Ecosystem consequences of plant life form changes at three sites in the semiarid United States

Many semiarid rangelands have recently experienced changes in dominant plant life form. Both woody plant expansion into grasslands and the invasion of annual grasses into shrublands have potential influence on regional carbon cycling. Soil carbon content, chemistry, and distribution may change following shifts in dominant plant life form because plant life forms differ in litter chemistry and patterns of detrital input. This study assesses the amount, quality, and distribution of soil C below woody vegetation and grasses at three rangelands in Texas, New Mexico, and Utah. At each of these sites there has been a well-documented shift in dominant plant life form. In Texas and New Mexico, woody plants have increased in grasslands, while grasses have invaded into former shrublands in Utah. We measured total soil carbon, particulate organic matter (POM) C, and the carbon isotopic composition of soil carbon beneath woody plants and grasses at each of these three sites. At the La Copita Research Area in south-central Texas there was significantly more soil C found beneath Prosopis glandulosa, the dominant woody plant, than was found beneath grasses. Mean soil C content to 1 m was 7.2 kg C m^–2 beneath P. glandulosa and 6.0 kg C m^–2 beneath grasses. There was also significantly more POM C beneath P. glandulosa than beneath grasses. Stable carbon isotopic composition indicated that the expansion of P. glandulosa in savannas in Texas first influences carbon cycling in surface soils, then deep soil C, and finally throughout the soil profile. At the Sevilleta National Wildlife Refuge in central New Mexico, we found that there was significantly more soil C in the upper 10 cm of the soil profile beneath Larrea tridentata than was found beneath Bouteloua spp. Stable carbon isotopic composition indicated that the expansion of L. tridentata influenced C cycling throughout the soil profile. At Curlew Valley in northern Utah, we found no significant differences in total profile soil C beneath different plant life forms. However, there was significantly more soil C found at the soil surface beneath woody plants than was observed beneath annual grasses. There was significantly less POM C beneath annual grasses than was found beneath woody plants or perennial grasses. Based on stable carbon isotopic analyses, we concluded that the invasion of grasses into shrublands influenced only the upper 30 cm of the soil profile. We determined that following changes in plant life form dominance, the most consistent change in soil C was an alteration in content and distribution of POM C, a slowly cycling pool of soil C. While we failed to find a consistent change in total profile soil C with plant life form across our sites, the change in soil C chemistry may have important implications for long-term soil C storage in semiarid systems where there have been shifts in plant life form. Received: 30 March 1999 / Accepted: 11 August 1999     

内蒙古半干旱草原土壤水分对降水格局变化的响应

在全球气候变化背景下, 未来我国北方半干旱地区的降水格局将呈现出季节与年际间降水波动增强和极端降水事件增加的趋势。水分是半干旱草原的主要限制因子, 降水格局变化导致的土壤水分状况的改变必然对生态系统的结构和功能产生显著的影响。该研究选取内蒙古多伦和锡林浩特两个典型半干旱草原群落, 通过分析2006-2013年的降水和多层次土壤(0-10 cm, 10 cm, 20 cm, 30 cm和50 cm)含水量连续观测数据, 研究降水格局变化对土壤水分状况及其垂直分布的影响, 特别是土壤水分对降水事件的脉冲响应过程。结果表明: 两个站点的土壤含水量均呈现显著的季节及年际间波动, 其中土壤表层 0-10 cm水分波动更剧烈。锡林浩特50 cm处土壤含水量波动较大, 主要由于春季融雪的影响。年际间多伦和锡林浩特生长季土壤表层0-10 cm土壤含水量与降水量存在显著的正相关关系, 下层(10-50 cm)土壤含水量与降水量相关性不显著。研究发现小至2 mm的降水事件就能够引起两个站点表层0-10 cm土壤含水量的升高, 即该地区有效降水为日降水量> 2 mm。表层0-10 cm土壤含水量对独立降水事件的脉冲响应可通过指数方程很好地拟合。降水事件的大小决定了降水后表层0-10 cm土壤含水量的最大增量和持续时间, 同时这个脉冲响应过程还受到降水前土壤含水量的影响, 但该过程中并未发现植被因子(叶面积指数)的显著影响。降水后水分下渗深度及该深度的土壤含水量增量主要由降水事件的大小主导, 同时受到降水前土壤含水量的影响。在多伦和锡林浩特, 平均每增加1 mm降水, 下渗深度分别增加1.06和0.79 cm。由此作者认为, 在内蒙古半干旱草原, 降水事件大小和降水前土壤干湿状况是影响土壤水分对降水响应的主要因素, 而植被因子的影响较小。     

