Burrowing activities of kangaroo rats and patterns in plant species dominance at a shortgrass steppe-desert grassland ecotone

Abstract. Our objective was to evaluate the effects of burrowing activities by banner-tail kangaroo rats (Dipodomys spectabilis Merriam) on plant community structure and species dominance for two patch types at the ecotone between shortgrass steppe and desert grassland in New Mexico, USA. 10 mounds produced by kangaroo rats were selected in patches dominated by Bouteloua gracilis (the dominant in shortgrass steppe communities) and 10 mounds were selected in patches dominated by B. eriopoda (the dominant in Chihuahuan desert grasslands). Plant cover and density by species were sampled from three locations associated with each mound: the mound proper, the edge of the mound in the transition area, and the off-mound vegetation. Similar cover of B. eriopoda for the edges of mounds in both patch types indicates the ability of this species to respond to animal disturbances regardless of the amount of cover in the surrounding undisturbed vegetation. By contrast, cover of B. gracilis was low for all mounds and mound edges in patches dominated by this species. Much higher cover of B. eriopoda on mound edges compared to the undisturbed vegetation in B. gracilis-dominated patches indicates that kangaroo rats have important positive effects on this species. Lower cover of perennial grasses and higher cover of forbs, shrubs, and succulents on the edges of mounds in B. eriopoda-dominated patches compared to patches dominated by B. gracilis indicate the importance of surrounding vegetation to plant responses on disturbed areas. Our results show that kangaroo rats have important effects on both species dominance and composition for different patch types, and may provide a mechanism for small-scale dominance patterns at an ecotone; thus providing further support for their role as keystone species in desert grasslands.     

Distribution of plant species at a biome transition zone in New Mexico

Question: Is there a difference in plant species and life form composition between two major patch types at a biome transition zone? Are subordinate species associated with different patch types at the shortgrass steppe — Chihuahuan desert grassland transition zone? Is this association related to differences in soil texture between patch types and the geographic range of associated species? Location: central New Mexico, USA. Methods: Patches dominated by either Bouteloua gracilis, the dominant species in the shortgrass steppe, or Bouteloua eriopoda, dominant species in the Chihuahuan desert grasslands, were sampled for the occurrence of subordinate species and soil texture within a 1500‐ha transitional mosaic of patches. Results: Of the 52 subordinate species analysed, 16 species were associated with B. gracilis‐dominated patches and 12 species with B. eriopoda‐dominated patches. Patches dominated by B. gracilis were richer in annual grasses and forbs, whereas patches dominated by B. eriopoda contained more perennials forbs and shrubs. Soils of B. gracilis‐dominated patches had higher clay and lower rock contents compared with soils of B. eriopoda‐dominated patches. Differences in species characteristics of the dominant species as well as differences in soil texture between patch types contribute to patch‐scale variation in composition. The association of species to patch types was not related to their geographic range and occurrence in the adjacent biomes. Conclusions: Patch types at this biome transition zone have characteristic life‐form and species composition, but species are associated to patch types due to local constraints, independently from their affinity to the adjacent biomes.     

Climatic variation and simulated patterns in seedling establishment of two dominant grasses at a semi‐arid‐arid grassland ecotone

Abstract. The objective of this study was to predict the effects of climatic variation at multiple temporal frequencies on seedling establishment by two congeneric C₄ perennial grasses (Bouteloua gracilis and B. eriopoda) at the ecotone between shortgrass steppe grassland and Chihuahuan desert grassland in central New Mexico, USA. The approach was to use a daily time‐step simulation model to determine the occurrence of a recruitment event in each year based upon the amount and timing of soil water required for establishment. Historical weather data were used to predict effects of seasonal and inter‐decadal variation in climate on establishment. A sensitivity analysis was used to predict effects of directional climate change on establishment. Bouteloua gracilis had a broad pattern of simulated establishment from May through September that included periods with high year‐to‐year variation in precipitation. B. eriopoda establishment events occurred primarily in July when precipitation amounts were most reliable. Climatic conditions from 1949 through 1968 were more favorable for B. eriopoda establishment compared to the cooler, wetter conditions from 1969 through 1988 that favored B. gracilis. Establishment of B. eriopoda was lowest in El Niño years whereas B. gracilis establishment was highest in La Niña years. Establishment of B. gracilis was most sensitive to temperature when precipitation was higher than current amounts. The greatest response to temperature by B. eriopoda for all precipitation amounts occurred at cooler temperatures than found currently. These results indicate that climatic variation at multiple frequencies has differential effects on seedling establishment for these two perennial grasses, and may account at least in part for patterns in dominance at this biome transition zone.     

