Environmental controls on nitric oxide emission from northern Chihuahuan desert soils

A survey of nitric oxide (NO) emission from Chihuahuan desert soils found mean NO fluxes <0.1 ng NO-N cm^–2h^–1 during the dry season. These fluxes were at thelower end of the range reported for temperate grassland and woodlandecosystems. NO fluxes from wet or watered soils were higher(0.1–35 ng NO-N cm^–2 h^–1).Watering of black grama grassland soils produced an initial pulse of 12ng cm^–2 h^–1 (12-h after 1-cm watering)with high fluxes sustained over 4 days with repeated watering. Initialpulses from shrubland soils were lower (maximum 5 ngcm^–2 h^–1), and fluxes declined withrepeated watering. Repeated watering of creosotebush soils depleted thesoil NH ₄ ⁺ pool, and NO emissions weredirectly related to soil NH ₄ ⁺ concentrationsat the end of the experiment. In watered andNH ₄ ⁺ -fertilized creosotebush soils, NO fluxeswere positively related to potential net nitrification rates.NH ₄ ⁺ -fertilization boosted the initial NOpulse 15 times in the shrubland and 5 times in black grama grasslandrelative to watered controls. These experimental results point towardgreater substrate limitation in shrublands. In this desert basin, NOemission averaged 0.12 kg N ha^–1 y^–1in untreated soil and 0.76 kg N ha^–1y^–1 in watered soil. We multiplied these averages bythe distribution of grassland and shrubland vegetation within a58,600-ha area of the Jornada del Muerto basin to estimate regionallosses of 0.15–0.38 kg NO-N ha^–1y^–1 for this area of the Chihuahuan desert.     

As above,not so below: Long-term dynamics of net primary production across a dryland transition zone

Drylands are key contributors to interannual variation in the terrestrial carbon sink, which has been attributed primarily to broad-scale climatic anomalies that disproportionately affect net primary production (NPP) in these ecosystems. Current knowledge around the patterns and controls of NPP is based largely on measurements of aboveground net primary production (ANPP), particularly in the context of altered precipitation regimes. Limited evidence suggests belowground net primary production (BNPP), a major input to the terrestrial carbon pool, may respond differently than ANPP to precipitation, as well as other drivers of environmental change, such as nitrogen deposition and fire. Yet long-term measurements of BNPP are rare, contributing to uncertainty in carbon cycle assessments. Here, we used 16 years of annual NPP measurements to investigate responses of ANPP and BNPP to several environmental change drivers across a grassland–shrubland transition zone in the northern Chihuahuan Desert. ANPP was positively correlated with annual precipitation across this landscape; however, this relationship was weaker within sites. BNPP, on the other hand, was weakly correlated with precipitation only in Chihuahuan Desert shrubland. Although NPP generally exhibited similar trends among sites, temporal correlations between ANPP and BNPP within sites were weak. We found chronic nitrogen enrichment stimulated ANPP, whereas a one-time prescribed burn reduced ANPP for nearly a decade. Surprisingly, BNPP was largely unaffected by these factors. Together, our results suggest that BNPP is driven by a different set of controls than ANPP. Furthermore, our findings imply belowground production cannot be inferred from aboveground measurements in dryland ecosystems. Improving understanding around the patterns and controls of dryland NPP at interannual to decadal scales is fundamentally important because of their measurable impact on the global carbon cycle. This study underscores the need for more long-term measurements of BNPP to improve assessments of the terrestrial carbon sink, particularly in the context of ongoing environmental change.     

Mechanisms for the keystone status of kangaroo rats: graminivory rather than granivory?

