Climate change may restrict dryland forest regeneration in the 21st century.

The persistence and geographic expansion of dryland forests in the 21st century will be influenced by how climate change supports the demographic processes associated with tree regeneration. Yet, the way that climate change may alter regeneration is unclear. We developed a quantitative framework that estimates forest regeneration potential (RP) as a function of key environmental conditions for ponderosa pine, a key dryland forest species. We integrated meteorological data and climate projections for 47 ponderosa pine forest sites across the western United States, and evaluated RP using an ecosystem water balance model. Our primary goal was to contrast conditions supporting regeneration among historical, mid-21st century and late-21st century time frames. Future climatic conditions supported 50% higher RP in 2020-2059 relative to 1910-2014. As temperatures increased more substantially in 2060-2099, seedling survival decreased, RP declined by 50%, and the frequency of years with very low RP increased from 25% to 58%. Thus, climate change may initially support higher RP and increase the likelihood of successful regeneration events, yet will ultimately reduce average RP and the frequency of years with moderate climate support of regeneration. Our results suggest that climate change alone may begin to restrict the persistence and expansion of dryland forests by limiting seedling survival in the late 21st century.

Impact of labile and recalcitrant carbon treatments on available nitrogen and plant communities in a semiarid ecosystem.

In a 10-year study, we assessed the influence of five carbon (C) treatments on the labile C and nitrogen (N) pools of historically N-enriched plots on the Shortgrass Steppe Long Term Ecological Research site located in northeastern Colorado. For eight years, we applied sawdust, sugar, industrial lignin, sawdust + sugar, and lignin + sugar to plots that had received N and water additions in the early 1970s. Previous work showed that past water and N additions altered plant species composition and enhanced rates of nutrient cycling; these effects were still apparent 25 years later. We hypothesized that labile C amendments would stimulate microbial activity and suppress rates of N mineralization, whereas complex forms of carbon (sawdust and lignin) could enhance humification and lead to longer-term reductions in N availability. Results indicated that, of the five carbon treatments, sugar, sawdust, and sawdust + sugar suppressed N availability, with sawdust + sugar being the most effective treatment to reduce N availability. The year after treatments stopped, N availability remained less in the sawdust + sugar treatment plots than in the high-N control plots. Three years after treatments ended, reductions in N availability were smaller (40-60%). Our results suggest that highly labile forms of carbon generate strong short-term N sinks, but these effects dissipate within one year of application, and that more recalcitrant forms reduce N longer. Sawdust + sugar was the most effective treatment to decrease exotic species canopy cover and increase native species density over the long term. Labile carbon had neither short- nor long-term effects on exotic species. Even though the organic amendments did not contribute to recovery of the dominant native species Bouteloua gracilis, they were effective in increasing another native species, Carex eleocharis. These results indicate that organic amendments may be a useful tool for restoring some native species in the shortgrass steppe, though not all.

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

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

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

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

Forage quality in relation to long-term grazing history, current-year defoliation, and water resource.

Forage nitrogen concentrations, nitrogen yields, and in vitro digestibilities were assessed in shortgrass steppe that had been ungrazed, lightly, or heavily grazed for 50 years. Caged plots were defoliated in amounts based upon removals observed in naturallygrazed reference plots or not defoliated. This was done in a year of average precipitation and with a supplemental water treatment to simulate a wet year. In general, current-year defoliation had positive effects, and longterm grazing and supplemental water had negative effects, on forage nitrogen concentrations and digestibilities. However, defoliation interacted with long-term grazing in determning forage nitrogen concentrations, and with grazing and with watering in determining digestibilities. Nitrogen concentration and digestibility increased with defoliation in lightly, but not in heavily, grazed treatments. The dilution effect of supplemental water an digestibilities through increased plant growth was offset by defoliation. The negative effects of long-term grazing on forage quality were small, equally or more than compensated for by defoliation in a year of average precipitation, but more pronounced in the simulated wet year. Nitrogen yields and digestible forage production were usually increased by defoliation, but this depended upon grazing and watering treatments. Increased nitrogen and digestible forage yields and concentrations in response to defoliation were greater than the biomass response in lightly grazed grassland. For both nitrogen and digestibility, yields were greater in grazed than ungrazed treatments in the year of average precipitation, but less in the simulated wet year. Optimizing quantity and year-to-year stability of nitrogen and digestible forage yield may best be achieved with light grazing rather than no or heavy grazing. Clipping was conducted in a manner closely resembling the natural pattern and intensity of defoliation by the cattle, and confirms the potential for a positive feedback of increased forage quality with defoliation observed in pot experiments. Long-term heavy grazing can diminish this response. Quantily (aboveground primary production, ANPP), quantity of quality (digestible and N yields), and quality (concentrations) do not necessarily respond similarly in interactions between current-year defoliation, long-term grazing history, and level of water resource.