Responses of soil moisture to precipitation pattern change in semiarid grasslands in Nei Mongol,China

Aims Under global climate change, precipitation patterns were predicted to change with larger seasonal and annual variations and more extreme events in the semiarid regions of northern China. Water availability is one of the key limited factors in semiarid grasslands. Changes in precipitation patterns will inevitably affect ecosystem structure and function through soil water condition. Our objective was to investigate the response of soil water content to changes of precipitation pattern, especially its pulse response to precipitation events.
Methods Two semiarid steppe sites (Duolun and Xilinhot) in Nei Mongol were chosen and meteorological stations were installed to monitor precipitation and soil volumetric water content (VWC) at five soil depths (0-10 cm, 10 cm, 20 cm, 30 cm, 50 cm) from 2006 to 2013. The pulse response of VWC at 0-10 cm to an individual precipitation event was simulated by an exponential equation.
Important findings Significant seasonal and inter-annual variations of VWC were observed at the Duolun and Xilinhot sites. VWC at 50 cm soil layer in Xilinhot showed an obvious increase during the early spring due to the influences of snow melting. Mean surface (0-10 cm soil layer) VWC was significantly correlated with annual precipitation across eight years, but VWC in the deeper soil layers (10-50 cm) were not impacted by precipitation. We also found that the precipitation event larger than 2 mm could induce a significant increase in surface (0-10 cm soil layer) VWC, and could be regarded as an effective precipitation in this region. The maximum increment of surface VWC after the events and lasting time (T_lasting) were determined by the event size, while showed negatively linear correlations with the initial soil water content before the events. Vegetation development (leaf area index) did not show significant impacts on the responses of surface soil moisture to precipitation pulses. The infiltration depth of rain water was also determined by rain size and pre-event soil moisture. In average, soil water can infiltrate 1.06 cm and 0.79 cm deeper in Duolun and Xilinhot with 1 mm more precipitation, respectively. Therefore, our results suggest that the event size and pre-event soil moisture were the most important factors affecting response patterns of soil moisture to rain events in semiarid ecosystems.     

Spatial and temporal soil moisture resource partitioning by trees and grasses in a temperate savanna, Arizona, USA

Stable isotope analysis was used to determine sources of water used by coexisting trees and grasses in a temperate savanna dominated by Quercus emoryi Torr. We predicted that (1) tree seedlings and bunchgrasses utilize shallow sources of soil water, (2) mature savanna trees use deeper sources of water, and (3) trees switch from shallow to deep water sources within 1 year of germination. We found that Q. emoryi trees, saplings, and seedlings (about 2 months, 1 year, and 2 years old), and the dominant bunchgrass [Trachypogon montufari (H.B.K.) Nees.] utilized seasonally available moisture from different depths within the soil profile depending on size/age relationships. Sapling and mature Q. emoryi acquired water from >50 cm deep, 2-month-old seedlings utilized water from <15 cm, and 1- and 2-year-old seedlings and grasses used water from between 20 cm and 35 cm. This suggests that very young seedlings are decoupled from grasses in this system, which may facilitate germination and early establishment of Q. emoryi within extant stands of native grasses. The potential for subsequent interaction between Q. emoryi and native grasses was evidenced by similar patterns of soil water use by 1- and 2-year-old seedlings and grasses. Q. emoryi seedlings did not switch from shallow to deep sources of soil water within 2 years of germination: water use by these seedlings apparently becomes independent of water use by grasses after 2 years of age. Finally, older trees (saplings, mature trees) use water from deeper soil layers than grasses, which may facilitate the stable coexistence of mature trees and grasses. Potential shifts in the seasonality of precipitation may alter interactions between woody plants and grasses within temperate savannas characterized by bimodal precipitation regimes: reductions in summer precipitation or soil moisture may be particularly detrimental to warm-season grasses and seedlings of Q. emoryi. Received: 21 November 1996 / Accepted: 2 May 1997     

