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.     

Does grazing mediate soil carbon and nitrogen accumulation beneath C4, perennial grasses along an environmental gradient?

An experiment was conducted to evaluate the influence of long-term (>25 yrs) grazing on soil organic carbon (SOC) and total soil nitrogen (N) accumulation beneath individual plants of three perennial grasses along an environmental gradient in the North American Great Plains. The zone of maximum SOC and N accumulation was restricted vertically to the upper soil depth (0-5 cm) and horizontally within the basal area occupied by individual caespitose grasses, which contributed to fine-scale patterning of soil heterogeneity. Long-term grazing mediated SOC and N accumulation in the tall-, mid- and shortgrass communities, but the responses were community specific. SOC and N were lower beneath Schizachyrium scoparium plants in long-term grazed sites of the tall- and midgrass communities, but higher beneath Bouteloua gracilis plants in the long-term grazed site of the shortgrass community. SOC, but not N, was greater in soils beneath compared to between S. scoparium plants in an abandoned field seeded in 1941, indicating that this caespitose grass accumulated SOC more rapidly than N. SOC and N were greater in the 0-5 cm soil depth beneath a caespitose grass (S. scoparium) compared to a rhizomatous grass (Panicum virgatum) in the tallgrass community, with no significant accumulation of either SOC or N beneath P. virgatum plants. Grazing appears to indirectly mediate nutrient accumulation beneath caespitose grasses along the environmental gradient by modifying the size class distribution of plants. Populations with a greater proportion of large plants have a greater potential for biomass incorporation into soils and may more effectively capture redistributed organic matter from between plant locations. Contrasting plant responses to grazing at various locations along the environmental gradient conform to the predictions of the generalized grazing model, as the selection pressures of grazing and aridity may have also influenced the ability of caespitose grasses to accumulate nutrients in soils beneath them by mediating grazing resistance, competitive ability and population structure.     

Rhizosphere microorganism effects on soluble amino acids,sugars and organic acids in the root zone ofAgropyron cristatum,A. smithii andBouteloua gracilis

Three axenic and rhizosphere microorganism-inoculated shortgrass steppe plant species were evaluated for possible differences in residual organic carbon and nitrogen present as sugars, organic acids and amino acids. IntroducedAgropyron cristatum was compared toA. smithii andBouteloua gracilis, which are dominant species in the native shortgrass steppe. These plants, grown for 90 days in root growth chambers, showed differences in residual organic carbon and nitrogen per gram of root, and rhizosphere microbe presence resulted in additional changes in these compounds. The root biomass ofB. gracilis was significantly increased with microbes present. TheAgropyron species had significantly lower amino acid levels with microbes present, while under the same conditions, theB. gracilis showed significant decreases in residual sugars. Based on the amino acids, sugars and organic acids, the C/N ratio of the sterileA. cristatum was higher than forB. gracilis. Rhizosphere microbe presence did not result in changes in these C/N ratios. These results suggest thatA. cristatum, with microbes present, will have lower levels of amino acids present, whileB. gracilis, with a lower C/N ratio, will have sugars used to a greater extent by the rhizosphere microbes. This resulted in the higher levels of residual soluble organic C and N in the rhizosphere ofB. gracilis, in comparison with the introducedA. cristatum. These differences may be critical in influencing the course of nutrient accumulation and plant competition in short-grass steppe communities, and in understanding basic aspects of plant-rhizosphere microorganism interactions.     

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.     

Conservation of nitrogen increases with precipitation across a major grassland gradient in the Central Great Plains of North America

Regional analyses and biogeochemical models predict that ecosystem N pools and N cycling rates must increase from the semi-arid shortgrass steppe to the sub-humid tallgrass prairie of the Central Great Plains, yet few field data exist to evaluate these predictions. In this paper, we measured rates of net N mineralization, N in above- and belowground primary production, total soil organic matter N pools, soil inorganic N pools and capture in resin bags, decomposition rates, foliar ¹⁵N, and N use efficiency (NUE) across a precipitation gradient. We found that net N mineralization did not increase across the gradient, despite more N generally being found in plant production, suggesting higher N uptake, in the wetter areas. NUE of plants increased with precipitation, and δ¹⁵N foliar values and resin-captured N in soils decreased, all of which are consistent with the hypothesis that N cycling is tighter at the wet end of the gradient. Litter decomposition appeared to play a role in maintaining this regional N cycling trend: litter decomposed more slowly and released less N at the wet end of the gradient. These results suggest that immobilization of N within the plant–soil system increases from semi-arid shortgrass steppe to sub-humid tallgrass prairie. Despite the fact that N pools increase along a bio-climatic gradient from shortgrass steppe to mixed grass and tallgrass prairie, this element becomes relatively more limiting and is therefore more tightly conserved at the wettest end of the gradient. Similar to findings from forested systems, our results suggest that grassland N cycling becomes more open to N loss with increasing aridity.     

