CSR strategy shifts under biotic resistance and grazing drive invasion success of Solanum rostratum in northern China
Data files
Dec 17, 2025 version files 7.12 MB
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Best_model_result.csv
2.02 KB
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boot.csv
4.58 KB
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code_CSR.R
25.65 KB
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CSR_strategies.csv
190.62 KB
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cv_dataresult.csv
5.27 KB
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cvdata.csv
188.82 KB
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LMMdata.csv
41.41 KB
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Mean_CSR_strategies.csv
210 B
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model_avg_full_estimates.csv
1.95 KB
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random_factors.csv
4.20 KB
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README.md
15.70 KB
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StrateFy.xlsx
6.40 MB
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trait_data.csv
235.57 KB
Abstract
Biotic resistance and environmental filter are the primary ecological barriers to successful invasions. Although the role of functional strategies in overcoming these barriers has been extensively investigated, whether and how the plasticity of functional strategies facilitates successful invasions remains poorly understood. In this study, we explored the invasion mechanisms of a widespread invasive plant, Solanum rostratum, across a 3,000 km spatial extent in northern China using Grime’s competitor, stress tolerator, and ruderal (CSR) framework. Our results showed that S. rostratum consistently invested more in C-strategy compared to native species, but it exhibited plastic shifts between S- and R-strategies. These strategic shifts were primarily influenced by the CSR strategy and the taxonomic and functional diversity of native communities, which selected for greater strategic divergence between S. rostratum and native species. Grazing further modulated these shifts by reducing native diversity. By increasing its strategic divergence from native competitors, S. rostratum may persist in species-diverse native communities and dominate in frequently disturbed and species-limited communities. Synthesis: Our findings reveal a dynamic strategy-environment matching process across space, underscoring the significance of functional plasticity in overcoming varying ecological barriers during large-scale invasions.
Dataset DOI: 10.5061/dryad.tqjq2bwcf
Description of the data and file structure
We conducted large-scale field surveys and trait measurements across40 invaded and native plant communities spanning a 3,000 km transect in northern China to investigate how the functional strategies of Solanum rostratum vary with ecological contexts. We recorded species composition, environmental conditions, and plant functional traits to calculate CSR strategies for both invasive and native species. These data were collected to assess how functional strategy plasticity facilitates invasion success across environmental and biotic gradients.
Files and variables
1. Best_model_result.csv
Description: Model selection results for candidate linear mixed-effects models.
Variables:
model_id— Model identifier (character)df— Degrees of freedomlogLik— Log-likelihood of the modelAICc— Corrected Akaike Information Criteriondelta— Difference in AICc relative to the best modelweight— AICc weightR2m— Marginal R² (variance explained by fixed effects)R2c— Conditional R² (variance explained by fixed + random effects)variables— Predictors included in the model (character)Moran_I— Moran’s I statistic for spatial autocorrelationExpected_I— Expected Moran’s I under the null modelSD— Standard deviation of Moran’s Ip_value— p-value for Moran’s I test
2. boot.csv
Description: Bootstrap results of CSR strategic dissimilarity indices (SDI).
Variables:
SDI_C— Strategic dissimilarity index for C-strategySDI_S— Strategic dissimilarity index for S-strategySDI_R— Strategic dissimilarity index for R-strategy
Units: Dimensionless indices (0–1)
3. cv_dataresult.csv / cvdata.csv
Description: Trait variability data.
Variables:
species— Species name (character)mean_value— Mean trait valuesd_value— Standard deviation of trait valuecv— Coefficient of variationC,S,R— CSR strategy scores for individual species (0–100%)
4. Mean_CSR_strategies.csv / CSR_strategies.csv
Description: Site-level and species-level CSR strategy scores.
Variables:
C— Competitor strategy score (%)S— Stress-tolerator strategy score (%)R— Ruderal strategy score (%)Group— Species group (e.g., S. rostratum vs. native community)
5. LMMdata.csv
Description: Predictor and response variables used in linear mixed-effects models.