Using GIS and remote sensing to explore the influence of physical environmental factors and historical land use on bushland structure

Bushland is a vegetation type with a dense cover of woody plants of low stature and a grass understory. We explored the influence of environmental factors (precipitation, topography, geology, hydrology and waterhole density) and historical land use in the Tsavo ecosystem bushland. We mapped vegetation patterns in the region using remote sensing, GIS and field data and tested relationships with physical factors. We used regression analysis to explore the interaction of physical factors and human influence by including a protected and a nonprotected area, representing contrasting historical land use policies, as factors in the regression model. Bushland vegetation had the largest proportion in the region as a whole, but its proportion was notably greater in the nonprotected area. Precipitation, elevation, geology and historical land use were significant predictors of vegetation patterns in the regression models. Higher precipitation and elevation make woody plants superior competitors over grasses, resulting in predominance of bushland in such areas. Geologically, marine deposits result in shallow calcareous soils that favour grasses over trees, hence, absence of bushland. Wildlife management policies such as providing water for wildlife, protective boundaries and controlling wildfires influence distribution of wildlife and result in conditions that prevent regeneration and recruitment of trees.     

Ecological strategies in a Patagonian arid steppe

The vegetation in the Coironal arid steppe consists of grasses and shrubs. The objective of this paper was to test Walter's hypothesis that woody vegetation and grasses compete for water in the upper layers of the soil, but woody vegetation has exclusive access to a source of water at deeper levels.Analysis of root profiles and patterns of leaf and soil water potential led us to accept the hypothesis for this arid steppe. Additional information on phenology and on the ability of the major grass species to respond to watering permitted to identify two ecological strategies corresponding to grasses and shrubs. Grasses behave as opportunists having always leaves ready to grow as soon as water becomes available. They have a shallow root system and are able to respond very rapidly to increases in soil water availability. In contrast, woody species have a clear-cut periodic pattern of growth and dormancy. They possess thick horizontal roots running below 35–40 cm and utilized water stored in lower layers of the soil.A diagrammatic model summarizes the role of periodic and opportunistic species upon water circulation in the ecosystem. The effect of changes in the proportion of the two groups upon water dynamics is also discussed.Nomenclature follows Nicora (1978) and Cabrera (1971). Acknowledgements. We would like to thank Instituto Nacional de Tecnología Agropecuaria for its valuable support. This work was also supported by Subsecretaría de Ciencia y Técnica and Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET). E. H. Satorre, A. Nuñez and M. Aguiar helped in data collection in the field and in laboratory sample processing.     

Windows of opportunity for Prosopis velutina seedling establishment and encroachment in a semiarid grassland

Large increases in the density of woody plants in former grasslands have been documented globally over the past two centuries. Prosopis velutina (velvet mesquite) has recently expanded on over 38 million ha in the grasslands of the American southwest.Mesquite establishment and expansion is potentially influenced by competitive interactions with grasses, particle size distribution (texture) of soil and changes in the amount and/or seasonality of precipitation, amongst other factors.To investigate the relative importance of precipitation seasonality, grass competition, and soil texture on mesquite seedling and establishment, we experimentally manipulated seasonality of precipitation across 72 1.5 m × 1.7 m plots that we planted with two grasses from contrasting functional groups (or left unvegetated) across two highly contrasting soil textures within a semi-desert grassland in Arizona, USA, wherein we followed the demography of 8640 planted mesquite seeds between 2002 and 2006. Up to 70% of the variance in seedling emergence was explained by soil water content, and seedling mortality was more dependent on summer than on winter precipitation.Mortality of less drought tolerant grasses after drought in coarse textured soils created conditions most favorable for seedling establishment, whereas lower rates of seedling establishment were observed in finer textured soils or in plots dominated by more drought tolerant grasses.We conclude that mesquite encroachment is episodic, based on discontinuous patterns of seedling establishment, favored on coarse textured soils and associated with unique conditions of high summer water availability when precipitation is not limiting and when competition from grasses is lowered.     

