Dynamic impact of alien plant invasion on plant diversity: a global meta-analysis

Dynamic impact of alien plant invasion on plant diversity: a global meta-analysis | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 26 September 2025 V1 Latest version Share on Dynamic impact of alien plant invasion on plant diversity: a global meta-analysis Authors : Zhaoqi Zhu 0009-0000-1364-7485 , Li Fan , Xue Fan , Haochen Yu , Miaomiao Cui 0000-0002-6578-9298 , Yijian Wang , Shoujiang Liu , Zhi-Cong Dai 0000-0002-0748-8059 , GuangQian Ren , and Daolin Du [email protected] Authors Info & Affiliations https://doi.org/10.22541/au.175888646.66598029/v1 293 views 194 downloads Contents Abstract Abstract： 1. Introduction Supplementary Material Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Invasive alien plants not only alter the structure and functioning of ecosystems but can also pose serious threats to native biodiversity. Understanding vegetation dynamics and succession across temporal gradients remains a central issue in plant ecology, yet the long-term effects of plant invasions on biodiversity lack a clear consensus. In this study, we conducted a global meta-analysis of 635 data sets drawn from 79 peer-reviewed publications to quantitatively assess the temporal impacts of invasive alien plants on plant diversity. Overall, invasions significantly reduced local plant diversity by 22.57% (P<0.05) and exhibited a gradual downward trend with increasing invasion duration. External disturbances showed a partial restorative effect: compared with invaded plots, disturbed plots had on average 13.71% higher diversity (P<0.05), but their diversity remained 11.07% lower than that of uninvaded plots (P<0.05). Differences among groups across varying invasion stages were relatively small, suggesting that invasion duration exerted only a minor influence on changes in diversity. Subgroup analyses further revealed that climatic zone, habitat type, and disturbance type are key moderators of invasion impacts on diversity, with their effects being more pronounced under disturbance conditions. These findings underscore the need to integrate climatic context, disturbance regimes, and habitat characteristics when predicting and managing plant invasions, and they provide quantitative evidence to inform ecological risk assessments and targeted management strategies for invasive alien plants worldwide. Dynamic impact of alien plant invasion on plant diversity: a global meta-analysis Abstract： Invasive alien plants not only alter the structure and functioning of ecosystems but can also pose serious threats to native biodiversity. Understanding vegetation dynamics and succession across temporal gradients remains a central issue in plant ecology, yet the long-term effects of plant invasions on biodiversity lack a clear consensus. In this study, we conducted a global meta-analysis of 635 data sets drawn from 79 peer-reviewed publications to quantitatively assess the temporal impacts of invasive alien plants on plant diversity. Overall, invasions significantly reduced local plant diversity by 22.57% ( P <0.05) and exhibited a gradual downward trend with increasing invasion duration. External disturbances showed a partial restorative effect: compared with invaded plots, disturbed plots had on average 13.71% higher diversity ( P <0.05), but their diversity remained 11.07% lower than that of uninvaded plots ( P <0.05). Differences among groups across varying invasion stages were relatively small, suggesting that invasion duration exerted only a minor influence on changes in diversity. Subgroup analyses further revealed that climatic zone, habitat type, and disturbance type are key moderators of invasion impacts on diversity, with their effects being more pronounced under disturbance conditions. These findings underscore the need to integrate climatic context, disturbance regimes, and habitat characteristics when predicting and managing plant invasions, and they provide quantitative evidence to inform ecological risk assessments and targeted management strategies for invasive alien plants worldwide. Keywords: Alien plant invasion; Plant diversity; Dynamic impact; Temporal scale; Invasion stage ¿p#1 1. Introduction With the accelerating process of globalization, the frequency and extent of cross-regional species dispersal have increased significantly, and plant invasions by alien species have been widely recognized as one of the major global ecological and environmental issues (Gaertner et al., 2014). At present, very few ecosystems remain unaffected by alien plants. These invasive species often overcome biogeographical barriers and expand rapidly, exerting profound impacts on native ecosystems (Essl et al., 2018; Richardson & Gaertner, 2013). Studies have shown that alien plant invasions not only reduce the diversity and abundance of native plant communities, but may also alter the genetic structure of native populations, disrupt the behavioral patterns of native animals, affect the phylogenetic diversity of communities, and reshape trophic network structures (Jones et al., 2020; Liao et al., 2008; Pyšek et al., 2020). Overall, alien plant invasions are generally considered to have negative impacts on biodiversity, most directly manifested in the decline of native plant diversity; however, their specific effects vary depending on ecosystem characteristics and invasion contexts (Xu et al., 2022; Zhu et al., 2025). In ecological research, plant diversity is one of the key indicators for assessing ecosystem health, as it effectively reflects changes in plant communities within a region and captures both species richness and evenness (Sanaei et al., 2018; Vellend et al., 2013). Plant diversity is not only directly linked to the structure and functioning of ecosystems, but also influences their productivity, stability, and capacity to respond to environmental change (Lange et al., 2015). Over the past few decades, the impact of invasive alien plants on plant diversity has remained a central focus of ecological studies, with