Evidence of local adaptation to aridity but not nitrogen deposition in invasive annuals
Data files
Jun 25, 2025 version files 58.28 KB
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BRODIA_data.csv
23.68 KB
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CENMEL_data.csv
22.94 KB
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README.md
10.60 KB
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site_data.csv
1.06 KB
Abstract
Rapid adaptation of traits and trait plasticity may contribute to plant invasiveness and confer fitness advantages in novel environments resulting from global change. The importance of trait differentiation in invasive plant populations is well-recognized, particularly in response to climate. However, it is largely unknown how invasive plant populations will respond evolutionarily to atmospheric nitrogen (N) deposition resulting from air pollution, which is a major contributor to invasion success in many ecosystems. Using a common garden experiment, a widely used method for testing local adaptation, we assessed potential differences in plant functional traits and nitrogen plasticity across populations of two widespread invasive annuals from sites spanning a range of N deposition and aridity throughout southern California. These species exhibited clear population-level differences in traits and N responses, but these were unrelated to N deposition. Instead, we detected significant relationships between several traits and aridity, and populations from more arid sites exhibited reduced N plasticity for multiple traits. Multivariate plasticity indices also showed a strong negative relationship with aridity across populations for both species. However, trait responses to N addition also appeared be influenced by species’ drought-coping strategies. In Bromus diandrus, a drought-escaping early-season annual grass, populations from less arid sites showed increased plasticity in shoot growth and more rapid flowering in response to N addition. In contrast, Centaurea melitensis, a drought-tolerant late-season forb, showed climate-driven shifts in biomass allocation in response to N; populations from more arid sites invested more in roots, while populations from less arid sites allocated more to leaves. These contrasting N responses strongly suggest distinct growth strategies and ecophysiological tradeoffs shaped by adaptation to local climate conditions. While elevated N availability may indeed promote invasion, climate stress might exert an overriding influence on local adaptation of plant invaders in dryland ecosystems subject to N deposition.
Dataset DOI: 10.5061/dryad.r4xgxd2r3
Description of the data and file structure
Data collected from a common garden experiment in which multiple populations of two invasive plant species, Bromus diandrus and Centaurea melitensis, were grown under high and low nitrogen. Plant traits and responses were then related to aridity and nitrogen deposition of the source populations.
Files and variables
File: BRODIA_data.csv
Description: Plant trait data for Bromus diandrus populations grown under high and low nitrogen availability. For some plant traits (i.e., Amax, Cond, WUE, SLA, perc_N, CN, and d13C), data were collected only on a subset of individuals in each treatment, so these columns contain blank cells.
Variables
Variable | Description | Units |
---|---|---|
site_number |
Site identification number | unitless |
rep |
Replicate number | unitless |
Maternal_ID |
Identifier for maternal plant source | unitless (ID code) |
M_seed_ave |
Average mass of maternal seeds | g (grams) |
N |
Nitrogen treatment (high N or low N) | unitless (treatment) |
root |
Root biomass | g (grams) |
Infl_count |
Inflorescence count | count |
leaf |
Leaf biomass | g (grams) |
repr_mass |
Reproductive biomass (flowers and/or seeds) | g (grams) |
total_mass |
Total plant biomass (root + shoot) | g (grams) |
shoot_mass |
Total above-ground biomass (stem + leaf + repr) | g (grams) |
LMR |
Leaf mass ratio (leaf_mass / shoot_mass) | unitless (proportion) |
RMR |
Root mass ratio (root_mass / total_mass) | unitless (proportion) |
seed_mass |
Average seed mass | g (grams) |
DTF |
Days to flowering | days |
Amax |
Maximum photosynthetic rate | μmol CO₂ m⁻² s⁻¹ |
Cond |
Stomatal conductance | mol H₂O m⁻² s⁻¹ |
WUE |
Water use efficiency (Amax / transpiration) | μmol CO₂ / mmol H₂O |
SLA |
Specific leaf area | cm² g⁻¹ |
perc_N |
Leaf nitrogen content | % dry weight |
CN |
Carbon-to-nitrogen ratio | unitless (C/N ratio) |
d13C |
Carbon isotope discrimination | ‰ (per mil) |
File: CENMEL_data.csv
Description: Plant trait data for Centaurea melitensis populations grown under high and low nitrogen availability. For some plant traits (i.e., Amax, Cond, WUE, SLA, perc_N, CN, and d13C), data were collected only on a subset of individuals in each treatment, so these columns contain blank cells.
