Interspecific competition negates CO₂ benefits for most plant species
Data files
Feb 16, 2026 version files 469.75 KB
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MetaAnalysis_Data5.csv
98.74 KB
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MetaAnalysis_Data6.csv
103.34 KB
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RCode.R
127.54 KB
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README_DRYAD.Rmd
17.19 KB
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README.md
11.83 KB
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Sum_Of_Mix.csv
70.92 KB
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Traits2.csv
40.18 KB
Abstract
Elevated CO₂ (eCO₂) frequently increases plant growth and primary productivity, and is considered a driver of increased land carbon sinks. This evidence contributes to a widespread but surprisingly untested assumption in global change science and policy, and society at large, that eCO₂ consistently stimulates the growth of plant species within communities. However, species and community responses depend on species’ growth strategies and abiotic and biotic contexts, and among these, the role of competition remains underexplored. We therefore ask whether eCO2 enhances growth broadly across species, or primarily in a dominant few? To address this question of how species interactions are shaped by eCO₂ response and competitive ability, we analysed responses from 97 species in 20 grassland CO₂ enrichment experiments. Herein, we show that interspecific competition markedly reduced eCO₂ stimulation: while most species responded positively to eCO₂ in isolation, over half responded negatively in mixed communities. Despite this, community biomass was often and on average was increased under eCO₂, driven by dominant species contributions. eCO₂ did not enhance competition tolerance, and acquisitive traits—including fast growth and invasion success—conferred a consistent advantage under both CO₂ enrichment and competitive stress. Our findings suggest that in species mixtures, which represent the majority of natural ecosystems, competition causes both positive and negative growth responses to eCO2 to be ubiquitous and pervasive. We show that eCO₂ does not consistently reduce competitive vulnerability, but instead, trait-based filtering in a competitive context determines both which species benefit from eCO2. These results challenge assumptions built into ecosystem models; and incorporating these dynamics will improve forecasts of vegetation response to rising CO₂.
Overview
This repository contains the R code and datasets used to generate Figures 2, 3, and 4 for the manuscript Interspecific competition negates CO₂ benefits for most plant species (Raubenheimer et al., 2026, Nature Plants revision).
All analyses are run from a single master script and reproduce all figures and supplementary tables.
Repository structure
/Data/
MetaAnalysis_Data5.csv
MetaAnalysis_Data6.csv
Sum_Of_Mix.csv
Traits2.csv
/Code/
RCode.R
README_DRYAD.Rmd
README.md
Dataset variable definitions
MetaAnalysis_Data5.csv
Description: Species-level biomass. Cells with 'NA' represent where a value is not applicable (rather than just missing). Note where 'CO2' is abbreviated in the variables; this refers to carbon dioxide.
Variables:
| Column | Description | Units / Notes |
|---|---|---|
| X | Row index | — |
| Species | Scientific name | matches Traits2.csv |
| Species.Status.FunctMetab | Functional type | character |
| Neighbours.FunctMetab | Functional type of neighbours | character |
| Competition.Treatment | Competition treatment | Isolated / Monoculture / Inter-Functional Mixture / Intra-Functional Mixture |
| CO2.Treatment | CO₂ treatment | High_CO2 / Low_CO2 |
| CO2.Treatment.Value | CO₂ ppm | numeric |
| Plant.age..days. | Plant age | days |
| Length.of.CO2.exposure..days. | Duration of CO₂ exposure | days |
| Method.of.CO2.Treatment | CO₂ enrichment method | categorical |
| Paper | Study name | short citation |
| Biome.Category | Biome type | categorical |
| Location | Study location | character |
| Continent | Continent | character |
| Dominance.ratio | Dominance metric | unitless |
| Sum.of.mixture | Total mixture biomass | g |
| Biomass | Mean biomass of focal species | g |
| SE | Standard error | g |
| RCV.RR | Response ratio of RCV | unitless |
| yi_RCV | Effect size for RCV | unitless |
| vi_RCV | Variance of yi_RCV | unitless |
| yi_CO2 | Effect size for CO₂ | unitless |
| vi_CO2 | Variance of yi_CO₂ | unitless |
Variable names that are created in the code:
| Variable | Description | Units | Notes |
|---|---|---|---|
| Method | Experimental method | categorical | (this was Method.of.CO2.Treatment in the dataframe) |
| Duration | Duration of experiment | days | (this was Length.of.CO2.exposure..days. in the dataframe) |
| Comp | Harmonised competition variable | used in modelling | (this was Competition.Treatment in the dataframe) |
| FT | Harmonised functional type | used in modelling | (this was Species.Status.FunctMetab in the dataframe) |
Sum_of_Mix.csv
Description: Community-level biomass. Values are in the row of one species per community (i.e., NAs for the other species in the same community. Species names are not relevant in the code - values are taken according to each study and treatment). Cells with 'NA' represent where a value is not applicable (rather than just missing). Note where 'CO2' is abbreviated in the variables; this refers to carbon dioxide.
