Climate models predict more frequent, prolonged, and extreme droughts in the future. Therefore, drought experiments varying in amount and duration across a range of biogeographical scenarios provide a powerful tool for estimating how drought will affect future ecosystems. Past experimental work has been focused on the manipulation of meteorological drought: Rainout shelters are used to reduce precipitation inputs into the soil. This work has been instrumental in our ability to predict the expected effects of altered rainfall. But what about the nonrainfall components of drought? We review recent literature on the co-occurring and sometimes divergent impacts of atmospheric drying and meteorological drying. We discuss how manipulating meteorological drought or rainfall alone may not predict future changes in plant productivity, composition, or species interactions that result from climate change induced droughts. We make recommendations for how to improve these experiments using manipulations of relative humidity.
Note: When clicking on a Digital Object Identifier (DOI) number, you will be taken to an external site maintained by the publisher.
Some full text articles may not yet be available without a charge during the embargo (administrative interval).
What is a DOI Number?
Some links on this page may take you to non-federal websites. Their policies may differ from this site.
-
Abstract -
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.
Free, publicly-accessible full text available February 1, 2025 -
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.Free, publicly-accessible full text available February 1, 2025 -
Abstract The predicted intensification of the North American Monsoon is expected to alter growing season rainfall patterns in the southwestern United States. These patterns, which have historically been characterized by frequent small rain events, are anticipated to shift towards a more extreme precipitation regime consisting of fewer, but larger rain events. Furthermore, human activities are contributing to increased atmospheric nitrogen deposition throughout this dryland region.
Alterations in rainfall size and frequency, along with changes in nitrogen availability, are likely to have significant consequences for above‐ground net primary production (ANPP) and plant community dynamics in drylands. The conceptual bucket model predicts that a shift towards fewer, but larger rain events could promote greater rates of ANPP in these regions by maintaining soil moisture availability above drought stress thresholds for longer periods during the growing season. However, only a few short‐term studies have tested this hypothesis, and none have explored the interaction between altered rainfall patterns and nitrogen enrichment.
To address this knowledge gap, we conducted a 14‐year rainfall addition and nitrogen fertilization experiment in a northern Chihuahuan Desert grassland to explore the long‐term impacts of changes in monsoon rainfall size and frequency, along with chronic nitrogen enrichment, on ANPP (measured as peak biomass) and plant community dynamics.
Contrary to bucket model predictions, small frequent rain events promoted comparable rates of ANPP to large infrequent rain events in the absence of nitrogen enrichment. It was only when nitrogen limitation was alleviated that large infrequent rain events resulted in the greatest ANPP. Furthermore, we found that nitrogen enrichment had the greatest impact on plant community composition under the small frequent rainfall regime.
Synthesis . Our long‐term field experiment highlights limitations of the bucket model by demonstrating that water and nitrogen availability sequentially limit dryland ecological processes. Specifically, our findings suggest that while water availability is the primary limiting factor for above‐ground net primary production in these ecosystems, nitrogen limitation becomes increasingly important when water is not limiting. Moreover, our findings reveal that small frequent rain events play an important but underappreciated role in driving dryland ecosystem dynamics. -
Abstract Drylands are key contributors to interannual variation in the terrestrial carbon sink, which has been attributed primarily to broad‐scale climatic anomalies that disproportionately affect net primary production (NPP) in these ecosystems. Current knowledge around the patterns and controls of NPP is based largely on measurements of aboveground net primary production (ANPP), particularly in the context of altered precipitation regimes. Limited evidence suggests belowground net primary production (BNPP), a major input to the terrestrial carbon pool, may respond differently than ANPP to precipitation, as well as other drivers of environmental change, such as nitrogen deposition and fire. Yet long‐term measurements of BNPP are rare, contributing to uncertainty in carbon cycle assessments. Here, we used 16 years of annual NPP measurements to investigate responses of ANPP and BNPP to several environmental change drivers across a grassland–shrubland transition zone in the northern Chihuahuan Desert. ANPP was positively correlated with annual precipitation across this landscape; however, this relationship was weaker within sites. BNPP, on the other hand, was weakly correlated with precipitation only in Chihuahuan Desert shrubland. Although NPP generally exhibited similar trends among sites, temporal correlations between ANPP and BNPP within sites were weak. We found chronic nitrogen enrichment stimulated ANPP, whereas a one‐time prescribed burn reduced ANPP for nearly a decade. Surprisingly, BNPP was largely unaffected by these factors. Together, our results suggest that BNPP is driven by a different set of controls than ANPP. Furthermore, our findings imply belowground production cannot be inferred from aboveground measurements in dryland ecosystems. Improving understanding around the patterns and controls of dryland NPP at interannual to decadal scales is fundamentally important because of their measurable impact on the global carbon cycle. This study underscores the need for more long‐term measurements of BNPP to improve assessments of the terrestrial carbon sink, particularly in the context of ongoing environmental change.
