Pastures and Climate Extremes: Impacts of Cool Season Warming and Drought on the Productivity of Key Pasture Species in a Field Experiment.

Shifts in the timing, intensity and/or frequency of climate extremes, such as severe drought and heatwaves, can generate sustained shifts in ecosystem function with important ecological and economic impacts for rangelands and managed pastures. The Pastures and Climate Extremes experiment (PACE) in Southeast Australia was designed to investigate the impacts of a severe winter/spring drought (60% rainfall reduction) and, for a subset of species, a factorial combination of drought and elevated temperature (ambient +3°C) on pasture productivity. The experiment included nine common pasture and Australian rangeland species from three plant functional groups (C3 grasses, C4 grasses and legumes) planted in monoculture. Winter/spring drought resulted in productivity declines of 45% on average and up to 74% for the most affected species (Digitaria eriantha) during the 6-month treatment period, with eight of the nine species exhibiting significant yield reductions. Despite considerable variation in species' sensitivity to drought, C4 grasses were more strongly affected by this treatment than C3 grasses or legumes. Warming also had negative effects on cool-season productivity, associated at least partially with exceedance of optimum growth temperatures in spring and indirect effects on soil water content. The combination of winter/spring drought and year-round warming resulted in the greatest yield reductions. We identified responses that were either additive (Festuca), or less-than-additive (Medicago), where warming reduced the magnitude of drought effects. Results from this study highlight the sensitivity of diverse pasture species to increases in winter and spring drought severity similar to those predicted for this region, and that anticipated benefits of cool-season warming are unlikely to be realized. Overall, the substantial negative impacts on productivity suggest that future, warmer, drier climates will result in shortfalls in cool-season forage availability, with profound implications for the livestock industry and natural grazer communities.

High safety margins to drought-induced hydraulic failure found in five pasture grasses.

Determining the relationship between reductions in stomatal conductance (gs ) and leaf water transport during dehydration is key to understanding plant drought responses. While numerous studies have analysed the hydraulic function of woody species, minimal research has been conducted on grasses. Here, we sought to characterize hydraulic vulnerability in five widely-occurring pasture grasses (including both C3 and C4 grasses) and determine whether reductions in gs and leaf hydraulic conductance (Kleaf ) during dehydration could be attributed to xylem embolism. Using the optical vulnerability (OV) technique, we found that all species were highly resistant to xylem embolism when compared to other herbaceous angiosperms, with 50% xylem embolism (PX50 ) occurring at xylem pressures ranging from -4.4 to -6.1 MPa. We observed similar reductions in gs and Kleaf under mild water stress for all species, occurring well before PX50 . The onset of xylem embolism (PX12 ) occurred consistently after stomatal closure and 90% reduction of Kleaf . Our results suggest that factors other than xylem embolism are responsible for the majority of reductions in gs and Kleaf during drought and reductions in the productivity of pasture species under moderate drought may not be driven by embolism.

Semiarid grasslands and extreme precipitation events: do experimental results scale to the landscape?

The frequency and magnitude of deluges (extremely large rain events) are increasing globally as the atmosphere warms. Small-scale experiments suggest that semiarid grasslands are particularly sensitive to both the timing and size of deluge events. However, the assumption that plot-scale results can be extrapolated across landscapes with variable soil textures, plant communities, and grazing regimes has seldom been tested, despite being key to forecasting regional consequences of precipitation extremes. We used precipitation data from an extensive rain gauge network to identify natural deluges (mean size = 60 ± 31 mm, 1984-2012) that occurred across a ˜60-km2 heterogeneous native shortgrass steppe landscape in Colorado. We then related spatial variation in deluge precipitation to postdeluge responses in canopy greenness (normalized difference vegetation index, NDVI) via satellite imagery. Consistent with results from experiments, this semiarid grassland was most sensitive to mid-growing-season deluges, and postdeluge canopy greenness usually increased linearly (67% of the time) with increasing deluge size. This suggests that aboveground productivity in these semiarid systems will likely increase, rather than asymptote, with forecasted increases in deluge size. Importantly, differences in grazing regime did not significantly alter deluge responses, indicating that these patterns are robust to this widespread management practice.

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.

How big is big enough? Surprising responses of a semiarid grassland to increasing deluge size.

Climate change has intensified the hydrologic cycle globally, increasing the magnitude and frequency of large precipitation events, or deluges. Dryland ecosystems are expected to be particularly responsive to increases in deluge size, as their ecological processes are largely dependent on distinct soil moisture pulses. To better understand how increasing deluge size will affect ecosystem function, we conducted a field experiment in a native semiarid shortgrass steppe (Colorado, USA). We quantified ecological responses to a range of deluge sizes, from moderate to extreme, with the goal of identifying response patterns and thresholds beyond which ecological processes would not increase further (saturate). Using a replicated regression approach, we imposed single deluges that ranged in size from 20 to 120 mm (82.3rd to >99.9th percentile of historical event size) on undisturbed grassland plots. We quantified pre- and postdeluge responses in soil moisture, soil respiration, and canopy greenness, as well as leaf water potential, growth, and flowering of the dominant grass species (Bouteloua gracilis). We also measured end of season above- and belowground net primary production (ANPP, BNPP). As expected, this water-limited ecosystem responded strongly to the applied deluges, but surprisingly, most variables increased linearly with deluge size. We found little evidence for response thresholds within the range of deluge sizes imposed, at least during this dry year. Instead, response patterns reflected the linear increase in the duration of elevated soil moisture (2-22 days) with increasing event size. Flowering of B. gracilis and soil respiration responded particularly strongly to deluge size (14- and 4-fold increases, respectively), as did ANPP and BNPP (~60% increase for both). Overall, our results suggest that this semiarid grassland will respond positively and linearly to predicted increases in deluge size, and that event sizes may need to exceed historical magnitudes, or occur during wet years, before responses saturate.

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.

Plant growth and aboveground production respond differently to late-season deluges in a semi-arid grassland.

Semi-arid ecosystems are strongly water-limited and typically quite responsive to changes in precipitation amount and event size. In the C4-dominated shortgrass steppe of the Central US, previous experiments suggest that large rain events more effectively stimulate plant growth and aboveground net primary production (ANPP) than an equal amount of precipitation from smaller events. Responses to naturally occurring large events have generally been consistent with experimental results, with the exception of large events occurring later in the growing season (e.g., August). These have been reported as less effective at increasing net C uptake, despite temperatures optimal for C4 plant growth. Since atmospheric warming is increasing the frequency of statistically extreme rain events (deluges) throughout the growing season, how late-season deluges affect semi-arid ecosystems remains to be resolved. We applied deluges in August of three sizes (1.0-2.5 times average August precipitation) to assess the potential for late-season deluges to stimulate plant growth and ANPP. These late-season deluges led to significant "green-up" of this grassland, with new leaf production, and an increase in flowering of the dominant grass species. Further, these responses increased as deluge size increased, suggesting that larger or multiple deluges may lead to even greater growth responses. However, despite strong plant-level responses, no increase in ANPP was measured. Our results confirm that aboveground plant growth in the C4-dominated shortgrass steppe does respond to late-season deluges; however, if there is an increase in plant biomass, net accumulation aboveground is minimal at this time of year.

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.