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Soil water availability shapes species richness in mid-latitude shrub steppe plant communities

Questions Ecological communities are controlled by multiple, interacting abiotic and biotic factors that influence the distribution, abundance, and diversity of species. These processes jointly determine resource availability, resource competition, and ultimately species richness. For many terrestrial ecosystems in dryland climates, soil water availability is the most frequent limiting resource for plant species. We used field sampling coupled with process‐based soil water balance modeling to explore the relative importance of multiple macroclimatic, ecohydrological, and biotic variables on plant species and functional type richness at the landscape scale in dryland plant communities. Location Dryland plant communities dominated by big sagebrush ( Artemisia tridentata ) that span climatic and elevational gradients in Wyoming, USA. Methods We quantified species richness at 1,000 m 2 and used multiple regression to determine whether mean climatic conditions, multiple metrics of soil moisture from a soil water balance model (SOILWAT2), soil physical and chemical properties, and shrub stand structure (biotic) variables were related to species and functional type richness. Results Species richness varied between 16 and 54 across sites. We found that species and functional type richness were related to both macroclimate and ecohydrology, but ecohydrology explained slightly more variation than climate. Biotic variables were always secondary to macroclimate and ecohydrology in our models. Variance partitioning revealed that large portions of variability in species (~54%), forb (~47%), and grass (~40%) richness were explained by ecohydrological variables. Conclusions Our results highlight the importance of the spatial and temporal distribution of soil water for dryland plant species richness and suggest that documenting the ways in which climate, vegetation, and soil properties interact to determine soil water availability is critical for understanding biodiversity patterns in dryland plant communities. This work has relevance for other mid‐latitude, shrub‐dominated dryland plant communities where soil water availability strongly influences ecosystem structure and function.

Wyoming

Biocrusts mediate a new mechanism for land degradation under a changing climate

Global concerns for desertification have focused on the slow recovery of extensive and expanding drylands following disturbance, which may be exacerbated by climate change. Biological soil crusts (biocrusts) are photosynthetic soil communities found in drylands worldwide, which are central to the stability and resilience of dryland ecosystems, but vulnerable to global change. Here we use multiple decade-long experiments to investigate the consequences of climate and land-use change on biocrusts and soil stability. Biocrusts recovered rapidly under ambient temperatures but warming interacted with the precipitation disturbance to halt recovery. Moreover, warming alone caused losses of mosses, lichens and soil stability. Our results present a new mechanism contributing to land degradation in drylands whereby warming drives a state shift in biocrust communities, which degrades soil stability. The synergistic effects of climate and land-use change co-occur globally and our results support projections of increased desertification and lowered dryland resilience under warming.

Nature Climate Change

Water from air: An overlooked source of moisture in arid and semiarid regions

Water drives the functioning of Earth’s arid and semiarid lands. Drylands can obtain water from sources other than precipitation, yet little is known about how non-rainfall water inputs influence dryland communities and their activity. In particular, water vapor adsorption – movement of atmospheric water vapor into soil when soil air is drier than the overlying air – likely occurs often in drylands, yet its effects on ecosystem processes are not known. By adding 18 O-enriched water vapor to the atmosphere of a closed system, we documented the conversion of water vapor to soil liquid water across a temperature range typical of arid ecosystems. This phenomenon rapidly increased soil moisture and stimulated microbial carbon (C) cycling, and the flux of water vapor to soil had a stronger impact than temperature on microbial activity. In a semiarid grassland, we also observed that non-rainfall water inputs stimulated microbial activity and C cycling. Together these data suggest that, during rain-free periods, atmospheric moisture in drylands may significantly contribute to variation in soil water content, thereby influencing ecosystem processes. The simple physical process of adsorption of water vapor to soil particles, forming liquid water, represents an overlooked but potentially important contributor to C cycling in drylands.

