Small effects of livestock grazing intensification on diversity, abundance, and composition in a dryland plant community.

Livestock grazing is a globally important land use and has the potential to significantly influence plant community structure and ecosystem function, yet several critical knowledge gaps remain on the direction and magnitude of grazing impacts. Furthermore, much of our understanding of the long-term effects on plant community composition and structure are based on grazer exclusion experiments, which explicitly avoid characterizing effects along grazing intensity gradients. We sampled big sagebrush plant communities using 68 plots located along grazing intensity gradients to determine how grazing intensity influences multiple aspects of plant community structure over time. This was accomplished by sampling plant communities at different distances from 17 artificial watering sources, using distance from water and cow dung density as proxies for grazing intensity at individual plots. Total vegetation cover and total grass cover were negatively related to grazing intensity, and cover of annual forbs, exotic cover, and exotic richness were positively related to grazing intensity. In contrast, species richness and composition, bunchgrass biomass, shrub density and size, percentage cover of bare ground, litter, and biological soil crusts did not vary along our grazing intensity gradients, in spite of our expectations to the contrary. Our results suggest that the effects of livestock grazing over multiple decades (mean = 46 years) in our sites are relatively small, especially for native perennial species, and that the big sagebrush plant communities we sampled are somewhat resistant to livestock grazing. Collectively, our findings are consistent with existing evidence that indicates the stability of the big sagebrush plant functional type composition under current grazing management regimes.

Divergent climate change effects on widespread dryland plant communities driven by climatic and ecohydrological gradients.

Plant community response to climate change will be influenced by individual plant responses that emerge from competition for limiting resources that fluctuate through time and vary across space. Projecting these responses requires an approach that integrates environmental conditions and species interactions that result from future climatic variability. Dryland plant communities are being substantially affected by climate change because their structure and function are closely tied to precipitation and temperature, yet impacts vary substantially due to environmental heterogeneity, especially in topographically complex regions. Here, we quantified the effects of climate change on big sagebrush (Artemisia tridentata Nutt.) plant communities that span 76 million ha in the western United States. We used an individual-based plant simulation model that represents intra- and inter-specific competition for water availability, which is represented by a process-based soil water balance model. For dominant plant functional types, we quantified changes in biomass and characterized agreement among 52 future climate scenarios. We then used a multivariate matching algorithm to generate fine-scale interpolated surfaces of functional type biomass for our study area. Results suggest geographically divergent responses of big sagebrush to climate change (changes in biomass of -20% to +27%), declines in perennial C3 grass and perennial forb biomass in most sites, and widespread, consistent, and sometimes large increases in perennial C4 grasses. The largest declines in big sagebrush, perennial C3 grass and perennial forb biomass were simulated in warm, dry sites. In contrast, we simulated no change or increases in functional type biomass in cold, moist sites. There was high agreement among climate scenarios on climate change impacts to functional type biomass, except for big sagebrush. Collectively, these results suggest divergent responses to warming in moisture-limited versus temperature-limited sites and potential shifts in the relative importance of some of the dominant functional types that result from competition for limiting resources.

Robust ecological drought projections for drylands in the 21st century.

Dryland ecosystems may be especially vulnerable to expected 21st century increases in temperature and aridity because they are tightly controlled by moisture availability. However, climate impact assessments in drylands are difficult because ecological dynamics are dictated by drought conditions that are difficult to define and complex to estimate from climate conditions alone. In addition, precipitation projections vary substantially among climate models, enhancing variation in overall trajectories for aridity. Here, we constrain this uncertainty by utilizing an ecosystem water balance model to quantify drought conditions with recognized ecological importance, and by identifying changes in ecological drought conditions that are robust among climate models, defined here as when >90% of models agree in the direction of change. Despite limited evidence for robust changes in precipitation, changes in ecological drought are robust over large portions of drylands in the United States and Canada. Our results suggest strong regional differences in long-term drought trajectories, epitomized by chronic drought increases in southern areas, notably the Upper Gila Mountains and South-Central Semi-arid Prairies, and decreases in the north, particularly portions of the Temperate and West-Central Semi-arid Prairies. However, we also found that exposure to hot-dry stress is increasing faster than mean annual temperature over most of these drylands, and those increases are greatest in northern areas. Robust shifts in seasonal drought are most apparent during the cool season; when soil water availability is projected to increase in northern regions and decrease in southern regions. The implications of these robust drought trajectories for ecosystems will vary geographically, and these results provide useful insights about the impact of climate change on these dryland ecosystems. More broadly, this approach of identifying robust changes in ecological drought may be useful for other assessments of climate impacts in drylands and provide a more rigorous foundation for making long-term strategic resource management decisions.

