Dryland soil recovery after disturbance across soil and climate gradients of the Colorado Plateau.

Drylands impacted by energy development often require costly reclamation activities to reconstruct damaged soils and vegetation, yet little is known about the effectiveness of reclamation practices in promoting recovery of soil quality due to a lack of long-term and cross-site studies. Here, we examined paired on-pad and adjacent undisturbed off-pad soil properties over a 22-year chronosequence of 91 reclaimed oil or gas well pads across soil and climate gradients of the Colorado Plateau in the southwestern United States. Our goals were to estimate the time required for soil properties to reach undisturbed conditions, examine the multivariate nature of soil quality following reclamation, and identify environmental factors that affect reclamation outcomes. Soil samples, collected in 2020 and 2021, were analyzed for biogeochemical pools (total nitrogen, and total organic and inorganic carbon), chemical characteristics (salinity, sodicity, pH), and texture. Predicted time to recovery across all sites was 29 years for biogeochemical soil properties, 31 years for soil chemical properties, and 6 years for soil texture. Ordination of soil properties revealed differences between on- and off-pad soils, while site aridity explained variability in on-pad recovery. The predicted time to total soil recovery (distance between on- and off-pad in ordination space) was 96 years, which was longer than any individual soil property. No site reached total recovery, indicating that individual soil properties alone may not fully indicate recovery in soil quality as soil recovery does not equal the sum of its parts. Site aridity was the largest predictor of reclamation outcomes, but the effects differed depending on soil type. Taken together, results suggest the recovery of soil quality - which reflects soil fertility, carbon sequestration potential, and other ecosystem functions - was influenced primarily by site setting, with soil type and aridity major mediators of on-pad carbon, salinity, and total soil recovery following reclamation.

Ecosystem resilience to invasion and drought: Insights after 24 years in a rare never-grazed grassland.

Understanding the resilience of ecosystems globally is hampered by the complex and interacting drivers of change characteristic of the Anthropocene. This is true for drylands of the western US, where widespread alteration of disturbance regimes and spread of invasive non-native species occurred with westward expansion during the 1800s, including the introduction of domestic livestock and spread of Bromus tectorum, an invasive non-native annual grass. In addition, this region has experienced a multi-decadal drought not seen for at least 1200 years with potentially large and interacting impacts on native plant communities. Here, we present 24 years of twice-annual plant cover monitoring (1997-2021) from a semiarid grassland never grazed by domestic livestock but subject to a patchy invasion of B. tectorum beginning in ~1994, compare our findings to surveys done in 1967, and examine potential climate drivers of plant community changes. We found a significant warming trend in the study area, with more than 75% of study year temperatures being warmer than average (1966-2021). We observed a native perennial grass community with high resilience to climate forcings with cover values like those in 1967. In invaded patches, B. tectorum cover was greatest in the early years of this study (1997-2001; ~20%-40%) but was subsequently constrained by climate and subtle variation in soils, with limited evidence of long-term impacts to native vegetation, contradicting earlier studies. Our ability to predict year-to-year variation in functional group and species cover with climate metrics varied, with a 12-month integrated index and fall and winter patterns appearing most important. However, declines to near zero live cover in recent years in response to regional drought intensification leave questions regarding the resiliency of intact grasslands to ongoing aridification and whether the vegetation observations reported here may be a leading indicator of impending change in this protected ecosystem.

Synergistic soil, land use, and climate influences on wind erosion on the Colorado Plateau: Implications for management.