Scientific Reports

Earlier plant growth helps compensate for reduced carbon fixation after 13 years of warming

1. Drylands play a dominant role in global carbon cycling and are particularly vulnerable to increasing temperatures, but our understanding of how dryland ecosystems will respond to climatic change remains notably poor. Considering that the area of drylands is projected to increase 11–23% by 2100, understanding the impacts of warming on the functions and services furnished by these arid and semiarid ecosystems has numerous implications. 2.In a unique 13‐year ecosystem warming experiment in a southwestern U.S. dryland, we investigated the consequences of rising temperature on Achnatherum hymenoides , a widespread, keystone grass species on the Colorado Plateau. We tracked individual‐ and population‐level responses to identify optimal strategies that may have been masked if considering only one level of plant response. 3.We found several factors combined to affect the timing and magnitude of plant responses during the 13 th year of warming. These included large warming‐induced biomass increases for individual plants, an 8.5‐day advancement in the growing season, and strong reductions in photosynthetic rates and population cover. 4.Importantly, we observed a lack of photosynthetic acclimation and, thus, a warming‐induced down regulation of photosynthetic rates. However, these physiological responses were concurrent with warmed‐plant increases in growing season length and investment in photosynthetic surfaces, demonstrating the species' ability to balance carbon fixation limitations with warming. 5.These results, which bring together ecophysiological, phenological, reproductive, and morphological assessments of plant responses to warming, suggest that the extent of change in A. hymenoides populations will be based upon numerous adaptive responses that vary in their direction and magnitude. Plant population responses to climatic warming remain poorly resolved, particularly for Earth's drylands, and our in situ experiment assessing multiple strategies offers a novel look into a warmer world.

Utah

Water movement through the unsaturated zone of the High Plains Aquifer in the Central Platte Natural Resources District, Nebraska, 2008-12

Uncertainty about the effects of land use and climate on water movement in the unsaturated zone and on groundwater recharge rates can lead to uncertainty in water budgets used for groundwater-flow models. To better understand these effects, a cooperative study between the U.S. Geological Survey and the Central Platte Natural Resources District was initiated in 2007 to determine field-based estimates of recharge rates in selected land-use areas of the Central Platte Natural Resources District in Nebraska. Measured total water potential and unsaturated-zone profiles of tritium, chloride, nitrate as nitrogen, and bromide, along with groundwater-age dates, were used to evaluate water movement in the unsaturated zone and groundwater recharge rates in the central Platte River study area. Eight study sites represented an east-west precipitation contrast across the study area—four beneath groundwater-irrigated cropland (sites 2, 5, and 6 were irrigated corn and site 7 was irrigated alfalfa/corn rotation), three beneath rangeland (sites 1, 4, and 8), and one beneath nonirrigated cropland, or dryland (site 3). Measurements of transient vertical gradients in total water potential indicated that periodic wetting fronts reached greater mean maximum depths beneath the irrigated sites than beneath the rangeland sites, in part, because of the presence of greater and constant antecedent moisture. Beneath the rangeland sites, greater temporal variation in antecedent moisture and total water potential existed and was, in part, likely a result of local precipitation and evapotranspiration. Moreover, greater variability was noticed in the total water potential profiles beneath the western sites than the corresponding eastern sites, which was attributed to less mean annual precipitation in the west. The depth of the peak post-bomb tritium concentration or the interface between the pre-bomb/post-bomb tritium, along with a tritium mass balance, within sampled soil profiles were used to estimate water fluxes in the unsaturated zone at three of the eight study sites: site 2 (irrigated), site 3 (dryland), and site 8 (rangeland). Estimates for recharge were about 68 millimeters per year [(mm/yr), post-bomb peak], 133 to 159 mm/yr (tritium interface), and 137 mm/yr (mass balance) at site 2 (irrigated); about 63 mm/yr (tritium interface) and 12 mm/yr (mass balance) at site 3 (dryland); and about 53 mm/yr (tritium interface) and 10 mm/yr (mass balance) at site 8 (rangeland). Recharge values from the mass balance at site 2 were more than an order of magnitude greater than recharge values at site 3, suggesting irrigation is an important control on water movement through the unsaturated zone. For the remaining five sites, the post-bomb tritium had flushed through the system and recharge was considered modern (within 10 years of sampling). The chloride mass-balance method was used to determine water fluxes below the root zone (less than 2 meters below land surface) at the rangeland sites: sites 1, 4, and 8. At these rangeland sites, water fluxes ranged from 1.8 to 96 mm/yr at site 1, 1.1 to 9.6 mm/yr at site 4, and 1.1 to 68 mm/yr at site 8, with mean rates of 21, 4.3, and 13 mm/yr, respectively. Site 1 had a greater mean water flux, which was consistent with the greater precipitation in the east than at site 8 in the west. Chloride mass balance was not calculated at the irrigated and dryland sites because of uncertainty about additional sources of chloride. Concentrations of nitrate as nitrogen in pore water in the unsaturated zone were larger beneath the irrigated and dryland (agricultural) sites compared with the rangeland sites. The larger concentrations at the agricultural sites are consistent with the application of nitrogen fertilizer at the agricultural sites and no substantial accumulation at the rangeland sites. The shape of the nitrate as nitrogen and chloride concentration profiles at site 1 (rangeland) indicate a reasonably larger and more consistent water flux in the UZ than beneath the other two rangeland sites (sites 4 or 8). Excluding site 7, the general shape of the nitrate as nitrogen profiles was similar beneath the agricultural sites and supports the estimates of water movement and recharge rates determined from the tritium and chloride methods. Movement of bromide through the unsaturated zone indicated greater water fluxes are found beneath irrigated lands than beneath rangeland. Bromide profiles in the unsaturated zone, determined from center of mass and peak displacement methods, document water fluxes ranged from 58 to 394 mm/yr beneath irrigated sites and 9 to 201 mm/yr beneath rangeland sites. Water-flux estimates from the potassium bromide tests at most sites did not represent overall recharge rates because the bromide remained primarily in the root zone. Apparent groundwater age was used to determine the groundwater residence time at the eight sites and to estimate recharge rates. Groundwater ages in the study area ranged from old water (defined here as groundwater that was recharged more than 50 years ago) to modern (defined here as groundwater that has recharged within the past 10 years). Groundwater ages indicated that the shallow monitoring wells generally had younger residence times, whereas the deeper monitoring wells generally had the older residence times. Groundwater dates from the shallowest monitoring wells were used to determine recharge rates at the water table. These rates generally were similar to recharge rates determined from tritium and chloride mass-balance methods. Groundwater recharge rates generally increased with well depth, and the deeper monitoring wells likely do not represent local recharge conditions but recharge from a regional flow system that receives recharge from distant sources. Overall, these data generally indicate that water movement within the unsaturated zone primarily is affected by spatial contrasts in mean annual precipitation and by the land use or land cover. The eight unsaturated-zone sites each generated unique, valuable datasets that likely will improve the understanding of water movement and recharge rates in the central Platte River valley.