numerous empirical and synthetic investigations conducted in this field. Statistics suggest that approximately 63% of studies have reported significant alterations in species, communities, or ecosystem attributes following invasion events (Pyšek et al., 2020). For example, a meta-analysis by Vilà et al. (2011), which synthesized 1,041 field studies from 199 publications, demonstrated that invasive plants typically reduce the abundance and diversity of native species and reshape the community structure of invaded regions. However, some scholars have put forward different perspectives. Vellend et al. (2017), for instance, argued that the introduction of alien plants has substantially increased species richness in many regions worldwide and may even facilitate the formation of novel hybrid polyploid species. Overall, changes in regional species diversity are heterogeneous, with impacts that may be negative in some cases but potentially beneficial in others. However, plant invasions are a continuous and dynamic process rather than a static phenomenon (Gioria et al., 2023; Jandova et al., 2014). During the course of invasion, the colonization, replacement, and loss of both alien and native plant species often occur, thereby exerting profound impacts on the community structure and functioning of the invaded ecosystems (Liu et al., 2022). At the same time, plant communities undergoing succession are frequently subject to natural disturbances (e.g., natural disasters or herbivory) or anthropogenic disturbances (e.g., herbicide application, manual removal, or prescribed burning), which lead to divergent outcomes in plant diversity (Guido & Pillar, 2017; Meffin et al., 2010). Previous studies on invasion dynamics and their ecological impacts have mostly adopted a ”one-time snapshot” approach, often neglecting the invasion starting point as well as its temporal dynamics (D’Antonio & Flory, 2017; Strayer et al., 2006). Among the hundreds of studies assessing the dynamic impacts of alien plant invasions, the vast majority lasted only one year, while research on long-term invasion impacts remains relatively limited (D’Antonio & Flory, 2017; Karatayev et al., 2023). Nevertheless, over the past decade, research on invasion dynamics has increased, with growing attention directed toward the long-term effects of invasions on plant communities (Stricker et al., 2015). For example, a 40-year study on the invasion of Ramnus cathartica demonstrated that the species’ cover increased significantly with invasion duration, while the cover of native species declined, leading to reduced community diversity over time (Ortega et al., 2019). Conversely, a study on Acacia longifolia found that with increasing invasion duration, species richness gradually declined, and the cover, diversity, and turnover rate of native plants decreased. These changes were more pronounced during the early stages of invasion but tended to stabilize after several decades (Marchante et al., 2015). In addition, under invasion scenarios involving external disturbances, plant diversity showed significant short-term recovery (6 years), but the effectiveness of recovery was limited under long-term interventions (25 years) (Chikowore et al., 2025; Marchante et al., 2011). To date, the long-term dynamic impacts of alien plants on the plant communities they invade remain controversial, and no consensus has been reached within the academic community. Furthermore, when delineating invasion stages, most researchers rely on ecological characteristics of the invasion process, while few studies explicitly use invasion duration as the primary criterion. To our knowledge, no review has systematically analyzed changes in plant diversity from the perspective of invasion duration. In this context, we conducted a meta-analysis of 635 data sets extracted from 79 published studies to provide a comprehensive assessment of plant diversity changes across different invasion stages. Our objectives were to: (1) quantify the overall and temporal impacts of alien plant invasions on plant diversity, (2) determine the differentiated outcomes of alien plant invasion on plant diversity under different contexts, and (3) further analyze the causes of these differentiated outcomes. We hypothesized that: (1) alien plant invasions exert negative impacts on plant diversity, and that these impacts intensify over time; (2) external disturbances can facilitate the recovery of plant diversity in invaded areas; and (3) the effects of alien plant invasions on local plant diversity may also be influenced by climate type, plot size, habitat type, and disturbance type. 2. Materials and methods 2.1. Data collection To comprehensively investigate the dynamic impacts of alien plant invasions on plant diversity, we conducted a systematic literature search on May 23, 2025 using the Web of Science and China National Knowledge Infrastructure (CNKI) databases. The search strategy employed the terms TS = (invasive plants OR plant invasions) AND TS = (plant diversity OR plant richness), with no restriction on publication year. To meet the requirements of a meta-analysis, the specific inclusion criteria were set as follows: (1) The study must clearly specify the starting year of the alien plant invasion and provide the corresponding research year(s) or time interval; (2) The study must include both a treatment group and a control group, with no fewer than three replicates in each group (n≥3); (3) The study must report either a species richness index (R) or a Shannon-Wiener index (H), presented in numerical form or as figures/graphs; (4) Results presented in graphical form must allow for the extraction of specific values using image processing software; (5) The study must provide the mean values and sample sizes for the extracted metrics. If the standard deviation (SD) is not reported, it should be derived from the standard error (SE) using the formula . If neither SD nor SE is provided, the SD should be approximated as one-tenth of the mean (Zhou et al., 2024). Based on the PRISMA