Graminivory by kangaroo rats (Dipodomys spp.) was investigated as a potential mechanism for the keystone role of these rodents in the dynamics of desert grasslands. Experiments confirmed that Ord's kangaroo rats (Dipodomys ordii) cut and consumed a large proportion of the tillers of three Chihuahuan Desert tussock-forming grass species. Field observations indicated that the characteristically cut grass tillers were absent from all-rodent and medium-sized kangaroo rat exclosures, but were frequent in large-sized kangaroo rat and rabbit exclosures, indicating that the medium-sized kangaroo rats (D. ordii, D. merriami) were responsible for grass cutting. Tiller waste as a percentage of peak standing crop ranged from 7% in grassland habitats to 0.7% in Flourensia cernua shrubland. Of the 13 species of perennial, tussock-forming grasses measured, only one, Muhlenbergia porteri, had no tillers cut by kangaroo rats. This study demonstrates that the keystone role of kangaroo rats in Chihuahuan Desert grassland ecosystems is probably the result of their graminivory. Received: 28 October 1996 / Accepted: 26 February 1997     

Nutrient losses in runoff from grasslandand shrubland habitats in southern New Mexico: II. Field plots

Losses of dissolved nutrients (N, P, K, Ca, Mg, Na,Cl, and SO₄) in runoff were measured on grasslandand shrubland plots in the Chihuahuan desert ofsouthern New Mexico. Runoff began at a lowerthreshold of rainfall in shrublands than ingrasslands, and the runoff coefficient averaged 18.6%in shrubland plots over a 7-year period. In contrast,grassland plots lost 5.0 to 6.3% of incidentprecipitation in runoff during a 5.5-year period. Nutrient losses from shrubland plots were greater thanfrom grassland plots, with nitrogen losses averaging0.33 kg ha^–1 yr^–1 vs0.15 kg ha^–1 yr^–1, respectively, during a 3-year period. Thegreater nutrient losses in shrublands were due tohigher runoff, rather than higher nutrientconcentrations in runoff. In spite of these nutrientlosses in runoff, all plots showed net accumulationsof most elements due to inputs from atmosphericdeposition. Therefore, loss of soil nutrients byhillslope runoff cannot, by itself, account for thedepletion of soil fertility associated withdesertification in the Chihuahuan desert.     

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.     

Effects of an increase in summer precipitation on leaf, soil, and ecosystem fluxes of CO2 and H2O in a sotol grassland in Big Bend National Park, Texas

Global climate models predict that in the next century precipitation in desert regions of the USA will increase, which is anticipated to affect biosphere/atmosphere exchanges of both CO₂ and H₂O. In a sotol grassland ecosystem in the Chihuahuan Desert at Big Bend National Park, we measured the response of leaf-level fluxes of CO₂ and H₂O 1 day before and up to 7 days after three supplemental precipitation pulses in the summer (June, July, and August 2004). In addition, the responses of leaf, soil, and ecosystem fluxes of CO₂ and H₂O to these precipitation pulses were also evaluated in September, 1 month after the final seasonal supplemental watering event. We found that plant carbon fixation responded positively to supplemental precipitation throughout the summer. Both shrubs and grasses in watered plots had increased rates of photosynthesis following pulses in June and July. In September, only grasses in watered plots had higher rates of photosynthesis than plants in the control plots. Soil respiration decreased in supplementally watered plots at the end of the summer. Due to these increased rates of photosynthesis in grasses and decreased rates of daytime soil respiration, watered ecosystems were a sink for carbon in September, assimilating on average 31 mmol CO₂ m⁻² s⁻¹ ground area day⁻¹. As a result of a 25% increase in summer precipitation, watered plots fixed eightfold more CO₂ during a 24-h period than control plots. In June and July, there were greater rates of transpiration for both grasses and shrubs in the watered plots. In September, similar rates of transpiration and soil water evaporation led to no observed treatment differences in ecosystem evapotranspiration, even though grasses transpired significantly more than shrubs. In summary, greater amounts of summer precipitation may lead to short-term increased carbon uptake by this sotol grassland ecosystem.     