Local and regional differences in abundance of co-dominant grasses in the shortgrass steppe: a modeling analysis of potential causes

The shortgrass steppe is co-dominated by two C₄perennial grasses, Bouteloua gracilis andBuchloë dactyloides. At our site and throughouteastern Colorado Bouteloua gracilis has higher cover andbiomass than Buchloë dactyloides. We hypothesizedthatthe interaction between seedling recruitment differences and disturbance regimeand tolerance to drought conditions were the most likely causes of the observeddifferences in relative abundances. We used a simulation model to investigatethe relative importance of the three factors. We studied plant biomass of thesetwo species in 18 simulated treatments that resulted from a factorialcombination of seedling recruitment differences, disturbance regime, andtolerance to drought conditions. Analysis of the simulation outputs with ANOVAindicated that biomass of each species responded positively to increases inrecruitment probability. Increasing disturbance frequency favoredBuchloë dactyloides over Boutelouagracills, whereas the susceptibility Buchloëdactyloides to drought favored Boutelouagracilis. The results indicated that differences in droughttoleranceand seedling recruitment probabilities along with their interactions withdisturbance regimes exert a major control on the biomass of the species. In theabsence of or with intermediate disturbance, a higher recruitment probabilityand greater tolerance to drought of Bouteloua gracilisthanBuchloë dactyloides yielded patterns of relativebiomass similar to the patterns observed in the shortgrass steppe.     

Longevity and turnover of roots in the shortgrass steppe: influence of diameter and depth

We used minirhizotrons to determine patterns of root longevity andturnover for the perennial bunchgrass Bouteloua gracilisinthe shortgrass steppe of eastern Colorado, USA. We hypothesized that rootlongevity would be partially controlled by root diameter, following previouslyobserved patterns in woody plants. In addition, we hypothesized that rootturnover would be greatest in surface soil horizons and decrease with depth dueto variation in soil moisture availability and temperature. Root longevity wascorrelated with root diameter. Median life span of roots > 0.4mm was approximately 320 days, while roots < 0.2mmhad a median life span of 180 days. There was approximately a 6%decreasein the likelihood of mortality with a 0.10-mm increase inroot diameter, controlling for the effect of depth in the soil profile. Rootlength production and mortality were highest in the upper20 cm of the soil profile and decreased with depth.However,because root length density also decreased with depth, there were nosignificantdifferences in turnover rate of root length among sampling intervals. Turnoverwas approximately 0.86 yr^–1 based on root length production,while turnover was 0.35 yr^–1 using root length mortality as ameasurement of flux. The imbalance between turnover estimates may be aconsequence of the time the minirhizotrons were in place prior to imaging or mayresult from our lack of over-winter measures of mortality. Our worksuggests that Bouteloua gracilis roots have complex lifehistory strategies, similar to woody species. Some portion of the root systemishighly ephemeral, while slightly larger roots persist much longer. Thesedifferences have implications for belowground carbon and nitrogen cycles in theshortgrass steppe.     

Plant population and community responses to removal of dominant species in the shortgrass steppe

Question: What are the plant population‐ and community‐level effects of removal of dominant plant species in the shortgrass steppe? Location: The Shortgrass Steppe Long‐Term Ecological Research site in northern Colorado, USA. Methods: We annually measured plant cover and density by species for 10 years after a one‐time aboveground removal of the dominant perennial grass, Bouteloua gracilis. Removal and control plots (3 m × 3 m) were within grazed and ungrazed locations to assess the influence of grazing on recovery dynamics. Our analyses examined plant species, functional type, and community responses to removal, paying special attention to the dynamics of subdominant and rare species. Results: Basal cover of B. gracilis increased by an average of 1% per year, but there was significantly less plant cover in treatment compared to control plots for 5 years following removal. In contrast to the lower cover in treatment plots, the plant density (number of plants m^?2) of certain subdominant perennial grasses, herbaceous perennial and annual forbs, a dwarf shrub, and cactus increased after removal of the dominant species, with no major change in species richness (number of species per 1 m × 1 m) or diversity. Subdominant species were more similar between years than rare species, but dominant removal resulted in significantly lower similarity of the subdominant species in the short term and increased the similarity of rare species in the long term. Conclusions: Removal of B. gracilis, the dominant perennial grass in the shortgrass steppe, increased the absolute density of subdominant plants, but caused little compensation of plant cover by other plants in the community and changes in species diversity.     

