Data from: Trait responses to competition are mostly parallel among competitor plants but can cause convergence or divergence in key traits
Data files
Jul 01, 2026 version files 859.21 KB
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biomassdatafinal.csv
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eigenvalues_9_30.csv
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eigenvaluesplast.csv
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eigenvectors_9_30.csv
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eigenvectorsplast.csv
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finalplotbio.csv
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finalplots.csv
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finalrootlength.csv
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floweringfinal.csv
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heightfinal2024.csv
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individualbiomassgrassesfinal.csv
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isotope1.csv
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isotope10.csv
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isotope11.csv
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isotope12.csv
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isotope13.csv
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isotope14.csv
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isotope15.csv
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isotope16.csv
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isotope17.csv
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isotope18.csv
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isotope19.csv
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isotope2.csv
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isotope20.csv
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isotope21.csv
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isotope22.csv
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isotope23.csv
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isotope24.csv
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isotope25.csv
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isotope26.csv
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isotope27.csv
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isotope28.csv
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isotope3.csv
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isotope4.csv
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isotope5.csv
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isotope6.csv
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isotope7.csv
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isotope8.csv
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isotope9.csv
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PCAscores_9_30.csv
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PCAscoresplast.csv
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plasticitycoderevised.R
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predawnwaterpotentialfinal.csv
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pwpleafweights.csv
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README.md
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rootmassfinal.csv
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SeedCounts.csv
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SIMPERCOLOR(2).txt
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SIMPERCOLORSPPBC(2).txt
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SIMPERCOLORSPPBH(2).txt
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SIMPERCOLORSPPFM(2).txt
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SIMPERCOLORSPPFO(2).txt
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SIMPERCOLORSPPLP(2).txt
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SIMPERCOLORSPPOP(2).txt
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SIMPERINTRA(2).txt
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SIMPERINTRABC(2).txt
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SIMPERINTRABH(2).txt
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SIMPERINTRAFM(2).txt
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SIMPERINTRAFO(2).txt
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SIMPERINTRALP(2).txt
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SIMPERINTRAOP(2).txt
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SIMPERSPP(2).txt
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Abstract
Plants competing for resources exhibit different traits than plants which are not. Phenotypic plasticity can shape competition by altering trait differences between competitors. Studies show that plastic traits may converge or diverge among competitors as competition intensifies. However, another possibility is that interacting species could exhibit similar trait shifts as competition intensifies (trait parallelism). We used a field experiment to assess trait responses and performance outcomes for six co-occurring native and invasive annual California grassland species when grown together at various densities. We grew one focal individual of each species in each plot and manipulated the identity and density of competitors, from bare ground to a dense lawn. Species displayed many trait responses to competitor density which depended on competitor identity more than focal species identity. Most trait responses were parallel among competing plants with no effect on trait differences between competitors. The exceptions were specific root length, which diverged among competitors with increased competitor density, root mass fraction and leaf dry matter content, which both converged among competitors with increased competitor density. Competitors’ effects on performance were not correlated with greater responses in any trait. A few traits displayed non-parallel responses among competing plants, altering trait differences between competitors. These traits may be of interest for future studies of plasticity's effects on competition or species coexistence.
Dataset DOI: 10.5061/dryad.4qrfj6qqw
Description of the data and file structure
When plants compete for resources, they grow with different traits than plants which are not experiencing competition. Phenotypic plasticity can shape competition by altering trait differences among competitors. While there are empirical examples of both plastic trait divergence and convergence among competitors, the commonality of parallel versus non-parallel plasticity among competitors remains unclear.
We used a field experiment to assess plastic trait shifts and fitness outcomes for six annual species when grown together at a range of densities. These data describe the traits and fitness of the annual plants grown along a gradient of competitor density.
We found that species displayed plasticity in a wide variety of traits which depended more on competitor species than focal species identity.
Contrary to expectations, most plasticity was parallel among competing plants. The exceptions were specific root length, which diverged among competitors with increased competitor density, root mass fraction and leaf dry matter content, which both converged among competitors with increased competitor density.
Competitors’ effects on fitness were not correlated with greater plasticity of any trait.