Nebraska

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.

Ecological Applications

C 3 and C 4 plant responses to increased temperatures and altered monsoonal precipitation in a cool desert on the Colorado Plateau, USA

Dryland ecosystems represent >40 % of the terrestrial landscape and support over two billion people; consequently, it is vital to understand how drylands will respond to climatic change. However, while arid and semiarid ecosystems commonly experience extremely hot and dry conditions, our understanding of how further temperature increases or altered precipitation will affect dryland plant communities remains poor. To address this question, we assessed plant physiology and growth at a long-term (7-year) climate experiment on the Colorado Plateau , USA, where the community is a mix of shallow-rooted C 3 and C 4 grasses and deep-rooted C 4 shrubs. The experiment maintained elevated-temperature treatments (+2 or +4 °C) in combination with altered summer monsoonal precipitation (+small frequent precipitation events or +large infrequent events). Increased temperature negatively affected photosynthesis and growth of the C 3 and C 4 grasses, but effects varied in their timing: +4 °C treatments negatively affected the C 3 grass early in the growing season of both years, while the negative effects of temperature on the C 4 grass were seen in the +2 and +4 °C treatments, but only during the late growing season of the drier year. Increased summer precipitation did not affect photosynthesis or biomass for any species, either in the year the precipitation was applied or the following year. Although previous research suggests dryland plants, and C 4 grasses in particular, may respond positively to elevated temperature, our findings from a cool desert show marked declines in C 3 and C 4 photosynthesis and growth, with temperature effects dependent on the degree of warming and growing-season precipitation.

Utah

Spectrally monitoring the response of the biocrust moss Syntrichia caninervis to altered precipitation regimes