Effects of monsoon precipitation variability on the physiological response of two dominant C4 grasses across a semiarid ecotone

For the southwestern United States, climate models project an increase in extreme precipitation events and prolonged dry periods. While most studies emphasize plant functional type response to precipitation variability, it is also important to understand the physiological characteristics of dominant plant species that define plant community composition and, in part, regulate ecosystem response to climate change. We utilized rainout shelters to alter the magnitude and frequency of rainfall and measured the physiological response of the dominant C₄ grasses, Bouteloua eriopoda and Bouteloua gracilis. We hypothesized that: (1) the more drought-adapted B. eriopoda would exhibit faster recovery and higher rates of leaf-level photosynthesis (A _net) than B. gracilis, (2) A _net would be greater under the higher average soil water content in plots receiving 30-mm rainfall events, (3) co-dominance of B. eriopoda and B. gracilis in the ecotone would lead to intra-specific differences from the performance of each species at the site where it was dominant. Throughout the study, soil moisture explained 40–70 % of the variation in A _net. Consequently, differences in rainfall treatments were not evident from intra-specific physiological function without sufficient divergence in soil moisture. Under low frequency, larger rainfall events B. gracilis exhibited improved water status and longer periods of C gain than B. eriopoda. Results from this study indicate that less frequent and larger rainfall events could provide a competitive advantage to B. gracilis and influence species composition across this arid–semiarid grassland ecotone.     

Influence of soil depth on the decomposition of Bouteloua gracilis roots in the shortgrass steppe

The distribution and turnover of plant litter contribute to soil structure, the availability of plant nutrients, and regional budgets of greenhouse gasses. Traditionally, studies of decomposition have focused on the upper soil profile. Other work has shown that temperature, precipitation, and soil texture are important determinates of patterns of decomposition. Since these factors all vary through a soil profile, it has been suggested that decomposition rates may vary with depth in a soil profile. In this work, we examine patterns of root decomposition through a shortgrass steppe soil profile. We buried fresh root litter from Bouteloua gracilis plants in litterbags at 10, 40, 70, and 100 cm. Litterbags were retrieved six times between July 1996 and May 1999. We found that the decomposition rate for fresh root litter was approximately 50% slower at 1 m than it was at 10 cm. After 33 months, 55% of the root mass buried at 10 cm remained, while 72% of the root mass buried at 1 m was still present. This corresponds to a 19-year residence time for roots at 10 cm and a 36-year residence time for roots at 1 m. Mass loss rates decreased linearly from 10 cm to 1 m. Patterns of total carbon and cellulose loss rates followed those of mass loss rates. Roots at 1 m tended to accumulate lignin-like compounds over the course of the experiment. Differences in the stabilization of lignin may be a consequence of differences in microbial community through a shortgrass steppe soil profile.     

Spatial Variability of Soil Properties in the Shortgrass Steppe: The Relative Importance of Topography, Grazing, Microsite, and Plant Species in Controlling Spatial Patterns