Plant biodiversity on shortgrass steppe after 55 years of zero,light, moderate,or heavy cattle grazing

Shortgrass steppe rangeland near Nunn, Colorado, USA, has been lightly,moderately, or heavily grazed by cattle, or protected from grazing inexclosures, for 55 years. Plant species biodiversity and evenness were greatestin lightly- and moderately-grazed pastures. Both pastures weredominated by the warm-season shortgrass Boutelouagracilis, but the cool-season midgrasses Pascopyrumsmithii and Stipa comata contributedsignificantly to biomass production on the lightly-grazed pasture, asthey did in the exclosures. Diversity was least in the exclosures, which werestrongly dominated by the cactus Opuntia polyacantha.Buchloë dactyloides, another warm-seasonshortgrass, and Bouteloua gracilis were co-dominantsunder heavy grazing, and diversity was intermediate. Plant community structureand diversity were controlled by selective grazing by cattle and soildisturbance by cattle and rodents. Shortgrass steppe moderately or heavilygrazed by cattle was similar to and probably as sustainable as steppe grazed formillenia by bison and other wild ungulates.     

SPATIAL VARIABILITY IN SEED PRODUCTION OF THE PERENNIAL BUNCHGRASS BOUTELOUA GRACILIS (GRAMINEAE)

The effects of soil texture and grazing by cattle on the production of seeds of Bouteloua gracilis were evaluated for a semiarid grassland in northeastern Colorado. Ten locations were chosen to represent the range in soil textures and grazing intensities found at the Central Plains Experimental Range research site. Number of flowering culms, inflorescences and seeds, length of each flowering culm, total biomass of reproductive structures (culms, inflorescences, and seeds), and basal area were assessed for 96 B. gracilis plants at each location. Community-level estimates of density of flowering culms and density of viable seeds were made for each location. Both soil texture and grazing by cattle were important to spatial variability in seed production and other indicators of reproductive effort by B. gracilis. Grazing was important in mediating effects of soil texture. On locations protected from grazing, soil texture had significant effects on seed production; the largest number of seeds was produced on the coarsest-textured soil and the fewest number on the finest-textured soil. Relationships between seed production and clay content and between seed production and other indicators of reproductive effort by B. gracilis were different for grazed and ungrazed locations. Spatial variability in seed production of B. gracilis as a result of spatial variability in soil texture and grazing may be important to the continued dominance of this species in the presence of disturbances that vary in time and space.     

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.     

Grazing changes topography-controlled topsoil properties and their interaction on different spatial scales in a semi-arid grassland of Inner Mongolia, P.R. China

Semiarid steppe ecosystems account for large terrestrial areas and are considered as large carbon (C) sinks. However, fundamental information on topsoil sensitivity to grazing is lacking across different spatial scales including the effects of topography. Our interdisciplinary approach considering soil chemical, physical, and vegetation properties included investigations on pit scale (square-metre scale), plot scale (hectare scale), and the scale of a landscape section (several hectares). Five different sites, representing a grazing intensity gradient, ranging from a long-term grazing exclosure to a heavily grazed site were used. On the pit scale, data about aggregate size distribution, quantity of different soil organic carbon (SOC) pools, SOC mineralisation, hydraulic conductivity and shear strength was available for topsoil samples from representative soil profiles. Spatial variability of topographical parameters, topsoil texture, bulk density, SOC, water repellency, and vegetation cover was analysed on the basis of regular, orthogonal grids in differently grazed treatments by using two different grid sizes on the plot scale and landscape section. On the pit scale, intensive grazing clearly decreased soil aggregation and the amount of fresh, litter-like particulate organic matter (POM). The weak aggregation in combination with animal trampling led to an enhanced mineralisation of SOC, higher topsoil bulk densities, lower infiltration rates, and subsequently to a higher risk of soil erosion. On the plot scale, the effects of soil structure disruption due to grazing are enhanced by the degradation of vegetation patches and resulted in a texture-controlled wettability of the soil surface. In contrast, topsoils of grazing exclosures were characterised by advantageous mechanical topsoil characteristics and SOC-controlled wettability due to higher POM contents. A combined geostatistical and General Linear Model approach identified topography as the fundamental factor creating the spatial distribution of texture fractions and related soil parameters on the scale of a landscape section. Grazing strongly interfered with the topography-controlled particle relocation processes in the landscape and showed strongest effects on the aboveground biomass production and biomass-related soil properties like SOC stocks. We conclude that interdisciplinary multi-scale analyses are essential (i) to differentiate between topography- and grazing-controlled spatial patterns of topsoil and vegetation properties, and (ii) to identify the main grazing-sensitive processes on small scales that are interacting with the spatial distribution and relocation processes on larger 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.     

