DOI: 10.5061/dryad.hhmgqnkq5
title: Plant invasions alter soil biota and microbial activities: A global meta-analysis

description: Plant invasion is a major component of global environmental change that
 can significantly alter soil biota, which are essential for organic matter
 decomposition and nutrient cycling in terrestrial ecosystems. However, the
 particular linkage between invasive plants, soil biotas, and enzymatic
 activities remains unclear. Here, we conducted a comprehensive
 meta-analysis using 688 paired observations from 107 studies to evaluate
 to the impact of plant invasion on soil biota, enzymatic activities and
 nutrient cycling. Our results showed that plant invasion significantly
 reduced herbivores (45%), detritivores (27%) and omnivores (45%)
 abundance, while allelopathy, woody invaders and forest ecosystems
 significantly reduced predator abundance by 47, 46, and 32%, respectively.
 Invasive plants also slightly reduced bacterial and fungal biomass, but
 significantly increased arbuscular mycorrhizal fungal (AMF) biomass, MBC
 and MBN by 29, 19 and 32%, respectively. Moreover, invasive plants had
 inconsistent impact on the activities of C-decomposing enzymes, but
 invertase, phenol oxidase and β-glucosidase were found significantly
 higher in invaded than uninvaded sites.  Invaded sites had
 significantly higher activities of N- and P-releasing soil enzymes ranged
 from 18 to 27% than uninvaded sites. Soil microbial biomass,
 N-mineralization, soil respiration, available (N, P) nutrients, NH4+-N and
 nutrient stocks were all higher in invasive than native plants rhizosphere
 soils. Our findings suggest that invasive plants had negative impact on
 soil functional groups, however, they enhanced soil nutrient-releasing
 enzymes and soil available nutrients. This in turn accelerates nutrient
 cycling and promote the persistence and success of invasive plants.
description: We selected peer-reviewed journal articles using Web of Science
 and Google Scholars and screened the identified records using Preferred
 Reporting Items for Systematic Review and Meta-analysis procedure
 (PRISMA). Primary studies that satisfied the following selection criteria
 were included: (1) studies specifically described the identities of
 invasive and native plants and investigated either in the field
 observational or experimental studies, (2) at least one of the soil
 functional groups, or microbial composition or enzymatic activity was
 measured simultaneously in the invasive and native plants using the PLFA
 method (Kong et al., 2011), (3) invaded sites and control or native plants
 were found in the same biotic and abiotic conditions, (4) the means and
 sample sizes of the selected variables were available or could be
 calculated from the related publications. In our meta-analysis, we only
 used the soil microbial community composition and structure, soil
 microbial biomass and their enzymatic activities measured with PLFAs, the
 most used method for microbial measurements (G. Zhang et al., 2019). We
 applied the following criteria to identify independent case studies:  (1)
 If multiple independent pairs of invasive and native plants from different
 ecosystems were reported in the same article, we treated each pair as a
 separate case study that complied with other meta-analyses' criteria
 and (2) if the study article reported one native plant and two more
 invasive plants, each invasive-native combination was considered as a
 separate case study. The data presented in the form of figures were
 extracted by using webplotdigitizer 4 https://automeris.io/WebPlotDigitizer.      For each pair of response variables in invasive (i) and native (n) plants, the log response ratio (LRR) was calculated based on mean values (X) as a measurement of effect size (Bakbergenuly et al., 2020; Lajeunesse, 2011). The LRR is a unit-free index that estimates the size of the impact and its directions. The values of LRR = 0, showed no difference in the measured response variable between invasive and native plants, a negative value indicates that the invaded site has a lower value than the noninvaded sites and a positive value presents that the invaded site has a greater value than the noninvaded site (Luo et al., 2006). The effect of plant invasion was estimated (Eq. 1) using the succeeding natural LRR (Bakbergenuly et al., 2020).  ---------------------------------------------------- (1) Where  and <sub>, </sub>are the results of the means in the treatment and control group, respectively. We calculated the variance (<em>VR</em>) that related with the effects computed by Eq. 2: - -------------------------------------------------------------------------- (2) Where S<sub>t</sub> and S<sub>c</sub> are the standard deviations of the experimental and control treatments, respectively, where as <sub> </sub>and  are the data points of the experimental and control treatments, respectively. The standard error of the log response ratio calculated (Eq. 3) as follows: - SE (lnR)=  ----------------------------------------------------------------------------- (3) We used Eq. 4, the weighted mean response ratio (R++) to identify the overall effects of the experimental versus the control group treatments, respectively (Hedges et al., 1999).   ---------------------------------------------------------------------- (4) Where  is the weighting factor,  and  are th data points in the th dataset in each category group, ,  and  are the number of datasets and data points in each dataset, respectively, in the category groups. The SE of R<sub>++</sub> was calculated by Eq. 5: - ------------------------------------------------------------------ (5)
description: # Plant invasions alter soil biota and microbial activities: A global
 meta-analysis [Data for plant invasion effects on soil biota functional
 groups and microbial activities research
 project. ](https://doi.org/10.5061/dryad.hhmgqnkq5)The dataset includes
 authors name and year, invasive species, native plant species, growth
 forms, allelopathy and ecosystem types. In addition, response variables
 for plant invasion with control and treated data with mean, standard
 deviation and number of samples, geographic coordinates, continent and
 country included. Meta-analysis was conducted on soil biota functional
 groups, microbial activities and nutrient cycling to assess the
 effect-sizes of plant invasion on the response variables.  Contact: Zebene
 Negesse ([zedalem12@gmail.com](mailto:hahnp@ufl.edu)) with any questions.