guidelines and the screening criteria described above, this meta-analysis ultimately included 79 publications (see details in Appendix: Figure S1), from which a total of 635 valid data sets were extracted. According to the selected studies, the data fall into two main categories: alien plant invasions under natural conditions and those under experimental or disturbance conditions. The geographic distribution of sampling sites is shown in Figure 1. All data were obtained from figures and tables in the literature; for data presented in graphical form, values were extracted using GetData Graph Digitizer version 2.24. Figure 1. Global distribution of studies included in this review 2.2 Heterogeneity and publication bias tests This study employed the Q test and the I² test to assess heterogeneity in plant diversity under alien plant invasion (Table S1). The results showed that, under natural conditions, the Q value was 216.192 ( P = 0.00050%, and under disturbed conditions, the Q value was 4672.219 ( P = 0.00050%. These findings indicate significant heterogeneity among the study samples. Consequently, a random-effects model was applied in subsequent analyses, and the extracted data required further refinement. Publication bias was evaluated using a funnel plot and the fail-safe number (Figure S2). The funnel plot revealed that most effect sizes were symmetrically distributed on both sides of the mean effect size and concentrated toward the upper part of the funnel. Moreover, the fail-safe numbers were 677.2 and 18,336.4, both exceeding the threshold of 5n+10, indicating no evidence of publication bias in the selected literature. Therefore, the final results are considered robust and reliable. 2.3 Data categorization Based on the research objectives, and drawing on previous studies of invasion duration as well as the data we extracted, we classified the duration of alien plant invasion into four stages: < 3 years (Ⅰ), 3-10 years (Ⅱ), 11-30 years (Ⅲ), and 31-100 years (Ⅳ) (Strayer et al., 2006). The time‐scale classification for disturbance under invasion followed the same scheme, representing the length of time the invaded sites had been subject to disturbance. This approach provides a sound rationale and facilitates subsequent processing and analysis. In addition, to account for differences between control groups, plant invasion under disturbance was divided into two categories: sites invaded by alien plants serving as the control (invaded) and non-invaded sites serving as the control (non-invaded). Finally, based on the results of the preceding heterogeneity tests, we conducted further subgroup analyses and grouped the extracted data accordingly (Table 1). Table 1 Data grouping ¿p#1 R NT SS Grassland (GL) Herbicide application (HA) H T MS Shrubland (SL) Physical removal (PR) ST LS Woodland (WL) Grazing (GR) Dune system (DS) Fire treatment (FT) Wetland (WT) Multiple disturbances (MD) Riparian zone (RZ) Other disturbances (OD) Note: NT represents the Northern Temperate Zone (23°26′-66°34′ N); T represents the Tropics (23°26′ S-23°26′ N); ST represents the Southern Temperate Zone (23°26′-66°34′ S). SS denotes small-scale plots (1000 m²). Multiple disturbances refer to the simultaneous application of two or more disturbance treatments (e.g., a combination of herbicide application and grazing). Other disturbances include urbanization, tarp shading, fertilization, biological control, wild boar disturbance, logging treatments, and seeding restoration (a total of 40 data groups). 2.4 Data analysis To improve the accuracy of the effect sizes in the meta-analysis, we used the logarithm response ratio () as the effect size to evaluate the impact of alien plant invasions on plant diversity across temporal scales. The variance corresponding to is denoted as, The formulas for calculating and are as follows (Hedges et al., 1999; Shangguan et al., 2024): （1） （2） where and are the means for the treatment and control groups, respectively; and are the standard deviations for the treatment and control groups, respectively; and are the sample sizes for the treatment and control groups, respectively. In order to improve the accuracy of our research, we employed a random effects model to calculate the combined effect size/mean weighted response ratio (), with studies of higher precision being assigned greater weight. The calculation formulas for () and its weighting factor (), standard error (), and 95% bootstrap confidence interval () are as follows(Xu et al., 2022): （3） （4） （5） （6） Where represents the weighting factor of the th variable in the th group, indicates the number of groups, and denotes the pairwise quantity of target variables in the th group. The is used to test whether the weighted response ratio of the target variable to the invasive plant treatment is significant. If overlaps with 0, it indicates that the response ratio of the variable is not significant ( P >0.05); otherwise, it indicates that the response ratio of the variable is significant ( P <0.05). To facilitate better understanding, we log-transformed the weighted response ratio and its corresponding confidence interval limits and expressed them as percentage change (). The transformation formula is as follows: （7） In this meta-analysis, the database was constructed and managed using Microsoft Excel 2016. The meta-analysis itself was performed with MetaWin 2.1 and Comprehensive Meta-Analysis 3.0, while figures were generated using ArcMap 10.7 and GraphPad Prism 9. 