Grazing and Ecosystem Carbon Storage in the North American Great Plains

Isotopic signatures of ¹³C were used to quantify the relative contributions of C₃ and C₄ plants to whole-ecosystem C storage (soil+plant) in grazed and ungrazed sites at three distinct locations (short-, mid- and tallgrass communities) along an east–west environmental gradient in the North American Great Plains. Functional group composition of plant communities, the source and magnitude of carbon inputs, and total ecosystem carbon storage displayed inconsistent responses to long-term livestock grazing along this gradient. C₄ plants [primarily Bouteloua gracilis (H.B.K.) Lag ex Steud.] dominated the long-term grazed site in the shortgrass community, whereas the ungrazed site was co-dominated by C₃ and C₄ species; functional group composition did not differ between grazed and ungrazed sites in the mid- and tallgrass communities. Above-ground biomass was lower, but the relative proportion of fine root biomass was greater, in grazed compared to ungrazed sites at all three locations. The grazed site of the shortgrass community had 24% more whole-ecosystem carbon storage compared to the ungrazed site (4022 vs. 3236 g C m⁻²). In contrast, grazed sites at the mid- and tallgrass communities had slightly lower (8%) whole-ecosystem carbon storage compared to ungrazed sites (midgrass: 7970 vs. 8683 g C m⁻²; tallgrass: 8273 vs. 8997 g C m⁻²). Differential responses between the shortgrass and the mid- and tallgrass communities with respect to grazing and whole-ecosystem carbon storage are likely a result of: (1) maintenance of larger soil organic carbon (SOC) pools in the mid- and tallgrass communities (7476–8280 g C m⁻²) than the shortgrass community (2517–3307 g C m⁻²) that could potentially buffer ecosystem carbon fluxes, (2) lower root carbon/soil carbon ratios in the mid- and tallgrass communities (0.06–0.10) compared to the shortgrass community (0.20–0.27) suggesting that variation in root organic matter inputs would have relatively smaller effects on the size of the SOC pool, and (3) the absence of grazing-induced variation in the relative proportion of C₃ and C₄ functional groups in the mid- and tallgrass communities. We hypothesize that the magnitude and proportion of fine root mass within the upper soil profile is a principal driver mediating the effect of community composition on the biogeochemistry of these grassland ecosystems.     

Spatial heterogeneity of soil properties in areas under elephant-grass short-duration grazing system

Soil properties can be changed by several factors, such as plant roots and animal trampling. The identification of spatial heterogeneity of these properties depends on the sampling scale. This study was developed to test the hypothesis that soil chemical and physical properties beneath elephant-grass plants are different from those between them. The research was carried out in a soil classified as Kandiudalfic Eutrudox. Forty-eight soil samples were collected from 0-10 cm depth (24 beneath plants and 24 between plants). The following properties were measured: pH, organic matter, S, available P, K, Ca and Mg exchangeable, sum of bases, cation exchange capacity, base saturation percentage, dry-aggregate distribution, bulk density and soil penetration resistance. Statistical analyses (t test) indicated that there were no significant differences in soil chemical properties in relation to spatial position. However, significant differences were observed in soil physical properties, with higher values of bulk density and soil resistance to penetration between the plants than beneath the plants.     

Relationship Between Root Biomass and Soil Organic Matter Pools in the Shortgrass Steppe of Eastern Colorado