The traits which display non-parallel trait shifts among competing plants may be of particular interest for future studies of plasticity's role shaping community dynamics.
The data are in a collection of files. Some of these are raw data for fitness or traits of focal plants. Some of these are outputs from multivariate analyses conducted in PRIMER-e v7.
Files and variables
Experimental treatment data:
finalplotbio.csv:
Competitor biomass at end of growing season
block= replicate block, 1-3
plot= plots 1-48 within each block
biomass= g dry competitor aboveground biomass at end of growing season from a 7.6 cm x 80 cm strip down the center of the plot
finalplots.csv:
Plot species and treatment information. Each row is a plot.
spp= Competitor species (fo= Festuca octoflora, fm= Festuca myuros, op= Oncosiphon pilulifer, lp= Layia platyglossa, Bh= Bromus hordeaceus, Bc= Bromus sitchensis var. carinatus). NA means no competitor— omitted from analyses due to persistent flooding and poor growth.
vdensity= 8 evenly spaced levels of viable competitor seed density from 1-8 g/m^2 for all species except op. Due to seed limitations and small seed mass, op seed levels are 0.29–4.05 g/m². NA means no competitor— omitted from analyses due to persistent flooding and poor growth.
viab= seed viability rate for each competitor species
pdensity= plot total competitor seed density
pmass= plot total competitor seed mass
name= treatment name, competitor species + density level, NC is no competitor— omitted from analyses due to persistent flooding and poor growth.
block= replicate block, 1-3
plot= plots 1-48 within each block
sp1-6= focal species in each evenly spaced focal individual ring
gopher= record of gopher activity in plot, Y, slight (<25% of plot affected), or blank
Fitness outcomes data:
biomassdatafinal.csv:
Data for focal individuals’ weights and some trait measures. Each row is a focal individual.
spp= Competitor species (fo= Festuca octoflora, fm= Festuca myuros, op= Oncosiphon pilulifer, lp= Layia platyglossa, Bh= Bromus hordeaceus, Bc= Bromus sitchensis var. carinatus). NA means no competitor— omitted from analyses due to persistent flooding and poor growth.
vdensity= 8 evenly spaced levels of viable competitor seed density from 1-8 g/m^2 for all species except op. Due to seed limitations and small seed mass, op seed levels are 0.29–4.05 g/m². NA means no competitor— omitted from analyses due to persistent flooding and poor growth.
name= treatment name, competitor species + density level, NC is no competitor— omitted from analyses due to persistent flooding and poor growth.
block= replicate block, 1-3
plot= plots 1-48 within each block
color= Focal individual species in color coded rings (green= Festuca octoflora, purple= Festuca myuros, red= Oncosiphon pilulifer, orange= Layia platyglossa, blue= Bromus hordeaceus, yellow= Bromus sitchensis var. carinatus)
area= leaf area cm^2 obtained by analysis in ImageJ of photographs of leaves using the Analyze Particles function
perimeter= leaf perimeter cm obtained by analysis in ImageJ of photographs of leaves using the Analyze Particles function
fweight= fresh weight of leaf g
dweight= dry weight of leaf g
biomass= dry weight of individual g
biomass scale= coarse or fine balance used, depended on size of plant
flowerheads= number of flower heads on individual
basalshoots= number of shoots branching in the first cm from the soil surface for forb individuals (op and lp only)
herbivory= 0 for no sign of herbivory, 1 for signs of herbivory
notes= other information about focal individual
pwpleafweights.csv:
Data for mass of leaves taken for predawn water potential, to be added back to aboveground biomass. Each row is a leaf from a focal individual.
block= replicate block, 1-3
plot= plots 1-48 within each block
color= focal individual species (fo= Festuca octoflora, fm= Festuca myuros, op= Oncosiphon pilulifer, lp= Layia platyglossa, Bh= Bromus hordeaceus, Bc= Bromus sitchensis var. carinatus)
pwpdweight= dry weight of leaf taken for predawn water potential g
notes= other information about leaf
SeedCounts.csv:
Seed counts for seed heads collected from competitor individuals at each density for each species. Each row is a sample of seedheads from competitor plants at a particular density.