Climate change is expected to impact drylands worldwide by increasing temperatures and changing precipitation patterns. These effects have known feedbacks to the functional roles of dryland biological soil crust communities (biocrusts), which are expected to undergo significant climate-induced changes in community structure and function. Nevertheless, our ability to monitor the status and physiology of biocrusts with remote sensing is limited due to the heterogeneous nature of dryland landscapes and the desiccation tolerance of biocrusts, which leaves them frequently photosynthetically inactive and difficult to assess. To address this critical limitation, we subjected a dominant biocrust species Syntrichia caninervis to climate-induced stress in the form of small, frequent watering events, and spectrally monitored the dry mosses’ progression towards mortality. We found points of spectral sensitivity responding to experimentally-induced stress in desiccated mosses, indicating that spectral imaging is an effective tool to monitor photosynthetically inactive biocrusts. Comparing the Normalized Difference Vegetation Index (NDVI), the Simple Ratio (SR), and the Normalized Pigment Chlorophyll Index (NPCI), we found NDVI minimally effective at capturing stress in precipitation-stressed dry mosses, while the SR and NPCI were highly effective. Our results suggest the strong potential for utilizing spectroscopy and chlorophyll-derived indices to monitor biocrust ecophysiological status, even when biocrusts are dry, with important implications for improving our understanding of dryland functioning.

Scientific Reports

Biocrust and seed placement influence emergence rates of perennial grass Elymus elymoides across five North American deserts

1. Dryland vascular plant emergence is often limited by water availability especially with projected climate and precipitation changes. Biological soil crusts (biocrusts) can promote water retention and nutrient availability that benefit germination, and emergence yet can also act as a surface barrier preventing critical seed soil contact and hindering emergence. Alongside these factors, dryland fire frequency has increased with the inclusion of invasive species and vegetation structural changes. With enhanced continuous fine fuel distribution there is a high potential to disrupt biocrust-plant interactions and influence associated management actions. 2. This study explores the dynamic relationship between biocrusts and fire-related heating effects on seedling emergence across five North American deserts: the Chihuahuan Desert, Colorado Plateau, Great Basin, Mojave Desert and Sonoran Desert. We conducted a greenhouse-based seedling emergence experiment using Elymus elymoides (bottlebrush squirreltail), a common perennial grass, with biocrust and bare soil mesocosms in which half were heated to mimic the effects of wildfire temperature. 3. The variables that had the greatest influence on germination rate and germination timing were the presence of biocrust and seed placement (on top of vs within the biocrust/soil matrix). Emergence rate was greatest atop bare soil followed by seeds inserted into biocrust. Emergence timing was faster with biocrust present, but only when seeds were inserted into it. Desert origin of biocrust and soil collection also influenced germination where the probability of any one seed emerging was highest in the Chihuahuan and Mojave desert sites relative to the Sonoran desert site which showed the lowest probability. Heating had mixed effects whereby it increased the likelihood of emergence but did not affect the overall rate or timing. 4. Synthesis and applications . This study underscores the importance of healthy and impaired biocrusts in dryland systems and suggests ways in which the combination of biocrust and seed placement can influence plant establishment, in addition to providing insight into seeding strategies for managers and restoration practitioners working in dryland sites.

Arizona, California, Colorado, Nevada, New Mexico

Biological soil crusts in ecological restoration: Emerging research and perspectives

Drylands encompass over 40% of terrestrial ecosystems and face significant anthropogenic degradation causing a loss of ecosystem integrity, services, and deterioration of social‐ecological systems. To combat this degradation, some dryland restoration efforts have focused on the use of biological soil crusts (biocrusts): complex communities of cyanobacteria, algae, lichens, bryophytes, and other organisms living in association with the top millimeters of soil. Biocrusts are common in many ecosystems and especially drylands. They perform a suite of ecosystem functions: stabilizing soil surfaces to prevent erosion, contributing carbon through photosynthesis, fixing nitrogen, and mediating the hydrological cycle in drylands. Biocrusts have emerged as a potential tool in restoration; developing methods to implement effective biocrust restoration has the potential to return many ecosystem functions and services. Although culture‐based approaches have allowed researchers to learn about the biology, physiology, and cultivation of biocrusts, transferring this knowledge to field implementation has been more challenging. A large amount of research has amassed to improve our understanding of biocrust restoration, leaving us at an opportune time to learn from one another and to join approaches for maximum efficacy. The articles in this special issue improve the state of our current knowledge in biocrust restoration, highlighting efforts to effectively restore biocrusts through a variety of different ecosystems, across scales and utilizing a variety of lab and field methods. This collective work provides a useful resource for the scientific community as well as land managers.