Two decades of drought in the southwestern USA are spurring concerns about increases in wind erosion, dust emissions, and associated impacts on ecosystems, agriculture, human health, and water supply. Different avenues of investigation into primary drivers of wind erosion and dust have yielded mixed results depending on the spatial and temporal sensitivity of the evidence. We monitored passive aeolian sediment traps from 2017 to 2020 across eighty-one sites near Moab, Utah to understand patterns of sediment flux. At measurement sites, we collated climate, soil, topography and vegetation spatial layers to better understand the context of wind erosion and then combined these data with field observations of land use in models to characterize the influence of cattle grazing, oil and gas well pads, and vehicle/heavy equipment disturbance that potentially drive both exposure of bare soil and increases in erodible sediment supply that increase vulnerability to erosion. Disturbed areas with low soil calcium carbonate content yielded high sediment transport in dry years, but notably areas with little disturbance and low bare soil exposure had much less activity. Cattle grazing had the largest land use association with erosional activity with analyses suggesting that both herbivory and trampling from cattle could be drivers. The amount and distribution of bare soil exposure from new sub-annual fractional cover remote sensing products proved very helpful in mapping erosion, and new predictive maps informed by field data are presented to help depict spatial patterns of wind erosion activity. Our results suggest that despite the magnitude of current droughts, minimizing surface disturbance in vulnerable soils can mitigate a large portion of dust emissions. Results can help land managers identify eroding areas where disturbance reduction and soil surface protection measures can be prioritized.

Soil surface treatments and precipitation timing determine seedling development across southwestern US restoration sites.

Restoration in dryland ecosystems often has poor success due to low and variable water availability, degraded soil conditions, and slow plant community recovery rates. Restoration treatments can mitigate these constraints but, because treatments and subsequent monitoring are typically limited in space and time, our understanding of their applicability across broader environmental gradients remains limited. To address this limitation, we implemented and monitored a standardized set of seeding and soil surface treatments (pits, mulch, and ConMod artificial nurse plants) designed to enhance soil moisture and seedling establishment across RestoreNet, a growing network of 21 diverse dryland restoration sites in the southwestern USA over 3 years. Generally, we found that the timing of precipitation relative to seeding and the use of soil surface treatments were more important in determining seeded species emergence, survival, and growth than site-specific characteristics. Using soil surface treatments in tandem with seeding promoted up to 3× greater seedling emergence densities compared with seeding alone. The positive effect of soil surface treatments became more prominent with increased cumulative precipitation since seeding. The seed mix type with species currently found within or near a site and adapted to the historical climate promoted greater seedling emergence densities compared with the seed mix type with species from warmer, drier conditions expected to perform well under climate change. Seed mix and soil surface treatments had a diminishing effect as plants developed beyond the first season of establishment. However, we found strong effects of the initial period seeded and of the precipitation leading up to each monitoring date on seedling survival over time, especially for annual and perennial forbs. The presence of exotic species exerted a negative influence on seedling survival and growth, but not initial emergence. Our findings suggest that seeded species recruitment across drylands can generally be promoted, regardless of location, by (1) incorporation of soil surface treatments, (2) employment of near-term seasonal climate forecasts, (3) suppression of exotic species, and (4) seeding at multiple times. Taken together, these results point to a multifaceted approach to ameliorate harsh environmental conditions for improved seeding success in drylands, both now and under expected aridification.

Droughting a megadrought: Ecological consequences of a decade of experimental drought atop aridification on the Colorado Plateau.

Global dryland vegetation communities will likely change as ongoing drought conditions shift regional climates towards a more arid future. Additional aridification of drylands can impact plant and ground cover, biogeochemical cycles, and plant-soil feedbacks, yet how and when these crucial ecosystem components will respond to drought intensification requires further investigation. Using a long-term precipitation reduction experiment (35% reduction) conducted across the Colorado Plateau and spanning 10 years into a 20+ year regional megadrought, we explored how vegetation cover, soil conditions, and growing season nitrogen (N) availability are impacted by drying climate conditions. We observed large declines for all dominant plant functional types (C3 and C4 grasses and C3 and C4 shrubs) across measurement period, both in the drought treatment and control plots, likely due to ongoing regional megadrought conditions. In experimental drought plots, we observed less plant cover, less biological soil crust cover, warmer and drier soil conditions, and more soil resin-extractable N compared to the control plots. Observed increases in soil N availability were best explained by a negative correlation with plant cover regardless of treatment, suggesting that declines in vegetation N uptake may be driving increases in available soil N. However, in ecosystems experiencing long-term aridification, increased N availability may ultimately result in N losses if soil moisture is consistently too dry to support plant and microbial N immobilization and ecosystem recovery. These results show dramatic, worrisome declines in plant cover with long-term drought. Additionally, this study highlights that more plant cover losses are possible with further drought intensification and underscore that, in addition to large drought effects on aboveground communities, drying trends drive significant changes to critical soil resources such as N availability, all of which could have long-term ecosystem impacts for drylands.

