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Variation in the competitive ability of plant species is suggested to determine their persistence and abundance in communities. In this study, we examine competition between grasses and woody plants, a phenomenon that occurs word-wide in savannas and along the margins between grasslands and forests. Specifically, we quantified the per- gram competitive effects of grasses and woody plants in native mixed-grass prairie by manipulating the natural vegetation. We separated the competitive effects of grasses, shrubs, and intact vegetation using selective herbicides to create four competition treatments: no neighbours (NN), no shrubs (NS), no grasses (NG), and all neighbours (AN). Competition treatments were applied to 2m x 2m experimental plots located in either prairie- or shrub- dominated habitats. Herbicides were applied starting one year prior to the study to create the four competition treatments. The effects of grasses and shrubs on resource availability (light, water, nitrogen) and the growth of two transplant species, Bouteloua gracilis, a perennial tussock grass, and Elaeagnus commutata, a common shrub, were measured over two growing seasons. Resource availability (i.e. light, soil moisture, soil available nitrogen) was two- to tenfold higher in no neighbour (NN) plots than in vegetated plots (NS, NG, AN). Both grasses and shrubs reduced resource levels to about the same extent. Light penetration declined linearly with increasing grass or shrub biomass, to a minimum of about 30% incident light at 500 g/m² neighbour shoot mass. Soil resources (i.e. water, nitrogen) did not decline with increasing neighbour shoot or root mass for either grasses or shrubs. Thus, for soil resources, the presence of neighbours was more important than their mass (i.e. abundance). Transplant growth did not decline with increasing neighbour biomass; the only exception to this pattern was a linear decline in Bouteloua growth with increasing neighbour shoot mass in plots containing only shrubs (NG). Competition was equally intense in prairie-dominated and shrub-dominated habitats for transplants of Bouteloua, but was less intense in shrub-dominated habitats for the shrub Elaeagnus. These results raise the interesting possibility that variation in the persistence and abundance of plants in communities is more strongly controlled by variation in competitive effects than by differences in competitive response ability.
1 Atmospheric nitrogen (N) deposition has become one of the most important agents of vegetation change in densely populated regions, It may also contribute to forest expansion into grasslands at the northern edge of the North American Great Plains. 2 We measured N deposition and available soil N with ion-exchange resin over 2 years in six national parks in areas varying in population density and industrialization. N deposition was significantly higher in four parks in densely populated regions than in two remote parks. 3 Available soil N increased significantly with N deposition across all parks. 4 We measured N mass and N-15 abundance (delta N-15) in vegetation and soil in two parks: Elk Island, receiving 22 kg N ha(-1) year(-1), and Jasper, receiving 8 kg N ha(-1) year(-1). Differences between parks in tissue N concentrations were small, but forest expansion over five decades resulted in the mass of N in vegetation increasing by 74% in Elk Island but by only 26% in Jasper. delta N-15 in forest vegetation was significantly lower in Elk Island than in Jasper, suggesting that anthropogenic sources contribute significantly to the high rates of N entering that ecosystem. 5 We determined the rate of forest expansion within parks using six decades of aerial photographs, Parks in aspen parkland and boreal forest showed a strong positive relationship between forest expansion and N deposition. The relationships found between N deposition, available soil N and forest expansion suggest that even comparatively low rates of N deposition may accelerate the expansion of forest into temperate grasslands.
Standing crop and species composition in semiarid grassland are linked to long-term patterns of water availability, but grasslands are characterized by large single-season variability in rainfall. We tested whether a single season of altered water availability influenced the proportions of grasses and shrubs in a semiarid grassland near the northern edge of the North American Great Plains. We studied stands of the clonal shrub snowberry (Symphoricarpos occidentalis) and adjacent grassland dominated by the native grasses Stipa spartea and Bouteloua gracilis. Rain was excluded and water supplied in amounts corresponding to years of low, medium, and high rainfall, producing a 2 - 4-fold range in monthly precipitation among water supply treatments. There were ten replicate plots of each water treatment in both snowberry stands and grassland. Grass standing crop increased significantly with water availability in grassland but not inside snowberry stands. Total standing crop and shrub stem density increased significantly with water supply, averaged across both communities. In contrast, water had no effect on shrub standing crop or light penetration. In summary, our finding that water has significant effects on a subset of components of grassland vegetation is consistent with long-term, correlational studies, but we also found that a single season of altered water supply had no effect on other important aspects of the ecosystem.