Neighborhood interactions in a natural population of the perennial bunchgrassBouteloua gracilis.

We analyzed neighborhood interactions in a natural population of the perennial bunchgrass blue grama (Bouteloua gracilis). Space occupation by individual plants was characterized in terms of neighborhood size. Neighborhood size was defined as the area potentially 'available' to an individual, which included the basal area of the plant and the bare area closer to the edges of the plant than to any others. Geographic Information Systems (GIS) were used to describe space partitioning. Growing season performance was evaluated as a function of neighborhood area and neighbor size, controlling for focal plant size. The area of the neighborhood was significant in explaining the remaining variation of allometric relationships between basal area and current vegetative and reproductive performance. In contrast, current performance of focal individuals was not related to the average basal area or the sum of basal areas of adjacent neighbors. Growing season performance was apparently affected by plant spacing, suggesting that competition for spatially distributed resources occurs. The presence of relatively small plants in neighborhoods with a high proportion of bare soil is consistent with the view of a community composed of patches undergoing their own successional dynamics. Competition and disturbances seem to play an important role in this semiarid grassland.

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.

Plant competition, abiotic, and long- and short-term effects of large herbivores on demography of opportunistic species in a semiarid grassland.

The emergence and subsequent survival and growth of five opportunistic "weeds" were monitored after seed additions to long-term grazing treatments with or without current-year grazing, long-term ungrazed treatments, and removal treatments designed to eliminate plant competition from existing perennials while either leaving vegetation and soil structure unaltered or disturbed. The treatments were applied on both uplands and lowlands to assess the relative influence of macroabiotic environment versus plant competition. The long-term effects of large herbivores on the initial emergence of seedlings were greater than the effects of removing competition. Very few individuals emerged on the long-term grazed treatments that were either grazed or ungrazed during the experiment. Numbers of individuals emerging on the long-term ungrazed treatments were greater or equal to those emerging on the no-competition-undisturbed treatments, but numbers were greatest on no-competition-disturbed treatments. None of the seeded individuals on the long-term grazed, currently grazed treatments survived to the end of the growing season. There was a slightly greater end-of-season biomass of seeded species and percentage of the total population reaching reproductive status on the long-term ungrazed compared with grazed-nondefoliated treatments, and very high survival, biomass, and proportions of reproductives on both no-competition treatments. Cover types in the immediate vicinity of seedlings influenced both germination and survival, but the effects differed between species and treatments. Equal compensation to current-year herbivory occurred on long-term heavily grazed treatments even though above-ground production was much greater on long-term protected sites. Productivity varied with topography, but very few topographic main effects or interactions occurred with demographic variables of seeded species, suggesting that macroabiotic effects were of minor importance compared with grazing and plant competition.

Resource partitioning between shrubs and grasses in the Patagonian steppe.

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.

Biomass dynamics and water use efficiencies of five plant communities in the shortgrass steppe.

Standing crop biomass and water-use efficiency were estimated for five plant communities of the Central Plains Experimental Range in north central Colorado. Aboveground biomass by functional groups, surface litter amounts, and standing dead biomass were compared, as were vertical and size-class distributions of belowground biomass. Greater production and water-use efficiency values were found: (1) at coarse-textured sites, indicating the importance of the inverse texture effect, and (2) wherever site characteristics favored the establishment of lifeforms other than grasses, e.g., succulents, and shrubs. Seasonal aboveground biomass and water-use efficiencies for the grass component were similar among sites, even though the mixes of C3 and C4 grass species were different. Similar grass biomass values in very different communities suggested that high biomass and high water-use efficiencies were related less to grass types than to the abundance of non-grass life-forms.

Small rainfall events: An ecological role in semiarid regions.

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

Water status of soil and vegetation in a shortgrass steppe.

