Historical shrub–grass transitions in the northern Chihuahuan Desert: modeling the effects of shifting rainfall seasonality and event size over a landscape gradient

We use a spatially explicit landscape model to investigate the potential role of rainfall on shrub–grass transitions in the Jornada Basin of southern New Mexico during the past century. In long‐term simulations (1915–1998) along a 2700 m transect running from a dry lake bed to the foothills of a small mountain, we test two hypotheses: (i) that wetter winters and drier summers may have facilitated shrub encroachment in grasslands, and (ii) that increases in large precipitation events may have increased soil water recharge at deeper layers, thus favoring shrub establishment and growth. Our model simulations generally support the hypothesis that wetter winters and drier summers may have played a key role, but we are unable to reproduce the major shifts from grass‐ to shrub‐domination that occurred in this landscape during the early part of the 1900s; furthermore, the positive shrub response to wetter winters and drier summers was only realized subsequent to the drought of 1951–1956, which was a relatively short ‘window of opportunity’ for increased shrub establishment and growth. Our simulations also generally support the hypothesis that an increase in the number of large precipitation events may also have favored shrub establishment and growth, although these results are equivocal, depending upon what constitutes a ‘large’ event and the timing of such events. We found complex interactions among (i) the amount/seasonality of rainfall, (ii) its redistribution in the landscape via run‐on and runoff, (iii) the depth of the soil water recharge, and (iv) subsequent water availability for the growth and reproduction of shrubs vs. herbaceous plants at various landscape positions. Our results suggest that only a mechanistic understanding of these interactions, plus the role of domestic cattle grazing, will enable us to elucidate fully the relative importance of biotic vs. abiotic factors in vegetation dynamics in this semiarid landscape.     

Heterogeneity of soil and plant N and C associated with individual plants and openings in North American shortgrass steppe

Small-scale spatial heterogeneity of soil organic matter (SOM) associated with patterns of plant cover can strongly influence population and ecosystem dynamics in dry regions but is not well characterized for semiarid grasslands. We evaluated differences in plant and soil N and C between soil from under individual grass plants and from small openings in shortgrass steppe. In samples from 0 to 5 cm depth, root biomass, root N, total and mineralizable soil N, total and respirable organic C, C:N ratio, fraction of organic C respired, and ratio of respiration to N mineralization were significantly greater for soil under plants than soil from openings. These differences, which were consistent for two sites with contrasting soil textures, indicate strong differentiation of surface soil at the scale of individual plants, with relative enrichment of soil under plants in total and active SOM. Between-microsite differences were substantial relative to previously reported differences associated with landscape position and grazing intensity in shortgrass steppe. We conclude that microscale heterogeneity in shortgrass steppe deserves attention in investigation of controls on ecosystem and population processes and when sampling to estimate properties at plot or site scales.     

Nitrogen mineralization in native cultivated and abandoned fields in shortgrass steppe

We conducted a set of in situ incubations to evaluate patterns of N availability among dominant land uses in the shortgrass steppe region of Colorado, USA, and to assess recovery of soil fertility in abandoned fields. Replicated 30 d incubations were performed in 3 sets of native (never cultivated), abandoned (cultivated until 1937), and currently cultivated, fallow fields. Net N mineralization and the percentage of total N that was mineralized increased in the order: native, abandoned, cultivated. Higher soil water content in fallow fields is the most likely reason for greater mineralization in cultivated fields, while higher total organic C and C/N ratios in native and abandoned fields may explain differences in mineralization between these land uses. Recovery of soil organic matter in abandoned fields appears to involve accumulation of soil C and N under perennial plants, but probable methodological artifacts complicate evaluation of the role of individual plants in recovery of N availability. Higher N mineralization and turnover in cultivated fields may make them more susceptible to N losses; recovery of N cycling in abandoned fields appears to involve a return to slower N turnover and tighter N cycling similar to native shortgrass steppe.     