Decline in biological soil crust N-fixing lichens linked to increasing summertime temperatures.

Biological soil crusts (biocrusts), comprised of mosses, lichens, and cyanobacteria, are key components to many dryland communities. Climate change and other anthropogenic disturbances are thought to cause a decline in mosses and lichens, yet few long-term studies exist to track potential shifts in these sensitive soil-surface communities. Using a unique long-term observational dataset from a temperate dryland with initial observations dating back to 1967, we examine the effects of 53 y of observed environmental variation and Bromus tectorum invasion on biocrust communities in a grassland never grazed by domestic livestock. Annual observations show a steep decline in N-fixing lichen cover (dominated by Collema species) from 1996 to 2002, coinciding with a period of extended drought, with Collema communities never able to recover. Declines in other lichen species were also observed, both in number of species present and by total cover, which were attributed to increasing summertime temperatures. Conversely, moss species gradually gained in cover over the survey years, especially following a large Bromus tectorum invasion at the study onset (ca. 1996 to 2001). These results support a growing body of studies that suggests climate change is a key driver in changes to certain components of late-successional biocrust communities. Results here suggest that warming may partially negate decades of protection from disturbance, with biocrust communities reaching a vital tipping point. The accelerated rate of ongoing warming observed in this study may have resulted in the loss of lichen cover and diversity, which could have long-term implications for global temperate dryland ecosystems.

Assessing vegetation recovery from energy development using a dynamic reference approach.

Ecologically relevant references are useful for evaluating ecosystem recovery, but references that are temporally static may be less useful when environmental conditions and disturbances are spatially and temporally heterogeneous. This challenge is particularly acute for ecosystems dominated by sagebrush (Artemisia spp.), where communities may require decades to recover from disturbance. We demonstrated application of a dynamic reference approach to studying sagebrush recovery using three decades of sagebrush cover estimates from remote sensing (1985-2018). We modelled recovery on former oil and gas well pads (n = 1200) across southwestern Wyoming, USA, relative to paired references identified by the Disturbance Automated Reference Toolset. We also used quantile regression to account for unmodelled heterogeneity in recovery, and projected recovery from similar disturbance across the landscape. Responses to weather and site-level factors often differed among quantiles, and sagebrush recovery on former well pads increased more when paired reference sites had greater sagebrush cover. Little (<5%) of the landscape was projected to recover within 100 years for low to mid quantiles, and recovery often occurred at higher elevations with cool and moist annual conditions. Conversely, 48%-78% of the landscape recovered quickly (within 25 years) for high quantiles of sagebrush cover. Our study demonstrates advantages of using dynamic reference sites when studying vegetation recovery, as well as how additional inferences obtained from quantile regression can inform management.

Evaluating natural experiments in ecology: using synthetic controls in assessments of remotely sensed land treatments.