3. Results 3.1 Overall effects of alien plant invasion on plant diversity Alien plant invasions significantly affect plant diversity in the invaded habitats and are further influenced by external factors during this ecological process. According to Figures 2 and 3, under natural conditions, 77.08% of the plots exhibited negative effects, with plant diversity decreasing by 22.57% ( P <0.05). Under disturbed conditions, when invaded plots were used as the control group, 58.32% of the plots showed positive effects, and plant diversity increased by 13.71% ( P <0.05). However, when non-invaded plots were used as the control group, 68.42% of the plots showed negative effects, with plant diversity decreasing by 11.07% ( P <0.05). Moreover, there were significant differences among groups, with a Qb value of 122.01 ( P <0.01). Figure 2. Frequency distribution of lnR for plant diversity. Note: (non-invaded) indicates the group compared with non-invaded plots as the control. (invaded) indicates the group compared with invaded plots as the control. The meanings of (non-invaded) and (invaded) in Figures 3-7 are the same. ¿p#1 Figure 3. Overall percentage change in plant diversity in response to alien plant invasion. Note: dots with error bars denote overall means and 95% confidence intervals (95%CI). Numbers in parentheses indicate sample sizes. Asterisks (*) indicates that invasive alien plants have a significant impact ( P <0.05). The Qb statistic represents the chi-square measure of between-group heterogeneity; a larger value indicates greater differences among groups. The meanings of error bars, parentheses, and asterisks (*) in Figures 4-7 are the same. 3.2 Dynamic effects of alien plant invasion on plant diversity Under natural conditions, plant diversity declined significantly across all four invasion stages, and the magnitude of this decline increased with the duration of invasion. However, differences among stages were not statistically significant (Qb = 3.11, P >0.05). By Stage Ⅳ, diversity had decreased markedly by 34.79% ( P <0.05). Under disturbed conditions, the two control-group comparisons produced opposite patterns. When invaded plots were used as the control group, diversity increased significantly during Stages I, Ⅱ, and Ⅲ ( P <0.05), with gains ranging from 11.81% to 24.37%, but declined in Stage Ⅳ without statistical significance. In contrast, when uninvaded plots were used as the control, no significant differences were observed in Stages I and Ⅳ, whereas diversity decreased by 21.76% and 11.32% in Stages Ⅱ and Ⅲ, respectively ( P <0.05). Between these two disturbance subgroups, a significant difference was detected, primarily driven by Stage Ⅳ (Qb = 10.58, P <0.05 and Qb = 53.09, P <0.01), while the first three stages remained relatively stable. Figure 4. Percentage change in plant diversity in response to invasive species across invasion stages. Note: Since only a small number of cases with invasion stage Ⅳ were included in the study (n = 2-4), the effect size estimates are unstable and the 95% confidence intervals are wide. In view of the insufficient sample size and its impact on statistical reliability and graphical readability, stage Ⅳ will not be displayed in subsequent subgroup analysis visualizations. 3. 3 Subgroup analysis of the dynamic effects of alien plant invasion on plant diversity 3.3.1 Subgroup analysis under the natural (non-invaded) condition Combining the subgroup analysis (Figure 5) with the results of the between-group heterogeneity test (Table 2), climate zone and habitat type emerged as the main moderating factors of plant diversity change under natural invasion conditions, with clear differences across invasion stages. Variations among these factors were minimal and nonsignificant during Stage I. In terms of climate zones, the change patterns of NT and ST were opposite in Stages Ⅱ and Ⅲ, and the difference between them was significant ( P <0.05). Regarding habitat types, the decline in diversity was smaller for SL in Stage Ⅱ and for GL and WL in Stage Ⅲ, while SL in Stage Ⅲ showed a particularly sharp decrease of 89.14% ( P <0.05). Diversity index results indicated that the impact of plant invasion on R was greater than on H, although neither effect was significant. From the limited data on plot size, all effects were significantly negative ( P <0.05), with MS showing a relatively larger magnitude of change. Figure 5. Subgroup analysis results under the natural (non-invaded) condition Table 2 Results of between-group heterogeneity tests for variables at the same invasion stage (random-effects model). Qb P-value Qb P-value Qb P-value Invasion stage Ⅰ Diversity index 0.53 0.47 2.66 0.10 5.18 0.02 Climatic zone 1.88 0.17 10.90 < 0.01 0.96 0.33 Plot size 1.88 0.17 3.39 0.18 28.98 < 0.01 Habitat type 1.88 0.17 11.84 0.04 18.25 < 0.01 Disturbance type — — 25.32 < 0.01 19.73 < 0.01 Invasion stage Ⅱ Diversity index 2.64 0.10 14.76 < 0.01 0.46 0.50 Climatic zone 35.15 < 0.01 15.89 < 0.01 15.82 < 0.01 Plot size 12.05 < 0.01 4.75 0.09 1.34 0.25 Habitat type 6.01 0.04 133.74 < 0.01 20.14 < 0.01 Disturbance type — — 95.72 < 0.01 4.11 0.13 Invasion stage Ⅲ Diversity index 2.64 0.10 22.22 < 0.01 — — Climatic zone 153.10 < 0.01 20.55 < 0.01 40.86 < 0.01 Plot size 2.08 0.15 59.80 < 0.01 40.86 < 0.01 Habitat type 41.81 < 0.01 199.85 < 0.01 40.86 < 0.01 Disturbance type — — 52.84 < 0.01 13.33 < 0.01 Note：”—” indicates that the heterogeneity test was not applicable due to no or only one category under this variable. Significances ( P < 0.05) are highlight in bold. 3.3.2 Subgroup analysis under the disturbed (invaded) condition Based on the subgroup analysis (Figure 6) and the results of the between-group heterogeneity test (Table 2), diversity index, climatic zone, habitat type, and disturbance type were identified as the primary moderators under invaded conditions. External disturbances significantly increased plant community R (17.54 % to 50.1 %, P <0.05), whereas their effects on H were minor and not significant. During the invasion process, disturbance exerted a significant positive influence on the diversity of NT and T plants (8.7 % to 38.44 %, P 0.05). Plot size showed a significant reduction in MS only in Stage Ⅲ (−17.91 %, P <0.05), while all other categories increased significantly (10.07 % to 76.63 %, P <0.05), with the largest increase observed in LS. Among the six habitat types, only DS in Stage I and GL in Stages Ⅱ and Ⅲ exhibited significant decreases (−2.78 %, −5.69 %, and −14.84 %, respectively; P <0.05). All other habitat types showed positive effects, and DS in Stage Ⅲ increased markedly by 221.68 % ( P <0.05). Most disturbance measures had a promotive effect, with the exception of HA and FT in Stage Ⅱ (−7.69 % and −3.46 %, respectively) and FT and GR in Stage Ⅲ (−4.36 % and −14.75 %, respectively), which showed negative effects. Figure 6 Subgroup analysis results under the disturbed (invaded) condition 3.3.3 Subgroup analysis under the disturbed ( non-invaded) condition Based on the subgroup analysis (Figure 7) and the results of the between-group heterogeneity test (Table 2), climatic zone, plot size, habitat type, and disturbance type were the primary moderators under non-invaded conditions. Across all three invasion stages, R showed a consistently significant negative effect (−9.48 % to −20.31 %, P 0.05). Climatic zone analysis revealed that differences in diversity change widened over time: T in Stage Ⅱ and both T and ST in Stage Ⅲ exhibited significant negative effects ( P <0.05), whereas NT showed positive effects in Stages Ⅱ and Ⅲ. Regarding plot size, only SS shifted to a significantly positive effect in Stage Ⅲ (21.62 %, P <0.05), while most other categories declined significantly (−12.87 % to −26.9 %, P <0.05). Among habitat types, only DS, RZ, and WL showed relatively good diversity recovery, all displaying positive effects (1.17 % to 39.43 %). Analysis of disturbance type indicated that only FT in Stage I and PR in Stage Ⅲ exhibited positive effects (0.18 % and 8.2 %, respectively; P >0.05). ¿p#1 Figure 7 Subgroup analysis results under the disturbed (non-invaded) condition 4. Discussion 4.1 Analysis of the effects of alien plant invasion on plant diversity In the field of invasion ecology, the long-term effects of alien plant invasions on the plant diversity of recipient ecosystems remain a matter of debate. Our meta-analysis indicates that alien plant invasions generally exert a negative impact on local plant diversity, although the magnitude of this impact varies across contexts. Under natural invasion conditions, plant diversity declined significantly by 22.57%, with negative effects observed in 77.08% of the cases (Figures 2 and 3). Another related study reported an even greater overall decline of 36.97% (Xu et al., 2022). This discrepancy may stem from differences in case-selection criteria: our study emphasized invasion duration and excluded cases lacking information on the time since invasion. Further synthesis revealed a downward trend in plant diversity with increasing invasion time, although this trend was not statistically significant (Figure 4). This result largely aligns with our initial hypothesis. However, some scholars have suggested that observed declines in diversity may primarily reflect early- to mid-stage invasion effects, while long-term impacts may weaken or even disappear over time (Flory et al., 2017; Gruntman et al., 2017; Karatayev et al., 2023; Vellend et al., 2013). Several classic invasion hypotheses, such as the Enemy Release Hypothesis, the Evolution of Increased Competitive Ability Hypothesis, and the Shifting Defense Hypothesis, propose that alien plants often enjoy a competitive advantage during the early stages of invasion. Yet this advantage may diminish as time progresses, as native communities adapt through herbivory, parasitism, and disease, thereby reducing the invaders’ influence (Diez et al., 2010; Iacarella et al., 2015; Liu et al., 2022). Nevertheless, in many situations the ecological effects of alien plant invasions remain profound and persistent. Short-term impacts typically involve direct resource competition, space occupation, and the initial displacement of native species, whereas long-term effects may drive deeper alterations in ecosystem structure, such as changes in soil properties, nutrient cycling, and the complete replacement of native plant assemblages (He et al., 2025; Richardson & Gaertner, 2013; Zhang et al., 2020; Zhou & Staver, 2019). A well-documented example is Melaleuca quinquenervia , whose litter is gradually transforming the soils and topography of Florida wetlands, shifting them toward upland habitats (Gordon, 1998; Strayer et al., 2006). This case illustrates that the long-term consequences of alien plant invasions can extend beyond the community level to reshape entire ecosystems and landscapes. Such alterations often further facilitate the sustained expansion of invasive species, thereby continually constricting the ecological niches of native plants and reducing overall plant diversity. To more accurately assess the impact of alien plant invasions on plant diversity under disturbed conditions, this meta-analysis divided invasion data from disturbed sites into two comparative categories: one using naturally uninvaded plots as the control and the other using invaded plots as the control. Our analysis revealed that, relative to invaded sites, disturbance significantly increased plant diversity. However, compared with naturally uninvaded plots, plant diversity still showed a significant decline (Figure 3). This indicates that external disturbance can promote the recovery of plant diversity to some extent, but full restoration to natural levels remains difficult, and community recovery requires a longer timeframe (Guido & Pillar, 2017). When invaded sites are subjected to external disturbances, changes in resource availability and ecological conditions may create niches more favorable to native plants, enhancing their resilience and constraining invasive species (Khattak et al., 2024; Zhao et al., 2025). For example, certain native plants (e.g., Raphanus raphanistrum and Setaria faberi ) have developed tolerance to multiple herbicides, including glyphosate, through extensive genetic variation, allowing them to maintain a competitive advantage in human-disturbed environments (Khattak et al., 2024; Norsworthy et al., 2012). Other researchers argue that in the presence of external disturbances, invasive species may gain a greater competitive advantage over native communities and better adapt to disturbed landscapes (Urziceanu et al., 2024). Together, these findings highlight the inherent uncertainty of