and community stability of plant communities in heterogeneous landscapes in urban ecosystems in East China. Ecological Engineering 127, 504-509.\\u003c/li\\u003e\\n\\u003cli\\u003eWang, R., Dai, T., Quan, G., Zhang, J., 2015. Changes in soil physico-chemical properties, enzyme activities and soil microbial communities under Mimosa pudica invasion. Allelopathy J 36, 15-24.\\u003c/li\\u003e\\n\\u003cli\\u003eWu, X., Duan, C., Fu, D., Peng, P., Zhao, L., Jones, D.L., 2020. Effects of Ageratina adenophora invasion on the understory community and soil phosphorus characteristics of different forest types in southwest China. Forests 11, 806.\\u003c/li\\u003e\\n\\u003cli\\u003eWu, X., Xing, H., Wang, X., Yang, J., Chen, J., Liu, X., Dai, D., Zhang, M., Yang, Q., Dong, S., 2023. Changes in soil microbial communities are linked to metal elements in a subtropical forest. Applied Soil Ecology 188, 104919.\\u003c/li\\u003e\\n\\u003cli\\u003eXia, H., Riaz, M., Liu, B., Li, Y., El-Desouki, Z., Jiang, C., 2022. Over two years study: Peanut biochar promoted potassium availability by mediating the relationship between bacterial community and soil properties. Applied Soil Ecology 176, 104485.\\u003c/li\\u003e\\n\\u003cli\\u003eXiao, H., Schaefer, D.A., Yang, X., 2017. pH drives ammonia oxidizing bacteria rather than archaea thereby stimulate nitrification under Ageratina adenophora colonization. Soil Biology and Biochemistry 114, 12-19.\\u003c/li\\u003e\\n\\u003cli\\u003eXiao, L., Min, X., Liu, G., Li, P., Xue, S., 2023. Effect of plant-plant interactions and drought stress on the response of soil nutrient contents, enzyme activities and microbial metabolic limitations. Applied Soil Ecology 181, 104666.\\u003c/li\\u003e\\n\\u003cli\\u003eXu, C.-W., Yang, M.-Z., Chen, Y.-J., Chen, L.-M., Zhang, D.-Z., Mei, L., Shi, Y.-T., Zhang, H.-B., 2012. Changes in non-symbiotic nitrogen-fixing bacteria inhabiting rhizosphere soils of an invasive plant Ageratina adenophora. Applied Soil Ecology 54, 32-38.\\u003c/li\\u003e\\n\\u003cli\\u003eXu, Z., Xu, J., Chen, P., Zhong, S., Xu, Z., Yu, Y., Wang, C., Du, D., 2023. Heavy metal pollution is more conducive to the independent invasion of Solidago canadensis L. than the co-invasion of two Asteraceae invasive plants. Acta Oecologica 120, 103934.\\u003c/li\\u003e\\n\\u003cli\\u003eYang, Q., Carrillo, J., Jin, H., Shang, L., Hovick, S.M., Nijjer, S., Gabler, C.A., Li, B., Siemann, E., 2013. Plant-soil biota interactions of an invasive species in its native and introduced ranges: Implications for invasion success. Soil Biology and Biochemistry 65, 78-85.\\u003c/li\\u003e\\n\\u003cli\\u003eZhang, M., Li, X., Qiu, Z., Shi, C., Wang, K., Fukuda, K., Shi, F., 2022. Effects of Amaranthus palmeri invasion on soil extracellular enzyme activities and enzymatic stoichiometry. Journal of Soil Science and Plant Nutrition 22, 5183-5194.\\u003c/li\\u003e\\n\\u003cli\\u003eZhang, P., Nie, M., Li, B., Wu, J., 2017. The transfer and allocation of newly fixed C by invasive Spartina alterniflora and native Phragmites australis to soil microbiota. Soil Biology and Biochemistry 113, 231-239.