Small rainfall events: An ecological role in semiarid regions

Summary Small precipitation events account for a large proportion of the precipitation received in semiarid regions, and their potential ecological importance has previously been ignored. We investigated the effect of a small rainfall event (5 mm) upon Bouteloua gracilis, the dominant grass species of the central and southern Great Plains of North America. An effect of a small event on leaf water potential and leaf conductance to water vapor was observed in less than 12 h and lasted for up to two days.The remarkable short response time of Bouteloua gracilis to a rainfall stimulus enables this species to utilize small events and, therefore, may influence its persistence as a dominant species in the steppe region.We proposed response times to be one of the major species characteristics determining capacity for utilizing different portions of the water resource of the region. We suggest that small precipitation events are ecologically significant and a qualitatively distinct resource for ecosystems in semiarid regions.     

Combined effects of precipitation and nitrogen deposition on native and invasive winter annual production in California deserts

Primary production in deserts is limited by soil moisture and N availability, and thus is likely to be influenced by both anthropogenic N deposition and precipitation regimes altered as a consequence of climate change. Invasive annual grasses are particularly responsive to increases in N and water availabilities, which may result in competition with native forb communities. Additionally, conditions favoring increased invasive grass production in arid and semi-arid regions can increase fire risk, negatively impacting woody vegetation that is not adapted to fire. We conducted a seeded garden experiment and a 5-year field fertilization experiment to investigate how winter annual production is altered by increasing N supply under a range of water availabilities. The greatest production of invasive grasses and native forbs in the garden experiment occurred under the highest soil N (inorganic N after fertilization = 2.99 g m^?2) and highest watering regime, indicating these species are limited by both water and N. A classification and regression tree (CART) analysis on the multi-year field fertilization study showed that winter annual biomass was primarily limited by November–December precipitation. Biomass exceeded the threshold capable of carrying fire when inorganic soil N availability was at least 3.2 g m^?2 in piñon-juniper woodland. Due to water limitation in creosote bush scrub, biomass exceeded the fire threshold only under very wet conditions regardless of soil N status. The CART analyses also revealed that percent cover of invasive grasses and native forbs is primarily dependent on the timing and amount of precipitation and secondarily dependent on soil N and site-specific characteristics. In total, our results indicate that areas of high N deposition will be susceptible to grass invasion, particularly in wet years, potentially reducing native species cover and increasing the risk of fire.     

Carbon and Nitrogen Decoupling Under an 11-Year Drought in the Shortgrass Steppe

The frequency and magnitude of drought is expected to increase in the US Great Plains under future climate regimes. Although semiarid systems are considered highly resistant to water limitation, novel drought events could alter linkages among biogeochemical processes, and result in new feedbacks that influence the timescale of ecosystem recovery. We examined changes in carbon and nitrogen cycling in the last 2 years of an 11-year drought manipulation in the shortgrass steppe, and under the first 2 years of recovery from drought. We measured plant production, plant tissue chemistry, soil trace gas flux, and soil inorganic nitrogen dynamics to test the extent that this magnitude of drought altered carbon and nitrogen fluxes and how these changes affected post-drought dynamics. We found that soil inorganic nitrogen was up to five times higher under severe drought than under control conditions, but that this nitrogen may not have been accessible to plants and microbial communities during drought due to diffusion limitations. Drought plots had higher N₂O flux when they received equal rainfall pulses, showing that this accumulated N may be vulnerable to loss. In addition, plants in drought plots had higher tissue nitrogen for 2 years following drought. These results show that decadal-length droughts that may occur under future precipitation regimes are likely to alter ecosystem properties through interactions among precipitation, vegetation, and N cycling. Shifts in plant N, vulnerability of nitrogen to loss, and rainfall use efficiency that we observed are likely to affect the recovery time of semiarid systems subject to droughts of this magnitude.     

