Abstract. Multiple regression analyses were performed on a worldwide 236-site data set compiled from studies that compared species composition, aboveground net primary production (ANPP), root biomass, and soil nutrients of grazed vs. protected, ungrazed sites. The objective was to quantitatively assess factors relating to differential sensitivities of ecosystems to grazing by large herbivores. A key question in this assessment was: Do empirically based, broad-scale relationships correspond to ecological theories of plantanimal interactions and conceptual frameworks for management of the world's grazing lands?Changes in species composition with grazing were primarily a function of ANPP and the evolutionary history of grazing of the site, with level of consumption third in importance. Changes in species composition increased with increasing productivity and with longer, more intense evolutionary histories of grazing. These three variables explained >50% of the variance in the species response of grasslands or grasslands-plus-shrublands to grazing, even though methods of measurement and grazing systems varied among studies. Years of protection from grazing was a significant variable only in the model for shrublands.Similar variables entered models of change in the dominant species with grazing. As with species composition, sensitivities of change in dominant species were greater to varying ecosystem-environmental variables than to varying grazing variables, from low to high values. Increases of the dominant species under grazing were predicted under some conditions, and decreases were more likely among bunch grasses than other life-forms and more likely among perennials than annuals. The response of shrublands was different from that of grasslands, both in terms of species composition and the dominant species. Our analyses support the perception of grazing as a factor in the conversion of grasslands to less desirable shrublands, but also suggest that we may be inadvertently grazing shrublands more intensively than grasslands.Percentage differences in ANPP between grazed and ungrazed sites decreased with increasingly long evolutionary histories of grazing and increased with increasing ANPP, levels of consumption, or years of treatment. Although most effects of grazing on ANPP were negative, some were not, and the statistical models predicted increases in ANPP with grazing under conditions of long evolutionary history, low consumption, few years of treatment, and low ANPP for grasslands-plus-shrublands. The data and the models support the controversial hypothesis that grazing can increase ANPP in some situations. Similar to species variables, percentage differences in ANPP between grazed and ungrazed treatments were more sensitive to varying ecosystem-environmental variables than to varying grazing variables. Within levels not considered to be abusive "overgrazing," the geographical location where grazing occurs may be more important than how many animals are grazed or how intensively an area is grazed.Counter to the...

Abstract. Aboveground net primary production of grasslands is strongly influenced by the amount and distribution of annual precipitation. Analysis of data collected at 9 500 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. Variability 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, sandy soils with low water-holding capacity are more productive than loamy soils with high waterholding capacity, while the opposite pattern occurs when precipitation is > 370 mm/yr.

Shifts in the timing of spring phenology are a central feature of global change research. Long-term observations of plant phenology have been used to track vegetation responses to climate variability but are often limited to particular species and locations and may not represent synoptic patterns. Satellite remote sensing is instead used for continental to global monitoring. Although numerous methods exist to extract phenological timing, in particular start-of-spring (SOS), from time series of reflectance data, a comprehensive intercomparison and interpretation of SOS methods has not been conducted. Here, we assess 10 SOS methods for North America between 1982 and 2006. The techniques include consistent inputs from the 8 km Global Inventory Modeling and Mapping Studies Advanced Very High Resolution Radiometer NDVIg dataset, independent data for snow cover, soil thaw, lake ice dynamics, spring streamflow timing, over 16 000 individual measurements of ground-based phenology, and two temperature-driven models of spring phenology. Compared with an ensemble of the 10 SOS methods, we found that individual methods differed in average day-of-year estimates by AE 60 days and in standard deviation by AE 20 days. The ability of the satellite methods to retrieve SOS estimates was highest in northern latitudes and lowest in arid, tropical, and Mediterranean ecoregions. The ordinal rank of SOS methods varied geographically, as did the relationships between SOS estimates and the cryospheric/hydrologic metrics. Compared with ground observations, SOS estimates were more related to the first leaf and first flowers expanding phenological stages. We found no evidence for time trends in spring arrival from ground-or model-based data; using an ensemble estimate from two methods that were more closely related to ground observations than other methods, SOS Correspondence: Michael A. White, tel. 1 1 435 797 3794, fax 1 1 435 797 187, trends could be detected for only 12% of North America and were divided between trends towards both earlier and later spring.

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.

Abstract. 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.

In dryland ecosystems, the timing and magnitude of precipitation pulses drive many key ecological processes, notably soil water availability for plants and soil microbiota. Plant available water has frequently been viewed simply as incoming precipitation, yet processes at larger scales drive precipitation pulses, and the subsequent transformation of precipitation pulses to plant available water are complex. We provide an overview of the factors that influence the spatial and temporal availability of water to plants and soil biota using examples from western USA drylands. Large spatial- and temporal-scale drivers of regional precipitation patterns include the position of the jet streams and frontal boundaries, the North American Monsoon, El Niño Southern Oscillation events, and the Pacific Decadal Oscillation. Topography and orography modify the patterns set up by the larger-scale drivers, resulting in regional patterns (10(2)-10(6) km2) of precipitation magnitude, timing, and variation. Together, the large-scale and regional drivers impose important pulsed patterns on long-term precipitation trends at landscape scales, in which most site precipitation is received as small events (< 5 mm) and with most of the intervals between events being short (< 10 days). The drivers also influence the translation of precipitation events into available water via linkages between soil water content and components of the water budget, including interception, infiltration and runoff, soil evaporation, plant water use and hydraulic redistribution, and seepage below the rooting zone. Soil water content varies not only vertically with depth but also horizontally beneath versus between plants and/or soil crusts in ways that are ecologically important to different plant and crust types. We highlight the importance of considering larger-scale drivers, and their effects on regional patterns; small, frequent precipitation events; and spatio-temporal heterogeneity in soil water content in translating from climatology to precipitation pulses to the dryland ecohydrology of water availability for plants and soil biota.

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Experiments 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.