We conducted a study to evaluate the relative importance of topography, grazing, the location of individual plants (microsite), and plant species in controlling the spatial variability of soil organic matter in shortgrass steppe ecosystems. We found that the largest spatial variation occurs in concert with topography and with microsite-scale heterogeneity, with relatively little spatial variability due to grazing or to plant species. Total soil C and N, coarse and fine particulate organic matter C and N, and potentially mineralizable C were significantly affected by topography, with higher levels in toeslope positions than in midslopes or summits. Soils beneath individual plants (Bouteloua gracilis and Opuntia polyacantha) were elevated by 2–3 cm relative to surrounding soils. All pools of soil organic matter were significantly higher in the raised hummocks directly beneath plants than in the soil surface of interspaces or this layer under plants. High levels of mineral material in the hummocks suggest that erosion is an important process in their formation, perhaps in addition to biotic accumulation of litter beneath individual plants. Over 50 y of heavy grazing by cattle did not have a significant effect on most of the soil organic matter pools we studied. This result was consistent with our hypothesis that this system, with its strong dominance of belowground organic matter, is minimally influenced by aboveground herbivory. In addition, soils beneath two of the important plant species of the shortgrass steppe, B. gracilis and O. polyacantha, differed little from one another. The processes that create spatial variability in shortgrass steppe ecosystems do not affect all soil organic matter pools equally. Topographic variability, developing over pedogenic time scales (centuries to thousands of years), has the largest effect on the most stable pools of soil organic matter. The influence of microsite is most evident in the pools of organic matter that turn over at time scales that approximate the life span of individual plants (years to decades and centuries).     

Soil water dynamics,transpiration, and water losses in a crested wheatgrass and native shortgrass ecosystem

The status of water in soil and vegetation was monitored in a stand of crested wheatgrass (Agropyron cristatum) and a nearby shortgrass steppe during a growing season. This was done to determine if water use and losses were similar among two very different communities in a similar climate. Precipitation was similar throughout the study period for both the crested wheatgrass and native shortgrass communities. However, the native shortgrass community with greater root biomass had consistently greater soil water depletion in the deeper soil horizons than was found in the crested wheatgrass community. Greater depletion of soil water by native shortgrass species suggests that they might be more competitive than crested wheatgrass in a water-limited environment.Crested wheatgrass maintained high leaf water potential early in the season, but lower water potential during the latter part of the growing season as compared with the major species of the shortgrass steppe, blue grama (Bouteloua gracilis) and western wheatgrass (Agropyron smithii). Leaf conductance was lower for crested wheatgrass than for the native grasses during the later part of the growing season. Consequently, seasonal transpiration for crested wheatgrass was lower when compared with blue grama or western wheatgrass. Lower conductance allowed crested wheatgrass to maintain relatively high internal water potential and may have accounted for less soil water use at deeper soil depths during the latter part of the growing season.Water loss through transpiration was less for western wheatgrass than for either blue grama or crested wheatgrass because western wheatgrass had less leaf area. However, western wheatgrass was as efficient as the other species in its use of water. Crested wheatgrass transpired more water than blue grama early in the growing season, but less than either native species for the remainder of the growing season. Estimated seasonal transpiration loss was greater in the shortgrass ecosystem than in the established crested wheatgrass stand.     

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.     

气候变暖与荒漠-草原过渡带植物动态响应灰色分析--以美国新墨西哥州Sevilleta生态过渡带为例

在当前全球气候变暖的形势下,地表生态系统表现出不同的响应。干旱-半干旱地区的生态过渡带的生物反应更为敏感。美国新墨西哥州中部的荒漠-草原生态过渡带是监测全球气候变化和人类活动对生物影响的重要区域之一。本文以Sevilleta荒漠．草原生态过渡带为例,运用灰色系统分析方法．研究在当前全球气候变暖的条件下,1989～1998年10年间该生态过渡带的两种共存的植物优势种(Bouteloua eripoda)和(Bouteloua grncilis)历年的种群密度、组合形式和变化趋势,分析引起这些变化的主要气候因子。结果表明,Sevilleta生态过渡带中来自荒漠草原的优势物种Bouteloua eripoda的密度有上升的趋势,来自大草原的优势物种Bouteloua gracilis的密度有下降的趋势,它们的密度比大于1且有上升的趋势;在影响种群密度变化的降水、最高温度、最低温度、湿度的4个气候因子中,温度因子起着重要的作用。由此可以认为,随着气候的变暖,本区来自荒漠地区的优势植物种在荒漠-草原群落过渡带中将逐渐处于强优势地位,生态过渡带将有可能被荒漠草原所替代。     

Successional dynamics of a semiarid grassland: effects of soil texture and disturbance size