 This manuscript has been published (14 Feb 2025):  Zebene Negesse, Kaiwen
 Pan, Awoke Guadie, Meta Francis Justine, Belayneh Azene, Bikram Pandey,
 Xiaogang Wu, Xiaoming Sun, Lin Zhang. Plant invasions alter soil biota and
 microbial activities: A global meta-analysis. Plant and Soil (2025).
  Datasets and associated with this data files can be found here:
 [http://doi](http://doi):10.5061/dryad.hhmgqnkq5.   Dataset This file
 contains data on invasive and native plants. All data were collected by
 the authors. Blank cells represent missing data. | Variable | Unit |
 Description | | :------------------------------ |
 :---------------------------------------- |
 :-------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------- | | Continent | Categorical | representing different continents or global regions (e.g., Europe, Latin America, Africa, Asia, North America, Oceania). | | Study site | Categorical | Location of the study, see detail in Table S1 | | Ecosystem type | Categorical | Forest and non-forest ecosystems invaded by invasive plants | | Allelopathic | Categorical | Plants with allelopathy = Yes and Without allelopathy = No, see in detail which invasive plants have allelopathy, and which is not. | | Growth form | Categorical | Woody = (Trees, shrubs); and non-woody = (herbs, grasses)  | | Land use types | Categorical | Representing different ways land is utilized by humans or nature, such as agriculture, forests, urban areas, and wetlands. | | Country | Categorical | Representing different nations where the study is conducted | | ln (LRR) | Dimensionless | Natural logarithm (Ln) of the response ratio (RR). | | In (V) | Dimensionless | Natural logarithm (Ln) of the variance | | Microbial biomass | µg biomass C/g soil | The total mass of living microorganisms (bacteria, fungi, archaea) in a given amount of soil. | | Bacteria biomass | CFU/g | Single-celled microorganisms found in various environments, including soil, water, air, and living organisms | | Fungal biomass | µg C/g soil | The total mass of living fungi (including mycorrhizal fungi and decomposers) in a given amount of soil. | | AMF biomass | µg C/g soil | The total biomass of Arbuscular Mycorrhizal Fungi (AMF) in soil | | Soil respiration | g CO₂/m²/day | The process by which soil microorganisms (bacteria, fungi, and other soil organisms) and plant roots break down organic material, releasing carbon dioxide (CO₂) into the atmosphere | | Acid phosphatase | µmol p-NP/g soil/hr | An enzyme produced by soil microorganisms, plants, and fungi that catalyzes the hydrolysis of phosphate esters, releasing inorganic phosphate | | Alkaline phosphatase | µmol p-NP/g soil/hr | An enzyme that catalyzes the hydrolysis of organic phosphate esters, releasing inorganic phosphate | | Urease | µmol NH₃/g soil/hr | an enzyme that catalyzes the hydrolysis of urea into ammonia (NH₃) and carbon dioxide (CO₂) | | Peroxidase | µmol phenol/g soil/hr | an enzyme involved in the breakdown of hydrogen peroxide (H₂O₂) and the oxidation of a variety of organic substrates in soil | | Phenol oxidase | µmol guaiacol/g soil/hr | an enzyme that catalyzes the oxidation of phenolic compounds to quinones and involved in the decomposition of organic matter | | N-nitrification | µmol NO₃⁻/g soil/hr | Nitrogen nitrification: Ammonia (NH₃) is converted to nitrite (NO₂⁻) through the action of ammonia-oxidizing bacteria (AOB) or ammonia-oxidizing archaea (AOA) | | N-mineralization | µmol NH₄⁺/g soil/hr | Nitrogen mineralization: microbial process by which organic nitrogen (found in plant residues, animal remains, and other organic matter) is converted into inorganic forms such as ammonium (NH₄⁺), which is available for plant uptake | | β-glucosidase | µmol glucose/g soil/hr | an enzyme produced by soil microorganisms (bacteria, fungi) that plays a key role in the breakdown of complex organic matter in soils, especially in cellulose and lignocellulosic compounds | | Abundance of soil invertebrates | mg/m² | organisms, which include earthworms, nematodes, arthropods (e.g., beetles, ants), mollusks (e.g., snails), and microarthropods (e.g., mites) | | Actinomycete biomass | µg/g | the total mass of actinomycetes present in the soil | | MBC | mg C/kg soil | Microbial biomass carbon (MC): - the total amount of carbon contained in the living microbial biomass (bacteria, fungi, actinomycetes) within the soil | | MBN | mg N/kg soil | Microbial biomass nitrogen (MBN): - the total amount of nitrogen contained in the living microbial biomass (bacteria, fungi, actinomycetes) within the soil | | Cellulase | µmol glucose/g soil/hr | An enzyme that breaks down cellulose, the primary structural component of plant cell walls, into simpler sugars like glucose | | Catalase | µmol O₂/g soil/min | An enzyme that breaks down hydrogen peroxide (H₂O₂) into water (H₂O) and oxygen (O₂) | | Invertase | µmol glucose/g soil/hr | An enzyme that catalyzes the hydrolysis of sucrose into glucose and fructose | | Acetyl Glucosamidase | µmol NAG/g soil/hr | An enzyme involved in the decomposition of chitin, a major component of fungal cell walls, insect exoskeletons, and some microbial structures | | Polyphenol oxidase | µmol quinone/g soil/hr | An enzyme that catalyzes the oxidation of phenolic compounds into quinones, which subsequently polymerize into humic substances | | Glycine amino peptidase | µmol glycine/g soil/hr | An enzyme involved in protein degradation and nitrogen cycling in soil | | Abundance of Herbivores |  g/m² | The population density or biomass of herbivorous soil organisms, including invertebrates such as nematodes, collembolans, mites, and insect larvae that feed on plant roots, fungi, or microbial biomass | | Abundance of Omnivores | g/m² | The population density or biomass of omnivorous soil organisms, which feed on a mix of plant material, microbes, fungi, and other small invertebrates | | Abundance of predators | g/m² | The number or biomass of organisms that primarily feed on other organisms (e.g., herbivores, omnivores, or other predators) in an ecosystem | | Abundance of Decomposers | g/m² | The number or biomass of organisms that break down organic matter in the soil, recycling nutrients and playing a key role in nutrient cycling | | Protease | µmol amino acid/g soil/hr | Enzymes that catalyze the breakdown of proteins into smaller peptides and amino acids | | abundance of microbivores | g/m² | soil organisms that feed primarily on microorganisms, including bacteria, fungi, and algae | | SD\_Control | SI unit of the measured variable | Standard deviation of the mean in the control groups | | N\_Control | Dimensionless | Number of sample size (N) in the control groups | | SD\_Treatment | SI unit of the measured variable | Standard deviation of the mean in the treatment groups | | N\_Treatment | Dimensionless | Number of sample size (N) in the treatment groups | | Native plant species | Categorical | Plant species native to the region and listed by their scientific names | | Soil response variables | Categorical | Soil response variables = soil parameters used in the effect size calculations  | | Mean\_Treatment | SI unit of each soil response variable | The mean value in the treatment groups | | Mean\_Control | SI unit of each soil response variable | The mean value in the control groups | | Geographic coordinates | degrees (°), minutes ('), and seconds (") | Shows the precise location of the studies on Earth's surface using latitude (Northing, Southing) and longitude (Easting, Westing). | | Invader plant species | Categorical | Plant species non-native to the area, invading and displacing the native plant communities | | Land use type | Categorical | Describe how land is utilized by humans and natural processes | | Forest vs non-forest | Categorical | Ecosystem types invaded by exotic plants are classified as forest (temperate, tropical, plantation) or non-forest (grasslands, agricultural lands) etc | | | | | | Woody vs non-woody | Categorical | Plant growth forms are categorized as woody plants (trees, shrubs) or non-woody plants (grasses, herbs). | | ES | Dimensionless | ES = represents effect sizes | | Northing | degrees (°), minutes ('), and seconds (") | Shows the location of