Restoration Ecology

Monitoring the water-quality response of agricultural conservation practices in the Bucks Branch watershed, Sussex County, Delaware, 2014–16

The purpose of this study was to evaluate the effects of irrigation and cover crops as conservation practices on water quality in groundwater and streams. Bucks Branch, a stream in the Nanticoke River watershed in southwestern Delaware, was identified as having one of the highest concentrations of nitrate in all surface-water sites sampled by the Delaware Department of Natural Resources and Environmental Control (DNREC). The study site is on two adjacent fields bordering Bucks Branch, one that has used irrigation since 2000 and one with dryland farming; both under conservation tillage and long-term rotation of corn, soybean, and small grain crops. A streamgage was installed near the study site fields to measure streamflow and water quality. The study area is typical of farming practices and environmental conditions throughout much of the intensively farmed agricultural land of the Coastal Plain of Delaware and surrounding parts of Maryland. Monitoring was conducted from January 2014 through June 2016. Corn was grown on both fields during the two growing seasons of the study period, and cover crops were planted before or shortly after harvest on both fields. During the second year of data collection, the effects of radish and rye grass cover crops on nutrient transport were studied. The combined results from data collected for this study show that water and nitrate moved below the root zone year round when soil moisture was high, especially after significant rainfall and frequently after irrigation. Soil water sampled 2 to 3 weeks after nutrients were applied had nitrate concentrations greater than 50 milligrams per liter as nitrogen (mg/L as N) and may be a significant source of nitrate to groundwater. Whereas recharge containing elevated nitrate concentrations also occurred under the dryland field, it was less frequent and of lower concentration than recharge under the irrigated field. Nitrate was present in all groundwater samples from these sites. Groundwater estimated to have recharged within 10 years or less had higher median concentrations of nitrate than in older water samples. The oldest groundwater encountered was over 30 years old, and had traveled along the longest, deep flowpaths from upland fields to the stream. The median nitrate concentration was 18 mg/L as N in younger water (less than 10 years old) beneath the irrigated field, compared to about 10 mg/L as N in younger water beneath the dryland field. Samples from the shallow upland wells in both study fields showed little, if any, evidence of denitrification. Several samples from deeper wells and from wells near forested riparian zone wetlands that border both fields did show partial denitrification. A mixing model estimated that between 12 and 22 percent of the nitrate discharging to the stream was lost through uptake and denitrification upstream of the streamgage on Bucks Branch. Continuous data collected at this site and evidence of denitrification in the surface-water samples showed a greater potential for loss of nitrate during the warmer months than the colder months. This pattern was similar to that seen below the streamgage at the most downstream site in the watershed. A mixed cover crop of radishes and rye was planted prior to removal (radishes) and just after harvest (rye) of the corn crop on the irrigated field. Rye grass was planted shortly after crop harvest on the dryland field. Cover crop biomass samples collected while radishes were growing and after they were killed by freezing temperatures indicates that the early planted radish crop effectively scavenged available nitrogen from the soil. Whereas radish biomass initially held more nitrogen than rye, at 55 to 8 pounds per acre, respectively, leaching of inorganic nitrate following radish die-off was minimal. Soil-water nitrate concentrations during the cover-crop growing period were lower than during the growing season prior to planting of the cover crop. There also was an increase in soil fertility and dissolved organic nitrate in samples of soil water that was likely related to increased soil microbial metabolism. Results indicate that cover crops stored plant nutrients over the winter and did not increase shallow groundwater concentrations of nitrate. Although conservation practices such as cover crops and nutrient management have been applied to these fields, there was still significant leaching of nitrate to groundwater, especially under the irrigated field. This will likely continue to be a challenge in this area and other parts of the Coastal Plain where soil moisture capacity is relatively low and managing irrigation around rainfall is difficult. Cover crops, when planted in standing corn, are one practice that can effectively pull nitrate from below the root zone to the top layer of soil, thus limiting the amount of potential nitrate leaching to groundwater. Irrigation management that would lower average soil moisture conditions during the growing season also could potentially limit nitrogen transport.

Delaware

Hydrogeology, water use, and simulation of flow in the High Plains aquifer in northwestern Oklahoma, southeastern Colorado, southwestern Kansas, northeastern New Mexico, and northwestern Texas