Differential water resource use by herbaceous and woody plant life-forms in a shortgrass steppe community

We conducted a study to test the predictions of Walter's two-layer model in the shortgrass steppe of northeastern Colorado. The model suggests that grasses and woody plants use water resources from different layers of the soil profile. Four plant removal treatments were applied in the spring of 1996 within a plant community codominated by Atriplex canescens (a C₄ shrub) and Bouteloua gracilis (a C₄ grass). During the subsequent growing season, soil water content was monitored to a depth of 180 cm. In addition, stem and leaf tissue of Atriplex, Bouteloua and the streamside tree Populus sargentii were collected monthly during the growing seasons of 1995 and 1996 for analysis of the δ¹⁸O value of plant stem water (for comparison with potential water sources) and the δ¹³C value of leaves (as an indicator of plant water status). Selective removal of shrubs did not significantly increase water storage at any depth in the measured soil profile. Selective removal of the herbaceous understory (mainly grasses) increased water storage in the top 60 cm of the soil. Some of this water gradually percolated to lower layers, where it was utilized by the shrubs. Based on stem water δ¹⁸O values, grasses were exclusively using spring and summer rain extracted from the uppermost soil layers. In contrast, trees were exclusively using groundwater, and the consistent δ¹³C values of tree leaves over the course of the summer indicated no seasonal changes in gas exchange and therefore minimal water stress in this life-form. Based on anecdotal rooting-depth information and initial measurements of stem water δ¹⁸O, shrubs may have also had access to groundwater. However, their overall δ¹⁸O values indicated that they mainly used water from spring and summer precipitation events, extracted from subsurface soil layers. These findings indicate that the diversity of life-forms found in this shortgrass steppe community may be a function of the spatial partitioning of soil water resources, and their differential use by grasses, shrubs, and trees. Consequently, our findings support the two-layer model in a broad sense, but indicate a relatively flexible strategy of water acquisition by shrubs. Received: 23 December 1997 / Accepted: 16 September 1998     

免耕对北方旱作玉米土壤水分及物理性质的影响

通过2006-2008年的田间试验,研究了北京地区免耕与传统翻耕条件下土壤水分的时空动态及其对土壤相关物理性质的影响.结果表明：研究期间,免耕和翻耕地0～100 cm土层的土壤贮水量随时间及降水变化的趋势一致,但免耕地的土壤贮水量在不同时刻以及不同降雨条件下均高于翻耕地,增幅在2.7%～30.3%,降雨相对充分条件下土壤贮水量的增量有所增加,但免耕在相对缺水的地区更值得推广;免耕地0～100 cm土层的平均土壤含水量明显高于传统翻耕,增幅在3.4%～12.8%,其中表层（0～20 cm）和底层（80～100 cm）的水分增量明显高于中间层次,最高增量可达22.2%;免耕覆盖的耕作方式可通过降低土壤容重、提高土壤孔隙度、促进土壤水稳性团聚体的形成等作用来提高土壤的保水能力和水分利用效率,进而促进作物增收.与对照相比,免耕3年后土壤水分利用率和春玉米产量分别较传统翻耕提高13.3%和16.4%.     

Aboveground net primary production dynamics in a northern Chihuahuan Desert ecosystem

Aboveground net primary production (ANPP) dynamics are a key element in the understanding of ecosystem processes. For semiarid environments, the pulse-reserve framework links ANPP to variable and unpredictable precipitation events contingent on surficial hydrology, soil moisture dynamics, biodiversity structure, trophic dynamics, and landscape context. Consequently, ANPP may be decoupled periodically from processes such as decomposition and may be subjected to complex feedbacks and thresholds at broader scales. As currently formulated, the pulse-reserve framework may not encompass the breadth of ANPP response to seasonal patterns of precipitation and heat inputs. Accordingly, we examined a 6-year (1999–2004), seasonal record of ANPP with respect to precipitation, soil moisture dynamics, and functional groups in a black grama (Bouteloua eriopoda) grassland and a creosotebush (Larrea tridentata) shrubland in the northern Chihuahuan Desert. Annual ANPP was similar in the grassland (51.1 g/m²) and shrubland (59.2 g/m²) and positively correlated with annual precipitation. ANPP differed among communities with respect to life forms and functional groups and responses to abiotic drivers. In keeping with the pulse-reserve model, ANPP in black grama grassland was dominated by warm-season C₄ grasses and subshrubs that responded to large, transient summer storms and associated soil moisture in the upper 30 cm. In contrast, ANPP in creosotebush shrubland occasionally responded to summer moisture, but the predominant pattern was slower, non-pulsed growth of cool-season C₃ shrubs during spring, in response to winter soil moisture accumulation and the breaking of cold dormancy. Overall, production in this Chihuahuan Desert ecosystem reflected a mix of warm-temperate arid land pulse dynamics during the summer monsoon and non-pulsed dynamics in spring driven by winter soil moisture accumulation similar to that of cool-temperate regions. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Semiarid grassland responses to short-term variation in water availability