color= focal individual species (fo= Festuca octoflora, fm= Festuca myuros, op= Oncosiphon pilulifer, lp= Layia platyglossa, Bh= Bromus hordeaceus, Bc= Bromus sitchensis var. carinatus)
vdensity= 8 evenly spaced levels of viable competitor seed density from 1-8 g/m^2 for all species except op. Due to seed limitations and small seed mass, op seed levels are 0.29–4.05 g/m².
seedheads= number of seed heads in sample
seeds= number of seeds in sample
individualbiomassgrassesfinal.csv:
Mass of late flowering grasses and number of seed heads for estimate of seed head production at end of growing season for individuals harvested prior to flowering
color= focal individual species (fm= Festuca myuros, Bh= Bromus hordeaceus, Bc= Bromus sitchensis var. carinatus)
biomass= g dry aboveground biomass competitor individual at end of growing season
no_sd= number of seed heads on competitor individual at end of growing season
Trait outcomes data:
finalrootlength.csv:
Root length of focal individuals collected via neural network from images of cleaned roots in a shallow tray of water on a scanner
block= replicate block, 1-3
plot= plots 1-48 within each block
species= focal individual species (fo= Festuca octoflora, fm= Festuca myuros, op= Oncosiphon pilulifer, lp= Layia platyglossa, Bh= Bromus hordeaceus, Bc= Bromus sitchensis var. carinatus)
length_cm= root length in cm
floweringfinal.csv:
Phenology data for flower and bud timing of focal individuals and competitors (out)
date= date of record, month/day/year
days_since_planting= days since planting
block= replicate block, 1-3
plot= plots 1-48 within each block
yellow_bc, red_op, orange_lp, green_fo, blue_bh, purple_fm= phenological stage of focal individual of each species (n= no buds or flowers, b= at least 1 bud, f= at least 1 open flower, NA= plant missing)
out= phenological stage of competitor lawn with percentage of plants at that stage 0-25%, 25-50%, 50-75%, 75-100%, or 100% (n= no buds or flowers, b= at least 1 bud, f= at least 1 open flower)
out_ctd= phenological stage of competitor lawn with percentage of plants at that stage 0-25%, 25-50%, 50-75%, 75-100%, or 100% (n= no buds or flowers, b= at least 1 bud, f= at least 1 open flower), equivalent out column since plots can have some % of competitor plants budding and some % of competitor plants flowering
heightfinal2024.csv:
Height of focal and competitor (out) plants during growing season. Each row is a height of a plant on a date.
date= date of record, day/month/year
day_since_planting= days since planting
block= replicate block, 1-3
plot= plots 1-48 within each block
color_or_out= focal individual’s species (green= Festuca octoflora, purple= Festuca myuros, red= Oncosiphon pilulifer, orange= Layia platyglossa, blue= Bromus hordeaceus, yellow= Bromus sitchensis var. carinatus) out means tallest competitor in one half of plot
height_cm= height of plant from soil surface to highest point in cm, NA means plant missing
predawnwaterpotentialfinal.csv:
Predawn water potential data for newest fully expanded leaves of focal individuals in 1 block, filled in with same treatment from next block when plant missing (due to herbivory)
name= treatment name, competitor species + density level, NC is no competitor— omitted from analyses due to persistent flooding and poor growth.
block= replicate block, 1-3
plot= plots 1-48 within each block
color= Focal individual species in color coded rings(green= Festuca octoflora, purple= Festuca myuros, red= Oncosiphon pilulifer, orange= Layia platyglossa, blue= Bromus hordeaceus, yellow= Bromus sitchensis var. carinatus)
water potential= - leaf predawn water potential in bars
notes= other information
date= date of sampling, month/day/year
rootmassfinal.csv:
Dry root mass of focal individuals. Each row is a root from a focal individual.