Variation in nitrogen deposition and available soil nitrogen in a forest–grassland ecotone in Canada
(2004)
Regional variation in nitrogen (N) deposition increases plant productivity and decreases species diversity, but landscape- or local-scale influences on N deposition are less well-known. Using ion-exchange resin, we measured variation of N deposition and soil N availability within Elk Island National Park in the ecotone between grassland and boreal forest in western Canada. The park receives regionally high amounts of atmospheric N deposition (22 kg ha⁻¹ yr⁻¹). N deposition was on average higher ton clayrich luvisols than on brunisols, and areas burned 1 – 15 years previously received more atmospheric N than unburned sites. We suggest that the effects of previous fires and soil type on deposition rate act through differences in canopy structure. The magnitude of these effects varied with the presence of ungulate grazers (bison, moose, elk) and vegetation type (forest, shrubland, grassland). Available soil N (ammonium and nitrate) was higher in burned than unburned sites in the absence of grazing, suggesting an effect of deposition. On grazed sites, differences between fire treatments were small, presumably because the removal of biomass by grazers reduced the effect of fire. Aspen invades native grassland in this region, and our results suggest that fire without grazing might reinforce the expansion of forest into grassland facilitated by N deposition.
Regional variation in nitrogen (N) deposition increases plant productivity and decreases species diversity, but landscape- or local -scale influences on N deposition are less well-known. Using ion-exchange resin, we measured variation of N deposition and soil N availability within Elk Island National Park in the ecotone between grassland and boreal forest in western Canada. The park receives regionally high amounts of atmospheric N deposition (22 kg ha(-1) yr(- 1)). N deposition was on average higher ton clay-rich luvisols than on brunisols, and areas burned 1-15 years previously received more atmospheric N than unburned sites. We suggest that the effects of previous fires and soil type on deposition rate act through differences in canopy structure. The magnitude of these effects varied with the presence of ungulate grazers (bison, moose, elk) and vegetation type (forest, shrubland, grassland). Available soil N (ammonium and nitrate) was higher in burned than unburned sites in the absence of grazing, suggesting an effect of deposition. On grazed sites, differences between fire treatments were small, presumably because the removal of biomass by grazers reduced the effect of fire. Aspen invades native grassland in this region, and our results suggest that fire without grazing might reinforce the expansion of forest into grassland facilitated by N deposition
Global Circulation Models of climate predict not only a change of annual precipitation amounts but also a shift in the daily distribution. To improve the understanding of the importance of daily rain pattern for annual plant communities, which represent a large portion of semi-natural vegetation in the Middle East, I used a detailed, spatially explicit model. The model explicitly considers water storage in the soil and has been parameterized and validated with data collected in field experiments in Israel and data from the literature. I manipulated daily rainfall variability by increasing the mean daily rain intensity on rainy days (MDI, rain volume/day) and decreasing intervals between rainy days while keeping the mean annual amount constant. In factorial combination, I also increased mean annual precipitation (MAP). I considered five climatic regions characterized by 100, 300, 450, 600, and 800 mm MAP. Increasing MDI decreased establishment when MAP was >250 mm but increased establishment at more arid sites. The negative effect of increasing MDI was compensated by increasing mortality with increasing MDI in dry and typical Mediterranean regions (c. 360-720 mm MAP). These effects were strongly tied to water availability in upper and lower soil layers and modified by competition among seedlings and adults. Increasing MAP generally increased water availability, establishment, and density. The order of magnitudes of MDI and MAP effects overlapped partially so that their combined effect is important for projections of climate change effects on annual vegetation. The effect size of MAP and MDI followed a sigmoid curve along the MAP gradient indicating that the semi-arid region (?300 mm MAP) is the most sensitive to precipitation change with regard to annual communities.