Antibiotic-resistantArthrobacter sp. andPseudomonas sp. responses in the rhizosphere of blue grama after herbage removal

Summary Streptomycin-resistant Pseudomonas and Arthrobacter were isolated from semi-arid grassland soil and their relative responses in the rhizosphere of blue grama (Bouteloua gracilis) subjected to herbage removal were evaluated. Using plants grown in normal soil, the two bacteria showed differential responses to herbage removal, which were most marked in the rhizoplane, where the Pseudomonas showed a two-log unit increase over a 60 hour period, while Arthrobacter, in contrast, exhibited a one-log unit decrease in viable counts for at least 48 hours after defoliation, responses which are similar to those observed in root exudate medium experiments by earlier workers. These results suggest that the rhizoplane may be a critical environment for interaction of these two types of microorganisms, and that sequential responses of the root-associated soil microorganisms may occur after herbage removal from this important rangeland plant. These responses are most likely associated with increased exudate release following herbage removal, which has been best documented using blue grama grown under sterile conditions.     

Environmental variation shapes genetic variation in Bouteloua gracilis: Implications for restoration management of natural populations and cultivated varieties in the southwestern United States

With the increasing frequency of large‐scale restoration efforts, the need to understand the adaptive genetic structure of natural plant populations and their relation to heavily utilized cultivars is critical. Bouteloua gracilis (blue grama) is a wind‐dispersed, perennial grass consisting of several cytotypes (2n = 2×–6×) with a widespread distribution in western North America. The species is locally dominant and used regularly in restoration treatments. Using amplified fragment length polymorphism (AFLP) and cpDNA analyses, we assessed the genetic variability and adaptive genetic structure of blue grama within and among 44 sampling sites that are representative of the species’ environmental and habitat diversity in the southwestern United States. Five cultivars were also included to investigate genetic diversity and differentiation in natural versus cultivated populations. Three main findings resulted from this study: (a) Ninety‐four polymorphic AFLP markers distinguished two population clusters defined largely by samples on and off the Colorado Plateau; (b) substructure of samples on the Colorado Plateau was indicated by genetic divergence between boundary and interior regions, and was supported by cytotype distribution and cpDNA analysis; and (c) six AFLP markers were identified as “outliers,” consistent with being under selection. These loci were significantly correlated to mean annual temperature, mean annual precipitation, precipitation of driest quarter, and precipitation of wettest quarter in natural populations, but not in cultivated samples. Marker × environment relationships were found to be largely influenced by cytotype and cultivar development. Our results demonstrate that blue grama is genetically variable, and exhibits genetic structure, which is shaped, in part, by environmental variability across the Colorado Plateau. Information from our study can be used to guide the selection of seed source populations for commercial development and long‐term conservation management of B. gracilis, which could include genetic assessments of diversity and the adaptive potential of both natural and cultivated populations for wildland restoration.     

Partitioning evapotranspiration fluxes from a Colorado grassland using stable isotopes: Seasonal variations and ecosystem implications of elevated atmospheric CO2

The stable isotopic composition of soil water is controlled by precipitation inputs, antecedent conditions, and evaporative losses. Because transpiration does not fractionate soil water isotopes, the relative proportions of evaporation and transpiration can be estimated using a simple isotopic mass balance approach. At our site in the shortgrass steppe in semi-arid northeastern Colorado, ¹⁸O values of soil water were almost always more enriched than those of precipitation inputs, owing to evaporative losses. The proportion of water lost by evaporation (E/ET) during the growing season ranged from nil to about 40% (to >90% in the dormant season), and was related to the timing of precipitation inputs. The sum of transpiration plus evaporation losses estimated by isotopic mass balance were similar to actual evapotranspiration measured from a nearby Bowen ratio system. We also investigated the evapotranspiration response of this mixed C₃/C₄ grassland to doubled atmospheric [CO₂] using Open-Top Chambers (OTC). Elevated atmospheric [CO₂] led to increased soil-water conservation via reduced stomatal conductance, despite greater biomass growth. We used a non-invasive method to measure the ¹⁸O of soil CO₂ as a proxy for soil water, after establishing a strong relationship between ¹⁸O of soil CO₂ from non-chambered control (NC) plots and ¹⁸O of soil–water from an adjacent area of native grassland. Soil–CO₂ ¹⁸O values showed significant treatment effects, particularly during a dry summer: values in ambient chambers (AC) were more enriched than in NC and elevated chamber (EC) plots. During the dry growing season of 2000, transpiration from the EC treatment was higher than from AC and lower than from NC treatments, but during 2001, transpiration was similar on all three treatments. Slightly higher evaporation rates from AC than either EC or NC treatments in 2000 may have resulted from increased convection across the soil surface from the OTC blowers, combined with lower biomass and litter cover on the AC treatment. Transpiration-use efficiency, or the amount of above-ground biomass produced per mm water transpired, was always greatest on EC and lowest on NC treatments.     