\\u003c/li\\u003e\\n\\u003cli\\u003eZhang, X., Wang, G., Peng, P., Zhou, Y., Chen, Z., Feng, Y., Wang, Y., Shi, S., Li, J., 2023. Influences of environment, human activity, and climate on the invasion of Ageratina adenophora (Spreng.) in Southwest China. PeerJ 11, e14902.\\u003c/li\\u003e\\n\\u003cli\\u003eZhang, Z., Suseela, V., 2021. Nitrogen availability modulates the impacts of plant invasion on the chemical composition of soil organic matter. Soil Biology and Biochemistry 156, 108195.\\u003c/li\\u003e\\n\\u003cli\\u003eZhao, M., Lu, X., Zhao, H., Yang, Y., Hale, L., Gao, Q., Liu, W., Guo, J., Li, Q., Zhou, J., 2019. Ageratina adenophora invasions are associated with microbially mediated differences in biogeochemical cycles. Science of the Total Environment 677, 47-56.\\u003c/li\\u003e\\n\\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\":\"info@researchsquare.com\",\"identity\":\"biological-invasions\",\"isNatureJournal\":false,\"hasQc\":true,\"allowDirectSubmit\":false,\"externalIdentity\":\"binv\",\"sideBox\":\"Learn more about [Biological Invasions](https://www.springer.com/journal/10530)\",\"snPcode\":\"10530\",\"submissionUrl\":\"https://submission.nature.com/new-submission/10530/3\",\"title\":\"Biological Invasions\",\"twitterHandle\":\"\",\"acdcEnabled\":true,\"dfaEnabled\":true,\"editorialSystem\":\"em\",\"reportingPortfolio\":\"Springer Hybrid\",\"inReviewEnabled\":true,\"inReviewRevisionsEnabled\":false},\"keywords\":\"Level of invasion, Ageratina adenophora, soil pH, soil nutrients, soil enzyme activity, structural equation model\",\"lastPublishedDoi\":\"10.21203/rs.3.rs-8012262/v1\",\"lastPublishedDoiUrl\":\"https://doi.org/10.21203/rs.3.rs-8012262/v1\",\"license\":{\"name\":\"CC BY 4.0\",\"url\":\"https://creativecommons.org/licenses/by/4.0/\"},\"manuscriptAbstract\":\"\\u003cp\\u003e\\u003cem\\u003eAgeratina adenophora\\u003c/em\\u003e poses significant threats to agricultural and forestry production and biodiversity conservation worldwide. However, its expansion mechanism in highly acidic environments has not been studied in depth. To address this issue, we investigated the impacts of \\u003cem\\u003eA. adenophora\\u003c/em\\u003einvasion on soil nutrient content and enzyme activity levels across 24 samples from Yunnan Province, China. The sampling sites were categorized into four invasion levels, namely, noninvaded (C), lightly invaded (L), moderately invaded (M), and severely invaded (S). Our findings revealed that the soil pH in severely invaded areas was 8.37% greater than that in noninvaded areas, with pH values ranging from 4.8 to 5.3. Notably, severely invaded soils demonstrated relatively high levels of soil organic carbon (SOC), available potassium (AK), aluminum (Al), available iron (Fe), available zinc (Zn), available copper (Cu), available manganese (Mn), exchangeable calcium (Ca), and exchangeable magnesium (Mg). However, the levels of available Al, boron (B), and phosphorus were significantly lower in these areas. Additionally, variations in the total nitrogen (TN), total potassium (TK), sucrase, and nitrate reductase activity levels were observed across the areas with different invasion levels. Correlation analysis underscored the pivotal role of pH in regulating soil nutrient availability and microbial activity levels. The random forest model (RFM) and structural equation modeling (SEM) results