Variables:
C_score_SR,S_score_SR,R_score_SR— CSR scores for S. rostratumC_score_CWM,S_score_CWM,R_score_CWM— Community-weighted mean CSR scores of native communitiesC_score_Dominant,S_score_Dominant,R_score_Dominant— CSR scores of dominant native speciesGrazing— Grazing intensity (categorical: grazed/ungrazed)Aridity— Aridity index (dimensionless)MAT— Mean annual temperature (°C)Species_richness— Number of species per plotSimpson— Simpson diversity indexPD— Faith’s phylogenetic diversity (dimensionless)Plant_density— Plant density (individuals m⁻²)Biomass— Aboveground biomass (g m⁻²)Vegetation_coverage— Vegetation cover (%)MPD_phy,MNTD_phy— Mean pairwise / nearest taxon distance (phylogenetic)NRI,NTI— Net Relatedness Index and Nearest Taxon Index (standardized effect sizes)MPD_trait,MNTD_trait— Mean pairwise / nearest trait distanceFD— Functional diversity indexSDI_C_score,SDI_S_score,SDI_R_score— Strategic dissimilarity indices for C, S, and RIII_SR— Invasion intensity index for S. rostratumLon,Lat— Longitude and latitude of sampling site (decimal degrees)
6. random_factors.csv
Description: Site-level random effects used in LMMs.
Variables:
Site— Site identifier (character)random_effect— Random effect estimateLon,Lat— Site coordinates (decimal degrees)
7. model_avg_full_estimates.csv
Description: Model-averaged parameter estimates and importance values.
Variables:
Variable— Predictor variable nameEstimate— Model-averaged estimateSE— Standard errorAdjusted_SE— Adjusted standard errorz value— z statisticp— p-valueLowerCI,UpperCI— 95% confidence interval boundsSignificance— Significance level (e.g., ‘*’, ‘**’)Importance— Variable importance weightFrequency— Frequency of occurrence in candidate modelsGroup— Variable group/category
8. StrateFy.xlsx
Description: StrateFy.xlsx is an external analytical tool originally published by Pierce et al. (2017, Functional ecology) for calculating CSR (Competitor–Stress tolerator–Ruderal) strategy scores from plant leaf traits.
This file is included in the repository solely to ensure full reproducibility of the CSR calculations performed in our study.
Important:
This Excel file is not raw data from our research.
All raw trait data used in our analyses are provided separately in unformatted CSV files.
Variables: Species ID, trait values (e.g., SLA, LDMC, LA), and site information.
Units: Trait-specific (e.g., LA in mm², LDMC in %, SLA in mm2 mg-1)).
Reference:
Pierce, S., Negreiros, D., Cerabolini, B. E. L., Kattge, J., Díaz, S., Kleyer, M., Shipley, B., Wright, S. J., Soudzilovskaia, N. A., Onipchenko, V. G., van Bodegom, P. M., Frenette-Dussault, C., Weiher, E., Pinho, B. X., Cornelissen, J. H. C., Grime, J. P., Thompson, K., Hunt, R., Wilson, P. J., … Tampucci, D. (2017). A global method for calculating plant CSR ecological strategies applied across biomes world-wide. Functional Ecology, 31(2), 444–457. https://doi.org/10.1111/1365-2435.12722
9. code_CSR.R
Description: R script containing code for trait data processing, CSR strategy calculation, SDI computation, and linear mixed-effects model fitting.
9. trait_data.csv
Description: raw leaf trait input file for matted for StrateFy
| Variable Description: | ||
|---|---|---|
| Column name | Description | Unit |
| Species binomial | Full species scientific name | — |
| Site | Sampling site identifier | — |
| Sample ID | Individual sample identifier | — |
| leaf area (mm²) | Leaf lamina area | mm² |
| leaf dry matter content (%) | LDMC = dry mass / fresh mass × 100 | % |
| specific leaf area (mm² mg⁻¹) | SLA = leaf area per dry mass | mm² mg⁻¹ |
Code/software
All data processing and analyses were conducted in the free and open-source statistical environment R (version 4.3.2). The workflow is fully contained in the provided R script code_CSR.R. This script allows users to reproduce the entire analytical pipeline from raw trait data to final statistical models and figures.