The U.S. Geological Survey, in cooperation with the Oklahoma Water Resources Board, began a three-year study of the High Plains aquifer in northwestern Oklahoma in 1996. The primary purpose of this study was to develop a ground-water flow model to provide the Water Board with the information it needs to manage the quantity of water withdrawn from the aquifer. The study area consists of about 7,100 square miles in Oklahoma and about 20,800 square miles in adjacent states to provide appropriate hydrologic boundaries for the flow model. The High Plains aquifer includes all sediments from the base of the Ogallala Formation to the potentiometric surface. The saturated thickness in Oklahoma ranges from more than 400 feet to less than 50 feet. Natural recharge to the aquifer from precipitation occurs throughout the area but is extremely variable. Dryland agricultural practices appear to enhance recharge from precipitation, and part of the water pumped for irrigation also recharges the aquifer. Natural discharge occurs as discharge to streams, evapotranspiration where the depth to water is shallow, and diffuse ground-water flow across the eastern boundary. Artificial discharge occurs as discharge to wells. Irrigation accounted for 96 percent of all use of water from the High Plains aquifer in the Oklahoma portion of the study area in 1992 and 93 percent in 1997. Total estimated water use in 1992 for the Oklahoma portion of the study area was 396,000 acre-feet and was about 3.2 million acre-feet for the entire study area. Since development of the aquifer, water levels have declined more than 100 feet in small areas of Texas County, Oklahoma, and more than 50 feet in areas of Cimarron County. Only a small area of Beaver County had declines of more than 10 feet, and Ellis County had rises of more than 10 feet. A flow model constructed using the MODFLOW computer code had 21,073 active cells in one layer and had a 6,000- foot grid in both the north-south and east-west directions. The model was used to simulate the period before major development of the aquifer and the period of development. The model was calibrated using observed conditions available as of 1998. The predevelopment-period model integrated data or estimates on the base of aquifer, hydraulic conductivity, streambed and drain conductances, and recharge from precipitation to calculate the predevelopment altitude of the water table, discharge to the rivers and streams, and other discharges. Hydraulic conductivity, recharge, and streambed conductance were varied during calibration so that the model produced a reasonable representation of the observed water table altitude and the estimated discharge to streams. Hydraulic conductivity was reduced in the area of salt dissolution in underlying Permianage rocks. Recharge from precipitation was estimated to be 4.0 percent of precipitation in greater recharge zones and 0.37 percent in lesser recharge zones. Within Oklahoma, the mean difference between water levels simulated by the model and measured water levels at 86 observation points is -2.8 feet, the mean absolute difference is 44.1 feet, and the root mean square difference is 52.0 feet. The simulated discharge is much larger than the estimated discharge for the Beaver River, is somewhat larger for Cimarron River and Wolf Creek, and is about the same for Crooked Creek. The development-period model added specific yield, pumpage, and recharge due to irrigation and dryland cultivation to simulate the period 1946 through 1997. During calibration, estimated specific yield was reduced by 15 percent in Oklahoma east of the Cimarron-Texas County line. Simulated recharge due to irrigation ranges from 24 percent for the 1940s and 1950s to 2 percent for the 1990s. Estimated recharge due to dryland cultivation is about 3.9 percent of precipitation. The mean difference between the simulated and observed waterlevel changes from predevelopment to 1998 at 162 observation points in Oklahoma is less than 0.01 foot, the mean absolute difference is 13.1 feet, and the root mean square difference is 17.9 feet. The model simulates 7.8 cubic feet per second discharge to the Beaver River above Optima Reservoir at the end of 1997, whereas the river was actually dry to this point by this time. The model simulates a decrease in discharge to the Cimarron River and Wolf Creek that appears to be reasonable. The sensitivity of the predevelopment-period model to changes in recharge and hydraulic conductivity was tested. Simulated water levels are sensitive to changes in both recharge and hydraulic conductivity. Simulated discharge to streams is sensitive to recharge but is insensitive to hydraulic conductivity. Recharge and hydraulic conductivity are closely related with respect to water levels but are not related with respect to discharge to streams. The sensitivity of the development-period model to changes in specific yield and recharge due to dryland cultivation was tested. The model appears more sensitive to specific yield, but the two inputs were not varied over the same range in percent change. The calibrated development-period model was used to simulate water-level changes from 1998 to 2020 using mean 1996-97 pumpage. The largest simulated water-level changes in Oklahoma occur in Texas County where water levels are simulated to decline 25 to 50 additional feet over a large area. Water levels also are simulated to decline 10 to 25 additional feet in two large areas of Cimarron County, two areas in Beaver County, and one area in Ellis County, all in Oklahoma. Water levels are simulated to decline more than 100 additional feet in several areas in Kansas and were simulated to decline 50 to 100 additional feet in several areas in Texas.