We examined the distribution of soil organic carbon (SOC) fractions and roots with depth to improve our understanding of belowground carbon dynamics in the shortgrass steppe of northern Colorado. Weaver and others (1935) found that the surface 15 cm of soil contained over 70% of the total roots found in a tallgrass prairie soil profile, while only accounting for 40% of the profile soil organic matter. We asked whether the relationship between roots and SOC that Weaver and others (1935) found in the tallgrass prairie was also found in the shortgrass steppe. Weaver and others (1935) suggested that the dissimilarity between belowground biomass and SOC with depth is the result of variability in decomposition rates. In an effort to determine whether patterns of SOC are the result of short-term plant input patterns or decomposition, we measured the ¹⁴C content of potentially mineralizable C and particulate organic matter (POM) C ten years after pulse labeling shortgrass steppe vegetation. We also estimated the mass specific decomposition rate constant (kPOM) for POM C through a shortgrass steppe soil profile. We found that the distribution of roots and SOM in the shortgrass steppe were similar to those observed in tallgrass prairie (Weaver and others 1935), with a higher proportion of total root biomass in the surface soils than total soil organic matter. Fifty-seven percent of root biomass was found in the surface 15-cm, while this same soil layer contained 23 percent of profile soil organic C. We measured the highest accumulation of ¹⁴C at the soil surface (12.0 ng ¹⁴C·m^-2·cm^-1 depth), with the least accumulation from 75-100 cm (0.724 ng ¹⁴C·m^-2·cm^-1 depth). The highest values of potentially mineralizable C were at the soil surface, with no significant differences in total mineralizable C among the 10-100 cm soil depths. The contribution of POM C to total C reached a profile minimum at the 15-20 cm depth increment, with profile maxima in the surface 5 cm and from 75-100 cm. We estimated that the proportion of particulate organic matter lost annually (kPOM) reached a profile maximum of 0.097 yr^-1 within the 10-15 cm depth increment. The 75-100 cm depth increment had the lowest kPOM value at 0.058 yr^-1. Thus, within the same physical fraction of SOC, decomposition rates vary with depth by nearly twofold. This pattern of high decomposition rates from 10-15 cm with lower decomposition rates at the soil surface and deeper in the soil profile may be the result of higher water availability in sub-surface soils in the shortgrass steppe.     

放牧强度对内蒙古荒漠草原土壤有机碳及其空间异质性的影响

放牧是内蒙古荒漠草原主要利用方式之一,研究不同放牧强度下土壤有机碳分布规律对退化草原恢复以及推广精准放牧技术具有重要的指导意义。基于不同放牧强度长期放牧样地（0、0.93、1.82、2.71羊单位hm^-2（a/2）^-1）,采用高样本数量的取样设计并结合地统计学分析方法,研究荒漠草原土壤有机碳及其空间异质性。结果表明：中度放牧会显著降低0-30 cm土层全氮含量（P<0.05）,全磷含量随放牧强度增强出现先降低后升高趋势;放牧样地土壤有机碳含量均显著低于对照样地（P<0.05）,不同放牧强度处理土壤有机碳含量没有显著差异;土壤有机碳密度受放牧影响在0-20 cm土层出现显著下降（P<0.05）,变化趋势同有机碳含量相似,碳氮比在重度放牧区0-10 cm土层显著降低（P<0.05）。土壤有机碳空间异质性和异质性斑块的破碎程度随放牧强度增加而增大;土壤有机碳含量与海拔高度在对照、轻度放牧和中度放牧区均呈极显著负相关（P<0.01）,在重度放牧区土壤有机碳含量和海拔无显著相关性;土壤有机碳含量与土壤全氮、全磷含量均呈极显著正相关（P<0.01）。综上所述,放牧降低土壤有机碳含量,提高土壤有机碳空间异质性,土壤有机碳含量的空间变异受海拔和土壤养分含量等因素的共同影响。     

Geographic patterns of simulated establishment of two Bouteloua species: implications for distributions of dominants and ecotones

Abstract. Our overall objective was to use a soil water model to predict spatial patterns in germination and establishment of two important perennial C₄-bunchgrasses across the North American shortgrass steppe and desert grassland regions. We also predicted changes in establishment patterns under climate change scenarios. Bouteloua gracilis dominates the shortgrass steppe from northeastern Colorado to southeastern New Mexico. Bouteloua eriopoda dominates desert grasslands in central and southern New Mexico. Germination and establishment for each species were predicted at 16 sites along the gradient using a daily time step, multi-layer soil water model (SOILWAT) to determine the percentage of years that temperature and soil water criteria for germination and establishment were met. Percentage of years with predicted establishment decreased from north to south for B. gracilis, but increased from north to south for B. eriopoda, comparable to observed dominance patterns. The 95 % confidence interval around the point at which simulated establishment were equal for the two species was near the location of the shortgrass steppe-desert grassland ecotone where both species are abundant. The intersection in percentage of years with establishment for the two species was predicted to move further north when climate was scaled using three Global Circulation Models (GCMs), indicating a possible northward expansion of B. eriopoda. Our results suggest that recruitment by seed may be an important process in determining, at least in part, the geographic distribution of these two species. Changes in climate that affect establishment constraints could result in shifts of species dominance that may or may not be accompanied by changes in species composition.     

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.     