Standing crop and species composition in semiarid grassland are linked to long-term patterns of water availability, but grasslands are characterized by large single-season variability in rainfall. We tested whether a single season of altered water availability influenced the proportions of grasses and shrubs in a semiarid grassland near the northern edge of the North American Great Plains. We studied stands of the clonal shrub snowberry (Symphoricarpos occidentalis) and adjacent grassland dominated by the native grasses Stipa spartea and Bouteloua gracilis. Rain was excluded and water supplied in amounts corresponding to years of low, medium, and high rainfall, producing a 2 − 4-fold range in monthly precipitation among water supply treatments. There were ten replicate plots of each water treatment in both snowberry stands and grassland. Grass standing crop increased significantly with water availability in grassland but not inside snowberry stands. Total standing crop and shrub stem density increased significantly with water supply, averaged across both communities. In contrast, water had no effect on shrub standing crop or light penetration. In summary, our finding that water has significant effects on a subset of components of grassland vegetation is consistent with long-term, correlational studies, but we also found that a single season of altered water supply had no effect on other important aspects of the ecosystem. This revised version was published online in June 2006 with corrections to the Cover Date.     

Sensitivity of plant species to warming and altered precipitation dominates the community productivity in a semiarid grassland on the Loess Plateau

Global warming and changes in precipitation patterns can critically influence the structure and productivity of terrestrial ecosystems. However, the underlying mechanisms are not fully understood. We conducted two independent but complementary experiments (one with warming and precipitation manipulation (+ or – 30%) and another with selective plant removal) in a semiarid grassland on the Loess Plateau, northwestern China, to assess how warming and altered precipitation affect plant community. Our results showed that warming and altered precipitation affected community aboveground net primary productivity (ANPP) through impacting soil moisture. Results of the removal experiment showed competitive relationships among dominant grasses, the dominant subshrub and nondominant species, which played a more important role than soil moisture in the response of plant community to warming and altered precipitation. Precipitation addition intensified the competition but primarily benefited the dominant subshrub. Warming and precipitation reduction enhanced water stresses but increased ANPP of the dominant subshrub and grasses, indicating that plant tolerance to drought critically meditated the community responses. These findings suggest that specie competitivity for water resources as well as tolerance to environmental stresses may dominate the responses of plant communities on the Loess Plateaus to future climate change factors.     

Inorganic Nitrogen Supply and Dissolved Organic Nitrogen Abundance across the US Great Plains

Across US Great Plains grasslands, a gradient of increasing mean annual precipitation from west to east corresponds to increasing aboveground net primary productivity (ANPP) and increasing N-limitation. Previous work has shown that there is no increase in net N mineralization rates across this gradient, leading to the question of where eastern prairie grasses obtain the nitrogen to support production. One as-yet unexamined source is soil organic N, despite abundant literature from other ecosystems showing that plants take up dissolved soil organic N. This study measured KCl-extractable dissolved organic N (DON) in surface soils across the grassland productivity gradient. We found that KCl-extractable DON pools increased from west to east. If available to and used by plants, this DON may help explain the high ANPP in the eastern Great Plains. These results suggest a need for future research to determine whether, in what quantities, and in what forms prairie grasses use organic N to support primary production.     

