Traits that distinguish dominant species across aridity gradients differ from those that respond to soil moisture.

Many plant traits respond to changes in water availability and might be useful for understanding ecosystem properties such as net primary production (NPP). This is especially evident in grasslands where NPP is water-limited and primarily determined by the traits of dominant species. We measured root and shoot morphology, leaf hydraulic traits, and NPP of four dominant North American prairie grasses in response to four levels of soil moisture in a greenhouse experiment. We expected that traits of species from drier regions would be more responsive to reduced water availability and that this would make these species more resistant to low soil moisture than species from wetter regions. All four species grew taller, produced more biomass, and increased total root length in wetter treatments. Each species reduced its leaf turgor loss point (TLP) in drier conditions, but only two species (one xeric, one mesic) maintained leaf water potential above TLP. We identified a suite of traits that clearly distinguished species from one another, but, surprisingly, these traits were relatively unresponsive to reduced soil moisture. Specifically, more xeric species produced thinner roots with higher specific root length and had a lower root mass fraction. This suggest that root traits are critical for distinguishing species from one another but might not respond strongly to changing water availability, though this warrants further investigation in the field. Overall, we found that NPP of these dominant grass species responded similarly to varying levels of soil moisture despite differences in species morphology, physiology, and habitat of origin.

Rapid evolution of a coastal marsh ecosystem engineer in response to global change.

There is increasing evidence that global change can alter ecosystems by eliciting rapid evolution of foundational plants capable of shaping vital attributes and processes. Here we describe results of a field-scale exposure experiment and multilocus assays illustrating that elevated CO2 (eCO2) and nitrogen (N) enrichment can result in rapid shifts in genetic and genotypic variation in Phragmites australis, an ecologically dominant plant that acts as an ecosystem engineer in coastal marshes worldwide. Compared to control treatments, genotypic diversity declined over three years of exposure, especially to N enrichment. The magnitude of loss also increased over time under conditions of N enrichment. Comparisons of genotype frequencies revealed that proportional abundances shifted with exposure to eCO2 and N in a manner consistent with expected responses to selection. Comparisons also revealed evidence of tradeoffs that constrained exposure responses, where any particular genotype responded favorably to one factor rather than to different factors or to combinations of factors. These findings challenge the prevailing view that plant-mediated ecosystem outcomes of global change are governed primarily by differences in species responses to shifting environmental pressures and highlight the value of accounting for organismal evolution in predictive models to improve forecasts of ecosystem responses to global change.

Rising ecosystem water demand exacerbates the lengthening of tropical dry seasons.

Precipitation-based assessments show a lengthening of tropical dry seasons under climate change, without considering simultaneous changes in ecosystem water demand. Here, we compare changes in tropical dry season length and timing when dry season is defined as the period when precipitation is less than: its climatological average, potential evapotranspiration, or actual evapotranspiration. While all definitions show more widespread tropical drying than wetting for 1983-2016, we find the largest fraction (48.7%) of tropical land probably experiencing longer dry seasons when dry season is defined as the period when precipitation cannot meet the need of actual evapotranspiration. Southern Amazonia (due to delayed end) and central Africa (due to earlier onset and delayed end) are hotspots of dry season lengthening, with greater certainty when accounting for water demand changes. Therefore, it is necessary to account for changing water demand when characterizing changes in tropical dry periods and ecosystem water deficits.

Compound hydroclimatic extremes in a semi-arid grassland: Drought, deluge, and the carbon cycle.

Climate change is predicted to increase the frequency and intensity of extreme events including droughts and large precipitation events or "deluges." While many studies have focused on the ecological impacts of individual events (e.g., a heat wave), there is growing recognition that when extreme events co-occur as compound extremes, (e.g., a heatwave during a drought), the additive effects on ecosystems are often greater than either extreme alone. In this study, we assessed a unique type of extreme-a contrasting compound extreme-where the extremes may have offsetting, rather than additive ecological effects, by examining how a deluge during a drought impacts productivity and carbon cycling in a semi-arid grassland. The experiment consisted of four treatments: a control (average precipitation), an extreme drought (<5th percentile), an extreme drought interrupted by a single deluge (>95th percentile), or an extreme drought interrupted by the equivalent amount of precipitation added in several smaller events. We highlight three key results. First, extreme drought resulted in early senescence, reduced carbon uptake, and a decline in net primary productivity relative to the control treatment. Second, the deluge imposed during extreme drought stimulated carbon fluxes and plant growth well above the levels of both the control and the drought treatment with several additional smaller rainfall events, emphasizing the importance of precipitation amount, event size, and timing. Third, while the deluge's positive effects on carbon fluxes and plant growth persisted for 1 month, the deluge did not completely offset the negative effects of extreme drought on end-of-season productivity. Thus, in the case of these contrasting hydroclimatic extremes, a deluge during a drought can stimulate temporally dynamic ecosystem processes (e.g., net ecosystem exchange) while only partially compensating for reductions in ecosystem functions over longer time scales (e.g., aboveground net primary productivity).