The impact of grazing intensity on soil characteristics of Stipa grandis and Stipa bungeana steppe in northern China (autonomous region of Ningxia)

The effects of grazing intensity on selected soil characteristics in the feather-grass steppes of the autonomous region of Ningxia (northern China) were investigated by a comparison of non-grazed areas (grazing intensity 0), slightly grazed areas (grazing intensity I), moderately grazed areas (II), intensively grazed areas (III) and over-grazed areas (IV). Even in areas used only minimally for grazing activities (I), a serious increase (doubling) in soil hardness was apparent in the upper soil layer. A continual decrease in organic matter in the surface soil can be correlated directly to soil compaction. The content of organic matter in soil of degree IV amounts to only a third of the organic matter found in non-grazed areas. This decrease can be attributed partly to the poor living conditions for soil organisms in compacted soils, but also to a significant reduction in litter. This is because intensive grazing causes reduced vegetation cover leading to litter being blown away by wind or washed away by heavy rainfall. Thus in level III hardly any plant litter remained to be incorporated into the soil as humus. Likewise root density also suffered its largest decrease in areas with a grazing intensity level III. With regard to the content of nitrogen and phosphorous (total and available) hardly any difference between soils of grazing intensity 0 and I was observed, whereas a noticeable decrease was apparent between levels I and II. Available Potassium was similar for all grazing levels. The pH-value of the soil solution is not significantly affected by grazing. We did not observe differences in the soils of the two main types of steppe vegetation (Stipa grandis and Stipa bungeana steppe) in response to grazing. Only the amount of litter in the S. grandis-steppe in non-grazed or slightly grazed areas is noticeably higher than in the S. bungeana steppe.     

The spatial distribution of termites in shortgrass steppe: a geostatistical approach

The broad-scale distribution of subterranean termites (Reticulitermestibialis) was studied in a shortgrass-steppe ecosystem in northern Colorado, United States. Termite occurrence and abundance was measured over 4 months at 10-m intervals along a 900-m transect that spanned a topographic gradient. Geostatistics were used to model the probability of termite occurrence along the transect, and to identify the distributional extent and potential roles of termites in shortgrass steppe. Semivariance was calculated between sample pairs of differing distances and kriging was used to interpolate the probability of termite occurrence along the transect. The semivariogram showed spatial dependence in termite distribution between samples 10–330 m apart and converged on the population variance at distances >330 m, which suggested that spatial dependence explained much of the broad-scale variation in termite distribution. A relatively large nugget variance, however, indicated there was spatial dependence below the 10-m sampling resolution. Termites were most frequently found on a south-facing slope and in a lowland swale. Four-wing saltbush (Atriplexcanescens) was also common in these areas and is important in the production of woody litter. The distribution of termites was significantly associated with proximity to saltbush, which showed a strong spatial dependence at scales <500 m. Kriged probabilities of occurrence and cross-correlation between termites and shrubs showed that peak termite occurrence was shifted upslope 100 m from areas of closest shrub proximity. Other factors, such as soil temperature, texture, or organic matter, are therefore likely to influence termite distributions in shortgrass steppe. The geostatistical approach used here provides a basis for further study on termites in shortgrass steppe, where their roles in decomposition and nutrient cycling are unknown. Geostatistics could also be used to describe distribution patterns on other soil arthropods sampled from traps or soil cores along transects that span topographic or land-use changes. Received: 4 April 1997 / Accepted: 4 November 1997     

Effects of simulated grazing on different genotypes of Bouteloua gracilis: how important is morphology?

Populations of grasses exposed to grazing by vertebrates often exhibit reduced stature, increased tillering, reduced flowering, and other morphological differences which distinguish them from ungrazed populations. These differences frequently are interpreted as an adaptive response that reduces grazing damage; however, there are few experimental tests of this hypothesis. This paper describes a field experiment designed to determine whether morphological variation among genotypes of the grass Bouteloua gracilis is related to variation in their responses to grazing. Eleven genotypes differing in morphological and reproductive characters were transplanted into a shortgrass steppe community near Fort Collins, Colorado. Replicates of each genotype were subjected to clipping treatments intended to realistically simulate three grazing intensities. After two growing seasons, different genotypes still maintained significant differences in a wide range of morphological and demographic characters. However, there were few significant effects of grazing treatment, and no significant genotype×treatment interactions. These results suggest that for B. gracilis clipped in simulation of natural grazing, defoliation has few short-term effects on fitness components, and intrapopulation morphological variation has few consequences for defoliation resistance. Received: 22 March 1999 / Accepted: 26 October 1999     

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.