Colorado, Kansas, New Mexico, Oklahoma, Texas

Interactions of microhabitat and time control grassland bacterial and fungal composition

Dryland grasslands are vast and globally important and, as in all terrestrial ecosystems, soil microbial communities play fundamental roles in regulating dryland ecosystem function. A typical characteristic of drylands is the spatial mosaic of vascular plant cover surrounded by interspace soils, where biological soil crusts (biocrusts)—a complex community of organisms including bacteria, fungi, algae, mosses, and lichens—are common. The implications of this heterogeneity, where plants and biocrust cover co-occur, are often explored in the context of soil fertility and hydrology, but rarely has the impact of these multiple microhabitat types been simultaneously explored to determine the influence on bacterial and fungal communities, key biological players in these ecosystems. Further, our understanding of the temporal dynamics of bacterial and fungal communities in grasslands, and of how these dynamics depend on the microhabitat within the ecosystem, is notably poor. Here we used a temporally and spatially explicit approach to assess bacterial and fungal communities in a grassland on the Colorado Plateau, and to link variation in these communities to edaphic characteristics. We found that microhabitat (e.g., vascular plant rhizosphere, biocrust, and below biocrust) was the strongest driver of differences in bacterial and fungal community richness, diversity, and composition. Microhabitat type also significantly mediated the impact of temporal change in shaping community composition. Taken together, 29% of the variation in bacterial community composition could be explained by microhabitat, date, and microhabitat-by-date interactions, while only 11% of the variation in fungal community composition could be explained by the same factors, suggesting important differences in community assembly processes. Soil microbial communities dictate myriad critical ecosystem functions, thus understanding the factors that control their compostition is crucial to considering and forecasting how terrestrial ecosystems work. Overall, this case study provides insights for future studies on the spatial and temporal dynamics of bacterial and fungal communities in dryland grasslands.

Frontiers in Ecology and Evolution

Influence of atmospheric rivers on vegetation productivity and fire patterns in the southwestern U.S.

In the southwestern U.S., the meteorological phenomenon known as atmospheric rivers (ARs) has gained increasing attention due to its strong connections to floods, snowpacks, and water supplies in the West Coast states. Relatively less is known about the ecological implications of ARs, particularly in the interior Southwest, where AR storms are less common. To address this gap, we compared a chronology of AR landfalls on the west coast between 1989 and 2011 and between 25°N and 42.5°N to annual metrics of the normalized difference vegetation index (NDVI; an indicator of vegetation productivity) and daily resolution precipitation data to assess influences of AR-fed winter precipitation on vegetation productivity across the southwestern U.S. We mapped correlations between winter AR precipitation during landfalling ARs and (1) annual maximum NDVI and (2) area burned by large wildfires summarized by ecoregion during the same year as the landfalls and during the following year. Interannual variations of AR precipitation strongly influenced both NDVI and area burned by wildfire in some dryland ecoregions. The influence of ARs on dryland vegetation varied significantly depending on the latitude of landfall, with those ARs making landfall below 35°N latitude more strongly influencing these systems, and with effects observed as far as 1300 km from the landfall location. As climatologists' understanding of the synoptic patterns associated with the occurrence of ARs continues to evolve, an increased understanding of how AR landfalls, in aggregate, influence vegetation productivity and associated wildfire activity in dryland ecosystems may provide opportunities to better predict ecological responses to climate and climate change.

Arizona, California, Nevada, Utah

Navigating the possibilities and pitfalls of biocrust recovery in a changing climate

Biological soil crusts are complex communities composed of lichens, mosses, bacteria, and cyanobacteria that create a living skin on the soil surface across drylands worldwide. Although small in size, the vast area that biocrusts cover and the critical functions they provide make them a cornerstone of dryland health and resiliency. In addition to being important, biocrusts are exceptionally vulnerable to certain types of disturbance. Although they can withstand a wide range of temperatures and long periods without precipitation, biocrusts are highly sensitive to land-use change and are vulnerable to physical and compressional disturbance (i.e., trampling, vehicles, cattle, heavy machinery). In the face of these disturbances, a critical, long-standing question of interest to dryland ecologists is: Can biocrusts recover following disturbance without active intervention. If so, how long does it take? Early estimates of biocrust recovery suggested recovery can be incredibly slow (on the order of thousands of years), with more modern studies finding potential for faster recovery, especially with intervention. Multiple lines of evidence agree that recovery is context dependent, differing across climates, soils, and with the types of disturbance and biocrust. Additionally, active restoration of biocrusts is becoming more common as tractable strategies are developed for facilitating the establishment of biocrusts after disturbance. Here, we add to the body of knowledge about biocrust recovery following disturbances by reviewing recovery patterns, their connection to climate change, considerations for recovery in changing climates, and the role of restoration.