external disturbance effects on plant diversity in invaded sites. In this meta-analysis, 85 % of the disturbance type were active human interventions, which is one of the primary reasons why plant diversity increased significantly compared with invaded plots. From the perspective of invasion duration, only Stage Ⅳ showed substantial differences, leading to significant among-group variation in plant diversity under disturbed conditions. The main cause of this result was the influence of a small sample size. For instance, among the Stage Ⅳ samples in disturbed (invaded) conditions, three datasets involved long-term logging in forests and another involved grassland grazing, both of which often exert negative impacts on plant diversity (Brown & Gurevitch, 2004; Porensky et al., 2020). From the perspective of diversity indices, the richness index primarily reflects the number of species within a community, whereas the Shannon-Wiener index integrates both species richness and the evenness of their distribution (Shannon, 1948; Zhu et al., 2021). Results from this meta-analysis indicate that invasive alien plants exert a stronger influence on the species number of invaded communities than on their evenness. Under conditions of natural invasion, the difference between these two effects is not significant; however, when external disturbances are present, the disparity becomes amplified and reaches a significant level. This finding is consistent with previous research: plant invasions often impose greater pressure on a few dominant or rare species, thereby reducing community species richness while exerting relatively limited effects on species evenness (Beaury et al., 2023; Mills et al., 2009). In the context of external disturbance, the impact on community structure becomes more pronounced, allowing some native species to gain living space and, to some extent, increasing species richness, though potentially reducing the evenness of species distribution (Valone & Weyers, 2019). In summary, plant invasion and external disturbance are two major drivers that significantly affect plant diversity. Invasive alien plants typically cause sustained and substantial declines in plant diversity, with particularly pronounced effects on community species numbers. Although external disturbances may, under certain conditions, partially facilitate the recovery of plant diversity in invaded areas, the overall negative impact of invasive alien plants on plant diversity remains dominant, while the moderating effect of invasion duration is relatively weak. ¿p#1 4.2 Analysis of external factors influencing the effects of alien plant invasion on plant diversity The impact of alien plant invasions on plant diversity is not a single process; rather, it is regulated and modified by multiple external factors. This meta-analysis shows that climatic zone, habitat type, and disturbance type are the principal moderators influencing how alien plant invasions affect diversity. Among these, climate, recognized as a key abiotic driver of invasion processes, has been widely acknowledged (Bradley et al., 2010; Chen et al., 2024; Ren et al., 2024; Thuiller et al., 2005; Zhou et al., 2019). Previous studies have likewise demonstrated strong correlations between climatic variables and all major indicators of alien plant invasion (Gonzalez-Moreno et al., 2014). Our findings further support the central role of climate in shaping invasion effects. Across climatic zones, we found that changes in plant diversity were smallest in the southern temperate zone, and this difference became more pronounced over time. Two factors may account for this pattern. First, southern temperate communities are inherently adapted to disturbances such as fire and grazing. When exposed to invasion or other disturbances, native plants may exhibit a degree of resilience, thereby reducing net changes in diversity (Foxcroft et al., 2010; Kitzberger et al., 2016). Second, relatively low population density and limited economic activity in southern temperate regions may prolong the lag phase of alien plant invasions, resulting in more gradual impacts over time (Robeck et al., 2024). By contrast, tropical regions exhibited the greatest changes in plant diversity, a finding consistent with previous research. The tropics, characterized by abundant light, heat, and water, harbor the highest global species richness and ecological niche saturation, so community structural shifts tend to be more dramatic (Pfadenhauer & Bradley, 2024; Solé et al., 2002; Waddell et al., 2020). The magnitude of diversity change in the northern temperate zone fell between these two extremes. This aligns well with its climatic characteristics: the northern temperate zone experiences pronounced seasonal variability, with environmental fluctuations intermediate between those of the tropics and the southern temperate zone, leading to a moderate level of invasion impact (Kreyling, 2010). Moreover, many major invasive plants originate from temperate or subtropical regions, and their life-history traits match northern temperate climates closely. As a result, they can establish populations successfully, but their impact on native diversity is generally less intense than the ”niche saturation–disturbance release” dynamic typical of the tropics (Bradley et al., 2012; Thuiller et al., 2005). Numerous studies have demonstrated that environmental context plays a critical role in the ecological processes of alien plant invasions (Catford et al., 2019; Cui et al., 2023; Ivison et al., 2024; Zeil-Rolfe et al., 2024). However, other research has found that, at broad spatial scales, invasion impacts depend more on the compositional traits of the invading species than on environmental factors (Beaury et al., 2023). Our integrative analysis shows that grasslands and shrublands are more sensitive to alien plant invasions, and this effect is significantly amplified under external disturbances. By contrast, woodlands and riparian zones exhibit stronger resistance to the combined effects of invasion and