Soil and stand structure explain shrub mortality patterns following global change-type drought and extreme precipitation.

The probability of extreme weather events is increasing, with the potential for widespread impacts to plants, plant communities, and ecosystems. Reports of drought-related tree mortality are becoming more frequent, and there is increasing evidence that drought accompanied by high temperatures is especially detrimental. Simultaneously, extreme large precipitation events have become more frequent over the past century. Water-limited ecosystems may be more vulnerable to these extreme events than other ecosystems, especially when pushed outside of their historical range of variability. However, drought-related mortality of shrubs-an important component of dryland vegetation-remains understudied relative to tree mortality. In 2014, a landscape-scale die-off of the widespread shrub, big sagebrush (Artemisia tridentata Nutt.), was reported in southwest Wyoming, following extreme hot and dry conditions in 2012 and extremely high precipitation in September of 2013. Here we examine how severe drought, extreme precipitation, soil texture and salinity, and shrub-stand characteristics contributed to this die-off event. At 98 plots within and around the die-off, we quantified big sagebrush mortality, characterized soil texture and salinity, and simulated soil-water conditions from 1916 to 2016 using an ecosystem water-balance model. We found that the extreme weather conditions alone did not explain patterns of big sagebrush mortality and did not result in extreme (historically unprecedented) soil-water conditions during the drought. Instead, plots with chronically dry soil conditions experienced greatest mortality following the global change-type (hot) drought in 2012. Furthermore, mortality was greater in locations with high potential run-on and low potential run-off where saturated soil conditions were simulated in September 2013, suggesting that extreme precipitation also played an important role in the die-off in these locations. In locations where drought alone contributed to mortality, stem density negatively impacted big sagebrush. In locations that may have been affected by both drought and saturation, however, mortality was greatest where stem density was lowest, suggesting that these locations may have already been less favorable to big sagebrush. Paradoxically, vulnerability to both extreme events (drought and saturation) was associated with finer-textured soils, and our results highlight the importance of soils in determining local variation of the vulnerability of dryland plants to extreme events.

Soil texture and precipitation seasonality influence plant community structure in North American temperate shrub steppe.

In drylands, the coexistence of grasses and woody plants has been attributed to soil-water resource partitioning. Soil texture and precipitation seasonality can influence the amount and distribution of water in the soil, and their interaction may play an important role in determining the relative importance of grasses and woody plants. We investigated the influence of this interaction on plant functional types across a broad range of precipitation regimes and soil textures in western North America by analyzing plant-cover data collected at 2,084 plots that included the widespread shrub big sagebrush (Artemisia tridentata Nutt.). We characterized how the significance of the inverse-texture effect varies across soil conditions by quantifying relationships between precipitation and foliar cover on finer- vs. coarser-textured soils across a range of potential texture divisions represented by sand content. We found evidence of the inverse-texture effect for every plant functional type (except for cheatgrass) that we examined with at least one component of precipitation (annual, warm, or cold season), and provide the first evidence for this effect in locations with cold-season-dominated precipitation regimes. The texture and precipitation combinations that exhibited the inverse-texture effect varied with plant functional type, presumably because of effects of soil texture on water availability at different soil depths with season. Furthermore, we found an inverse-texture effect that was remarkably similar for shrub cover with cold-season precipitation and grass cover with warm-season precipitation. These results provide new insight into how the inverse-texture effect interacts with precipitation seasonality to influence plant functional type composition in drylands, and further suggest that quantifying the soil-texture division at which the inverse-texture effect is relevant under a given set of environmental conditions may provide support for the effect across dryland plant communities.