the study in the north | | Easting | degrees (°), minutes ('), and seconds (") | Shows the location of the study in the east | | Case study | Discrete variable | It represents the number of studies within one article (Example: analyzing one invasive plant with two native plants results in two case studies because each native plant interacts separately with the invasive species)  | ## Description of the data and file structure The datasets structured by extracting from peer-reviewed journal articles as follows: We conducted a comprehensive search for peer-reviewed journal articles that assessed how soil functional groups, microbial compositions and enzymatic activities respond to plant invasion in terrestrial ecosystems using Web of Science and Google Scholars. For each pair of response variables in invasive (i) and native (n) plants, the log response ratio (LRR) was calculated based on mean values (X) as a measurement of effect size. lnV= natural logarism of variance. ln (RR) = ln (Xt/Xc), Where Xt and Xc , are the results of the means in the treatment and control group, respectively. We calculated the variance (*VR*) that related with the effects computed by: ln (V)= (Sc^2)/(nc*Xc^2)+(St^2)/(nt*Xt^2), where St and Sc are the standard deviations of the experimental and control treatments, respectively, where as nt and nc are the data points of the experimental and control treatments, respectively. ## Sharing/Access information Data was derived from the following sources: *Aguilera, Anna , Peter Alpert, Jeffrey Dukes, and Robin Harrington. 2009. 'Impacts of the invasive plant Fallopia japonica (Houtt.) on plant communities and ecosystem processes', Biological Invasions, 12: 1243-52. Allison, Steven D., Caroline Nielsen, and R. Flint Hughes. 2006. 'Elevated enzyme activities in soils under the invasive nitrogen-fixing tree Falcataria moluccana', Soil Biology and Biochemistry, 38: 1537-44. Anthony, M. A., S. D. Frey, and K. A. Stinson. 2017. 'Fungal community homogenization, shift in dominant trophic guild, and appearance of novel taxa with biotic invasion', Ecosphere, 8: 1-17. Aragón, Roxana, Jordi Sardans, and Josep Peñuelas. 2014. 'Soil enzymes associated with carbon and nitrogen cycling in invaded and native secondary forests of northwestern Argentina', Plant and Soil, 384: 169-83. Arthur, Mary A., Sarah R. Bray, Christina R. Kuchle, and Ryan W. McEwan. 2012. 'The influence of the invasive shrub, Lonicera maackii, on leaf decomposition and microbial community dynamics', Plant Ecology, 213: 1571-82. Ballard, M., J. Hough-Goldstein, and D. Tallamy. 2013. 'Arthropod communities on native and nonnative early successional plants', Environ Entomol, 42: 851-9. 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'Invasive plants decrease microbial capacity to nitrify and denitrify compared to native California grassland communities', Biological Invasions, 19: 2941-57. Castro-Díez, Pilar, Natalia Fierro-Brunnenmeister, Noelia González-Muñoz, and Antonio Gallardo. 2011. 'Effects of exotic and native tree leaf litter on soil properties of two contrasting sites in the Iberian Peninsula', Plant and Soil, 350: 179-91. Čerevková, Andrea, Kristina Ivashchenko, Dana Miklisová, Nadezhda Ananyeva, and Marek Renčo. 2020. 'Influence of invasion by Sosnowsky’s hogweed on nematode communities and microbial activity in forest and grassland ecosystems', Global Ecology and Conservation, 21. Chacón, Noemi, Ileana Herrera, Saúl Flores, José A. González, and Jafet M. Nassar. 2008. 'Chemical, physical, and biochemical soil properties and plant roots as affected by native and exotic plants in Neotropical arid zones', Biology and Fertility of Soils, 45: 321-28. Chang, Ed-Haun, and Chih-Yu Chiu. 2015. 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'Invasive plants affect arbuscular mycorrhizal fungi abundance and species richness as well as the performance of native plants grown in invaded soils', Biology and Fertility of Soils, 52: 879-93. ## Code/Software All the effect size calculations used Metawin Version 3 software. The forest plots produced using GraphPad Prism software.