Soil Microbial Community Response to Drought and Precipitation Variability in the Chihuahuan Desert

Increases in the magnitude and variability of precipitation events have been predicted for the Chihuahuan Desert region of West Texas. As patterns of moisture inputs and amounts change, soil microbial communities will respond to these alterations in soil moisture windows. In this study, we examined the soil microbial community structure within three vegetation zones along the Pine Canyon Watershed, an elevation and vegetation gradient in Big Bend National Park, Chihuahuan Desert. Soil samples at each site were obtained in mid-winter (January) and in mid-summer (August) for 2 years to capture a component of the variability in soil temperature and moisture that can occur seasonally and between years along this watershed. Precipitation patterns and amounts differed substantially between years with a drought characterizing most of the second year. Soils were collected during the drought period and following a large rainfall event and compared to soil samples collected during a relatively average season. Structural changes within microbial community in response to site, season, and precipitation patterns were evaluated using fatty acid methyl ester (FAME) and polymerase chain reaction-denaturing gradient gel electrophoresis (PCR-DGGE) analyses. Fungal FAME amounts differed significantly across seasons and sites and greatly outweighed the quantity of bacterial and actinomycete FAME levels for all sites and seasons. The highest fungal FAME levels were obtained in the low desert scrub site and not from the high elevation oak–pine forests. Total bacterial and actinomycete FAME levels did not differ significantly across season and year within any of the three locations along the watershed. Total bacterial and actinomycete FAME levels in the low elevation desert-shrub and grassland sites were slightly higher in the winter than in the summer. Microbial community structure at the high elevation oak–pine forest site was strongly correlated with levels of NH₄ ⁺–N, % soil moisture, and amounts of soil organic matter irrespective of season. Microbial community structure at the low elevation desert scrub and sotol grasslands sites was most strongly related to soil pH with bacterial and actinobacterial FAME levels accounting for site differences along the gradient. DGGE band counts of amplified soil bacterial DNA were found to differ significantly across sites and season with the highest band counts found in the mid-elevation grassland site. The least number of bands was observed in the high elevation oak–pine forest following the large summer-rain event that occurred after a prolonged drought. Microbial responses to changes in precipitation frequency and amount due to climate change will differ among vegetation zones along this Chihuahuan Desert watershed gradient. Soil bacterial communities at the mid-elevation grasslands site are the most vulnerable to changes in precipitation frequency and timing, while fungal community structure is most vulnerable in the low desert scrub site. The differential susceptibility of the microbial communities to changes in precipitation amounts along the elevation gradient reflects the interactive effects of the soil moisture window duration following a precipitation event and differences in soil heat loads. Amounts and types of carbon inputs may not be as important in regulating microbial structure among vegetation zones within in an arid environment as is the seasonal pattern of soil moisture and the soil heat load profile that characterizes the location.     

Intra-annual rainfall variability and grassland productivity: can the past predict the future?

Precipitation quantity has been shown to influence grassland aboveground net primary productivity (ANPP) positively whereas experimental increases in of temporal variability in water availability commonly exhibit a negative relationship with ANPP. We evaluated long term ANPP datasets from the Konza Prairie Long Term Ecological Research (LTER) program (1984–1999) to determine if similar relationships could be identified based on patterns of natural variability (magnitude and timing) in precipitation. ANPP data were analyzed from annually burned sites in native mesic grassland and productivity was partitioned into graminoid (principally C₄ grasses) and forb (C₃ herbaceous) components. Although growing season precipitation amount was the best single predictor of total and grass ANPP (r ²=0.62), several measures of precipitation variability were also significantly and positively correlated with productivity, independent of precipitation amount. These included soil moisture variability, expressed as CV, for June (r ²=0.45) and the mean change in soil moisture between weekly sampling periods in June and August (%wv) (r ²=0.27 and 0.32). In contrast, no significant relationships were found between forb productivity and any of the precipitation variables (p>0.05). A multiple regression model combining precipitation amount and both measures of soil moisture variability substantially increased the fit with productivity (r ²=0.82). These results were not entirely consistent with those of short-term manipulative experiments in the same grassland, however, because soil moisture variability was often positively, not negatively related to ANPP. Differences in results between long and short term experiments may be due to low variability in the historic precipitation record compared to that imposed experimentally as experimental levels of variability exceeded the natural variability of this dataset by a factor of two. Thus, forecasts of ecosystem responses to climate change (i.e. increased climatic variability), based on data constrained by natural and recent historical rainfall patterns may be inadequate for assessing climate change scenarios if precipitation variability in the future is expected to exceed current levels.     

