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Large mammalian herbivores may have positive, neutral, or negative effects on annual net aboveground plant production (NAP) in different ecosystems, depending on their indirect effects on availability of key nutrients such as soil N. In comparison, less is known about the corresponding influence of grazers, and nutrient dynamics, over annual net belowground plant production (NBP). In natural multi-species plant communities, it remains uncertain how grazing influences relative allocation in the above- and belowground compartments in relation to its effects on plant nutrients. We evaluated grazer impacts on NAP, NBP, and relative investment in the above- and belowground compartments, alongside their indirect effects on soil N availability in the multiple-use Trans-Himalayan grazing ecosystem with native grazers and livestock. Data show that a prevailing grazing intensity of 51% increases NAP (+61%), but reduces NBP (-35%). Grazing also reduced C:N ratio in shoots (-16%) and litter (-50%), but not in roots, and these changes coincided with increased plant-available inorganic soil N (+23%). Areas used by livestock and native grazers showed qualitatively similar responses since NAP was promoted, and NBP was reduced, in both cases. The preferential investment in the aboveground fraction, at the expense of the belowground fraction, was correlated positively with grazing intensity and with improvement in litter quality. These results are consistent with hypothesized herbivore-mediated positive feedbacks between soil nutrients and relative investment in above- and belowground compartments. Since potentially overlapping mechanisms, such as N mineralization rate, plant N uptake, compositional turnover, and soil microbial activity, may contribute towards these feedbacks, further studies may be able to discern their respective contributions.

Mineral nitrogen (N) has been suggested as a potential factor causing declines in amphibian populations, especially in agricultural landscapes; however, there is a question as to whether it remains in the water column long enough to be toxic. We explored the hypothesis that mineral N can cause both lethal and sublethal toxic effects in amphibian embryos and larvae in a manipulative field experiment. We sampled 12 ponds, fertilizing half with ammonium nitrate fertilizer early in the spring, and measured hatching, survival, development, growth, and the incidence of deformities in native populations of wood frog (Rana sylvatica) and eastern tiger salamander (Ambystoma tigrinum tigrinum) embryos and larvae held in in situ enclosures. We found that higher ammonium concentrations negatively affect R. sylvatica more strongly than A. tigrinum. R. sylvatica tended to have lower survival as embryos and young tadpoles, slowed embryonic development, and an increased proportion of hatchlings with deformities at experimentally elevated ammonium. A. tigrinum did not experience significantly reduced survival, but their larval development was slowed in response to elevated ammonium and the abundance of large invertebrate predators. Variable species susceptibility, such as that shown by R sylvatica and A. tigrinum, could have large indirect effects on aquatic community structure through modification of competitive or predator-prey relationships. Ammonium and nitrate + nitrite concentrations were not correlated with other measures that might have affected amphibians, such as pH, pond area, depth, or vegetation. Our results highlight the potential importance of elevated ammonium on the growth, development and survival of amphibians, especially those that breed in surface waters receiving anthropogenic N inputs.

Terrestrial plant community responses to herbivory depend on resource availability, but the separate influences of different resources are difficult to study because they often correlate across natural environmental gradients. We studied the effects of excluding ungulate herbivores on plant species richness and composition, as well as available soil nitrogen (N) and phosphorus (P), across eight grassland sites in Serengeti National Park (SNP), Tanzania. These sites varied independently in rainfall and available soil N and P. Excluding herbivores decreased plant species richness at all sites and by an average of 5.4 species across all plots. Although plant species richness was a unimodal function of rainfall in both grazed and ungrazed plots, fences caused a greater decrease in plant species richness at sites of intermediate rainfall compared to sites of high or low rainfall. In terms of the relative or proportional decreases in plant species richness, excluding herbivores caused the strongest relative decreases at lower rainfall and where exclusion of herbivores increased available soil P. Herbivore exclusion increased among-plot heterogeneity in species composition but decreased coexistence of congeneric grasses. Compositional similarity between grazed and ungrazed treatments decreased with increasing rainfall due to greater forb richness in exclosures and greater sedge richness outside exclosures and was not related to effects of excluding herbivores on soil nutrients. Our results show that plant resources, especially water and P, appear to modulate the effects of herbivores on tropical grassland plant diversity and composition. We show that herbivore effects on soil P may be an important and previously unappreciated mechanism by which herbivores influence plant diversity, at least in tropical grasslands.