Variables:
| Column | Description | Units / Notes |
|---|---|---|
| FT | Functional type | character |
| Neighbours.FT | Neighbour functional type | character |
| Comp | Competition treatment | Mono / Mix |
| CO2.Treatment | CO₂ treatment | High_CO2 / Low_CO2 |
| CO2.Treatment.Value | ppm | numeric |
| Plant.age..days. | Age | days |
| Duration | Duration | days |
| Method | Treatment method | categorical |
| Paper | Study name | character |
| Biome.Category | Biome | character |
| Location | Location | character |
| Continent | Continent | character |
| Sum.of.community | Community biomass | g |
| CO2.Effect | Log response ratio | unitless |
| n | Sample size | — |
| mean_SE | Mean SE | — |
| yi_Mix | Effect size | unitless |
| vi_Mix | Variance | unitless |
MetaAnalysis_Data6.csv
Same structure as MetaAnalysis_Data5, but includes:
Variable names that are created in the code:
| Column | Description |
|---|---|
| No.of.species | Number of species in community |
| X2 | Dominance ratio duplicate column |
| Dominance.ratio, Sum.of.mixture, Biomass, SE | As above |
Traits2.csv
Description: Trait values acquired from the TRY database, AusTraits, and references manuscripts. Unused variables are left in the dataset to avoid issues with column numbers in the accompanying code. Empty cells in the Traits2.csv data represent where trait data was not available for a given species. These empty cells are important for the code that imputes trait data.
Variables:
| Column | Description | Units |
|---|---|---|
| Species | Scientific name | — |
| Paper | Source study | — |
| Neighbours.Combo | Functional group of neighbouring species | categorical |
| Neighbour.Species | Name of neighbouring species | categorical |
| X2 | (unused column) | — |
| X3 | (unused column) | — |
| X4 | (unused column) | — |
| X5 | (unused column) | — |
| X6 | (unused column) | — |
| X7 | (unused column) | — |
| Species.Status | Functional type (e.g., C3 herbaceous) | categorical |
| Species.Status.Broad | Broad functional grouping | categorical |
| Leaf.area | Leaf area | cm² |
| Nmass | Leaf nitrogen concentration | mg g⁻¹ |
| LMA | Leaf mass per area | g m⁻² |
| Plant.height | Plant height | m |
| Diaspore.mass | Mass of dispersal unit (seed/fruit) | mg |
| LDMC | Leaf dry matter content | mg g⁻¹ |
| SSD | Stem specific density | g cm⁻³ |
| database | Data source / database | categorical |
| SLA | Specific leaf area | mm² mg⁻¹ or cm² g⁻¹ |
| LeafN | Leaf nitrogen | % |
| LeafC | Leaf carbon | % |
| LeafCN | Leaf carbon:nitrogen ratio | unitless |
| Root.Shoot | Root:shoot biomass ratio | unitless |
| Light.Rating | Shade / light tolerance rating | categorical (1–10) |
| Moisture.Rating | Soil moisture requirement | categorical (1–10) |
| Ellenberg.light | Ellenberg light indicator | categorical (1–12) |
| Ellenberg.moisture | Ellenberg moisture indicator | categorical (1–12) |
| Ellenberg.pH | Ellenberg soil pH indicator | categorical |
| Ellenberg.N | Ellenberg nutrient indicator | categorical |
| Ellenberg.salt | Ellenberg salinity indicator | categorical |
| Ellenberg.temperature | Ellenberg temperature indicator | categorical |
| Ellenberg.humidity | Ellenberg humidity indicator | categorical |
| Growth.Form | Growth form (herb, shrub, tree, etc.) | categorical |
| Lifespan.years | Lifespan | years |
| Growing.Speed | Growth rate class | categorical |
| Height | Maximum plant height | mm |
| Grime.strategy | CSR strategy (Competitor–Stress Tolerator–Ruderal) | categorical |
| Lifespan | Lifespan | years |
| Pollination | Pollination syndrome | categorical |
| Pollinator | Pollinator type | categorical |
| Propagation | Propagation type | categorical |
| Resprout.ability | Ability to resprout after disturbance | categorical (yes/no) |
| Shade.tolerant | Shade tolerance | categorical |
| Classified.as.invasive | Invasive status | categorical (yes/no) |
| Cold.Tolerant | Cold tolerance | categorical |
| Fire.tolerant | Fire tolerance | categorical |
| Drought.tolerant | Drought tolerance | categorical |
Figure Summaries
- Figure 2: Species-level responses to CO₂ × competition
- Figure 3: Community-level biomass responses and dominance ratios
- Figure 4: Trait-based DAPC and LMM analyses
Execution Instructions
- Place all data in
/Data/ - Open
RCode.R - Run figure sections sequentially
- All outputs saved to
/R_out/
6. Code and Software
All coding is done using the R software package (R Core Team (2024). _R: A Language and Environment for Statistical Computing_. R Foundation for Statistical Computing, Vienna, Austria. https://www.R-project.org/.)
'#' is used throughout the code to label actions.
Session Information
R version 4.5.0 (2025-04-11)
Platform: aarch64-apple-darwin20
Running under: macOS 26.2
Matrix products: default
BLAS: /System/Library/Frameworks/Accelerate.framework/...
LAPACK: /Library/Frameworks/R.framework/...
(Full session info retained exactly as provided.)
Data Provenance & Ethics
Data collection: Compiled from published experimental studies on plant CO₂ responses and trait measurements. Full study list in manuscript references.
Ethics: No human subject data. No sensitive location data for threatened species.
Special formatting: No merged cells, highlighting, or other special formatting are used in the provided tabular data files.
Related Resources
Manuscript DOI: to be added upon acceptance
Dryad DOI: to be assigned upon upload
Source datasets cited in manuscript reference list.
Reproducibility Notes
All figures can be reproduced using the RCode.R script provided, which contains self-contained code blocks for each figure.
Ensure all CSV files are placed in the /Data/ directory as expected by the script.
Outputs are saved automatically to /R_out/.
No manual steps are required between figures.