Grassland to shrubland state transitions enhance carbon sequestration in the northern Chihuahuan Desert

The replacement of native C₄‐dominated grassland by C₃‐dominated shrubland is considered an ecological state transition where different ecological communities can exist under similar environmental conditions. These state transitions are occurring globally, and may be exacerbated by climate change. One consequence of the global increase in woody vegetation may be enhanced ecosystem carbon sequestration, although the responses of arid and semiarid ecosystems may be highly variable. During a drier than average period from 2007 to 2011 in the northern Chihuahuan Desert, we found established shrubland to sequester 49 g C m^?2 yr^?1 on average, while nearby native C₄ grassland was a net source of 31 g C m^?2 yr^?1 over this same period. Differences in C exchange between these ecosystems were pronounced – grassland had similar productivity compared to shrubland but experienced higher C efflux via ecosystem respiration, while shrubland was a consistent C sink because of a longer growing season and lower ecosystem respiration. At daily timescales, rates of carbon exchange were more sensitive to soil moisture variation in grassland than shrubland, such that grassland had a net uptake of C when wet but lost C when dry. Thus, even under unfavorable, drier than average climate conditions, the state transition from grassland to shrubland resulted in a substantial increase in terrestrial C sequestration. These results illustrate the inherent tradeoffs in quantifying ecosystem services that result from ecological state transitions, such as shrub encroachment. In this case, the deleterious changes to ecosystem services often linked to grassland to shrubland state transitions may at least be partially offset by increased ecosystem carbon sequestration.     

Does shrub invasion indirectly limit grass establishment via seedling herbivory? A test at grassland‐shrubland ecotones

Question: Does shrub invasion at ecotones indirectly limit grass establishment by increasing mammalian seedling herbivory? Location: Chihuahuan Desert, New Mexico, USA. Methods: We tested the hypothesis that herbivore‐related mortality of seedlings of the dominant perennial grass Bouteloua eriopoda would be highest in shrub‐dominated portions of grassland‐shrubland ecotones. We tested the hypothesis in two Chihuahuan Desert sites featuring similar shrub encroachment patterns but different shrub species, grass cover, and different abundances of small mammals. Within each site we transplanted B. eriopoda seedlings to grass‐dominated, middle, and shrub‐dominated positions of replicate ecotones during the time of year (mid‐summer) when they would naturally appear and monitored seedling fates. We estimated population size/activity of putative small mammal herbivores. Results: Seedlings were killed by mammals in greater numbers in shrubland than in grassland or middle ecotone positions at the site with large herbivore numbers. At the site with low herbivore numbers, most seedlings were killed in middle ecotone positions. The abundance patterns of herbivores did not parallel patterns of seedling herbivory across the ecotones or between sites. Conclusions: Seedling herbivory is an important process and is related to vegetation composition, but the mechanisms underlying the relationship are not clear. We speculate that variation in small mammal foraging behavior may contribute to seedling herbivory patterns. Restoration strategies in the Chihuahuan Desert need to account for the abundance and/or behavior of native herbivores.     

