William K. Lauenroth

A Generalized Model of the Effects of Grazing by Large Herbivores on Grassland Community Structure

Current disturbance models do not adequately account for the wide range of responses by grassland plant communities to grazing by large generalist herbivores. The evolutionary history of grazing, an important factor in the response of grasslands to grazing, has not been explicitly addressed. Grazing history alone, however, is not a good predictor of plant-herbivore interactions. Interactions occur along gradients of convergent to divergent selection pressures with increasing environmental moisture and of intolerance to tolerance of grazing with increasingly long evolutionary histories of grazing. We suggest that feedback mechanisms between plants and grazing animals are well developed in grasslands with long evolutionary histories of grazing. Feedback mechanisms are manifest in the rapid switching capabilities (of plant species and modes of competition) of subhumid grasslands with long evolutionary histories of grazing and divergent selection pressures. Switching capabilities do not exist in semiarid grasslands with long evolutionary histories of grazing and convergent selection pressures. Rather, for heavily grazed dominant species dominance increases. Feedback mechanisms are not well developed in systems with short evolutionary histories of grazing. In these cases, the differences in response to grazing by semiarid and subhumid situations arise primarily from differences in the grazing tolerance of plants adapted to semiaridity or of plants adapted to competition for light and from the different effects of grazing on canopy structure.

Primary Production of the Central Grassland Region of the United States

Aboveground net primary production of grasslands is strongly influenced by the amount and distribution of annual precipitation. Analysis of data collected at 9500 sites throughout the central United States confirmed the overwhelming importance of water availability as a control on production. The regional spatial pattern of production reflected the east-west gradient in annual precipitation. Lowest values of aboveground net primary production were observed in the west and highest values in the east. This spatial pattern was shifted eastward during unfavorable years and westward during favorable years. Vari- ability in production among years was maximum in northern New Mexico and southwestern Kansas and decreased towards the north and south. The regional pattern of production was largely accounted for by annual precipitation. Production at the site level was explained by annual precipitation, soil water-holding capacity, and an interaction term. Our results support the inverse texture hypothesis. When precipitation is 370 mm/yr.

The effect of grazing on the spatial heterogeneity of vegetation

Grazing can alter the spatial heterogeneity of vegetation, influencing ecosystem processes and biodiversity. Our objective was to identify why grazing causes increases in the spatial heterogeneity of vegetation in some cases, but decreases in others. The immediate effect of grazing on heterogeneity depends on the interaction between the spatial pattern of grazing and the pre-existing spatial pattern of vegetation. Depending on the scale of observation and on the factors that determine animal distribution, grazing patterns may be stronger or weaker than vegetation patterns, or may mirror the spatial structure of vegetation. For each possible interaction between these patterns, we make a prediction about resulting changes in the spatial heterogeneity of vegetation. Case studies from the literature support our predictions, although ecosystems characterized by strong plant-soil interactions present important exceptions. While the processes by which grazing causes increases in heterogeneity are clear, how grazing leads to decreases in heterogeneity is less so. To explore how grazing can consistently dampen the fine-scale spatial patterns of competing plant species, we built a cell-based simulation model that features two competing plant species, different grazing patterns, and different sources of vegetation pattern. Only the simulations that included neighborhood interactions as a source of vegetation pattern produced results consistent with the predictions we derived from the literature review.

Long‐Term Forage Production of North American Shortgrass Steppe

We evaluated the relationship between annual forage production and annual and seasonal precipitation and temperature at a shortgrass steppe site in north-central Colorado using a long-term data set (52 yr). We also constructed a relationship between forage production and aboveground net primary production (ANPP). Precipitation fluctuated randomly, but temperature had clear warming and cooling trends including a 17-yr warming trend from 1974 to 1990. Forage production was significantly related to both annual and seasonal precipitation but not temperature. Precipitation events between 15 and 30 mm accounted for most of the variability in production because they accounted for most of the variability in precipitation and because they wetted the soil layers that have the largest effect on production. Forage production amplified variability in annual precipitation. Production showed time lags of several years in responding to increases in precipitation. Change in vegetation structure has a characteristic response time, which contrains production responses in wet years. Constraint caused by vegetation structure is the reason why regional ANPP-precipitation models have a steeper slope than long-term models and point out a weakness of exchanging space for time in predicting production patterns.

Small Rainfall Events: An Ecological Role in Semiarid Regions

SummarySmall 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.

Resource partitioning between shrubs and grasses in the Patagonian steppe

SummaryExperiments were conducted in the Patagonian steppe in southern South America to test the following hypotheses: (a) grasses take up most of the water from the upper layers of the soil and utilize frequent and short-duration pulses of water availability; (b) shrubs, on the contrary, take up most of the water from the lower layers of the soil and utilize infrequent and long-duration pulses of water availability. Grasses and shrubs were removed selectively and the performance of plants and the availability of soil resources were monitored. Results supported the overall hypothesis that grasses and shrubs in the Patagonian steppe use mainly different resources. Removal of shrubs did not alter grass production but removal of grasses resulted in a small increase in shrub production which was mediated by an increase in deep soil water and in shrub leaf water potential. The efficiency of utilization of resources freed by grass removal was approximately 25%. Shrubs used water exclusively from lower soil layers. Grasses took up most of the water from upper layers but they were also capable of absorbing water from deep layers. This pattern of water partitioning along with the lack of response in leaf nitrogen to the removal treatments suggested that shrubs may be at a disadvantage to grasses with respect to nutrient capture and led to questions about the role of nutrient recirculation, leaching, and nitrogen fixation in the steppe.