Variables
Variable | Description | Units |
---|---|---|
site_number | Site identification number (see site_data.csv) | unitless |
rep | Replicate number | unitless |
Maternal_ID | Identifier for maternal plant source | unitless (ID code) |
M_seed_ave | Average mass of maternal seeds | g (grams) |
N | Nitrogen treatment (high N or low N) | unitless (treatment) |
root_mass | Root biomass | g (grams) |
stem_mass | Stem biomass | g (grams) |
leaf_mass | Leaf biomass | g (grams) |
repr_mass | Reproductive biomass (flowers/seeds) | g (grams) |
flower# | Number of flowers produced | count |
seeds_per_flower | Average number of seeds per flower | count |
total_seed_mass | Total seed mass produced | g (grams) |
mean_seed_mass | Average mass per seed | g (grams) |
total_mass | Total above- and below-ground biomass | g (grams) |
shoot_mass | Total above-ground biomass (stem + leaf + repr) | g (grams) |
LMR | Leaf mass ratio (leaf_mass / shoot_mass) | unitless (proportion) |
RMR | Root mass ratio (root_mass / total_mass) | unitless (proportion) |
DTF | Days to flowering | days |
Amax | Maximum photosynthetic rate | μmol CO₂ m⁻² s⁻¹ |
Cond | Stomatal conductance | mol H₂O m⁻² s⁻¹ |
WUE | Water use efficiency (Amax / transpiration) | μmol CO₂ / mol H₂O |
SLA | Specific leaf area | cm² g⁻¹ |
perc_N | Leaf nitrogen content | % dry weight |
CN | Carbon-to-nitrogen ratio | unitless (C/N ratio) |
d13C | Stable carbon isotope ratio (carbon isotope discrimination) | ‰ (per mil) |
File: site_data.csv
Description: Site-level data for the locations from which seed of each species was collected for plant growth trials in the common garden experiment.
Variables
Variable | Description | Units |
---|---|---|
site | Site identification number | unitless |
site_name | Name of the site | text |
lat | Latitude of the site | decimal degrees (°N) |
lng | Longitude of the site | decimal degrees (°W) |
CMAQ_2002 | Modeled nitrogen deposition from CMAQ for the year 2002 | kg N ha⁻¹ yr⁻¹ |
CMAQ_2017 | Modeled nitrogen deposition from CMAQ for the year 2017 | kg N ha⁻¹ yr⁻¹ |
average_Ndep | Average nitrogen deposition across years (e.g., 2002 and 2017) | kg N ha⁻¹ yr⁻¹ |
elev | Elevation of the site | meters above sea level (m) |
mean_annual_ppt | Mean annual precipitation | mm yr⁻¹ |
map_CV | Coefficient of variation in mean annual precipitation | unitless |
mean_tmax | Mean maximum temperature (annual or seasonal, depending on dataset) | °C |
aridity_index | Aridity index | unitless (index) |
Software
All statistical analyses and data visualization were conducted using open-source software in R (R Core Team 2013), version 4.1.0, within the RStudio environment. Analyses were performed on a standard desktop computing setup. To view and reproduce the data analyses, users will need to install R and RStudio, both of which are freely available at https://www.r-project.org and https://www.rstudio.com, respectively.
The following R packages were used in the analysis:
psych
: for principal component analysis (PCA) and data explorationggplot2
: for data visualizationdplyr
,tidyr
: for data wrangling and transformationcar
: for model diagnostics and ANOVA toolsstats
(base): foraov
,lm
, andanova
functions used in model fitting- Custom functions were written for calculating plasticity indices such as log response ratios and the multivariate plasticity index (MVPi) following the method of Pennacchi et al. (2021)
Pennacchi, J. P., J. M. S. Lira, M. Rodrigues, F. H. S. Garcia, A. M. d. C. Mendonça, and J. P. R. A. D. Barbosa. 2021. A systemic approach to the quantification of the phenotypic plasticity of plant physiological traits: the multivariate plasticity index. Journal of Experimental Botany 72:1864-1878.
Access information
Climate data were acquired from the Oregon State University PRISM Group’s online tool (https://prism.oregonstate.edu/). All data retrieved from this website or otherwise provided on the website may be freely reproduced and distributed for non-commercial purposes
PRISM Group, Oregon State University, https://prism.oregonstate.edu, data accessed August 2020.
Nitrogen deposition data were acquired from the Environmental Protection Agency’s Community Multiscale Air Quality model. These data are publicly available.
Benish, S. E., J. O. Bash, K. M. Foley, K. W. Appel, C. Hogrefe, R. Gilliam, and G. Pouliot. 2022. Long-term Regional Trends of Nitrogen and Sulfur Deposition in the United States from 2002 to 2017. Atmospheric Chemistry and Physics Discussions:1-27.