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  1. Abstract

    Many plant species can exhibit remarkable variation in leaf characteristics, depending on their abiotic and biotic environment. Environmental changes therefore have the potential to alter leaf traits, which in turn scale up to influence ecosystem processes including net primary productivity, susceptibility to fire, and palatability to herbivores. It is not well understood how consistent trait–environment relationships are among species, across sites and over time. This presents a fundamental challenge for functional ecology, because no study can measure all relevant species in all places at all times. Thus, understanding the limits of transferability is critical. We collected leaf trait measurements on 13 species of grass (family: Poaceae) across 11 sites and five years (n = 3091 individuals). Sites were arrayed along a spatial precipitation gradient in coastal northern California (annual precipitation of 590–1350 mm) with substantial interannual precipitation variability (from 60% below the 30‐year average to 100% above average). Temporal and spatial linear relationships between precipitation and specific leaf area (SLA) appear at first idiosyncratic, with each species sometimes displaying positive and sometimes negative responses. However, this variation arises from sampling different portions of an underlying hump‐shaped relationship, which was shared across most species. This hump‐shaped relationship was driven primarily by changes in leaf tissue density. These results suggest the potential for transferability among species, as well as between space and time, as long as the gradients are sufficiently long to capture the nonlinear response. Future work could explore the physiological basis of the nonlinear SLA response, including the possibility that distinct physiological mechanisms are operating at the two extremes of the gradient.

     
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  2. Abstract

    Plant populations are limited by resource availability and exhibit physiological trade‐offs in resource acquisition strategies. These trade‐offs may constrain the ability of populations to exhibit fast growth rates under water limitation and high cover of neighbours. However, traits that confer drought tolerance may also confer resistance to competition. It remains unclear how fitness responses to these abiotic conditions and biotic interactions combine to structure grassland communities and how this relationship may change along a gradient of water availability.

    To address these knowledge gaps, we estimated the low‐density growth rates of populations in drought conditions with low neighbour cover and in ambient conditions with average neighbour cover for 82 species in six grassland communities across the Central Plains and Southwestern United States. We assessed the relationship between population tolerance to drought and resistance to competition and determined if this relationship was consistent across a precipitation gradient. We also tested whether population growth rates could be predicted using plant functional traits.

    Across six sites, we observed a positive correlation between low‐density population growth rates in drought and in the presence of interspecific neighbours. This positive relationship was particularly strong in the grasslands of the northern Great Plains but weak in the most xeric grasslands. High leaf dry matter content and a low (more negative) leaf turgor loss point were associated with high population growth rates in drought and with neighbours in most grassland communities.

    Synthesis: A better understanding of how both biotic and abiotic factors impact population fitness provides valuable insights into how grasslands will respond to extreme drought. Our results advance plant strategy theory by suggesting that drought tolerance increases population resistance to interspecific competition in grassland communities. However, this relationship is not evident in the driest grasslands, where above‐ground competition is likely less important. Leaf dry matter content and turgor loss point may help predict which populations will establish and persist based on local water availability and neighbour cover, and these predictions can be used to guide the conservation and restoration of biodiversity in grasslands.

     
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    Free, publicly-accessible full text available February 1, 2025
  3. Abstract

    Plant traits can be helpful for understanding grassland ecosystem responses to climate extremes, such as severe drought. However, intercontinental comparisons of how drought affects plant functional traits and ecosystem functioning are rare. The Extreme Drought in Grasslands experiment (EDGE) was established across the major grassland types in East Asia and North America (six sites on each continent) to measure variability in grassland ecosystem sensitivity to extreme, prolonged drought. At all sites, we quantified community‐weighted mean functional composition and functional diversity of two leaf economic traits, specific leaf area and leaf nitrogen content, in response to drought. We found that experimental drought significantly increased community‐weighted means of specific leaf area and leaf nitrogen content at all North American sites and at the wetter East Asian sites, but drought decreased community‐weighted means of these traits at moderate to dry East Asian sites. Drought significantly decreased functional richness but increased functional evenness and dispersion at most East Asian and North American sites. Ecosystem drought sensitivity (percentage reduction in aboveground net primary productivity) positively correlated with community‐weighted means of specific leaf area and leaf nitrogen content and negatively correlated with functional diversity (i.e., richness) on an intercontinental scale, but results differed within regions. These findings highlight both broad generalities but also unique responses to drought of community‐weighted trait means as well as their functional diversity across grassland ecosystems.

     
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    Free, publicly-accessible full text available February 1, 2025
  4. Plant traits are useful for predicting how species may respond to environmental change and/or influence ecosystem properties. Understanding the extent to which traits vary within species and across climatic gradients is particularly important for understanding how species may respond to climate change. We explored whether climate drives spatial patterns of intraspecific trait variation for three traits (specific leaf area (SLA), plant height, and leaf nitrogen content (Nmass)) across 122 grass species (family: Poaceae) with a combined distribution across six continents. We tested the hypothesis that the sensitivity (i.e. slope) of intraspecific trait responses to climate across space would be related to the species' typical form and function (e.g. leaf economics, stature and lifespan). We observed both positive and negative intraspecific trait responses to climate with the distribution of slope coefficients across species straddling zero for precipitation, temperature and climate seasonality. As hypothesized, variation in slope coefficients across species was partially explained by leaf economics and lifespan. For example, acquisitive species with nitrogen-rich leaves grew taller and produced leaves with higher SLA in warmer regions compared to species with low Nmass. Compared to perennials, annual grasses invested in leaves with higher SLA yet decreased height and Nmass in regions with high precipitation seasonality (PS). Thus, while the influence of climate on trait expression may at first appear idiosyncratic, variation in trait–climate slope coefficients is at least partially explained by the species' typical form and function. Overall, our results suggest that a species' mean location along one axis of trait variation (e.g. leaf economics) could influence how traits along a separate axis of variation (e.g. plant size) respond to spatial variation in climate. 
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    Free, publicly-accessible full text available May 16, 2024
  5. Abstract