Effect of precipitation variability on net primary production and soil respiration in a Chihuahuan Desert grassland

Precipitation regimes are predicted to become more variable with more extreme rainfall events punctuated by longer intervening dry periods. Water‐limited ecosystems are likely to be highly responsive to altered precipitation regimes. The bucket model predicts that increased precipitation variability will reduce soil moisture stress and increase primary productivity and soil respiration in aridland ecosystems. To test this hypothesis, we experimentally altered the size and frequency of precipitation events during the summer monsoon (July through September) in 2007 and 2008 in a northern Chihuahuan Desert grassland in central New Mexico, USA. Treatments included (1) ambient rain, (2) ambient rain plus one 20 mm rain event each month, and (3) ambient rain plus four 5 mm rain events each month. Throughout two monsoon seasons, we measured soil temperature, soil moisture content (θ), soil respiration (R_s), along with leaf‐level photosynthesis (A_net), predawn leaf water potential (Ψ_pd), and seasonal aboveground net primary productivity (ANPP) of the dominant C₄ grass, Bouteloua eriopoda. Treatment plots receiving a single large rainfall event each month maintained significantly higher seasonal soil θ which corresponded with a significant increase in R_s and ANPP of B. eriopoda when compared with plots receiving multiple small events. Because the strength of these patterns differed between years, we propose a modification of the bucket model in which both the mean and variance of soil water change as a consequence of interannual variability from 1 year to the next. Our results demonstrate that aridland ecosystems are highly sensitive to increased precipitation variability, and that more extreme precipitation events will likely have a positive impact on some aridland ecosystem processes important for the carbon cycle.     

Relationship between soil P fractions and microbial biomass in an oligotrophic grassland-desert scrub system

Phosphorus (P) is an essential element of the biosphere, both as a constituent of living organisms and as a regulator of biological processes. The Cuatro Ciénegas Basin in the central Chihuahuan Desert of Mexico is characterized by extreme P oligotrophy. The aim of this study was to quantify P distribution in soil P fractions, P sorption capacity, and P in microbial biomass in a desert scrub and grassland soil system in the Churince area of the Cuatro Ciénegas Basin over summer and winter seasons. Our objective, as part of an exploration of ecosystem functioning, was to ascertain the relationship between soil P fractions and P in microbial biomass. Our results demonstrate a scarcity of P, mainly in grassland, and also a higher P sorption capacity in grassland soil than in desert scrub. Desert scrub soil retained more P (228 ± 5 μg g^?1 dry soil) than grassland soil (87 ± 10 μg g^?1 dry soil), mainly in inorganic forms, but grassland soil retained more P in accessible organic forms. We suggest that biotic controls regulated by access to water shape the dynamics of soil P availability in the Churince grassland-desert scrub system.     

The distribution of vascular plant species and guilds in space and time along a desert gradient

Abstract. We studied distribution patterns of vascular plant species and environmental variables for three years along a permanent transect traversing a closed-drainage watershed in the northern Chihuahuan Desert of south-central New Mexico, USA. The transect extended for 2.7 km from a basin floor playa (1310 m elevation, fine-textured soil), across a piedmont slope, and onto the base of a granitic mountain (1410 m elevation, coarse-textured soil). The gradients in elevation and soils across our transect, along with variable seasonal rainfall, downslope redistribution of water and organic matter, and soil texture-related variation in infiltration, water holding capacity, and moisture release characteristics, interact to generate a complex spatial and temporal gradient of available soil water and nitrogen. We grouped plant species into guilds according to growth form and photo synthetic pathway type. These guilds are spatially and temporally differentiated along the transect such that particular groups utilize particular seasonal phases or spatial regions of the gradient. We identified six distinct plant communities along the transect. C₄ perennial grasses dominated the mesic/high nitrogen portion of the gradient, which occurred at the highest (upper piedmont grassland, dominated by Bouteloua eriopoda) and lowest (playa, dominated by Panicum obtusum) elevations along the transect. C₃ shrubs were dominant in the xeric/low nitrogen portion of the gradient located near the middle of the transect (bajada shrubland, dominated by Larrea tridentata). C₃ shrubs also dominated a narrow zone of vegetation located adjacent to the playa (playa fringe, dominated by Prosopisglandulosa). C₄ perennial grasses, C₃ subshrubs, and C₃ and C₄ perennial forbs and annuals were co-dominant in the intermediate locations along the gradient, which occurred below (mixed basin slopes) and above (lower piedmont grassland) the bajada shrubland. Life-form distribution patterns at the small scale of our study reflect some of the patterning that occurs at larger scales in response to climate gradients. The distributions of some species and guilds along the transect are apparently modified by competitive interactions.     