American Journal of Botany

Landscape genetic approaches to guide native plant restoration in the Mojave Desert

Restoring dryland ecosystems is a global challenge due to synergistic drivers of disturbance coupled with unpredictable environmental conditions. Dryland plant species have evolved complex life-history strategies to cope with fluctuating resources and climatic extremes. Although rarely quantified, local adaptation is likely widespread among these species and potentially influences restoration outcomes. The common practice of reintroducing propagules to restore dryland ecosystems, often across large spatial scales, compels evaluation of adaptive divergence within these species. Such evaluations are critical to understanding the consequences of large-scale manipulation of gene flow and to predicting success of restoration efforts. However, genetic information for species of interest can be difficult and expensive to obtain through traditional common garden experiments. Recent advances in landscape genetics offer marker-based approaches for identifying environmental drivers of adaptive genetic variability in non-model species, but tools are still needed to link these approaches with practical aspects of ecological restoration. Here, we combine spatially-explicit landscape genetics models with flexible visualization tools to demonstrate how cost-effective evaluations of adaptive genetic divergence can facilitate implementation of different seed sourcing strategies in ecological restoration. We apply these methods to Amplified Fragment Length Polymorphism (AFLP) markers genotyped in two Mojave Desert shrub species of high restoration importance: the long-lived, wind-pollinated gymnosperm Ephedra nevadensis , and the short-lived, insect-pollinated angiosperm Sphaeralcea ambigua . Mean annual temperature was identified as an important driver of adaptive genetic divergence for both species. Ephedra showed stronger adaptive divergence with respect to precipitation variability, while temperature variability and precipitation averages explained a larger fraction of adaptive divergence in Sphaeralcea . We describe multivariate statistical approaches for interpolating spatial patterns of adaptive divergence while accounting for potential bias due to neutral genetic structure. Through a spatial bootstrapping procedure, we also visualize patterns in the magnitude of model uncertainty. Finally, we introduce an interactive, distance-based mapping approach that explicitly links marker-based models of adaptive divergence with local or admixture seed sourcing strategies, promoting effective native plant restoration.

Mojave 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 semi-arid 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. The impact of plant and biocrust cover was tested on soil erosion potential in the piñon-juniper woodlands of Bandelier National Monument, NM. 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. In a full factorial design, plots were inoculated on eroding soils before the summer monsoon with greenhouse-cultured biocrusts, and administered the erosion intervention treatments of overland water flow barriers (flashing), slash placement, and seeding of vascular plants. Although significant and dynamic changes to soil stability, penetration resistance, and extractable soil nutrients were observed through time, no strong effects with the addition of inoculum, seeding, or erosion intervention treatments were seen. These results 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 important social-ecological dryland systems.

Ecological Applications

Flexible phenology of a C4 grass linked to resiliency to seasonal and multiyear drought events in the American southwest

Rising temperatures are predicted to further limit dryland water availability as droughts become more intense and frequent and seasonal precipitation patterns shift. Vegetation drought stress may increase mortality and cause declines and delays in phenological events, thereby impacting species' capacity to persist and recover from extreme drought conditions. We compare phenological responses of two common dryland perennial grass species, Achnatherum hymenoides (C 3 ) and Pleuraphis jamesii (C 4 ), to 4 years of experimentally imposed precipitation drought treatments (cool season, warm season, ambient), followed by 2 years of recovery on the Colorado Plateau, United States of America. Tagged individual grasses from both species were monitored biweekly and assessed for phenological metrics and mortality. The C 3 grass exhibited less phenological flexibility to both seasonal and interannual drought conditions and experienced high rates of mortality, thus reducing resiliency. Conversely, the C 4 grass showed more phenological plasticity during imposed drought treatments, with treatment effects diminishing in the two-year recovery period during a severe ambient drought. Synthesis: Results suggest that plant photosynthetic strategies may impact plant resistance and resiliency to drought. Here, C 3 grass populations may decline, potentially shifting cool dryland ecosystems into a system comprised predominantly of warm-season adapted species.

Utah