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  1. Abstract

    Multiyear periods (≥4 years) of extreme rainfall are increasing in frequency as climate continues to change, yet there is little understanding of how rainfall amount and heterogeneity in biophysical properties affect state changes in a sequence of wet and dry periods. Our objective was to examine the importance of rainfall periods, their legacies, and vegetation and soil properties to either the persistence of woody plants or a shift toward perennial grass dominance and a state reversal. We examined a 28‐year record of rainfall consisting of a sequence of multiyear periods (average, dry, wet, dry, average) for four ecosystem types in the Jornada Basin. We analyzed relationships between above ground net primary production (ANPP) and rainfall for three plant functional groups that characterize alternative states (perennial grasses, other herbaceous plants, dominant shrubs). A multimodel comparison was used to determine the relative importance of rainfall, soil, and vegetation properties. For perennial grasses, the greatest mean ANPP in mesquite‐ and tarbush‐dominated shrublands occurred in the wet period and in the dry period following the wet period in grasslands. Legacy effects in grasslands were asymmetric, where the lowest production was found in a dry period following an average period, and the greatest production occurred in a dry period following a wet period. For other herbaceous plants, in contrast, the greatest ANPP occurred in the wet period. Mesquite was the only dominant shrub species with a significant positive response in the wet period. Rainfall amount was a poor predictor of ANPP for each functional group when data from all periods were combined. Initial herbaceous biomass at the plant scale, patch‐scale biomass, and soil texture at the landscape scale improved the predictive relationships of ANPP compared with rainfall alone. Under future climate, perennial grass production is expected to benefit the most from wet periods compared with other functional groups with continued high grass production in subsequent dry periods that can shift (desertified) shrublands toward grasslands. The continued dominance by shrubs will depend on the effects that rainfall has on perennial grasses and the sequence of high‐ and low‐rainfall periods rather than the direct effects of rainfall on shrub production.

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  2. Abstract

    Prediction of abrupt ecosystem transitions resulting from climatic change will be an essential element of adaptation strategies in the coming decades. In the arid southwest USA, the collapse and recovery of long‐lived perennial grasses have important effects on ecosystem services, but the causes of these variations have been poorly understood. Here we use a quality‐controlled vegetation monitoring dataset initiated in 1915 to show that grass cover dynamics during the 20th century were closely correlated to the Pacific decadal oscillation (PDO) index. The relationship out‐performed models correlating grasses to yearly precipitation and drought indices, suggesting that ecosystem transitions attributed only to local disturbances were instead influenced by climate teleconnections. Shifts in PDO phase over time were associated with the persistent loss of core grass species and recovery of transient species, so recovery of grasses in aggregate concealed significant changes in species composition. However, the relationship between PDO and grass cover broke down after 1995; grass cover is consistently lower than PDO would predict. The decoupling of grass cover from the PDO suggests that a threshold had been crossed in which warming or land degradation overwhelmed the ability of any grass species to recover during favorable periods.

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  3. Abstract

    Janus is the Roman god of transitions. In many environments, state transitions are an important part of our understanding of ecological change. These transitions are controlled by the interactions between exogenous forcing factors and stabilizing endogenous feedbacks. Forcing factors and feedbacks are typically considered to consist of different processes. We argue that during extreme events, a process that usually forms part of a stabilizing feedback can behave as a forcing factor. And thus, like Janus, a single process can have two faces. The case explored here pertains to state change in drylands where interactions between wind erosion and vegetation form an important feedback that encourages grass‐to‐shrub state transitions. Wind concentrates soil resources in shrub‐centered fertile islands, removes resources through loss of fines to favor deep‐rooted shrubs, and abrades grasses' photosynthetic tissue, thus further favoring the shrub state that, in turn, experiences greater aeolian transport. This feedback is well documented but the potential of wind to act also as a forcing has yet to be examined. Extreme wind events have the potential to act like other drivers of state change, such as drought and grazing, to directly reduce grass cover. This study examines the responses of a grass‐shrub community after two extreme wind events in 2019 caused severe deflation. We measured grass cover and root exposure due to deflation, in addition to shrub height, grass patch size, and grass greenness along 50‐m transects across a wide range of grass cover. Root exposure was concentrated in the direction of erosive winds during the storms and sites with low grass cover were associated with increased root exposure and reduced greenness. We argue that differences between extreme, rare wind events and frequent, small wind events are significant enough to be differences in kind rather than differences in degree allowing extreme winds to behave as endogenous forcings and common winds to participate in an endogenous stabilizing feedback. Several types of state change in other ecological systems in are contextualized within this framework.