disturbance, in some cases even maintaining or enhancing community diversity. Diversity in dunes and wetlands remains relatively stable, suggesting that their community structures possess a certain degree of ecological resilience to invasion, likely because these habitats experience less human disturbance (Catford et al., 2019). Overall, current evidence supports the view that environmental context regulates, to some extent, the ecological consequences of alien plant invasions. Such differential responses are likely linked to factors such as vegetation structure, resource availability, and disturbance frequency. Grasslands and shrublands typically offer higher light levels and nutrient availability and are frequently disturbed by human activities, creating favorable conditions for the establishment and expansion of invasive species (Davis et al., 2000). For example, tropical savannas are highly susceptible to exogenous disturbances such as human activities, while invasive plant incursions and disturbances like wildfires can paradoxically facilitate vegetation recovery and help reestablish community structure (Buisson et al., 2019; Seastedt & Pyšek, 2011). Nevertheless, endemic species may struggle to reproduce and recover during this process, leading to greater fluctuations in species richness than in evenness. In comparison, the more closed canopies and resource-limited conditions of woodlands may impose environmental filters that reduce the impact of some invaders on community structure (Martin et al., 2009). Occasional increases in diversity observed in riparian zones following invasion may be related to their high dynamism and rapid species turnover, which confer a degree of recovery potential (Katz et al., 2005). Thus, environmental context does not operate in isolation; rather, it interacts with the traits of invading species (e.g., resource requirements, life-history strategies) and the disturbance regimes to jointly shape the ecological outcomes of invasion (Foster et al., 2022). These findings partly reconcile the macro-scale debate between ”environmental control” and ”species composition control,” suggesting that the relative importance of the two varies across spatial and ecological contexts. Despite the fact that climate and habitat largely determine the ecological consequences of plant invasions, their effects are often amplified or attenuated by the presence of external disturbances. As one of the most pervasive ecological processes under global change, disturbance regimes are regarded as key variables regulating the impacts of invasion (Jo et al., 2024; Marcolin et al., 2024). The results of this study likewise indicate that the ecological effects of the variables examined were intensified under disturbed conditions. Various types of disturbance mentioned in this work generally facilitated the recovery of plant diversity in invaded habitats, primarily because active human interventions accounted for the largest proportion. Within the disturbance categories of this meta-analysis, removal of invasive plants yielded the greatest improvement in the recovery of plant diversity, followed by combined or multiple disturbance measures, whereas herbicide application, grazing, and fire treatments were comparatively less effective. Removing invasive plants can directly reduce competitive pressure and restore resource availability for native species, thereby facilitating a marked recovery of community diversity (Kettenring & Adams, 2011). Some researchers, however, argue that the sole removal of invasive species is not ideal because the native seed bank in invaded sites is often severely depleted; thus, supplementing native seeds promptly after removal is recommended (Halassy et al., 2023; Orrock et al., 2023). By contrast, herbicide treatments, while capable of effectively suppressing invasive species in the short term, often have unintended negative effects on non-target species, limiting their restoration outcomes (Appah et al., 2020; Baucom et al., 2025). Moreover, long-term herbicide application can significantly disrupt soil microbial communities, which may also explain why plant diversity in disturbed (invaded) sites showed a marked negative effect during Phase Ⅱ (Wen et al., 2025). Grazing and fire treatments frequently create resource fluctuations and expand bare ground, yet their influence on invasion remains uncertain because the ultimate outcomes depend on the competitive balance between invasive and native species in exploiting these ecological opportunities (Beck et al., 2015; Cao et al., 2024). Importantly, the ecological consequences of different disturbance regimes do not occur in isolation; their effects often hinge on the surrounding environmental context and climatic conditions. For example, fire disturbance may exacerbate invasion in arid grasslands but promote the dominance of native species in more mesic habitats (Damasceno & Fidelis, 2020; Klinger & Brooks, 2017). Similarly, the impacts of grazing differ markedly between resource-rich temperate grasslands and resource-limited semi-arid ecosystems (Cipriotti et al., 2019). These findings underscore that disturbance type interact strongly with climate and habitat conditions, jointly determining both the direction and the magnitude of invasion impacts. Meanwhile, the results of field surveys on species diversity may also be affected by the size of the sampling plots. Based on the available data, the present meta-analysis indicates that, under the influence of invasion and external disturbance, there is no consistent pattern between plot scale and changes in plant diversity. In certain stages of invasion, however, there is a significant trend in which larger plot sizes are associated with greater fluctuations in diversity. The species-area relationship suggests that species richness increases with ecosystem area (Veresoglou et al., 2016). Because alien plant invasions and external disturbances often pose serious threats to a small number of species and to rare species, invaded sites with greater area and higher species richness tend to exhibit larger changes in diversity when disturbed. This finding is consistent with Solé et al. (2002), who argued that ecosystem complexity places systems near the edge of instability, such that even minor external perturbations can trigger dramatic shifts in state. Some scholars contend that biodiversity can enhance community stability, and that this stabilizing effect tends to strengthen moderately with increasing spatial scale (Liang et al., 2022; Liang et al., 2025). Others, however, point out that beyond the explanation of plot-scale differences, observed diversity outcomes also depend on researchers’ specific choices regarding study systems and experimental design (Tomasetto et al., 2019). Overall, the evidence shows that the effect of plot scale on detecting the impacts of invasion remains inconsistent. 