skip to main content


Title: Impact of rainfall seasonality on intraspecific trait variation in a shrub from a Mediterranean climate
Abstract

Selection pressures along climate gradients give rise to predictable variation in plant functional traits of individual species suggestive of local adaptation. Species whose ranges include winter rainfall, Mediterranean climates, or other strongly seasonal climates, may be exposed to divergent selection pressures at different ends of seasonality gradients.

Here, we evaluate how rainfall seasonality in conjunction with other key climatic variables impacts patterns of trait variation inPelargonium scabrum, a woody shrub from the Greater Cape Floristic Region of South Africa. This biodiversity hotspot encompasses a Mediterranean climate (wet winters and hot, dry summers) and displays steep gradients in temperature and water availability.

We used Bayesian regression models to evaluate leaf trait–trait and trait–climate relationships among 26 populations. Models included rainfall seasonality and its interaction with other climate variables (mean annual temperature, mean annual precipitation and potential evapotranspiration) as predictors to test for the impact of climate variation on three leaf traits: size, dissection and leaf mass per area (LMA). We evaluated model explanatory power by calculating BayesianR2values, and predictive power via leave‐one‐out cross‐validation.

Trait–trait associations were modulated by rainfall seasonality, including a reversal in the relationship between leaf size and dissection depending on the proportion of rain received in winter. Trait–climate models were improved by including rainfall seasonality as a predictor for both explanatory and predictive power. For leaf dissection and LMA, we detected significant interactions between rainfall seasonality and other environmental variables, leading to reversals in the relationships between these traits and the three environmental variables depending on the proportion of winter rainfall.

Differences in the timing of rainfall, coupled with strong differences in the covariation of climate variables, impose divergent selection pressures onP. scabrumpopulations resulting in divergence of trait values, trait integration and responses to climate gradients. These patterns are consistent with local adaptation ofP. scabrumpopulations mediated by the interactions between temperature and the amount and timing of rainfall. Species arrayed along broad climate gradients represent an excellent opportunity for investigating patterns of trait variation and abundances and distributions of species in relation to future changes in climate.

A freePlain Language Summarycan be found within the Supporting Information of this article.

 
more » « less
NSF-PAR ID:
10458086
Author(s) / Creator(s):
 ;  ;  ;  ;
Publisher / Repository:
Wiley-Blackwell
Date Published:
Journal Name:
Functional Ecology
Volume:
34
Issue:
4
ISSN:
0269-8463
Page Range / eLocation ID:
p. 865-876
Format(s):
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. Abstract

    Traits differentially adapt plant species to particular conditions generating compositional shifts along environmental gradients. As a result, community‐scale trait values show concomitant shifts, termed trait‒environment relationships. Trait‒environment relationships are often assessed by evaluating community‐weighted mean (CWM) traits observed along environmental gradients. Regression‐based approaches (CWMr) assume that local communities exhibit traits centred at a single optimum value and that traits do not covary meaningfully. Evidence suggests that the shape of trait‒abundance relationships can vary widely along environmental gradients—reflecting complex interactions—and traits are usually interrelated. We used a model that accounts for these factors to explore trait‒environment relationships in herbaceous forest plant communities in Wisconsin (USA).

    We built a generalized linear mixed model (GLMM) to analyse how abundances of 185 species distributed among 189 forested sites vary in response to four functional traits (vegetative height—VH, leaf size—LS, leaf mass per area—LMA and leaf carbon content), six environmental variables describing overstorey, soil and climate conditions, and their interactions. The GLMM allowed us to assess the nature and relative strength of the resulting 24 trait‒environment relationships. We also compared results between GLMM and CWMr to explore how conclusions differ between approaches.

    The GLMM identified five significant trait‒environment relationships that together explain ~40% of variation in species abundances across sites. Temperature appeared as a key environmental driver, with warmer and more seasonal sites favouring taller plants. Soil texture and temperature seasonality affected LS and LMA; seasonality effects on LS and LMA were nonlinear, declining at more seasonal sites. Although often assumed for CWMr, only some traits under certain conditions had centred optimum trait‒abundance relationships. CWMr more liberally identified (13) trait‒environment relationships as significant but failed to detect the temperature seasonality‒LMA relationship identified by the GLMM.

    Synthesis. Although GLMM represents a more methodologically complex approach than CWMr, it identified a reduced set of trait‒environment relationships still capable of accounting for the responses of forest understorey herbs to environmental gradients. It also identified separate effects of mean and seasonal temperature on LMA that appear important in these forests, generating useful insights and supporting broader application of GLMM approach to understand trait‒environment relationships.

     
    more » « less
  2. Summary

    Large intraspecific functional trait variation strongly impacts many aspects of communities and ecosystems, and is the medium upon which evolution works. Yet intraspecific trait variation is inconsistent and hard to predict across traits, species and locations.

    We measured within‐species variation in leaf mass per area (LMA), leaf dry matter content (LDMC), branch wood density (WD), and allocation to stem area vs leaf area in branches (branch Huber value (HV)) across the aridity range of seven Australian eucalypts and a co‐occurringAcaciaspecies to explore how traits and their variances change with aridity.

    Within species, we found consistent increases in LMA, LDMC and WD and HV with increasing aridity, resulting in consistent trait coordination across leaves and branches. However, this coordination only emerged across sites with large climate differences. Unlike trait means, patterns of trait variance with aridity were mixed across populations and species. Only LDMC showed constrained trait variation in more xeric species and drier populations that could indicate limits to plasticity or heritable trait variation.