Effects of grazing, topography, and precipitation on the structure of a semiarid grassland

Structural aspects of the shortgrass steppe plant community, functional groups, and species populations were examined in response to long-term heavy grazing and exclosure from grazing, contiguous wet or dry years, and an environmental gradient of topography. Of the three factors, relatively greater differences in community similarity were observed between catena positions, particularly on the ungrazed treatments. Grazing was intermediate between catena position and short-term weather in shaping plant community structure. Grazed treatments and ridgetops had a less variable species composition through fluctuations in weather. An increase with grazing of the dominant, heavily grazed species was observed. Basal cover and density of total species was also greater on grazed sites. The more uniform grazing lawn structure of the grazed plant communities had an influence on segregation of plant populations along topographical gradients. Segregation was less on grazed catenas, but diversity and the abundance of introduced and opportunistic-colonizer species was also less. Although the shortgrass steppe community was relatively invariant, less abundant species were dynamic and interactions occurred with respect to grazing, weather, and catena position. The effects of grazing may be mitigated by favorable growing seasons but magnified in unfavorable years in populations that are adapted to favorable sites. Grazing can be considered a disturbance at the level of the individual but it may or may not be a disturbance at the level of the population, and it is not a disturbance at the level of the community in this particular grassland.

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.

Long‐Term Soil Water Dynamics in the Shortgrass Steppe

To assess the temporal and spatial dynamics of soil water at a shortgrass steppe site in northcentral Colorado, we evaluated the precipitation regime for a 33-yr period and ran a simulation model for this period. Small precipitation events accounted for a large fraction of the total number of events and represented a source of water with small interannual variability. The difference between wet and dry years was related to the occurrence of a few large events. Average daily precipitation was concentrated during the warmest months of the year with a maximum in late spring. Water in the surface soil layers had a short residence time and no seasonal pattern. Intermediate layers reflected the seasonal pattern of precipitation. Maximum soil water availability occurred in the late spring, but this was also the period with the highest interannual variability. The wettest layer was at 4-15 cm of depth. The frequency of wet conditions decreased above this layer because of the strong influence of evaporation and below because recharge was infrequent. No deep percolation events were recorded. During dry years distribution of soil water was very shallow and during wet years wet conditions reached to depths of 120-135 cm. This shallow distribution of soil water matches the distribution of processes and struc- tural elements in the steppe suggesting that there is a cause-effect relationship between them. We speculate that the pattern of water availability interacts with biotic constraints and determines the rate of ecosystem processes. The depth distribution of water in dry and wet years is compared to the root distribution of grasses, shrubs, herbs, and succulents to suggest the response of each group to modal and extreme conditions. Comparison of long- term soil water patterns and traits of the major species allows us to suggest why Bouteloua graci/is is the dominant species in the shortgrass steppe.

Plant-Soil interactions in temperate grasslands

We present a conceptual model in which plant-soil interactions in grasslands are characterized by the extent to which water is limiting. Plant-soil interactions in dry grasslands, those dominated by water limitation (‘belowground-dominance’), are fundamentally different from plant-soil interactions in subhumid grasslands, where resource limitations vary in time and space among water, nitrogen, and light (‘indeterminate dominance’). In the belowground-dominance grasslands, the strong limitation of soil water leads to complete (though uneven) occupation of the soil by roots, but insufficient resources to support continuous aboveground plant cover. Discontinuous aboveground plant cover leads to strong biological and physical forces that result in the accumulation of soil materials beneath individual plants in resource islands. The degree of accumulation in these resource islands is strongly influenced by plant functional type (lifespan, growth form, root:shoot ratio, photosynthetic pathway), with the largest resource islands accumulating under perennial bunchgrasses. Resource islands develop over decadal time scales, but may be reduced to the level of bare ground following death of an individual plant in as little as 3 years. These resource islands may have a great deal of significance as an index of recovery from disturbance, an indicator of ecosystem stability or harbinger of desertification, or may be significant because of possible feedbacks to plant establishment. In the grasslands in which the dominant resource limiting plant community dynamics is indeterminate, plant cover is relatively continuous, and thus the major force in plant-soil interactions is related to the feedbacks among plant biomass production, litter quality and nutrient availability. With increasing precipitation, the over-riding importance of water as a limiting factor diminishes, and four other factors become important in determining plant community and ecosystem dynamics: soil nitrogen, herbivory, fire, and light. Thus, several different strategies for competing for resources are present in this portion of the gradient. These strategies are represented by different plant traits, for example root:shoot allocation, height and photosynthetic pathway type (C3 vs. C4) and nitrogen fixation, each of which has a different influence on litter quality and thus nutrient availability. Recent work has indicated that there are strong feedbacks between plant community structure, diversity, and soil attributes including nitrogen availability and carbon storage. Across both types of grasslands, there is strong evidence that human forces that alter plant community structure, such as invasions by nonnative annual plants or changes in grazing or fire regime, alters the pattern, quantity, and quality of soil organic matter in grassland ecosystems. The reverse influence of soils on plant communities is also strong; in turn, alterations of soil nutrient supply in grasslands can have major influences on plant species composition, plant diversity, and primary productivity.