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  4. Abstract

    In dryland soils, spatiotemporal variation in surface soils (0–10 cm) plays an important role in the function of the “critical zone” that extends from canopy to groundwater. Understanding connections between soil microbes and biogeochemical cycling in surface soils requires repeated multivariate measurements of nutrients, microbial abundance, and microbial function. We examined these processes in resource islands and interspaces over a two‐month period at a Chihuahuan Desert bajada shrubland site. We collected soil inProsopis glandulosa(honey mesquite),Larrea tridentata(creosote bush), and unvegetated (interspace) areas to measure soil nutrient concentrations, microbial biomass, and potential soil enzyme activity. We monitored the dynamics of these belowground processes as soil conditions dried and then rewetted due to rainfall. Most measured variables, including inorganic nutrients, microbial biomass, and soil enzyme activities, were greater under shrubs during both wet and dry periods, with the highest magnitudes under mesquite followed by creosote bush and then interspace. One exception was nitrate, which was highly variable and did not show resource island patterns. Temporally, rainfall pulses were associated with substantial changes in soil nutrient concentrations, though resource island patterns remained consistent during all phases of the soil moisture pulse. Microbial biomass was more consistent than nutrients, decreasing only when soils were driest. Potential enzyme activities were even more consistent and did not decline in dry periods, potentially helping to stimulate observed pulses in CO2efflux following rain events observed at a co‐located eddy flux tower. These results indicate a critical zone with organic matter cycling patterns consistently elevated in shrub resource islands (which varied by shrub species), high decomposition potential that limits soil organic matter accumulation across the landscape, and nitrate fluxes that are decoupled from the organic matter pathways.

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  5. Abstract Aims

    Grassland-to-shrubland transition is a common form of land degradation in drylands worldwide. It is often attributed to changes in disturbance regimes, particularly overgrazing. A myriad of direct and indirect effects (e.g., accelerated soil erosion) of grazing may favor shrubs over grasses, but their relative importance is unclear. We tested the hypothesis that topsoil “winnowing” by wind erosion would differentially affect grass and shrub seedling establishment to promote shrub recruitment over that of grass.


    We monitored germination and seedling growth of contrasting perennial grass (Bouteloua eriopoda,Sporobolus airoides, andAristida purpurea) and shrub (Prosopis glandulosa,Atriplex canescens, andLarrea tridentata) functional groups on field-collected non-winnowed and winnowed soils under well-watered greenhouse conditions.


    Non-winnowed soils were finer-textured and had higher nutrient contents than winnowed soils, but based on desorption curves, winnowed soils had more plant-available moisture. Contrary to expectations, seed germination and seedling growth on winnowed and non-winnowed soils were comparable within a given species. The N2-fixing deciduous shrubP. glandulosawas first to emerge and complete germination, and had the greatest biomass accumulation of all species.


    Germination and early seedling growth of grasses and shrubs on winnowed soils were not adversely nor differentially affected comparing with that observed on non-winnowed soils under well-watered greenhouse conditions. Early germination and rapid growth may giveP. glandulosaa competitive advantage over grasses and other shrub species at the establishment stage in grazed grasslands. Field establishment experiments are needed to confirm our findings in these controlled environment trials.

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  6. Abstract

    Grassland‐to‐shrubland state change has been widespread in arid lands globally. Long‐term records at the Jornada Basin USDA‐LTER site in the North American Chihuahuan Desert document the time series of transition from grassland dominance in the 1850s to shrubland dominance in the 1990s. This broadscale change ostensibly resulted from livestock overgrazing in conjunction with periodic drought and represents the classic “grassland‐to‐shrubland” regime shift. However, finer‐scale observations reveal a more nuanced view of this state change that includes transitions from dominance by one shrub functional type to another (e.g., based on leaf habit [evergreen vs. deciduous], N2fixation potential, and drought tolerance). We analyzed the Jornada Basin historic vegetation data using a fine‐scale grid and classified the dominant vegetation in the resulting 890 cells on each of four dates (1858, 1915, 1928, and 1998). This analysis allowed us to quantify on contrasting soil geomorphic units the rate and spatial distribution of: (1) state change from grasslands to shrublands across the Jornada Basin, (2) transitions between shrub functional groups, and (3) transitions from shrub‐to‐grass dominance. Results from our spatially explicit, decadal timescale perspective show that: (1) shrubland ecosystems developing on former grasslands were spatially and temporally more dynamic than has been generally presumed, (2) in some locations, shrublands initially developing on grasslands subsequently transitioned to ecosystems dominated by a different shrub functional type, with these changes in shrub composition likely involving changes in soil properties, and (3) some shrub‐dominated locations have reverted to grass dominance. Accordingly, traditional, broad characterizations of “grassland‐to‐shrubland” state change may be too simplistic. An accounting of these complexities and transitions from one shrub functional group to another is important for projecting state change consequences for ecosystem processes. Understanding the mechanisms, drivers, and influence of interactions between patterns and processes on transitions between shrub states defined by woody plant functional types will be germane to predicting future landscape change.