In an attempt to describe some major relationships between soil and plant compartments in a shortgrass steppe, the process of water loss from the system and plant water relations throughout a drying cycle were studied. The water supply was manipulated and some soil and plant variables monitored throughout a drying cycle. Leaf conductance and leaf water potential of blue grama (Bouteloua gracilis) were measured periodically at predawn and noon. Soil water content and water potential of different layers were also monitored.Three different periods were distinguished in the water loss process throughout a drying cycle. These distinctions were made taking into account the relative contribution of different soil layers. Leaf conductance and water potential at noon slowly declined throughout the first 50 days of plant growth. After that, they rapidly decreased, reaching values of 0.29 mm s-1 and-5.0 MPa, respectively. The predawn leaf water potential remained unchanged around-0.5 MPa during the first 45 days, then rapidly decreased. This occurred when soil water of the wettest soil layer was near depletion.Predawn leaf water potentials were highly correlated with water potentials of the wettest layer. Leaf conductance and water potential at noon were correlated with effective soil water potential (soil water potential weighted by the root distribution in the profile). We concluded that root surface area limited the water flow through an important part of the day in this semiarid ecosystem. Axial root resistance did not appear important in determining the equilibrium status between leaves and the wettest soil layer.

Partitioning of carbon an SO2-sulfur in a native grassland.

The seasonal assimilation and within-plant partitioning of 14CO2-carbon and 35SO2-sulfur in field plots of mixed-grass prairie was investigated, as was the dry deposition of 35SO2 onto surfaces of dead leaves, litter, and soil, and possible effects of continuous low-level SO2 fumigation on these processes. The proportion of total net-assimilated carbon found below-ground was 45% in May, 51% in July, and 17% in September. As the season progressed, greater proportions of assimilate were partitioned to 5-20 cm depths and less to the 0-5 cm depth. Rhizomes and crowns received greater proportions in late season. Significant fractions of total 34SO2-deposited sulfur were recovered on dead leaf surfaces as well as litter and soil, suggesting estimates of SO2 removal based on stomatal resistance alone are inadequate. Only 4% to 7% of total deposited sulfur was translocated belowground, with most going to 0-5 cm roots. In July much greater proportions of the total translocated SO2-sulfur were found in deeper depths than in September. On SO2-fumigated plots roots had lower total sulfur concentrations than controls. Furthermore, while on control plots total sulfur in roots at 5-20 cm increased from May to July and decreased from July to September, on fumigated plots there was a decrease followed by an increase suggesting that SO2 uptake by shoots interferes with the normal pattern of root sulfur uptake and redistribution within the plant. Continuous SO2 fumigation also seemed to stimulate root growth in July, possibly through a stimulation of photosynthesis.

The effects of water- and nitrogen-induced stresses on plant community structure in a semiarid grassland.

A replicated factorial experiment was designed to test the hypothesis that manipulating inputs of water and mineral nitrogen to a semiarid grassland would disrupt existing interactions resulting in alteration of the structure of the primary producer community. Alteration of community structure was measured as either changes in growing season average biomass of 6 functional groups of plants or their relative contribution to total biomass.Additions of water greatly increased total biomass and resulted in the replacement of one of the dominant functional groups by a subordinate group. The water plus nitrogen treatment resulted in large biomass increases in two of the dominant functional groups, elimination of succulents as an important component of community structure, and establishment of several introduced weedy species. Continuation of the experiment will likely result in complete dominance of the water plus nitrogen treatment by these introduced species.Despite the large changes in community structure observed as a result of water- and nitrogen-induced stresses we conclude that the shortgrass prairie in northcentral Colorado is asymptotically stable with respect to these influences.

Dynamics of dry matter production in a mixed-grass prairie in western North Dakota.

Above- and belowground biomass of primary producers were estimated by the harvest method on 10 dates in 1969 in a mixed-prairie grassland. A range of estimates of above- and belowground net primary production is established using several methods of calculation. The range for aboveground production is 240 to 302 g·m-2 and 931 to 1221 g·m-2 for belowground production. Correlation analysis indicated that above- and belowground biomass dynamics are significantly (α≦0.05) related to air and soil temperature, soil water, precipitation, and vapor pressure deficit. Analysis of energy flow through primary producers indicates a net storage of energy in the standing dead, litter, and belowground compartments. Accumulation in the standing dead was 63% of inputs, in the litter 8%, and belowground 37%. Belowground decomposition was 57% of belowground production and the same value aboveground was 50%.