We evaluated effects of soil texture and disturbance size on the successional dynamics of a semiarid grassland dominated by the perennial bunchgrass, Bouteloua gracilis (H.B.K.) Lag. ex Griffiths. A spatially-explicit gap dynamics simulation model was used to evaluate recovery patterns. The model simulates establishment, growth, and mortality of individual plants on an array of small plots through time at an annual time step. Each simulated disturbance consisted of a grid of plots of the same soil texture interconnected by processes associated with dispersal of B. gracilis seeds. Soil texture was incorporated into the model as effects on seed germination, seedling establishment, and subsequent growth of B. gracilis. Five soil texture classes and five disturbance sizes were simulated.Soil texture was more important to recovery of B. gracilis than either size of a disturbance or location of plots within a disturbance. Constraints on recruitment of seedlings had a greater effect on recovery than constraints associated with plant growth. Fastest recovery occurred on soils with the largest silt content, the variable that affects seedling establishment. Disturbances with slowest recovery were on soils with low silt contents, and either high or low water-holding capacity, the variable that affects plant growth. Biomass and recovery decreased as disturbance size increased, and as distance from a disturbed plot to the edge of the disturbance increased. In most cases, important interactions between soil texture and disturbance size on recovery were not found.     

Elevated CO2 enhances water relations and productivity and affects gas exchange in C3 and C4 grasses of the Colorado shortgrass steppe.

Six open‐top chambers were installed on the shortgrass steppe in north‐eastern Colorado, USA from late March until mid‐October in 1997 and 1998 to evaluate how this grassland will be affected by rising atmospheric CO₂. Three chambers were maintained at current CO₂ concentration (ambient treatment), three at twice ambient CO₂, or approximately 720 μmol mol^?1 (elevated treatment), and three nonchambered plots served as controls. Above‐ground phytomass was measured in summer and autumn during each growing season, soil water was monitored weekly, and leaf photosynthesis, conductance and water potential were measured periodically on important C₃ and C₄ grasses. Mid‐season and seasonal above‐ground productivity were enhanced from 26 to 47% at elevated CO₂, with no differences in the relative responses of C₃/C₄ grasses or forbs. Annual above‐ground phytomass accrual was greater on plots which were defoliated once in mid‐summer compared to plots which were not defoliated during the growing season, but there was no interactive effect of defoliation and CO₂ on growth. Leaf photosynthesis was often greater in Pascopyrum smithii (C₃) and Bouteloua gracilis (C₄) plants in the elevated chambers, due in large part to higher soil water contents and leaf water potentials. Persistent downward photosynthetic acclimation in P. smithii leaves prevented large photosynthetic enhancement for elevated CO₂‐grown plants. Shoot N concentrations tended to be lower in grasses under elevated CO₂, but only Stipa comata (C₃) plants exhibited significant reductions in N under elevated compared to ambient CO₂ chambers. Despite chamber warming of 2.6 °C and apparent drier chamber conditions compared to unchambered controls, above‐ground production in all chambers was always greater than in unchambered plots. Collectively, these results suggest increased productivity of the shortgrass steppe in future warmer, CO₂ enriched environments.     

Subdominant species distribution in microsites around two life forms at a desert grassland‐shrubland transition zone

Question: In the same landscape context — at a desert grassland‐shrubland transition zone, how does subdominant plant abundance vary in microsites around dominant grasses and shrubs? Location: Sevilleta LTER, New Mexico, USA (34°21’N; 106°53’W; 1650 m a.s.l.). Methods: We compared the distribution of subdominant plants in canopy, canopy edge and interspace microsites around individual shrubs (Larrea tridentata) and grasses (Bouteloua eriopoda) at a transition zone that has been encroached by shrubs within the past 50 ‐ 100 a. Plots of variable size according to microsite type and dominant plant size were sampled. Results: Subdominant abundance was higher in microsites around L. tridentata shrubs than in microsites around B. eriopoda. Furthermore, differences in species abundance and composition were higher among microsites around grasses than among microsites around shrubs. The distribution of subdominants was mostly explained by their phenological characteristics, which indicates the importance of temporal variation in resources to their persistence. Conclusions: This study of coexistence patterns around dominants revealed ecological contrasts between two dominant life forms, but other factors (such as disturbances) have to be taken into consideration to evaluate landscape‐scale diversity.     