Future soil moisture and temperature extremes imply expanding suitability for rainfed agriculture in temperate drylands.

The distribution of rainfed agriculture, which accounts for approximately ¾ of global croplands, is expected to respond to climate change and human population growth and these responses may be especially pronounced in water limited areas. Because the environmental conditions that support rainfed agriculture are determined by climate, weather, and soil conditions that affect overall and transient water availability, predicting this response has proven difficult, especially in temperate regions that support much of the world's agriculture. Here, we show that suitability to support rainfed agriculture in temperate dryland climates can be effectively represented by just two daily environmental variables: moist soils with warm conditions increase suitability while extreme high temperatures decrease suitability. 21st century projections based on daily ecohydrological modeling of downscaled climate forecasts indicate overall increases in the area suitable for rainfed agriculture in temperate dryland regions, especially at high latitudes. The regional exception to this trend was Europe, where suitability in temperate dryland portions will decline substantially. These results clarify how rising temperatures interact with other key drivers of moisture availability to determine the sustainability of rainfed agriculture and help policymakers, resource managers, and the agriculture industry anticipate shifts in areas suitable for rainfed cultivation.

Climate change reduces extent of temperate drylands and intensifies drought in deep soils.

Drylands cover 40% of the global terrestrial surface and provide important ecosystem services. While drylands as a whole are expected to increase in extent and aridity in coming decades, temperature and precipitation forecasts vary by latitude and geographic region suggesting different trajectories for tropical, subtropical, and temperate drylands. Uncertainty in the future of tropical and subtropical drylands is well constrained, whereas soil moisture and ecological droughts, which drive vegetation productivity and composition, remain poorly understood in temperate drylands. Here we show that, over the twenty first century, temperate drylands may contract by a third, primarily converting to subtropical drylands, and that deep soil layers could be increasingly dry during the growing season. These changes imply major shifts in vegetation and ecosystem service delivery. Our results illustrate the importance of appropriate drought measures and, as a global study that focuses on temperate drylands, highlight a distinct fate for these highly populated areas.

Climate change-induced vegetation shifts lead to more ecological droughts despite projected rainfall increases in many global temperate drylands.

Drylands occur worldwide and are particularly vulnerable to climate change because dryland ecosystems depend directly on soil water availability that may become increasingly limited as temperatures rise. Climate change will both directly impact soil water availability and change plant biomass, with resulting indirect feedbacks on soil moisture. Thus, the net impact of direct and indirect climate change effects on soil moisture requires better understanding. We used the ecohydrological simulation model SOILWAT at sites from temperate dryland ecosystems around the globe to disentangle the contributions of direct climate change effects and of additional indirect, climate change-induced changes in vegetation on soil water availability. We simulated current and future climate conditions projected by 16 GCMs under RCP 4.5 and RCP 8.5 for the end of the century. We determined shifts in water availability due to climate change alone and due to combined changes of climate and the growth form and biomass of vegetation. Vegetation change will mostly exacerbate low soil water availability in regions already expected to suffer from negative direct impacts of climate change (with the two RCP scenarios giving us qualitatively similar effects). By contrast, in regions that will likely experience increased water availability due to climate change alone, vegetation changes will counteract these increases due to increased water losses by interception. In only a small minority of locations, climate change-induced vegetation changes may lead to a net increase in water availability. These results suggest that changes in vegetation in response to climate change may exacerbate drought conditions and may dampen the effects of increased precipitation, that is, leading to more ecological droughts despite higher precipitation in some regions. Our results underscore the value of considering indirect effects of climate change on vegetation when assessing future soil moisture conditions in water-limited ecosystems.

Mid-latitude shrub steppe plant communities: climate change consequences for soil water resources.