Many important ecological phenomena occur on large spatial scales and/or are unplanned and thus do not easily fit within analytical frameworks that rely on randomization, replication, and interspersed a priori controls for statistical comparison. Analyses of such large-scale, natural experiments are common in the health and econometrics literature, where techniques have been developed to derive insight from large, noisy observational data sets. Here, we apply a technique from this literature, synthetic control, to assess landscape change with remote sensing data. The basic data requirements for synthetic control include (1) a discrete set of treated and untreated units, (2) a known date of treatment intervention, and (3) time series response data that include both pre- and post-treatment outcomes for all units. Synthetic control generates a response metric for treated units relative to a no-action alternative based on prior relationships between treated and unexposed groups. Using simulations and a case study involving a large-scale brush-clearing management event, we show how synthetic control can intuitively infer treatment effect sizes from satellite data, even in the presence of confounding noise from climate anomalies, long-term vegetation dynamics, or sensor errors. We find that accuracy depends on the number and quality of potential control units, highlighting the importance of selecting appropriate control populations. Although we consider the synthetic control approach in the context of natural experiments with remote sensing data, we expect the methodology to have wider utility in ecology, particularly for systems with large, complex, and poorly replicated experimental units.

Seasonal and individual event-responsiveness are key determinants of carbon exchange across plant functional types.

Differentiation in physiological activity is a critical component of resource partitioning in resource-limited environments. For example, it is crucial to understand how plant physiological performance varies through time for different functional groups to forecast how terrestrial ecosystems will respond to change. Here, we tracked the seasonal progress of 13 plant species representing C3 shrub, perennial C3 and C4 grass, and annual forb functional groups of the Colorado Plateau, USA. We tested for differences in carbon assimilation strategies and how photosynthetic rates related to recent, seasonal, and annual precipitation and temperature variables. Despite seasonal shifts in species presence and activity, we found small differences in seasonally weighted annual photosynthetic rates among groups. However, differences in the timing of maximum assimilation (Anet) were strongly functional group-dependent. C3 shrubs employed a relatively consistent, low carbon capture strategy and maintained activity year-round but switched to a rapid growth strategy in response to recent climate conditions. In contrast, grasses maintained higher carbon capture during spring months when all perennials had maximum photosynthetic rates, but grasses were dormant during months when shrubs remained active. Perennial grass Anet rates were explained in part by precipitation accumulated during the preceding year and average maximum temperatures during the past 48 h, a result opposite to shrubs. These results lend insight into diverse physiological strategies and their connections to climate, and also point to the potential for shrubs to increase in abundance in response to increased climatic variability in drylands, given shrubs' ability to respond rapidly to changing conditions.

Natural and anthropogenic processes affecting radon releases during mining and early stage reclamation activities, Pinenut uranium mine, Arizona, USA.

Radon (Rnair) was monitored in open air in publicly accessible areas surrounding the Pinenut uranium (U) mine during mining and reclamation activities in 2015-16 to address concerns about mining related effects to areas surrounding Grand Canyon National Park (GCNP) in Arizona, USA. During July 2015, Rnair concentrations associated with the ore storage pile monitoring site were larger than those at the mine vent monitoring site and likely resulted from the relatively large amount of ore stored on site during this period. Higher wind velocities at the ore pile monitoring site generally resulted in lower Rnair concentrations; however, wind velocity did not appear to be an important factor in controlling Rnair concentrations at the mine vent monitoring site. Physical disturbances of the ore pile by heavy equipment did not coincide with elevated Rnair concentrations at the ore storage pile or mine vent monitoring sites. The relative size of the ore storage pile showed a positive trend with the daily mean Rnair concentration measured at the ore pile monitoring site. Principal component analysis (PCA) was applied to the ore pile and mine vent multivariate data sets for simultaneous comparison of all measured variables during 230 days of the study period. A significant positive coefficient for Rnair was associated with a significant negative coefficient for wind speed for principal component (PC) 2ore pile. Significant, positive PC2mine vent coefficients included Rnair, wind direction, and relative ore pile size indicating that Rnair variations at the mine vent monitoring site may be affected by Rn sourced from the ore pile. The ore pile is located about 200 m south of the mine vent Rn monitor with the prevalent wind direction coming from the south. All data generated during the field study and laboratory verification tests were published by Naftz et al. (2018) and are available online at: https://doi.org/10.5066/F79Z946T.