indicated that available Mn, AK, and available Zn are the dominant factors in noninvaded areas (p \\u0026lt; 0.05), while Mg, B, and available Cu were the main factors in severely invaded areas (p \\u0026lt; 0.05). These findings collectively demonstrate that \\u003cem\\u003eA. adenophora\\u003c/em\\u003e invasion establishes favorable habitat conditions by altering soil pH, nutrient cycling, and enzyme activity levels, thereby suppressing the growth of native plants and ultimately displacing them from their ecological niches. This study provides a sound foundation for the formulation of stage-specific control strategies against \\u003cem\\u003eA. adenophora \\u003c/em\\u003einvasion.\\u003c/p\\u003e\",\"manuscriptTitle\":\"The Soil pH and Micronutrients Drive Ageratina Adenophora Invasion in Areas with Acidic and Nutrient-Poor Soils\",\"msid\":\"\",\"msnumber\":\"\",\"nonDraftVersions\":[{\"code\":1,\"date\":\"2025-11-28 14:57:13\",\"doi\":\"10.21203/rs.3.rs-8012262/v1\",\"editorialEvents\":[{\"type\":\"communityComments\",\"content\":0},{\"type\":\"reviewerAgreed\",\"content\":\"\",\"date\":\"2025-12-11T12:38:21+00:00\",\"index\":0,\"fulltext\":\"\"},{\"type\":\"reviewersInvited\",\"content\":\"\",\"date\":\"2025-11-25T06:40:28+00:00\",\"index\":\"\",\"fulltext\":\"\"},{\"type\":\"editorInvited\",\"content\":\"Biological Invasions\",\"date\":\"2025-11-15T20:31:22+00:00\",\"index\":\"\",\"fulltext\":\"\"},{\"type\":\"editorAssigned\",\"content\":\"\",\"date\":\"2025-11-03T12:57:04+00:00\",\"index\":\"\",\"fulltext\":\"\"},{\"type\":\"submitted\",\"content\":\"Biological Invasions\",\"date\":\"2025-11-02T11:32:15+00:00\",\"index\":\"\",\"fulltext\":\"\"}],\"status\":\"published\",\"journal\":{\"display\":true,\"email\":\"info@researchsquare.com\",\"identity\":\"biological-invasions\",\"isNatureJournal\":false,\"hasQc\":true,\"allowDirectSubmit\":false,\"externalIdentity\":\"binv\",\"sideBox\":\"Learn more about [Biological Invasions](https://www.springer.com/journal/10530)\",\"snPcode\":\"10530\",\"submissionUrl\":\"https://submission.nature.com/new-submission/10530/3\",\"title\":\"Biological Invasions\",\"twitterHandle\":\"\",\"acdcEnabled\":true,\"dfaEnabled\":true,\"editorialSystem\":\"em\",\"reportingPortfolio\":\"Springer Hybrid\",\"inReviewEnabled\":true,\"inReviewRevisionsEnabled\":false}}],\"origin\":\"\",\"ownerIdentity\":\"f149b169-2b0a-4ce0-8e3a-501886a0f7d4\",\"owner\":[],\"postedDate\":\"November 28th, 2025\",\"published\":true,\"recentEditorialEvents\":[],\"rejectedJournal\":[],\"revision\":\"\",\"amendment\":\"\",\"status\":\"under-review\",\"subjectAreas\":[],\"tags\":[],\"updatedAt\":\"2025-11-28T14:57:13+00:00\",\"versionOfRecord\":[],\"versionCreatedAt\":\"2025-11-28 14:57:13\",\"video\":\"\",\"vorDoi\":\"\",\"vorDoiUrl\":\"\",\"workflowStages\":[]},\"version\":\"v1\",\"identity\":\"rs-8012262\",\"journalConfig\":\"researchsquare\"},\"__N_SSP\":true},\"page\":\"/article/[identity]/[[...version]]\",\"query\":{\"redirect\":\"/article/rs-8012262\",\"identity\":\"rs-8012262\",\"version\":[\"v1\"]},\"buildId\":\"XKTyCvWXoU3ODBz1xrDgd\",\"isFallback\":false,\"isExperimentalCompile\":false,\"dynamicIds\":[84888],\"gssp\":true,\"scriptLoader\":[]}","source_license":"CC-BY-4.0","license_restricted":false}