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.     

Population and clonal level responses of a perennial grass following fire in the northern Chihuahuan Desert

Relationships involving fire and perennial grasses are controversial in Chihuahuan Desert grasslands of southern New Mexico, USA. Research suggests that fire delays the resprouting of perennial grasses well after two growing seasons. However, such results are confounded by livestock grazing, soil erosion, and drought. Additionally, post-fire grass responses may depend on initial clone size. We evaluated the effects of fire, grazing, and clone size on Bouteloua eriopoda (black grama) in southern New Mexico grasslands. Four 2-ha plots were established in each of four sites. Fire and grazing were applied or not applied in 1999 such that four treatment combinations were assigned randomly to plots within each site. Within each plot, small (0–10 cm² basal area), medium (10–30 cm²), and large ( > 30 cm²) clones were initially mapped in five 0.91-m² quadrats where grass attributes and litter cover were evaluated before and at the end of two growing seasons following fire. Maximum fire temperature was also measured. At a population level, canopy and litter cover were each approximately 50% less in burned than unburned areas. However, compared to initial levels, canopy height had increased by 10% at the end of the study, regardless of fire. At a clonal level, basal cover reductions were attributed mostly to large clones that survived fire. Smaller clone densities had decreased by as much as 19% in burned compared to unburned areas, and fire reduced the basal cover of medium clones. Basal and canopy cover, recruitment, and clone basal area decreased with increased fire temperatures. Almost all responses were independent of grazing, and interactive effects of grazing and fire were not detected. Fire did not kill all perennial grass clones, regardless of size. However, rapid responses were likely influenced by above-average precipitation after fire. Future studies in desert grasslands should examine how perennial grass dynamics are affected by fire, precipitation patterns, and interactions with grazing.     

Cost of resistance and tolerance under competition: the defense-stress benefit hypothesis

Defense costs provide a major explanation for why plants in nature have not evolved to be better defended against pathogens and herbivores; however, evidence for defense costs is often lacking. Plants defend by deploying resistance traits that reduce damage, and tolerance traits that reduce the fitness effects of damage. We first tested the defense-stress cost (DSC) hypothesis that costs of defenses increase and become important under competitive stress. In a greenhouse experiment, uniparental maternal families of the host plant Arabis perennans were grown in the presence and absence of the bunch grass Bouteloua gracilis and the herbivore Plutella xylostella. Costs of resistance and tolerance manifest as reduced growth in the absence of herbivory were significant when A. perennans grew alone, but not in the competitive environment, in contrast to the DSC hypothesis. We then tested the defense-stress benefit (DSB) hypothesis that plant defenses may benefit plants in competitive situations thereby reducing net costs. For example, chemical resistance agents and tolerance may also have functions in competitive interactions. To test the DSB hypothesis, we compared differentially competitive populations for defense costs, assuming that poorer competitors from less dense habitats were less likely to have evolved defenses that also function in competition. Without competitive benefits of defenses, poorer competitors were expected to have higher net costs of defenses under competition in accordance with DSB. Populations of A. perennans and A. drummondii that differed dramatically in competitiveness were compared for costs, and as the DSB hypothesis predicts, only the poor competitor population showed costs of resistance under competition. However, cost of tolerance under competition did not differ among populations, suggesting that the poor competitors might have evolved a general stress tolerance. Although the DSC hypothesis may explain cases where defense costs increase under stress, the DSB hypothesis may explain some cases where costs decrease under competitive stress.