One cornerstone of ecological theory is that nutrient availability limits the number of species that can inhabit a community. However, the relationship between the spatial distribution of limiting nutrients and species diversity is not well established because there is no single scale appropriate for measuring variation in resource distribution. Instead, the correct scale for analyzing resource variation depends on the range of species sizes within the community. To quantify the relationship between nutrient distribution and plant species diversity, we measured NO(3)(-) distribution and plant species diversity in 16 paired, modified Whittaker grassland plots in Serengeti National Park, Tanzania. Semivariograms were used to quantify the spatial structure of NO(3)(-) from scales of 0.4-26 m. Plant species diversity (Shannon-Weiner diversity index; H ') was quantified in 1-m(2) plots, while plant species richness was measured at multiple spatial scales between 1 and 1000 m(2). Small-scale variation in NO(3)(-) (<0.4 m) was positively correlated with 1-m(2) H ', while 1000-m(2) species richness was a log-normal function of average NO(3)(-) patch size. Nine of the 16 grassland plots had a fractal (self-similar across scales) NO(3)(-) spatial distribution; of the nine fractal plots, five were adjacent to plots that had a non-fractal distribution of NO(3)(-). This finding offered the unique opportunity to test predictions of Ritchie and Olff (1999): when the spatial distribution of limiting resources is fractal, communities should display a left-skewed log-size distribution and a log-normal relationship between net primary production and species richness. These predictions were supported by comparisons of plant size distributions and biomass-richness relationships in paired plots, one with a fractal and one with a non-fractal distribution of NO(3)(-). In addition, fractal plots had greater large-scale richness than paired non-fractal plots (1,0-1000 m(2)), but neither species diversity ( H') nor richness was significantly different at small scales (1 m(2)). This result is most likely explained by differences in the scale of resource variation among plots: fractal and non-fractal plots had equivalent NO(3)(-) variation at small scales but differed in NO(3)(-) variation at large scales (as measured by the fractal dimension). We propose that small-scale variation in NO(3)(-) is largely due to the direct effects of plants on soil, while patterns of species richness at large scales is controlled by the patch size and fractal dimension of NO(3)(-) in the landscape. This study provides an important empirical step in understanding the relationship between the spatial distribution of resources and patterns of species diversity across multiple spatial scales.

1 Herbivores can play a key role in affecting ecosystem function, but their direct and indirect effects are often confounded with each other and have rarely been dissected. Predictions for open systems, i.e. those with cross-habitat nutrient fluxes and dispersal, may differ from those expected in closed systems, where no such transfers occur, but these differences have only recently begun to be characterized. 2 We present a theoretical model of plant productivity and soil nitrogen (N) based on the Serengeti ecosystem in order to investigate the interplay among herbivore movement, nutrient transport across habitats (spatial subsidies), the fire regime and the effects of herbivores on N cycling. 3 Model results suggest that the fire regime and herbivore migration are key determinants of primary productivity and fertility, and that the impact of these factors depends on grazing intensity. At low intensity, high grass biomass leads to extensive fires and N volatilization, but this effect is reduced at high grazing intensity. Without migration, the model predicts that primary productivity and fertility initially increase, then decline with increasing grazing intensity. Conversely, seasonal migration decouples the growing and grazing seasons, leading to a monotonic increase in productivity with grazing intensity. Cross-habitat N transport has a relatively modest effect on N dynamics, and the magnitude of the seasonality effect outweighs the spatial subsidy effect. 4 Our model suggests that herbivores and fire may play key interactive roles in regulating producers and determining ecosystem functional properties in grazing systems, both through consumption and via indirect effects on nutrient availability. The direction and magnitude of these effects could potentially vary greatly among ecosystems with resident vs. migratory herbivores, suggesting that spatial coupling through animal movement can be important at both the population and ecosystem levels. 5 This study suggests that the degree to which trophic interactions influence ecosystem function may be strongly modulated by whether systems are open or closed. It also suggests that direct and indirect effects of consumers on their resources (top-down effects) may be far more important than nutrient subsidies in open terrestrial systems. [ABSTRACT FROM AUTHOR]

Abstract: We examined how elevated atmospheric [CO2] and higher rate of nitrogen (N) input may influence grasshopper growth by changing food plant quality and how such effects may be modified by species diversity of the plant community. We reared grasshopper nymphs (Melanoplus femurrubrum) on Poa pratensis from field-grown monocultures or polycultures (16 species) that were subjected to either ambient or elevated levels of CO2 and N. Grasshopper growth rate was higher on P. pratensis leaves grown in monocultures than in polycultures, higher on P. pratensis grown under elevated than under ambient [CO2], and higher on P. pratensis grown under elevated than under ambient [N]. The higher growth rate observed on P. pratensis exposed to elevated [CO2] was, however, less pronounced for polyculture- than monoculture-grown P. pratensis. Growth rate of the grasshoppers was positively correlated with leaf [N], [C], and concentration of soluble carbohydrates+lipids. Concentration of non-structural carbohydrates+lipids was higher in leaves grown under elevated than under ambient [CO2], and the difference between P. pratensis grown under ambient and elevated [CO2] was greater for monoculture- than polyculture-grown P. pratensis. In addition, leaf N concentration was higher in P. pratensis grown in monocultures than in polycultures, suggesting that plant species richness, indirectly, may influence insect performance by changed nutritional value of the plants. Because we found interactive effects between all factors included ([CO2], [N], and plant species diversity), our results suggest that these parameters may influence plant–insect interactions in a complex way that is not predictable from the sum of single factor manipulations. [Copyright &y& Elsevier]