In the coming century, climate change is projected to impact precipitation and temperature regimes worldwide, with especially large effects in drylands. We use big sagebrush ecosystems as a model dryland ecosystem to explore the impacts of altered climate on ecohydrology and the implications of those changes for big sagebrush plant communities using output from 10 Global Circulation Models (GCMs) for two representative concentration pathways (RCPs). We ask: (1) What is the magnitude of variability in future temperature and precipitation regimes among GCMs and RCPs for big sagebrush ecosystems, and (2) How will altered climate and uncertainty in climate forecasts influence key aspects of big sagebrush water balance? We explored these questions across 1980-2010, 2030-2060, and 2070-2100 to determine how changes in water balance might develop through the 21st century. We assessed ecohydrological variables at 898 sagebrush sites across the western US using a process-based soil water model, SOILWAT, to model all components of daily water balance using site-specific vegetation parameters and site-specific soil properties for multiple soil layers. Our modeling approach allowed for changes in vegetation based on climate. Temperature increased across all GCMs and RCPs, whereas changes in precipitation were more variable across GCMs. Winter and spring precipitation was predicted to increase in the future (7% by 2030-2060, 12% by 2070-2100), resulting in slight increases in soil water potential (SWP) in winter. Despite wetter winter soil conditions, SWP decreased in late spring and summer due to increased evapotranspiration (6% by 2030-2060, 10% by 2070-2100) and groundwater recharge (26% and 30% increase by 2030-2060 and 2070-2100). Thus, despite increased precipitation in the cold season, soils may dry out earlier in the year, resulting in potentially longer, drier summer conditions. If winter precipitation cannot offset drier summer conditions in the future, we expect big sagebrush regeneration and survival will be negatively impacted, potentially resulting in shifts in the relative abundance of big sagebrush plant functional groups. Our results also highlight the importance of assessing multiple GCMs to understand the range of climate change outcomes on ecohydrology, which was contingent on the GCM chosen.

Scale dependence of disease impacts on quaking aspen (Populus tremuloides) mortality in the southwestern United States.

Depending on how disease impacts tree exposure to risk, both the prevalence of disease and disease effects on survival may contribute to patterns of mortality risk across a species' range. Disease may accelerate tree species' declines in response to global change factors, such as drought, biotic interactions, such as competition, or functional traits, such as allometry. To assess the role of disease in mediating mortality risk in quaking aspen (Populus tremuloides), we developed hierarchical Bayesian models for both disease prevalence in live aspen stems and the resulting survival rates of healthy and diseased aspen near the species' southern range limit using 5088 individual trees on 281 United States Forest Service Forest Inventory and Analysis plots in the southwestern United States. We found that disease prevalence depended primarily on tree size, tree allometry, and spatial variation in precipitation, while mortality depended on tree size, allometry, competition, spatial variation in summer temperature, and both temporal and spatial variation in summer precipitation. Disease prevalence was highest in large trees with low slenderness found on dry sites. For healthy trees, mortality decreased with diameter, slenderness, and temporal variation in summer precipitation, but increased with competition and spatial variation in summer temperature. Mortality of diseased trees decreased with diameter and aspen relative basal area and increased with mean summer temperature and precipitation. Disease infection increased aspen mortality, especially in trees of intermediate size and trees on plots at climatic extremes (i.e., cool, wet and warm, dry climates). By examining variation in disease prevalence, mortality of healthy trees, and mortality of diseased trees, we showed that the role of disease in aspen tree mortality depended on the scale of inference. For variation among individuals in diameter, disease tended to expose intermediate-size trees experiencing moderate risk to greater risk. For spatial variation in summer temperature, disease exposed lower risk populations to greater mortality probabilities, but the magnitude of this exposure depended on summer precipitation. Furthermore, the importance of diameter and slenderness in mediating responses to climate supports the increasing emphasis on trait variation in studies of ecological responses to global change.

Forest stand structure, productivity, and age mediate climatic effects on aspen decline.