Soil C:N:P stoichiometry responds to vegetation change from grassland to woodland

Woody encroachment has been a major land cover change in dryland ecosystems during the past century. While numerous studies have demonstrated strong effects of woody encroachment on soil carbon (C), nitrogen (N), and phosphorus (P) storage, far less is known about the plasticity of soil C:N:P stoichiometry in response to woody encroachment. We assessed landscape-scale patterns of spatial heterogeneity in soil C:N:P ratios throughout a 1.2 m soil profile in a region where grassland is being replaced by a diverse assemblage of subtropical woody plants dominated by Prosopis glandulosa, an N₂-fixing tree. Woody species had leaf and fine root C:N:P ratios significantly different from grasses. Variation in soil C:N ratios in both horizontal and vertical planes was remarkably smaller than that of soil N:P and C:P ratios. Spatial patterns of soil C:N ratio throughout the profile were not strongly related to vegetation cover. In contrast, spatial patterns of soil N:P and C:P ratios displayed a strong resemblance to that of vegetation cover throughout the soil profile. Within the uppermost soil layer (0–5 cm), soil N:P and C:P ratios were higher underneath woody patches while lower within the grassland; however, this pattern was reversed in subsurface soils (15–120 cm). These results indicate a complex response of soil C:N:P stoichiometry to vegetation change, which could have important implications for understanding C, N, and P interactions and nutrient limitations in dryland ecosystems.     

Plant feedbacks increase the temporal heterogeneity of soil moisture

Plant feedbacks on resource levels are well-known, but feedbacks on resource variability have received little attention. Semi-arid grasslands have greater temporal heterogeneity of rainfall than mesic forests, leading to the possibility that grasses further enhance this variability as a mechanism for excluding woody plants originating in habitats with less heterogeneity. Here we test the hypothesis that grasses create greater levels of temporal heterogeneity of soil resources than do woody plants. We used monocultures of five replicate species of both growth forms. Daily soil moisture measurements taken 10 and 30 cm beneath monocultures over a growing season showed that temporal heterogeneity was significantly greater under grasses than under woody plants. This occurred during a dry period when plants are most likely to compete for moisture. Differences in temporal heterogeneity between growth forms were related to differences in their abilities to reduce soil moisture: during the dry period, the net effect of vegetation on moisture 10 cm deep was greatest under grasses. Although the rate of change of soil moisture was higher under grasses, the growth forms exploited different depths of soil moisture: soils 10 cm deep were driest under grasses, but soils 30 cm deep were driest under woody species. In summary, grasses increased within-season resource variability in a habitat already characterized by high among-year variability.     

Differential use of large summer rainfall events by shrubs and grasses: a manipulative experiment in the Patagonian steppe

In the Patagonian steppe, years with above-average precipitation (wet years) are characterized by the occurrence of large rainfall events. The objective of this paper was to analyze the ability of shrubs and grasses to use these large events. Shrubs absorb water from the lower layers, grasses from the upper layers, intercepting water that would otherwise reach the layers exploited by shrubs. We hypothesized that both life-forms could use the large rainfalls and that the response of shrubs could be more affected by the presence of grasses than vice versa. We performed a field experiment using a factorial combination of water addition and life-form removal, and repeated it during the warm season of three successive years. The response variables were leaf growth, and soil and plant water potential. Grasses always responded to experimental large rainfall events, and their response was greater in dry than in wet years. Shrubs only used large rainfalls in the driest year, when the soil water potential in the deep layers was low. The presence or absence of one life-form did not modify the response of the other. The magnitude of the increase in soil water potential was much higher in dry than in humid years, suggesting an explanation for the differences among years in the magnitude of the response of shrubs and grasses. We propose that the generally reported poor response of deep-rooted shrubs to summer rainfalls could be because (1) the water is insufficient to reach deep soil layers, (2) the plants are in a dormant phenological status, and/or (3) deep soil layers have a high water potential. The two last situations may result in high deep-drainage losses, one of the most likely explanations for the elsewhere-reported low response of aboveground net primary production to precipitation during wet years. Received: 23 January 1997 / Accepted: 19 November 1997