Transient population dynamics impede restoration and may promote ecosystem transformation after disturbance.

The apparent failure of ecosystems to recover from increasingly widespread disturbance is a global concern. Despite growing focus on factors inhibiting resilience and restoration, we still know very little about how demographic and population processes influence recovery. Using inverse and forward demographic modelling of 531 post-fire sagebrush populations across the western US, we show that demographic processes during recovery from seeds do not initially lead to population growth but rather to years of population decline, low density, and risk of extirpation after disturbance and restoration, even at sites with potential to support long-term, stable populations. Changes in population structure, and resulting transient population dynamics, lead to a > 50% decline in population growth rate after disturbance and significant reductions in population density. Our results indicate that demographic processes influence the recovery of ecosystems from disturbance and that demographic analyses can be used by resource managers to anticipate ecological transformation risk.

Shrub persistence and increased grass mortality in response to drought in dryland systems.

Droughts in the southwest United States have led to major forest and grassland die-off events in recent decades, suggesting plant community and ecosystem shifts are imminent as native perennial grass populations are replaced by shrub- and invasive plant-dominated systems. These patterns are similar to those observed in arid and semiarid systems around the globe, but our ability to predict which species will experience increased drought-induced mortality in response to climate change remains limited. We investigated meteorological drought-induced mortality of nine dominant plant species in the Colorado Plateau Desert by experimentally imposing a year-round 35% precipitation reduction for eight continuous years. We distributed experimental plots across numerous plant, soil, and parent material types, resulting in 40 distinct sites across a 4,500 km2 region of the Colorado Plateau Desert. For all 8 years, we tracked c. 400 individual plants and evaluated mortality responses to treatments within and across species, and through time. We also examined the influence of abiotic and biotic site factors in driving mortality responses. Overall, high mortality trends were driven by dominant grass species, including Achnatherum hymenoides, Pleuraphis jamesii, and Sporobolus cryptandrus. Responses varied widely from year to year and dominant shrub species were generally resistant to meteorological drought, likely due to their ability to access deeper soil water. Importantly, mortality increased in the presence of invasive species regardless of treatment, and native plant die-off occurred even under ambient conditions, suggesting that recent climate changes are already negatively impacting dominant species in these systems. Results from this long-term drought experiment suggest major shifts in community composition and, as a result, ecosystem function. Patterns also show that, across multiple soil and plant community types, native perennial grass species may be replaced by shrubs and invasive annuals in the Colorado Plateau Desert.

Temporal and abiotic fluctuations may be preventing successful rehabilitation of soil-stabilizing biocrust communities.

Land degradation is a persistent ecological problem in many arid and semiarid systems globally (drylands hereafter). Most instances of dryland degradation include some form of soil disturbance and/or soil erosion, which can hinder vegetation establishment and reduce ecosystem productivity. To combat soil erosion, researchers have identified a need for rehabilitation of biological soil crusts (biocrusts), a globally relevant community of organisms aggregating the soil surface and building soil fertility. Here, the impact of plant and biocrust cover was tested on soil erosion potential in the piñon-juniper woodlands of Bandelier National Monument, New Mexico, USA. Biocrusts were found to be similarly influential to vascular plants in reducing erosion, largely acting by promoting surface roughness. The potential to rehabilitate biocrusts within the Monument was also tested. Plots were inoculated on eroding soils before the summer monsoon with greenhouse-cultured biocrusts. In a full-factorial design, treatments to reduce or halt erosion were administered to the inoculated plots and their paired controls. These erosion-reduction treatments included barriers to overland flow (flashing), slash placement, and seeding of vascular plants. Dynamic changes to soil stability, penetration resistance, and extractable soil nutrients were observed through time, but no strong effects with the addition of biocrust inoculum, seeding, or erosion intervention treatments were seen. The results do suggest possible ways forward to successfully rehabilitate biocrust, including varying the timing of biocrust application, amending inoculum application with different types of soil stabilization techniques, and adding nutrients to soils. The insights gleaned from the lack of response brings us closer to developing effective techniques to arrest soil loss in these socially and ecologically important dryland systems.