Livestock exclusion increases the spatial heterogeneity of vegetation in Colorado shortgrass steppe

Abstract. Spatial heterogeneity, an important characteristic in semi‐arid grassland vegetation, may be altered through grazing by large herbivores. We used Moran's I, a measure of autocorrelation, to test the effect of livestock grazing on the fine scale spatial heterogeneity of dominant plant species in the shortgrass steppe of northeastern Colorado. Autocorrelation in ungrazed plots was significantly higher than in grazed plots for the cover of the dominant species Bouteloua gracilis, litter cover and density of other bunchgrasses. No species had higher autocorrelation in grazed compared to ungrazed sites. B. gracilis cover was significantly auto‐correlated in seven of eight 60‐yr ungrazed exclosures, four of six 8‐yr exclosures, and only three of eight grazed sites. Autocorrelograms showed that B. gracilis cover in ungrazed sites was frequently and positively spatially correlated at lag distances less than 5 m. B. gracilis cover was rarely autocorrelated at any sampled lag distance in grazed sites. The greater spatial heterogeneity in ungrazed sites appeared linked to patches characterized by uniformly low cover of B. gracilis and high cover of C₃ grasses. This interpretation was supported by simple simulations that modified data from grazed sites by reducing the cover of B. gracilis in patches of ca. 8 m diameter and produced patterns quite similar to those observed in ungrazed sites. In the one exclosure where we intensively sampled soil texture, autocorrelation coefficients for sand content and B. gracilis cover were similar at lag distances up to 12 m. We suggest that the negative effect of sand content on B. gracilis generates spatial heterogeneity, but only in the absence of grazing. An additional source of heterogeneity in ungrazed sites may be the negative interaction between livestock exclusion and B. gracilis recovery following patchy disturbance.     

Distribution of range and cultivated grassland plants in southern Alberta

We analysed the geographic distribution of native grassland and introduced grassland plants and the association of species abundance with climate and site characteristics in southern Alberta using a database of range and pasture assessments collected during 1991, 1992 and 1993. Distribution maps of native and introduced grassland plants could be inferred from association between site data recording botanical composition or presence and mean monthly climate surfaces for May and August. Introduced species formed three main geographical patterns with alfalfa, smooth bromegrass and crested wheatgrass most widespread, Russian wildrye restricted to the south eastern quarter and timothy, creeping red fescue, bluegrass and clovers restricted to the northern half. Native plants formed four main patterns characterised by Stipa comata and Bouteloua gracilis – south eastern; Agropyron spp. – widespread; Festuca spp. – northern and foothills; and Stipa viridula – sparse. Relative abundance of Stipa comata and Bouteloua gracilis were related primarily to latitude and longitude and variables describing temperature during the growing season. Biomass increased and range condition decreased with increasing latitude reflecting the decline in dominant native species, the increase in introduced and invader species and the better growing season moisture in northern areas. The maps provide a recent snapshot of the condition of commercially utilised native and cultivated grasslands in Alberta. They constitute a valuable reference point for comparison with future climate or management-induced change.     

Woody plant invasion at a semi‐arid/arid transition zone: importance of ecosystem type to colonization and patch expansion

Question: How do patterns in colonization and patch expansion of an invasive woody plant (Larrea tridentata, Zygo‐phyllaceae) differ between two grassland ecosystems at a biome transition zone? Location: Semi‐arid/arid transition zone in central New Mexico. Methods: Frequency of occurrence, height, and surface area of saplings (n= 134) and patches of adult plants (n= 247) of the invasive shrub, L. tridentata, were measured within a mosaic of ecosystems dominated either by the Chihuahuan Desert species, Bouteloua eriopoda (Poaceae), or the shortgrass steppe species, B. gracilis, located within 1 km of the L. tridentata‐dominated ecosystem. Distances between L. tridentata patches and patch area were used to estimate connectivity as a measure of propagule pressure. Sapling age (estimated from height using previously established relationships) and distance to the L. tridentata‐dominated ecosystem was used to evaluate patterns in dispersal. Cover by species or functional group inside each L. tridentata patch was compared with surrounding vegetation to estimate changes in species composition with patch expansion. Results: L. tridentata saplings (< 1%) and adult patches (15%) occurred less frequently in B. gracilis‐dominated ecosystems than expected based on areal extent of this ecosystem type. Propagule pressure did not differ with distance from the core ecosystem dominated by L. tridentata. Evidence for both local and long‐distance dispersal events was found. Similar relationships between number of plants and patch area in both grassland types indicate similar patterns in patch expansion. Cover of perennial forbs was higher and cover of dominant grasses was lower in L. tridentata patches compared with the surrounding vegetation for both ecosystem types. Conclusions Spatial variation in L. tridentata saplings and patches at this biome transition zone is related to the different susceptibilities to invasion by two grassland ecosystems. The persistence of grasslands at this site despite region‐wide expansion by L. tridentata may be related to the spatial distribution of B. gracilis‐dominated ecosystems that resist or deter invasion by this woody plant.     