Is a drought a drought in grasslands? Productivity responses to different types of drought.

Drought, defined as a marked deficiency of precipitation relative to normal, occurs as periods of below-average precipitation or complete failure of precipitation inputs, and can be limited to a single season or prolonged over multiple years. Grasslands are typically quite sensitive to drought, but there can be substantial variability in the magnitude of loss of ecosystem function. We hypothesized that differences in how drought occurs may contribute to this variability. In four native Great Plains grasslands (three C4- and one C3-dominated) spanning a ~ 500-mm precipitation gradient, we imposed drought for four consecutive years by (1) reducing each rainfall event by 66% during the growing season (chronic drought) or (2) completely excluding rainfall during a shorter portion of the growing season (intense drought). The drought treatments were similar in magnitude but differed in the following characteristics: event number, event size and length of dry periods. We observed consistent drought-induced reductions (28-37%) in aboveground net primary production (ANPP) only in the C4-dominated grasslands. In general, intense drought reduced ANPP more than chronic drought, with little evidence that drought duration altered this pattern. Conversely, belowground net primary production (BNPP) was reduced by drought in all grasslands (32-64%), with BNPP reductions greater in intense vs. chronic drought treatments in the most mesic grassland. We conclude that grassland productivity responses to drought did not strongly differ between these two types of drought, but when differences existed, intense drought consistently reduced function more than chronic drought.

Resolving the Dust Bowl paradox of grassland responses to extreme drought.

During the 1930s Dust Bowl drought in the central United States, species with the C3 photosynthetic pathway expanded throughout C4-dominated grasslands. This widespread increase in C3 grasses during a decade of low rainfall and high temperatures is inconsistent with well-known traits of C3 vs. C4 pathways. Indeed, water use efficiency is generally lower, and photosynthesis is more sensitive to high temperatures in C3 than C4 species, consistent with the predominant distribution of C3 grasslands in cooler environments and at higher latitudes globally. We experimentally imposed extreme drought for 4 y in mixed C3/C4 grasslands in Kansas and Wyoming and, similar to Dust Bowl observations, also documented three- to fivefold increases in C3/C4 biomass ratios. To explain these paradoxical responses, we first analyzed long-term climate records to show that under nominal conditions in the central United States, C4 grasses dominate where precipitation and air temperature are strongly related (warmest months are wettest months). In contrast, C3 grasses flourish where precipitation inputs are less strongly coupled to warm temperatures. We then show that during extreme drought years, precipitation-temperature relationships weaken, and the proportion of precipitation falling during cooler months increases. This shift in precipitation seasonality provides a mechanism for C3 grasses to respond positively to multiyear drought, resolving the Dust Bowl paradox. Grasslands are globally important biomes and increasingly vulnerable to direct effects of climate extremes. Our findings highlight how extreme drought can indirectly alter precipitation seasonality and shift ecosystem phenology, affecting function in ways not predictable from key traits of C3 and C4 species.

Precipitation amount and event size interact to reduce ecosystem functioning during dry years in a mesic grassland.

Ongoing intensification of the hydrological cycle is altering rainfall regimes by increasing the frequency of extreme wet and dry years and the size of individual rainfall events. Despite long-standing recognition of the importance of precipitation amount and variability for most terrestrial ecosystem processes, we lack understanding of their interactive effects on ecosystem functioning. We quantified this interaction in native grassland by experimentally eliminating temporal variability in growing season rainfall over a wide range of precipitation amounts, from extreme wet to dry conditions. We contrasted the rain use efficiency (RUE) of above-ground net primary productivity (ANPP) under conditions of experimentally reduced versus naturally high rainfall variability using a 32-year precipitation-ANPP dataset from the same site as our experiment. We found that increased growing season rainfall variability can reduce RUE and thus ecosystem functioning by as much as 42% during dry years, but that such impacts weaken as years become wetter. During low precipitation years, RUE is lowest when rainfall event sizes are relatively large, and when a larger proportion of total rainfall is derived from large events. Thus, a shift towards precipitation regimes dominated by fewer but larger rainfall events, already documented over much of the globe, can be expected to reduce the functioning of mesic ecosystems primarily during drought, when ecosystem processes are already compromised by low water availability.