Increasing temperature seasonality may overwhelm shifts in soil moisture to favor shrub over grass dominance in Colorado Plateau drylands.

Ecosystems in the southwestern U.S. are predicted to experience continued warming and drying trends of the early twenty-first century. Climate change can shift the balance between grass and woody plant abundance in these water-limited systems, which has large implications for biodiversity and ecosystem processes. However, variability in topo-edaphic conditions, notably soil texture and depth, confound efforts to quantify specific climatic controls over grass vs. shrub dominance. Here, we utilized weather records and a mechanistic soil water model to identify the timing and depth at which soil moisture related most strongly to the balance between grass and shrub dominance in the southern Colorado Plateau. Shrubs dominate where there is high soil moisture availability during winter, and where temperature is more seasonally variable, while grasses are favored where moisture is available during summer. Climate change projections indicate consistent increases in mean temperature and seasonal temperature variability for all sites, but predictions for summer and winter soil moisture vary across sites. Together, these changes in temperature and soil moisture are expected to shift the balance towards increasing shrub dominance across the region. These patterns are strongly driven by changes in temperature, which either enhance or overwhelm effects of changes in soil moisture across sites. This approach, which incorporates local, edaphic factors at sites protected from disturbance, improves understanding of climate change impacts on grass vs. shrub abundance and may be useful in other dryland regions with high edaphic and climatic heterogeneity.

Adapting management to a changing world: Warm temperatures, dry soil, and interannual variability limit restoration success of a dominant woody shrub in temperate drylands.

Restoration and rehabilitation of native vegetation in dryland ecosystems, which encompass over 40% of terrestrial ecosystems, is a common challenge that continues to grow as wildfire and biological invasions transform dryland plant communities. The difficulty in part stems from low and variable precipitation, combined with limited understanding about how weather conditions influence restoration outcomes, and increasing recognition that one-time seeding approaches can fail if they do not occur during appropriate plant establishment conditions. The sagebrush biome, which once covered over 620,000 km2 of western North America, is a prime example of a pressing dryland restoration challenge for which restoration success has been variable. We analyzed field data on Artemisia tridentata (big sagebrush) restoration collected at 771 plots in 177 wildfire sites across its western range, and used process-based ecohydrological modeling to identify factors leading to its establishment. Our results indicate big sagebrush occurrence is most strongly associated with relatively cool temperatures and wet soils in the first spring after seeding. In particular, the amount of winter snowpack, but not total precipitation, helped explain the availability of spring soil moisture and restoration success. We also find considerable interannual variability in the probability of sagebrush establishment. Adaptive management strategies that target seeding during cool, wet years or mitigate effects of variability through repeated seeding may improve the likelihood of successful restoration in dryland ecosystems. Given consistent projections of increasing temperatures, declining snowpack, and increasing weather variability throughout midlatitude drylands, weather-centric adaptive management approaches to restoration will be increasingly important for dryland restoration success.

Disturbance automated reference toolset (DART): Assessing patterns in ecological recovery from energy development on the Colorado Plateau.