    Our results highlight that climate can drive consistent within‐species trait patterns, but that patterns might often be obscured by the complex nature of morphological traits, sampling incomplete species ranges or sampling confounded stress gradients.

     
    more » « less
  3. Abstract

    In Mediterranean climates, the timing of seasonal rains determines germination, flowering phenology and fitness. As climate change alters seasonal precipitation patterns, it is important to ask how these changes will affect the phenology and fitness of plant populations. We addressed this question experimentally with the annual plant speciesArabidopsis thaliana.

    In a first experiment, we manipulated the date of rainfall onset and recorded germination phenology on sand and soil substrates. In a second experiment, we manipulated germination date, growing season length and mid‐season drought to measure their effects on flowering time and fitness. Within each experiment, we manipulated seed dormancy and flowering time using multilocus near‐isogenic lines segregating strong and weak alleles of the seed dormancy geneDOG1and the flowering time geneFRI. We synthesized germination phenology data from the first experiment with fitness functions from the second experiment to project population fitness under different seasonal rainfall scenarios.

    Germination phenology tracked rainfall onset but was slower and more variable on sand than on soil. Many seeds dispersed on sand in spring and summer delayed germination until the cooler temperatures of autumn. The high‐dormancyDOG1allele also prevented immediate germination in spring and summer. Germination timing strongly affected plant fitness. Fecundity was highest in the October germination cohort and declined in spring germinants. The late floweringFRIallele had lower fecundity, especially in early fall and spring cohorts. Projections of population fitness revealed that: (1) Later onset of autumn rains will negatively affect population fitness. (2) Slow, variable germination on sand buffers populations against fitness impacts of variable spring and summer rainfall. (3) Seasonal selection favours high dormancy and early flowering genotypes in a Mediterranean climate with hot dry summers. The high‐dormancyDOG1allele delayed germination of spring‐dispersed fresh seeds until more favourable early fall conditions, resulting in higher projected population fitness.

    These findings suggest that Mediterranean annual plant populations are vulnerable to changes in seasonal precipitation, especially in California where rainfall onset is already occurring later. The fitness advantage of highly dormant, early flowering genotypes helps explain the prevalence of this strategy in Mediterranean populations.

    Read the freePlain Language Summaryfor this article on the Journal blog.

     
    more » « less
  4. Abstract

    Environmental gradients act as potent filters on species distributions driving compositional shifts across communities. Compositional shifts may reflect differences in physiological tolerances to a limiting resource that result in broad distributions for tolerant species and restricted distributions for intolerant species (i.e. a nested pattern). Alternatively, trade‐offs in resource use or conflicting species' responses to multiple resources can result in complete turnover of species along gradients.

    We combined trait (leaf area, leaf mass per area, wood density and maximum height) and distribution data for 550 tree species to examine taxonomic and functional composition at 72 sites across strong gradients of soil phosphorus (P) and rainfall in central Panama.

    We determined whether functional and taxonomic composition was nested or turned over completely and whether community mean traits and species composition were more strongly driven by P or moisture.

    Turnover characterized the functional composition of tree communities. Leaf traits responded to both gradients, with species having larger and thinner leaves in drier and more fertile sites than in wetter and less fertile sites. These leaf trait–moisture relationships contradict predictions based on drought responses and suggest a greater role for differences in light availability than in moisture. Shifts in wood density and maximum height were weaker than for leaf traits with taller species dominating wet sites and low wood density species dominating P‐rich sites.

    Turnover characterized the taxonomic composition of tree communities. Geographic distances explained a larger fraction of variation for taxonomic composition than for functional composition, and community mean traits were more strongly driven by P than moisture.

    Synthesis. Our results offer weak support for the tolerance hypothesis for tree communities in central Panama. Instead, we observe functional and taxonomic turnover reflecting trade‐offs and conflicting species' responses to multiple abiotic factors including moisture, soil phosphorus and potentially other correlated variables (e.g. light).

     
    more » « less
  5. Abstract

    The coordination of traits from individual organs to whole plants is under strong selection because of environmental constraints on resource acquisition and use. However, the tight coordination of traits may provide underlying mechanisms of how locally adapted plant populations can become maladapted because of climate change.

    To better understand local adaptation in intraspecific trait coordination, we studied trait variability in the widely distributed foundation tree species,Populus fremontiiusing a common garden near the mid‐elevational point of this species distribution. We examined 28 traits encompassing four spectra: phenology, leaf economic spectrum (LES), whole‐tree architecture (Corner's Rule) and wood economic spectrum (WES).

    Based on adaptive syndrome theory, we hypothesized that trait expression would be coordinated among and within trait spectra, reflecting local adaptation to either exposure to freeze‐thaw conditions in genotypes sourced from high‐elevation populations or exposure to extreme thermal stress in genotypes sourced from low‐elevation populations.

    High‐elevation genotypes expressed traits within the phenology and WES that limit frost exposure and tissue damage. Specifically, genotypes sourced from high elevations had later mean budburst, earlier mean budset, higher wood densities, higher bark fractions and smaller xylem vessels than their low‐elevation counterparts. Conversely, genotypes sourced from low elevations expressed traits within the LES that prioritized hydraulic efficiency and canopy thermal regulation to cope with extreme heat exposure, including 40% smaller leaf areas, 67% higher stomatal densities and 34% higher mean theoretical maximum stomatal conductance. Low‐elevation genotypes also expressed a lower stomatal control over leaf water potentials that subsequently dropped to pressures that could induce hydraulic failure.

    Synthesis. Our results suggest thatPopulus fremontiiexpresses a high degree of coordination across multiple trait spectra to adapt to local climate constraints on photosynthetic gas exchange, growth and survival. These results, therefore, increase our mechanistic understanding of local adaptation and the potential effects of climate change that in turn, improves our capacity to identify genotypes that are best suited for future restoration efforts.

     
    more » « less