Because forest stand structure, age, and productivity can mediate the impacts of climate on quaking aspen (Populus tremuloides) mortality, ignoring stand-scale factors limits inference on the drivers of recent sudden aspen decline. Using the proportion of aspen trees that were dead as an index of recent mortality at 841 forest inventory plots, we examined the relationship of this mortality index to forest structure and climate in the Rocky Mountains and Intermountain Western United States. We found that forest structure explained most of the patterns in mortality indices, but that variation in growing-season vapor pressure deficit and winter precipitation over the last 20 years was important. Mortality index sensitivity to precipitation was highest in forests where aspen exhibited high densities, relative basal areas, quadratic mean diameters, and productivities, whereas sensitivity to vapor pressure deficit was highest in young forest stands. These results indicate that the effects of drought on mortality may be mediated by forest stand development, competition with encroaching conifers, and physiological vulnerabilities of large trees to drought. By examining mortality index responses to both forest structure and climate, we show that forest succession cannot be ignored in studies attempting to understand the causes and consequences of sudden aspen decline.

Community responses of arthropods to a range of traditional and manipulated grazing in shortgrass steppe.

Responses of plants to grazing are better understood, and more predictable, than those of consumers in North American grasslands. In 2003, we began a large-scale, replicated experiment that examined the effects of grazing on three important arthropod groups-beetles, spiders, and grasshoppers-in shortgrass steppe of north-central Colorado. We investigated whether modifications of the intensity and seasonality of livestock grazing alter the structure and diversity of macroarthropod communities compared with traditional grazing practices. Treatments represented a gradient of grazing intensity by cattle and native herbivores: long-term grazing exclosures; moderate summer grazing (the traditional regime); intensive spring grazing; intensive summer grazing; and moderately summer-grazed pastures also inhabited by black-tailed prairie dogs (Cynomys ludovicianus Ord). Beetles and spiders were the most common groups captured, comprising 60% and 21%, respectively, of 4,378 total pitfall captures. Grasshopper counts were generally low, with 3,799 individuals observed and densities <4 m(-2). Two years after treatments were applied, vegetation structure differed among grazing treatments, responding not only to long-term grazing conditions, but also to the short-term, more-intensive grazing manipulations. In response, arthropods were, in general, relatively insensitive to these grazing-induced structural changes. However, species-level analyses of one group (Tenebrionidae) revealed both positive and negative effects of grazing treatments on beetle richness and activity-density. Importantly, these responses to grazing were more pronounced in a year when spring-summer rainfall was low, suggesting that both grazing and precipitation-which together may create the greatest heterogeneity in vegetation structure-are drivers of consumer responses in this system.

Mountain landscapes offer few opportunities for high-elevation tree species migration.

Climate change is anticipated to alter plant species distributions. Regional context, notably the spatial complexity of climatic gradients, may influence species migration potential. While high-elevation species may benefit from steep climate gradients in mountain regions, their persistence may be threatened by limited suitable habitat as land area decreases with elevation. To untangle these apparently contradictory predictions for mountainous regions, we evaluated the climatic suitability of four coniferous forest tree species of the western United States based on species distribution modeling (SDM) and examined changes in climatically suitable areas under predicted climate change. We used forest structural information relating to tree species dominance, productivity, and demography from an extensive forest inventory system to assess the strength of inferences made with a SDM approach. We found that tree species dominance, productivity, and recruitment were highest where climatic suitability (i.e., probability of species occurrence under certain climate conditions) was high, supporting the use of predicted climatic suitability in examining species risk to climate change. By predicting changes in climatic suitability over the next century, we found that climatic suitability will likely decline, both in areas currently occupied by each tree species and in nearby unoccupied areas to which species might migrate in the future. These trends were most dramatic for high elevation species. Climatic changes predicted over the next century will dramatically reduce climatically suitable areas for high-elevation tree species while a lower elevation species, Pinus ponderosa, will be well positioned to shift upslope across the region. Reductions in suitable area for high-elevation species imply that even unlimited migration would be insufficient to offset predicted habitat loss, underscoring the vulnerability of these high-elevation species to climatic changes.

Dominant species, rather than diversity, regulates temporal stability of plant communities.