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  7. Premise of the Study

    The C3desert shrub ocotillo (Fouquieria splendens) completely lacks xeromorphic leaves but is uncommonly both stem succulent and repetitively drought deciduous (documented to have produced many foliation–defoliation cycles during a growing season). Both adaptations conserve water in this xerophyte, but are the roles of succulence and deciduousness merely redundant? The observation that year‐to‐year reproductive effort was relatively consistent while vegetative growth was not offered a critical clue that, coupled with long‐term precipitation data, helped answer this question.


    At two sites in the Chihuahuan Desert in southern New Mexico, United States, 22 ocotillos were studied annually for more than two decades to explore the relationships among reproductive effort, vegetative stem growth, and patterns of precipitation.

    Key Results

    Vegetative stem growth occurred in mid‐ to late summer (July–September), the season of maximum precipitation in the Chihuahuan Desert, and was significantly related to summer precipitation received in the year of growth. Reproductive effort occurred in early to late spring (April–June), which with winter account for minimum precipitation during the year, but was significantly related to summer precipitation received in the previous year, suggesting the importance of stem succulence and stored water.


    While highly variable summer precipitation was responsible for enormous fluctuations in annual ocotillo stem growth, stem succulence insulated reproductive effort from such immense variability. Stem‐stored water allowed the production of flowers and fruits to proceed relatively consistently during the driest years and during the driest time of year in the Chihuahuan Desert.

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  8. Abstract

    Regional long‐term monitoring can enhance the detection of biodiversity declines associated with climate change, improving future projections by reducing reliance on space‐for‐time substitution and increasing scalability. Rodents are diverse and important consumers in drylands, regions defined by the scarcity of water that cover 45% of Earth's land surface and face increasingly drier and more variable climates. We analyzed abundance data for 22 rodent species across grassland, shrubland, ecotone, and woodland ecosystems in the southwestern USA. Two time series (1995–2006 and 2004–2013) coincided with phases of the Pacific Decadal Oscillation (PDO), which influences drought in southwestern North America. Regionally, rodent species diversity declined 20%–35%, with greater losses during the later time period. Abundance also declined regionally, but only during 2004–2013, with losses of 5% of animals captured. During the first time series (wetter climate), plant productivity outranked climate variables as the best regional predictor of rodent abundance for 70% of taxa, whereas during the second period (drier climate), climate best explained variation in abundance for 60% of taxa. Temporal dynamics in diversity and abundance differed spatially among ecosystems, with the largest declines in woodlands and shrublands of central New Mexico and Colorado. Which species were winners or losers under increasing drought and amplified interannual variability in drought depended on ecosystem type and the phase of the PDO. Fewer taxa were significant winners (18%) than losers (30%) under drought, but the identities of winners and losers differed among ecosystems for 70% of taxa. Our results suggest that the sensitivities of rodent species to climate contributed to regional declines in diversity and abundance during 1995–2013. Whether these changes portend future declines in drought‐sensitive consumers in the southwestern USA will depend on the climate during the next major PDO cycle.

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  9. Abstract

    Woody plant encroachment is a global phenomenon whereby shrubs or trees replace grasses. The hydrological consequences of this ecological shift are of broad interest in ecohydrology, yet little is known of how plant and intercanopy patch dynamics, distributions, and connectivity influence catchment‐scale responses. To address this gap, we established research catchments in the Sonoran and Chihuahuan Deserts (near Green Valley, Arizona and near Las Cruces, New Mexico, respectively) that represent shrub encroachment in contrasting arid climates. Our main goals in the coordinated observations were to: (a) independently measure the components of the catchment water balance, (b) deploy sensors to quantify the spatial patterns of ecohydrological processes, (c) use novel methods for characterizing catchment properties, and (d) assess shrub encroachment impacts on ecohydrological processes through modelling studies. Datasets on meteorological variables; energy, radiation, and CO2fluxes; evapotranspiration; soil moisture and temperature; and runoff at various scales now extend to nearly 10 years of observations at each site, including both wet and dry periods. Here, we provide a brief overview of data collection efforts and offer suggestions for how the coordinated datasets can be exploited for ecohydrological inferences and modelling studies. Given the representative nature of the catchments, the available databases can be used to generalize findings to other catchments in desert landscapes.

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  10. Abstract

    Primary production, a key regulator of the global carbon cycle, is highly responsive to variations in climate. Yet, a detailed, continental‐scale risk assessment of climate‐related impacts on primary production is lacking. We combined 16 years of MODIS NDVI data, a remotely sensed proxy for primary production, with observations from 1218 climate stations to derive values of ecosystem sensitivity to precipitation and aridity. For the first time, we produced an empirically‐derived map of ecosystem sensitivity to climate across the conterminous United States. Over this 16‐year period, annual primary production values were most sensitive to precipitation and aridity in dryland and grassland ecosystems. Century‐long trends measured at the climate stations showed intensifying aridity and climatic variability in many of these sensitive regions. Dryland ecosystems in the western US may be particularly vulnerable to reductions in primary production and consequent degradation of ecosystem services as climate change and variability increase in the future.

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