The Influence of Spatial Patterns of Soil Moisture on the Grass and Shrub Responses to a Summer Rainstorm in a Chihuahuan Desert Ecotone

The cycling of surface water, energy, nutrients, and carbon is different between semiarid grassland and shrubland ecosystems. Although differences are evident when grasslands are compared to shrublands, the processes that contribute to this transition are more challenging to document. We evaluate how surface redistribution of precipitation and plant responses to the resulting infiltration patterns could contribute to the changes that occur during the transition from grassland to shrubland. We measured soil water potential under grasses (Bouteloua eriopoda), shrubs (Larrea tridentata) and bare soil and changes in plant water relations and gas exchange following a 15 mm summer storm in the grassland–shrubland ecotone at the Sevilleta National Wildlife Refuge in central New Mexico USA. Following the storm, soil water potential (Ψ_s) increased to 30 cm depth beneath both grass and shrub canopies, with the greatest change observed in the top 15 cm of the soil. The increase in Ψ_s was greater beneath grass canopies than beneath shrub canopies. Ψ_s under bare soil increased only to 5 cm depth. The substantial redistribution of rainfall and different rooting depths of the vegetation resulted in high Ψ_s throughout most of the rooting volume of the grasses whereas soil moisture was unchanged throughout a large portion of the shrub rooting volume. Consistent with this pattern, predawn water potential (Ψ_PD) of grasses increased more than 5 MPa to greater than −1 MPa whereas Ψ_PD of shrubs increased to −2.5 MPa, a change of less than 2 MPa. Transpiration increased roughly linearly with Ψ_PD in both grasses and shrubs. In grasses, assimilation was strongly correlated with Ψ_PD whereas there was no relationship in shrubs where assimilation showed no significant response to the pulse of soil moisture following the storm. These data show that preferential redistribution of water to grass canopies enhances transpiration and assimilation by grasses following large summer storms. This process may inhibit shrubland expansion at the ecotone during periods without extreme drought.     

Physiological responses of two contrasting desert plant species to precipitation variability are differentially regulated by soil moisture and nitrogen dynamics

Alterations in global and regional precipitation patterns are expected to affect plant and ecosystem productivity, especially in water‐limited ecosystems. This study examined the effects of natural and supplemental (25% increase) seasonal precipitation on a sotol grassland ecosystem in Big Bend National Park in the Chihuahuan Desert. Physiological responses – leaf photosynthesis at saturating light (A_sat), stomatal conductance (g_s), and leaf nitrogen [N] – of two species differing in their life form and physiological strategies (Dasylirion leiophyllum, a C₃ shrub; Bouteloua curtipendula, a C₄ grass) were measured over 3 years (2004–2006) that differed greatly in their annual and seasonal precipitation patterns (2004: wet, 2005: average, 2006: dry). Precipitation inputs are likely to affect leaf‐level physiology through the direct effects of altered soil water and soil nitrogen. Thus, the effects of precipitation, watering treatment, soil moisture, and nitrogen were quantified via multivariate hierarchical Bayesian models that explicitly linked the leaf and soil responses. The two species differed in their physiological responses to precipitation and were differentially controlled by soil water vs. soil nitrogen. In the relatively deeply rooted C₃ shrub, D. leiophyllum, A_sat was highest in moist periods and was primarily regulated by deep (16–30 cm) soil water. In the shallow‐rooted C₄ grass, B. curtipendula, A_sat was only coupled to leaf [N], both of which increased in dry periods when soil [N] was highest. Supplemental watering during the wet year generally decreased A_sat and leaf [N] in D. leiophyllum, perhaps due to nutrient limitation, and physiological responses in this species were influenced by the cumulative effects of 5 years of supplemental watering. Both species are common in this ecosystem and responded strongly, yet differently, to soil moisture and nitrogen, suggesting that changes in the timing and magnitude of precipitation may have consequences for plant carbon gain, with the potential to alter community composition.     