Required software and packages
The following R packages were used and must be installed prior to running the script:
StrateFy– calculation of CSR strategy scores from leaf traits.dplyr,tidyr– data cleaning and processing.ggplot2,ggtern– data visualization (e.g., ternary plots, effect plots).lme4– linear mixed-effects modeling.MuMIn– model selection and model averaging (AICc-based).DHARMa– model diagnostics.spdep– spatial autocorrelation testing (Moran’s I).
All required packages are open source and available through CRAN.
Analytical workflow
- CSR calculation – Leaf trait data from
StrateFy.xlsxare processed to calculate C, S, and R scores for each species. - Trait plasticity – Within-species variation is quantified by calculating the coefficient of variation (CV) of CSR scores (
cvdata.csv). - CSR dissimilarity – Functional divergence between invader and native communities is computed as CSR dissimilarity indices (SDI) (
boot.csv). - Modeling and selection – Linear mixed-effects models are fitted using
lme4to assess environmental and community predictors of invasion intensity. AICc-based model selection and averaging are performed usingMuMIn(Best_model_result.csv,model_avg_full_estimates.csv). - Model diagnostics and spatial analysis – Residual diagnostics are performed with
DHARMa, and spatial autocorrelation of model residuals is tested using Moran’s I (spdep). - Visualization – Ternary plots of CSR strategies, CV distributions, and model effect plots are generated with
ggplot2andggtern.
Variable Description Table
| Variable | Definition | Unit / Scale | Notes |
| C_score_SR | C-strategy score of S. rostratum | Unitless (0–1) | Calculated from CSR triangle method |
| S_score_SR | S-strategy score of S. rostratum | Unitless (0–1) | – |
| R_score_SR | R-strategy score of S. rostratum | Unitless (0–1) | – |
| C_score_CWM | Community-weighted mean C-strategy score of native communities | Unitless (0–1) | Weighted by species relative abundance |
| S_score_CWM | Community-weighted mean S-strategy score of native communities | Unitless (0–1) | – |
| R_score_CWM | Community-weighted mean R-strategy score of native communities | Unitless (0–1) | – |
| C_score_Dominant | C-strategy score of dominant native species | Unitless (0–1) | Dominant species identified by relative abundance |
| S_score_Dominant | S-strategy score of dominant native species | Unitless (0–1) | – |
| R_score_Dominant | R-strategy score of dominant native species | Unitless (0–1) | – |
| Grazing | Grazing intensity | Categorical (grazed / ungrazed) | Based on field observation & management records |
| Aridity | Aridity index | Unitless | Higher = drier |
| MAT | Mean annual temperature | °C | – |
| Species_richness | Number of species per plot | Count | – |
| Simpson | Simpson diversity index | Unitless (0–1) | Higher = greater diversity |
| PD | Faith’s phylogenetic diversity | Branch length units | – |
| Plant_density | Plant individual density | individuals·m⁻² | – |
| Biomass | Aboveground biomass | g·m⁻² (dry weight) | – |
| Vegetation_coverage | Vegetation cover | % | – |
| MPD_phy | Mean pairwise phylogenetic distance | Branch length units | – |
| MNTD_phy | Mean nearest-taxon phylogenetic distance | Branch length units | – |
| NRI | Net Relatedness Index | Unitless (SES) | Higher = stronger clustering |
| NTI | Nearest Taxon Index | Unitless (SES) | – |
| MPD_trait | Mean pairwise distance based on trait distance | Trait distance units | Gower distance |
| MNTD_trait | Mean nearest-taxon trait distance | Trait distance units | Gower distance |
| FD | Functional diversity index | Unitless | Higher = more functionally diverse |
| SDI_C_score | Strategic dissimilarity index for C-strategy | Unitless | Dissimilarity between S. rostratum CSR-C and community |
| SDI_S_score | Strategic dissimilarity index for S-strategy | Unitless | – |
| SDI_R_score | Strategic dissimilarity index for R-strategy | Unitless | – |
| III_SR | Invasion intensity index of S. rostratum | Unitless index (0–1) | Higher = stronger invasion |
| Lon | Longitude of sampling plot | Decimal degrees | GPS measured |
| Lat | Latitude of sampling plot | Decimal degrees | GPS measured |
Usage
Users can open and run the script code_CSR.R directly in R. The script loads the data files automatically, executes the above steps sequentially, and reproduces the main analyses and figures reported in the manuscript.