This contribution describes a generator of stochastic time series of daily precipitation for the interior of Israel from c. 90 to 900 mm mean annual precipitation (MAP) as a tool for studies of daily rain variability. The probability of rainfall on a given day of the year is described by a regular Gaussian peak curve function. The amount of rain is drawn randomly from an exponential distribution whose mean is the daily mean rain amount (averaged across years for each day of the year) described by a flattened Gaussian peak curve. Parameters for the curves have been calculated from monthly aggregated, long-term rain records from seven meteorological stations. Parameters for arbitrary points on the MAP gradient are calculated from a regression equation with MAP as the only independent variable. The simple structure of the generator allows it to produce time series with daily rain patterns that are projected under climate change scenarios and simultaneously control MAP. Increasing within-year variability of daily precipitation amounts also increases among-year variability of MAP as predicted by global circulation models. Thus, the time series incorporate important characteristics for climate change research and represent a flexible tool for simulations of daily vegetation or surface hydrology dynamics.
Global Circulation Models of climate predict not only a change of annual precipitation amounts but also a shift in the daily distribution. To improve the understanding of the importance of daily rain pattern for annual plant communities, which represent a large portion of semi-natural vegetation in the Middle East, I used a detailed, spatially explicit model. The model explicitly considers water storage in the soil and has been parameterized and validated with data collected in field experiments in Israel and data from the literature. I manipulated daily rainfall variability by increasing the mean daily rain intensity on rainy days (MDI, rain volume/day) and decreasing intervals between rainy days while keeping the mean annual amount constant. In factorial combination, I also increased mean annual precipitation (MAP). I considered five climatic regions characterized by 100, 300, 450, 600, and 800 mm MAP. Increasing MDI decreased establishment when MAP was >250 mm but increased establishment at more arid sites. The negative effect of increasing MDI was compensated by increasing mortality with increasing MDI in dry and typical Mediterranean regions (c. 360–720 mm MAP). These effects were strongly tied to water availability in upper and lower soil layers and modified by competition among seedlings and adults. Increasing MAP generally increased water availability, establishment, and density. The order of magnitudes of MDI and MAP effects overlapped partially so that their combined effect is important for projections of climate change effects on annual vegetation. The effect size of MAP and MDI followed a sigmoid curve along the MAP gradient indicating that the semi-arid region (≈300 mm MAP) is the most sensitive to precipitation change with regard to annual communitie
Germination rates and germination fractions of seeds can be predicted well by the hydrothermal time (HTT) model. Its four parameters hydrothermal time, minimum soil temperature, minimum soil moisture, and variation of minimum soil moisture, however, must be determined by lengthy germination experiments at combinations of several levels of soil temperature and moisture. For some applications of the HTT model it is more important to have approximate estimates for many species rather than exact values for only a few species. We suggest that minimum temperature and variation of minimum moisture can be estimated from literature data and expert knowledge. This allows to derive hydrothermal time and minimum moisture from existing data from germination experiments with one level of temperature and moisture. We applied our approach to a germination experiment comparing germination fractions of wild annual species along an aridity gradient in Israel. Using this simplified approach we estimated hydrothermal time and minimum moisture of 36 species. Comparison with exact data for three species shows that our method is a simple but effective method for obtaining parameters for the HTT model. Hydrothermal time and minimum moisture supposedly indicate climate related germination strategies. We tested whether these two parameters varied with the climate at the site where the seeds had been collected. We found no consistent variation with climate across species, suggesting that variation is more strongly controlled by site-specific factors.
In a recent contribution in Nature (vol. 442, pp. 555-558) Austin & Vivanco showed that sunlight is the dominant factor for decomposition of grass litter in a semi-arid grassland in Argentine. The quantification of this effect was portrayed as a novel finding. I put this result in the context of three other publications from as early as 1980 that quantified photodegradation. My synopsis shows that photodegradation is an important process in semi-arid grasslands in South America, North America and eastern Europe.