Soil fungal composition changes with shrub encroachment in the northern Chihuahuan Desert

Woody species encroachment of grasslands globally causes many socioecological impacts, including loss of grazing pastures and decreased biodiversity. Soil microbial communities may partially regulate the pace of shrub encroachment, as plant-microbial interactions can strongly influence plant success. We measured fungal composition and activity under dominant plant species across a grassland to shrubland transition to determine if shrubs cultivate soil microbial communities as they invade. Specifically, soil microbial communities, abiotic soil properties, and extracellular enzyme activities were quantified for soils under four common Chihuahuan Desert plant species (three grasses, one shrub) in central New Mexico, U.S.A. Extracellular enzyme activity levels were fairly consistent under different plant species across the grassland to shrubland transition. Activity levels of two enzymes (alkaline phosphatase and beta-N-acetyl-glucosaminidase) were lower in the ecotone, presumably because soil organic matter content was also lower in ecotone soils. Community composition of soil fungi mirrored patterns in the plant community, with distinct plant and fungal communities in the shrubland and grassland, while grassland-shrubland ecotone soils hosted a mix of taxa from both habitats. We show that shrubs cultivate a distinct microbial community on the leading edge of the invasion, which may be necessary for shrub colonization, establishment, and persistence.     

Do increased summer precipitation and N deposition alter fine root dynamics in a Mojave Desert ecosystem?

Climate change is expected to impact the amount and distribution of precipitation in the arid southwestern United States. In addition, nitrogen (N) deposition is increasing in these regions due to increased urbanization. Responses of belowground plant activity to increases in soil water content and N have shown inconsistent patterns between biomes. In arid lands, plant productivity is limited by water and N availability so it is expected that changes in these factors will affect fine root dynamics. The objectives of this study were to quantify the effects of increased summer precipitation and N deposition on fine root dynamics in a Mojave Desert ecosystem during a 2‐year field experiment using minirhizotron measurements. Root length density, production, and mortality were measured in field plots in the Mojave Desert receiving three 25 mm summer rain events and/or 40 kg N ha^?1 yr^?1. Increased summer precipitation and N additions did not have an overall significant effect on any of the measured root parameters. However, differences in winter precipitation resulting from interannual variability in rainfall appeared to affect root parameters with root production and turnover increasing following a wet winter most likely due to stimulation of annual grasses. In addition, roots were distributed more deeply in the soil following the wet winter. Root length density was initially higher under canopies compared to canopy interspaces, but converged toward the end of the study. In addition, roots tended to be distributed more deeply into the soil in canopy interspace areas. Results from this study indicated that increased summer precipitation and N deposition in response to climate change and urbanization are not likely to affect fine root dynamics in these Mojave Desert ecosystems, despite studies showing aboveground plant physiological responses to these environmental perturbations. However, changes in the amount and possibly distribution of winter precipitation may affect fine root dynamics.     

Response of dominant grass and shrub species to water manipulation: an ecophysiological basis for shrub invasion in a Chihuahuan Desert Grassland

Increases in woody vegetation and declines in grasses in arid and semi-arid ecosystems have occurred globally since the 1800s, but the mechanisms driving this major land-cover change remain uncertain and controversial. Working in a shrub-encroached grassland in the northern Chihuahuan Desert where grasses and shrubs typically differ in leaf-level nitrogen allocation, photosynthetic pathway, and root distribution, we asked if differences in leaf-level ecophysiology could help explain shrub proliferation. We predicted that the relative performance of grasses and shrubs would vary with soil moisture due to the different morphological and physiological characteristics of the two life-forms. In a 2-year experiment with ambient, reduced, and enhanced precipitation during the monsoon season, respectively, the encroaching C(3) shrub (honey mesquite Prosopis glandulosa) consistently and substantially outperformed the historically dominant C(4) grass (black grama Bouteloua eriopoda) in terms of photosynthetic rates while also maintaining a more favorable leaf water status. These differences persisted across a wide range of soil moisture conditions, across which mesquite photosynthesis was decoupled from leaf water status and moisture in the upper 50 cm of the soil profile. Mesquite's ability to maintain physiologically active leaves for a greater fraction of the growing season than black grama potentially amplifies and extends the importance of physiological differences. These physiological and phenological differences may help account for grass displacement by shrubs in drylands. Furthermore, the greater sensitivity of the grass to low soil moisture suggests that grasslands may be increasingly susceptible to shrub encroachment in the face of the predicted increases in drought intensity and frequency in the desert of the southwestern USA.     