How ecologists define drought, and why we should do better.

Drought, widely studied as an important driver of ecosystem dynamics, is predicted to increase in frequency and severity globally. To study drought, ecologists must define or at least operationalize what constitutes a drought. How this is accomplished in practice is unclear, particularly given that climatologists have long struggled to agree on definitions of drought, beyond general variants of "an abnormal deficiency of water." We conducted a literature review of ecological drought studies (564 papers) to assess how ecologists describe and study drought. We found that ecologists characterize drought in a wide variety of ways (reduced precipitation, low soil moisture, reduced streamflow, etc.), but relatively few publications (~32%) explicitly define what are, and are not, drought conditions. More troubling, a surprising number of papers (~30%) simply equated "dry conditions" with "drought" and provided little characterization of the drought conditions studied. For a subset of these, we calculated Standardized Precipitation Evapotranspiration Index values for the reported drought periods. We found that while almost 90% of the studies were conducted under conditions quantifiable as slightly to extremely drier than average, ~50% were within the range of normal climatic variability. We conclude that the current state of the ecological drought literature hinders synthesis and our ability to draw broad ecological inferences because drought is often declared but is not explicitly defined or well characterized. We suggest that future drought publications provide at least one of the following: (a) the climatic context of the drought period based on long-term records; (b) standardized climatic index values; (c) published metrics from drought-monitoring organizations; (d) a quantitative definition of what the authors consider to be drought conditions for their system. With more detailed and consistent quantification of drought conditions, comparisons among studies can be more rigorous, increasing our understanding of the ecological effects of drought.

A reality check for climate change experiments: Do they reflect the real world?

Experiments are widely used in ecology, particularly for assessing global change impacts on ecosystem function. However, results from experiments often are inconsistent with observations made under natural conditions, suggesting the need for rigorous comparisons of experimental and observational studies. We conducted such a "reality check" for a grassland ecosystem by compiling results from nine independently conducted climate change experiments. Each experiment manipulated growing season precipitation (GSP) and measured responses in aboveground net primary production (ANPP). We compared results from experiments with long-term (33-yr) annual precipitation and ANPP records to ask if collectively (n = 44 experiment-years) experiments yielded estimates of ANPP, rain-use efficiency (RUE, grams per square meter ANPP per mm precipitation), and the relationship between GSP and ANPP comparable to observations. We found that mean ANPP and RUE from experiments did not deviate from observations. Experiments and observational data also yielded similar functional relationships between ANPP and GSP, but only within the range of historically observed GSP. Fewer experiments imposed extreme levels of GSP (outside the observed 33-yr record), but when these were included, they altered the GSP-ANPP relationship. This result underscores the need for more experiments imposing extreme precipitation levels to resolve how forecast changes in climate regimes will affect ecosystem function in the future.

Pushing precipitation to the extremes in distributed experiments: recommendations for simulating wet and dry years.

Intensification of the global hydrological cycle, ranging from larger individual precipitation events to more extreme multiyear droughts, has the potential to cause widespread alterations in ecosystem structure and function. With evidence that the incidence of extreme precipitation years (defined statistically from historical precipitation records) is increasing, there is a clear need to identify ecosystems that are most vulnerable to these changes and understand why some ecosystems are more sensitive to extremes than others. To date, opportunistic studies of naturally occurring extreme precipitation years, combined with results from a relatively small number of experiments, have provided limited mechanistic understanding of differences in ecosystem sensitivity, suggesting that new approaches are needed. Coordinated distributed experiments (CDEs) arrayed across multiple ecosystem types and focused on water can enhance our understanding of differential ecosystem sensitivity to precipitation extremes, but there are many design challenges to overcome (e.g., cost, comparability, standardization). Here, we evaluate contemporary experimental approaches for manipulating precipitation under field conditions to inform the design of 'Drought-Net', a relatively low-cost CDE that simulates extreme precipitation years. A common method for imposing both dry and wet years is to alter each ambient precipitation event. We endorse this approach for imposing extreme precipitation years because it simultaneously alters other precipitation characteristics (i.e., event size) consistent with natural precipitation patterns. However, we do not advocate applying identical treatment levels at all sites - a common approach to standardization in CDEs. This is because precipitation variability varies >fivefold globally resulting in a wide range of ecosystem-specific thresholds for defining extreme precipitation years. For CDEs focused on precipitation extremes, treatments should be based on each site's past climatic characteristics. This approach, though not often used by ecologists, allows ecological responses to be directly compared across disparate ecosystems and climates, facilitating process-level understanding of ecosystem sensitivity to precipitation extremes.