Development of microtopography in a semi-arid grassland: Effects of disturbance size and soil texture

Our objective was to evaluate effects of disturbance size and soil texture on the development of microtopography for a shortgrass plant community in north central Colorado USA. Disturbances, defined as the death of individual plants, were created in 1984 and 1985 to evaluate development through time of the small-scale pattern of perennial bunchgrasses and bare soil openings that characterize this semiarid grassland. Disturbed plots of three sizes (50, 100, 150 cm-diameter) comparable in size to naturally-occurring disturbances were produced by killing plants at two sites differing in soil texture (sandy loam, clay loam). Disturbed plots were not manipulated after being created. In 1993, a laser surveying instrument was used to measure heights of crowns of individual plants of the dominant species, the perennial bunchgrass Bouteloua gracilis ([H.B.K.] Lag. ex Griffiths), and bare soil openings between plants for two locations: within each disturbance and in the surrounding undisturbed landscape.Differences between crown heights of plants and bare soil openings were comparable for both the undisturbed landscape and inside disturbances indicating that small-scale microtopography had recovered within nine years after disturbance occurred. However, complete recovery to the undisturbed state had not occurred since crown heights of plants relative to bare soil openings were significantly less on disturbed than undisturbed locations. Differences in height between plant crowns and bare soil openings on disturbed plots increased as disturbance size increased, indicating greater soil redistribution with increasing plot size. Larger differences in height were also found on plots on the fine- than the coarse-textured soil, indicating the importance of soil particle size and plant cover type to the development of microtopography. Differences in height between microsites on disturbed plots were positively related to total plant cover and negatively related to cover of B. gracilis indicating the importance of this species to reducing erosion on disturbed areas.In this semiarid grassland, patterns in microtopography were heterogeneous, likely as a result of the small-scale redistribution of soil between bare soil openings and B. gracilis plants through time. Our results indicate that this redistribution of soil is affected by disturbance size, soil texture, and patchy plant cover. The major effect of small-scale disturbances on patterns in microtopography of the shortgrass steppe are causing plant death and exposing soil to erosional and depositional processes.     

Topographic and soil surface effects on gypsophile plant community patterns in central Mexico

Abstract. Semiarid gypsum karstlands in north central Mexico are characterized by a mosaic of shrubland and endemic gypsophile grassland community types. Proximal physical factors affecting community patterns on a local scale were investigated at an undisturbed site south of Matehuala, San Luis Potosi. Soil surface characteristics associated with depth of calcareous alluvium overlying gypsum base material were strongly correlated with distribution of desert shrub and endemic grass species. Muhlenbergia purpusii was dominant on bare gypsum soils with indurated crusts, while Bouteloua chasei was dominant where a thin veneer of alluvium permitted development of a cryptogamic crust. A high-diversity mixed shrub-succulent community occupied gypsum overlain by 5 - 20 cm of alluvium, whereas Larrea tridentata was the principal species on gypsum overlain by > 20 cm of alluvium. Open sink walls were occupied mostly by species of the Muhlenbergia grassland, while closed sinks supported more mesophytic mixed shrubland vegetation. Physical factors operating mostly at the establishment stage are probably more important in mediating local species patterns on gypsum than factors associated with mineral nutrition. Gypsum material was much less fertile than alluvial material but did not exclude widely distributed species unless exposed at the surface.