Press–pulse interactions: effects of warming,N deposition,altered winter precipitation,and fire on desert grassland community structure and dynamics

Global environmental change is altering temperature, precipitation patterns, resource availability, and disturbance regimes. Theory predicts that ecological presses will interact with pulse events to alter ecosystem structure and function. In 2006, we established a long‐term, multifactor global change experiment to determine the interactive effects of nighttime warming, increased atmospheric nitrogen (N) deposition, and increased winter precipitation on plant community structure and aboveground net primary production (ANPP) in a northern Chihuahuan Desert grassland. In 2009, a lightning‐caused wildfire burned through the experiment. Here, we report on the interactive effects of these global change drivers on pre‐ and postfire grassland community structure and ANPP. Our nighttime warming treatment increased winter nighttime air temperatures by an average of 1.1 °C and summer nighttime air temperature by 1.5 °C. Soil N availability was 2.5 times higher in fertilized compared with control plots. Average soil volumetric water content (VWC) in winter was slightly but significantly higher (13.0% vs. 11.0%) in plots receiving added winter rain relative to controls, and VWC was slightly higher in warmed (14.5%) compared with control (13.5%) plots during the growing season even though surface soil temperatures were significantly higher in warmed plots. Despite these significant treatment effects, ANPP and plant community structure were highly resistant to these global change drivers prior to the fire. Burning reduced the cover of the dominant grasses by more than 75%. Following the fire, forb species richness and biomass increased significantly, particularly in warmed, fertilized plots that received additional winter precipitation. Thus, although unburned grassland showed little initial response to multiple ecological presses, our results demonstrate how a single pulse disturbance can interact with chronic alterations in resource availability to increase ecosystem sensitivity to multiple drivers of global environmental change.     

The role of vegetation–microclimate feedback in promoting shrub encroachment in the northern Chihuahuan desert

Many arid and semi‐arid landscapes around the world are affected by a shift from grassland to shrubland vegetation, presumably induced by climate warming, increasing atmospheric CO₂ concentrations, and/or changing land use. This major change in vegetation cover is likely sustained by positive feedbacks with the physical environment. Recent research has focused on a feedback with microclimate, whereby cold intolerant shrubs increase the minimum nocturnal temperatures in their surroundings. Despite the rich literature on the impact of land cover change on local climate conditions, changes in microclimate resulting from shrub expansion into desert grasslands have remained poorly investigated. It is unclear to what extent such a feedback can affect the maximum extent of shrub expansion and the configuration of a stable encroachment front. Here, we focus on the case of the northern Chihuahuan desert, where creosotebush (Larrea tridentata) has been replacing grasslands over the past 100–150 years. We use a process‐based coupled atmosphere‐vegetation model to investigate the role of this feedback in sustaining shrub encroachment in the region. Simulations indicate that the feedback allows juvenile shrubs to establish in the grassland during average years and, once established, reduce their vulnerability to freeze‐induced mortality by creating a warmer microclimate. Such a feedback is crucial in extreme cold winters as it may reduce shrub mortality. We identify the existence of a critical zone in the surroundings of the encroachment front, in which vegetation dynamics are bistable: in this zone, vegetation can be stable both as grassland and as shrubland. The existence of these alternative stable states explains why in most cases the shift from grass to shrub cover is found to be abrupt and often difficult to revert.