-
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 free
Plain Language Summary for this article on the Journal blog. -
Abstract The encroachment of woody plants into grasslands is a global phenomenon with implications for biodiversity and ecosystem function. Understanding and predicting the pace of expansion and the underlying processes that control it are key challenges in the study and management of woody encroachment. Theory from spatial population biology predicts that the occurrence and speed of expansion should depend sensitively on the nature of conspecific density dependence. If fitness is maximized at the low‐density encroachment edge, then shrub expansion should be “pulled” forward. However, encroaching shrubs have been shown to exhibit positive feedbacks, whereby shrub establishment modifies the environment in ways that facilitate further shrub recruitment and survival. In this case there may be a fitness cost to shrubs at low density causing expansion to be “pushed” from behind the leading edge. We studied the spatial dynamics of creosotebush (
Larrea tridentata ), which has a history of encroachment into Chihuahuan Desert grasslands over the past century. We used demographic data from observational censuses and seedling transplant experiments to test the strength and direction of density dependence in shrub fitness along a gradient of shrub density at the grass–shrub ecotone. We also used seed‐drop experiments and wind data to construct a mechanistic seed‐dispersal kernel, then connected demography and dispersal data within a spatial integral projection model (SIPM) to predict the dynamics of shrub expansion. Contrary to expectations based on potential for positive feedbacks, the shrub encroachment wave is “pulled” by maximum fitness at the low‐density front. However, the predicted pace of expansion was strikingly slow (ca. 8 cm/year), and this prediction was supported by independent resurveys of the ecotone showing little to no change in the spatial extent of shrub cover over 12 years. Encroachment speed was acutely sensitive to seedling recruitment, suggesting that this population may be primed for pulses of expansion under conditions that are favorable for recruitment. Our integration of observations, experiments, and modeling reveals not only that this ecotone is effectively stalled under current conditions but also why that is so and how that may change as the environment changes. -
Abstract Asexual reproduction plays a fundamental role in the structure, dynamics and persistence of perennial grasslands. Thus, assessing how asexual reproductive traits of plant communities respond to drought may be key for understanding grassland resistance to drought and recovery following drought.
Here, we quantified three asexual reproductive traits (i.e. above‐ground tiller abundance, below‐ground bud abundance and the ratio of tillers to buds) during a 4‐year severe drought and a 2‐year drought recovery period in four grasslands that spanned an aridity gradient in northern China. We also assessed the relationship between these traits and the resistance and recovery of above‐ground net primary productivity (ANPP).
We found that drought had limited and largely inconsistent effects on asexual reproduction among drought and recovery years and grasslands overall. Drought increased tiller abundance in the first treatment year and reduced bud banks by the fourth treatment year across grasslands. However, neither of the three asexual reproductive traits were correlated with drought resistance of ANPP. Drought legacies differed among the four grasslands with positive, negative and no legacies evident for the three asexual reproductive traits, and no clear relationship with aridity. Bud banks and tiller to bud ratio decreased and increased, respectively, in the first recovery year, but not in the second recovery year. In contrast to drought resistance, community bud abundance was strongly related to recovery, such that communities with higher bud abundance had greater ANPP recovery following drought.
Synthesis . These results suggest that asexual reproductive traits may be important drivers of ecosystem recovery after drought, but that variable responses of these asexual reproduction traits during drought complicates predictions of overall grassland responses. -
Abstract Grasslands are subject to climate change, such as severe drought, and an important aspect of their functioning is temporal stability in response to extreme climate events. Previous research has explored the impacts of extreme drought and post‐drought periods on grassland stability, yet the mechanistic pathways behind these changes have rarely been studied.
Here, we implemented an experiment with 4 years of drought and 3 years of recovery to assess the effects of drought and post‐drought on the temporal stability of above‐ground net primary productivity (ANPP) and its underlying mechanisms. To do so, we measured community‐weighted mean (CWM) of six plant growth and nine seed traits, functional diversity, population stability and species asynchrony across two cold, semiarid grasslands in northern China. We also performed piecewise structural equation models (SEMs) to assess the relationships between ANPP stability and its underlying mechanisms and how drought and post‐drought periods alter the relative contribution of these mechanisms to ANPP stability.
We found that temporal stability of ANPP was not reduced during drought due to grasses maintaining productivity, which compensated for increased variation of forb productivity. Moreover, ANPP recovered rapidly after drought, and both grasses and forbs contributed to community stability during the post‐drought period. Overall, ANPP stability decreased during the combined drought and post‐drought periods because of rapid changes in ANPP from drought to post‐drought. SEMs revealed that the temporal stability of ANPP during drought and post‐drought periods was modulated by functional diversity and community‐weighted mean traits directly and indirectly by altering species asynchrony and population stability. Specifically, the temporal stability of ANPP was positively correlated with functional divergence of plant communities. CWMs of seed traits (e.g. seed width and thickness), rather than plant growth traits (e.g. specific leaf area and leaf nutrient content), stabilized grassland ANPP. Productivity of plant communities with large and thick seeds was less sensitive to precipitation changes over time.
These results emphasize the importance of considering both the functional trait distribution among species and seed traits of dominant species since their combined effects can stabilize ecosystem functions under global climate change scenarios.
Read the free
Plain Language Summary for this article on the Journal blog. -
Abstract Future climates will alter the frequency and size of rain events in drylands, potentially affecting soil microbes that generate carbon feedbacks to climate, but field tests are rare. Topsoils in drylands are commonly colonized by biological soil crusts (biocrusts), photosynthesis‐based communities that provide services ranging from soil fertilization to stabilization against erosion. We quantified responses of biocrust microbial communities to 12 years of altered rainfall regimes, with 60 mm of additional rain per year delivered either as small (5 mm) weekly rains or large (20 mm) monthly rains during the summer monsoon season. Rain addition promoted microbial diversity, suppressed the dominant cyanobacterium,
Microcoleus vaginatus , and enhanced nitrogen‐fixing taxa, but did not consistently increase microbial biomass. The addition of many small rain events increased microbial biomass, whereas few, large events did not. These results alter the physiological paradigm that biocrusts are most limited by the amount of rainfall and instead predict that regimes enriched in small rain events will boost cyanobacterial biocrusts and enhance their beneficial services to drylands.