A new disturbance automated reference toolset (DART) was developed to monitor human land surface impacts using soil-type and ecological context. DART identifies reference areas with similar soils, topography, and geology; and compares the disturbance condition to the reference area condition using a quantile-based approach based on a satellite vegetation index. DART was able to represent 26-55% of variation of relative differences in bare ground and 26-41% of variation in total foliar cover when comparing sites with nearby ecological reference areas using the Soil Adjusted Total Vegetation Index (SATVI). Assessment of ecological recovery at oil and gas pads on the Colorado Plateau with DART revealed that more than half of well-pads were below the 25th percentile of reference areas. Machine learning trend analysis of poorly recovering well-pads (quantile<0.23) had out-of-bag error rates between 37 and 40% indicating moderate association with environmental and management variables hypothesized to influence recovery. Well-pads in grasslands (median quantile [MQ]=13%), blackbrush (Coleogyne ramosissima) shrublands (MQ=18%), arid canyon complexes (MQ=18%), warmer areas with more summer-dominated precipitation, and state administered areas (MQ=12%) had low recovery rates. Results showcase the usefulness of DART for assessing discrete surface land disturbances, and highlight the need for more targeted rehabilitation efforts at oil and gas well-pads in the arid southwest US.

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.

Pulse-drought atop press-drought: unexpected plant responses and implications for dryland ecosystems.

In drylands, climate change is predicted to cause chronic reductions in water availability (press-droughts) through reduced precipitation and increased temperatures as well as increase the frequency and intensity of short-term extreme droughts (pulse-droughts). These changes in precipitation patterns may have profound ecosystem effects, depending on the sensitivities of the dominant plant functional types (PFTs). Here we present the responses of four Colorado Plateau PFTs to an experimentally imposed, 4-year, press-drought during which a natural pulse-drought occurred. Our objectives were to (1) identify the drought sensitivities of the PFTs, (2) assess the additive effects of the press- and pulse-drought, and (3) examine the interactive effects of soils and drought. Our results revealed that the C3 grasses were the most sensitive PFT to drought, the C3 shrubs were the most resistant, and the C4 grasses and shrubs had intermediate drought sensitivities. Although we expected the C3 grasses would have the greatest response to drought, the higher resistance of C3 shrubs relative to the C4 shrubs was contrary to our predictions based on the higher water use efficiency of C4 photosynthesis. Also, the additive effects of press- and pulse-droughts caused high morality in C3 grasses, which has large ecological and economic ramifications for this region. Furthermore, despite predictions based on the inverse texture hypothesis, we observed no interactive effects of soils with the drought treatment on cover or mortality. These results suggest that plant responses to droughts in drylands may differ from expectations and have large ecological effects if press- and pulse-droughts push species beyond physiological and mortality thresholds.

Desert grassland responses to climate and soil moisture suggest divergent vulnerabilities across the southwestern United States.

Climate change predictions include warming and drying trends, which are expected to be particularly pronounced in the southwestern United States. In this region, grassland dynamics are tightly linked to available moisture, yet it has proven difficult to resolve what aspects of climate drive vegetation change. In part, this is because it is unclear how heterogeneity in soils affects plant responses to climate. Here, we combine climate and soil properties with a mechanistic soil water model to explain temporal fluctuations in perennial grass cover, quantify where and the degree to which incorporating soil water dynamics enhances our ability to understand temporal patterns, and explore the potential consequences of climate change by assessing future trajectories of important climate and soil water variables. Our analyses focused on long-term (20-56 years) perennial grass dynamics across the Colorado Plateau, Sonoran, and Chihuahuan Desert regions. Our results suggest that climate variability has negative effects on grass cover, and that precipitation subsidies that extend growing seasons are beneficial. Soil water metrics, including the number of dry days and availability of water from deeper (>30 cm) soil layers, explained additional grass cover variability. While individual climate variables were ranked as more important in explaining grass cover, collectively soil water accounted for 40-60% of the total explained variance. Soil water conditions were more useful for understanding the responses of C3 than C4 grass species. Projections of water balance variables under climate change indicate that conditions that currently support perennial grasses will be less common in the future, and these altered conditions will be more pronounced in the Chihuahuan Desert and Colorado Plateau. We conclude that incorporating multiple aspects of climate and accounting for soil variability can improve our ability to understand patterns, identify areas of vulnerability, and predict the future of desert grasslands.