    Plant traits are useful proxies of plant strategies and can influence community and ecosystem responses to climate extremes, such as severe drought. Few studies, however, have investigated both the immediate and lagged effects of drought on community‐weighted mean (CWM) plant traits, with even less research on the relative roles of interspecific vs. intraspecific trait variability in such responses.

    We experimentally reduced growing season precipitation by 66% in two cold‐semi‐arid grassland sites in northern China for four consecutive years to explore the drought resistance of CWM traits as well as their recovery 2 years following the drought. In addition, we isolated the effects of both interspecific and intraspecific trait variability on shifts in CWM traits.

    At both sites, we observed significant effects of drought on interspecific and intraspecific trait variability which, in some cases, led to significant changes in CWM traits. For example, drought led to reduced CWM plant height and leaf phosphorous content, but increased leaf carbon content at both sites, with responses primarily due to intraspecific trait shifts. Surprisingly, these CWM traits recovered completely 2 years after the extreme drought. Intraspecific trait variability influenced CWM traits via both positive and negative covariation with interspecific trait variability during drought and recovery phases.

    These findings highlight the important role of interspecific and intraspecific trait variability in driving the response and recovery of CWM traits following extreme, prolonged drought.

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

     
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  6. Abstract

    Seeds provide the basis of genetic diversity in perennial grassland communities and their traits may influence ecosystem resistance to extreme drought. However, we know little about how drought effects the community functional composition of seed traits and the corresponding implications for ecosystem resistance to drought.

    We experimentally removed 66% of growing season precipitation for 4 years across five arid and semi‐arid grasslands in northern China and assessed how this multi‐year drought impacted community‐weighted means (CWMs) of seed traits, seed trait functional diversity and above‐ground net primary productivity (ANPP).

    Experimental drought had limited effects on CWM traits and the few effects that did occur varied by site and year. For three separate sites, and in different years, drought reduced seed length and phosphorus content but increased both seed and seed‐coat thickness. Additionally, drought led to increased seed functional evenness, divergence, dispersion and richness, but only in some sites, and mostly in later years following cumulative effects of water limitation. However, we observed a strong negative relationship between drought‐induced reductions in ANPP and CWMs of seed‐coat thickness, indicating that a high abundance of dominant species with thick seeds may increase ecosystem resistance to drought. Seed trait functional diversity was not significantly predictive of ANPP, providing little evidence for a diversity effect.

    Our results suggest that monitoring community composition with a focus on seed traits may provide a valuable indicator of ecosystem resistance to future droughts despite inconsistent responses of seed trait composition overall. This highlights the importance of developing a comprehensive seed and reproductive traits database for arid and semi‐arid grassland biomes.

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

     
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  7. Abstract

    We review results from field experiments that simulate drought, an ecologically impactful global change threat that is predicted to increase in magnitude, extent, duration and frequency. Our goal is to address, from primarily an ecosystem perspective, the questions ‘What have we learned from drought experiments?’ and ‘Where do we go from here?’.

    Drought experiments are among the most numerous climate change manipulations and have been deployed across a wide range of biomes, although most are conducted in short‐statured, water‐limited ecosystems. Collectively, these experiments have enabled ecologists to quantify the negative responses to drought that occur for most aspects of ecosystem structure and function. Multiple meta‐analyses of responses have also enabled comparisons of relative effect sizes of drought across hundreds of sites, particularly for carbon cycle metrics. Overall, drought experiments have provided strong evidence that ecosystem sensitivity to drought increases with aridity, but that plant traits associated with aridity are not necessarily predictive of drought resistance. There is also intriguing evidence that as drought magnitude or duration increases to extreme levels, plant strategies may shift from drought tolerance to drought escape/avoidance.

    We highlight three areas where more drought experiments are needed to advance our understanding. First, because drought is intensifying in multiple ways, experiments are needed that address alterations in drought magnitude versus duration, timing and/or frequency (individually and interactively). Second, drivers of drought may be shifting—from precipitation deficits to rising atmospheric demand for water—and disentangling how ecosystems respond to changes in hydrological ‘supply versus demand’ is critical for understanding drought impacts in the future. Finally, more attention should be focussed on post‐drought recovery periods since legacies of drought can affect ecosystem functioning much longer than the drought itself.

    We conclude with a call for a fundamental shift in the focus of drought experiments from those designed primarily as ‘response experiments’, quantifying the magnitude of change in ecosystem structure and function, to more ‘mechanistic experiments’—those that explicitly manipulate ecological processes or attributes thought to underpin drought responses.

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

     
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