A growing body of empirical evidence suggests that the temporal stability of communities typically increases with diversity. The counterview to this is that dominant species, rather than diversity itself, might regulate temporal stability. However, empirical studies that have explicitly examined the relative importance of diversity and dominant species in maintaining community stability have yielded few clear-cut patterns. Here, using a long-term data set, we examined the relative importance of changes in diversity components and dominance hierarchy following the removal of a dominant C4 grass, Bouteloua gracilis, in stabilizing plant communities. We also examined the relationships between the variables of diversity and dominance hierarchy and the statistical components of temporal stability. We found a significant negative relationship between temporal stability and species richness, number of rare species, and relative abundance of rare species, whereas a significant positive relationship existed between temporal stability and relative abundance of the dominant species. Variances and covariances summed over all species significantly increased with increasing species richness, whereas they significantly decreased with increasing relative abundance of dominant species. We showed that temporal stability in a shortgrass steppe plant community was controlled by dominant species rather than by diversity itself. The generality of diversity-stability relationships might be restricted by the dynamics of dominant species, especially when they have characteristics that contribute to stability in highly stochastic systems. A clear implication is that dominance hierarchies and their changes might be among the most important ecological components to consider in managing communities to maintain ecosystem functioning.

Conservation of nitrogen increases with precipitation across a major grassland gradient in the Central Great Plains of North America.

Regional analyses and biogeochemical models predict that ecosystem N pools and N cycling rates must increase from the semi-arid shortgrass steppe to the sub-humid tallgrass prairie of the Central Great Plains, yet few field data exist to evaluate these predictions. In this paper, we measured rates of net N mineralization, N in above- and belowground primary production, total soil organic matter N pools, soil inorganic N pools and capture in resin bags, decomposition rates, foliar (15)N, and N use efficiency (NUE) across a precipitation gradient. We found that net N mineralization did not increase across the gradient, despite more N generally being found in plant production, suggesting higher N uptake, in the wetter areas. NUE of plants increased with precipitation, and delta(15)N foliar values and resin-captured N in soils decreased, all of which are consistent with the hypothesis that N cycling is tighter at the wet end of the gradient. Litter decomposition appeared to play a role in maintaining this regional N cycling trend: litter decomposed more slowly and released less N at the wet end of the gradient. These results suggest that immobilization of N within the plant-soil system increases from semi-arid shortgrass steppe to sub-humid tallgrass prairie. Despite the fact that N pools increase along a bio-climatic gradient from shortgrass steppe to mixed grass and tallgrass prairie, this element becomes relatively more limiting and is therefore more tightly conserved at the wettest end of the gradient. Similar to findings from forested systems, our results suggest that grassland N cycling becomes more open to N loss with increasing aridity.

A multi-scale perspective of water pulses in dryland ecosystems: climatology and ecohydrology of the western USA.

In dryland ecosystems, the timing and magnitude of precipitation pulses drive many key ecological processes, notably soil water availability for plants and soil microbiota. Plant available water has frequently been viewed simply as incoming precipitation, yet processes at larger scales drive precipitation pulses, and the subsequent transformation of precipitation pulses to plant available water are complex. We provide an overview of the factors that influence the spatial and temporal availability of water to plants and soil biota using examples from western USA drylands. Large spatial- and temporal-scale drivers of regional precipitation patterns include the position of the jet streams and frontal boundaries, the North American Monsoon, El Niño Southern Oscillation events, and the Pacific Decadal Oscillation. Topography and orography modify the patterns set up by the larger-scale drivers, resulting in regional patterns (10(2)-10(6) km2) of precipitation magnitude, timing, and variation. Together, the large-scale and regional drivers impose important pulsed patterns on long-term precipitation trends at landscape scales, in which most site precipitation is received as small events (< 5 mm) and with most of the intervals between events being short (< 10 days). The drivers also influence the translation of precipitation events into available water via linkages between soil water content and components of the water budget, including interception, infiltration and runoff, soil evaporation, plant water use and hydraulic redistribution, and seepage below the rooting zone. Soil water content varies not only vertically with depth but also horizontally beneath versus between plants and/or soil crusts in ways that are ecologically important to different plant and crust types. We highlight the importance of considering larger-scale drivers, and their effects on regional patterns; small, frequent precipitation events; and spatio-temporal heterogeneity in soil water content in translating from climatology to precipitation pulses to the dryland ecohydrology of water availability for plants and soil biota.