block= replicate block, 1-3
plot= plots 1-48 within each block
species= focal individual species (fo= Festuca octoflora, fm= Festuca myuros, op= Oncosiphon pilulifer, lp= Layia platyglossa, Bh= Bromus hordeaceus, Bc= Bromus sitchensis var. carinatus)
dry mass (g)= dry mass of focal individual’s root g
isotope1.csv through isotope28.csv:
C and N isotope data from mass spectrometry by UC Merced of leaf tissue from focal individuals
Identifier_1= unique ID for each sample or standard
sample= name of sample or standard, for leaf tissue samples the name is block-plotspecies
amount= mg dry sample or standard
Area 28= N2 gas content, nitrogen content
Area 44= CO2 gas content, carbon content
d13C_corrected= delta 13 C, a measure of the ratio of the two stable isotopes of carbon reported in parts per thousand, corrected for drift and standardized
d15N_corrected= delta 15 N, a measure of the ratio of the two stable isotopes of nitrogen, corrected for drift and standardized
N_wt_percent= percent nitrogen by mass
C_wt_percent= percent carbon by mass
C/N= carbon to nitrogen mass ratio
Outputs of analyses in PRIMER-e v7:
PCAscores_9_30.csv:
PCA of trait variation scores
Sample= unique number for sample, in same order as input file
SCORE1-SCORE5= scores for samples from first 5 PC axes
PCAscoresplast.csv:
PCA of plasticities (linear change in log normalized trait over log competitor biomass in each species pair) scores
Sample= unique number for sample, in same order as input file
SCORE1-SCORE5= scores for samples from first 5 principal component (PC) axes
eigenvalues_9_30.csv:
Corresponding eigenvalues, variation explained by axes
PC= PC number
Eigenvalues= amount of trait variation explained by PC
%Variation= % total trait variation explained by PC
Cum.%Variation= cumulative % of trait variation explained by PC
eigenvaluesplast.csv:
Corresponding eigenvalues, variation explained by axes
PC= PC number
Eigenvalues= amount of plasticity variation explained by PC
%Variation= % total plasticity variation explained by PC
Cum.%Variation= cumulative % of plasticity variation explained by PC
eigenvectors_9_30.csv:
Corresponding eigenvectors, trait loadings
Variable= trait, abbreviations in Table 1
PC1-PC5= direction of maximum variance in that trait
eigenvectorsplast.csv:
Corresponding eigenvectors, trait plasticity loadings
Variable= trait, abbreviations in Table 1
PC1-PC5= direction of maximum variance in that plasticity
SIMPER files:
Files of SIMPER outputs for comparisons related to all PERMANOVA factors that significantly explained trait variation. The identity of the groups being compared can be found in the header above the tables.
SIMPERCOLOR(2).txt compares traits of focal species. This comparison reveals the traits responsible for the significant PERMANOVA main effect of focal species.
SIMPERCOLORSPP(BC, BH, FM, FO, OP, LP)(2).txt compares traits of focal species in different competitor species contexts. This comparison reveals the traits responsible for the significant PERMANOVA effect of focal x competitor species.
SIMPERINTRA(2).txt compares traits of focal individuals in conspecific and heterospecific contexts. This comparison reveals the traits responsible for the significant PERMANOVA main effect of conspecific vs heterospecific context.
SIMPERINTRA(BC, BH, FM, FO, OP, LP)(2).txt compares traits of focal species in conspecific and heterospecific contexts. This comparison reveals the traits responsible for the significant PERMANOVA effect of conspecific vs heterospecific context x focal species.
SIMPERSPP(2).txt compares traits of focal individuals in each competitor species context. This comparison reveals the traits responsible for the significant PERMANOVA main effect of competitor species.
Variable= trait being evaluated for its contribution to between-group dissimilarity
Av.Value= average contribution of the trait to the overall Euclidean dissimilarity
Av.Sq.Dist= average squared contribution of that trait to the dissimilarity
Sq.Dist/SD= ratio of squared dissimilarity to its standard deviation. Higher values = more consistent contribution across samples
Contrib%= percentage of total dissimilarity explained by that trait
Cum.%= cumulative percentage of dissimilarity explained up to and including this trait
Code/software
plasticitycoderevised.R: All code to reproduce figures and tables is in this one R file, which uses the public data. This code was made in RStudio Version 2023.12.1+402 (2023.12.1+402) running R version 4.4.2 (2024-10-31).
Packages required: tidyverse, vegan, missMDA, FactoMineR, factoextra, ggrepel, car, emmeans, corrplot, grid, cowplot, betareg, smatr, ggforce)
Access information
All data was collected for this project in 2024. No previously published data was used.
