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The trade-offs between sexual reproduction and clonal propagation are of great significance in terms of ecology and evolution for clonal plants, and they vary with environmental change. Nitrogen (N) deposition can increase litter accumulation in grassland and promote litter decomposition and consequently increase nutrient availability. However, the response of plant reproduction to N and litter addition in grasslands remains unclear. We examined the combined effects of N addition and litter manipulation (i.e. initial litter, removal, addition) on sexual reproduction and clonal propagation of a perennial clonal grass, Leymus chinensis, at the population (total number) and individual (number per ramet) levels in an 11-year field experiment. Nitrogen addition and litter additionally decreased spike and seed number at the population level, and N addition reduced those at the individual level. Nitrogen addition and litter interactively affected bud number at the two levels, and N addition decreased bud number in the litter removal treatments and increased that in the litter addition treatments. The increased soil available N, rather than light and soil water content, explained the change in sexual reproduction and clonal propagation. The positive effects of litter addition on bud number in the treatments with N addition, suggested that the N deposition increased litter accumulation could intensify clonal propagation of perennial grasses and increase their dominance in grasslands. Grassland management that reduces litter accumulation, such as grazing and mowing, can therefore, help alleviate the negative effects of N deposition on plant diversity through decreasing clonal propagation of the dominant species.

Atmospheric nitrogen (N) deposition reduces plant diversity. However, it often remains unclear how dominant species and litter accumulation feedbacks mediate N-induced plant diversity declines. We tested mechanisms of N-induced diversity change through dominant grasses and litter in a 7-year field experiment. Nitrogen addition reduced species richness, Shannon-Wiener diversity (H') and evenness from the second to the fourth year, however, surprisingly, increased them in the sixth and seventh year. The reversal in the response of diversity to N addition was explained by changes in grass dominance and standing litter accumulation. The diversity recovery during later years in fertilized plots was attributed to a decrease in the dominant grass and an increase in standing litter: standing litter reduced bud numbers of the dominant grass by decreasing light availability. The decreased light availability by standing litter reduced completion from the dominant species, which resulted in diversity increase. The negative feedback between dominant grasses and standing litter led to transient N-induced diversity loss in the short-term, but recovery of plant diversity in the long-term. Grassland management that affects litter accumulation, such as firing, grazing and mowing, can therefore, have substantial effects on the long-term response of plant diversity to N deposition.

Close spatial relationships between plant species are often important for defense against herbivory. The associational plant defense may have important implications for plant community structure, species diversity, and species coexistence. An increasing number of studies have focused on associational plant defense against herbivory at the scale of the individual plant and its nearest neighbors. However, the average neighborhood effects between plant species at the scale of whole plant communities have received almost no attention. The aims of this study were to determine patterns of spatial relationship between different plant species that can provide effective defense against herbivory. We conducted a manipulative experiment using sheep and three native plant species with different palatability. Consumption of palatable plants by herbivores was largest when the three plant species were isolated in three patches and independent of each other. A homogenous and spatially equal neighbor relationship between the three species did not reduce the risk of herbivory of palatable species compared to isolation of these species, but it reduced the total intake of all plant species. The palatable species was subject to less herbivory in a complex spatial neighborhood of several plant species. High complexity of spatial neighborhood resulted in herbivores passively reducing selectivity, thereby reducing the probability of damage to palatable species in the community, or making inaccurate judgments in foraging selectivity between and within patches, thereby reducing the vulnerability of palatable plants and even the whole plant community. We conclude that compelling herbivores to passively reduce the magnitude of foraging selectivity by establishing spatially complex neighborhoods between plant species is a compromise and optimal spatial strategy by plants to defend themselves again herbivory. This may contribute not only to maintenance of plant species diversity but also to a stable coexistence between herbivores and plants in grassland ecosystems.