4.3 Limitations and uncertainties Previous studies have extensively investigated the effects of alien plant invasions on plant diversity(Beck et al., 2015; Brown & Gurevitch, 2004; Chikowore et al., 2025; De Abreua & Durigan, 2011; Liu et al., 2025). Building on this body of work, the present study conducted a comprehensive meta-analysis that integrates published findings on how plant diversity responds to alien plant invasions across different invasion stages. We further examined how plant diversity changes in the presence of external disturbances, comparing results across different control groups. In this meta-analysis, 45.57% of the studies were conducted in North America and 72.15% in the Northern Hemisphere, resulting in uneven spatial coverage and introducing uncertainty into the synthesis. Moreover, studies focusing on late-stage invasions (Stage Ⅳ: 31-100 years) were extremely scarce, limiting our ability to evaluate plant diversity under long-term invasion. Consequently, most of the available evidence reflects changes in plant diversity under short- to medium-term invasion. In addition, disturbance events reported in the analyzed literature lacked precise records of invasion duration, constraining our ability to interpret the combined effects of disturbance intensity, frequency, and invasion stage. Therefore, future research urgently needs to expand data coverage and observational depth across both spatial and temporal scales to enhance the representativeness and generalizability of conclusions. Furthermore, the scope of diversity metrics should be broadened. Current research has focused primarily on α-diversity, whereas β- and γ-diversity, as well as functional and phylogenetic diversity, remain insufficiently synthesized in the context of plant invasions. Such efforts will not only address current gaps in spatiotemporal coverage and diversity dimensions but will also provide critical insights into the ecological mechanisms of alien plant invasions and improve predictions of their future risks. Ultimately, this work will offer a stronger scientific foundation for global biodiversity conservation and ecosystem management strategies. 5. Conclusions This meta-analysis quantitatively evaluated the dynamic impacts of invasive alien plants on plant diversity and examined the moderating roles of climate zone, plot size, habitat type, and disturbance type. The main conclusions are as follows: (1) Invasive plant incursions significantly reduce local plant diversity, showing a gradual decline with increasing invasion duration. (2) The effect of invasion duration on changes in plant diversity is relatively limited. (3) External disturbances can promote partial recovery of diversity but are insufficient to restore it to pre-invasion levels. (4) Climate zone, habitat type, and disturbance type are key variables regulating invasion effects, and their influence is amplified under disturbance conditions. These findings provide a quantitative basis for assessing the ecological risks of plant invasions worldwide and offer scientific guidance for developing region-specific management and ecological restoration strategies. Funding information This work was supported by the State Key Research Development Program of China (2024YFF1307500), the National Natural Science Foundation of China (32271587, 32401311), the Natural Science Foundation of the Jiangsu Higher Education Institutions of China (23KJB180003), Carbon Peak and Carbon Neutrality Technology Innovation Foundation of Jiangsu Province (BK20220030), Graduate Research and Practice Innovation Program of Jiangsu Province (KYCX25_4258, SJCX25_2472), Priority Academic Program Development of Jiangsu Higher Education Institutions (PAPD), Jiangsu Collaborative Innovation Center of Technology and Material of Water Treatment and the Special Scientific Research Project of the School of Emergency Management, Jiangsu University(KY-A-07). Conflicts of interest The authors declare no conflict of interest. References Appah, S., Jia, W., Ou, M., Wang, P., & Asante, E. A. (2020). 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Keywords alien plant invasion dynamic impact invasion stage plant diversity temporal scale Authors Affiliations Zhaoqi Zhu 0009-0000-1364-7485 Jiangsu University View all articles by this author Li Fan Nanjing Normal University View all articles by this author Xue Fan Jiangsu University View all articles by this author Haochen Yu Nanjing Normal University View all articles by this author Miaomiao Cui 0000-0002-6578-9298 Nanjing Institute of Technology View all articles by this author Yijian Wang Jiangsu University View all articles by this author Shoujiang Liu China West Normal University School of Educational Science View all articles by this author Zhi-Cong Dai 0000-0002-0748-8059 Jiangsu University View all articles by this author GuangQian Ren Jiangsu University View all articles by this author Daolin Du [email protected] Jiangsu University Jingjiang College View all articles by this author Metrics & Citations Metrics Article Usage 293 views 194 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Zhaoqi Zhu, Li Fan, Xue Fan, et al. 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