Effects of dispersal, shrubs, and density-dependent mortality on seed and seedling distributions in temperate forests

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Abstract
Processes limiting recruitment of trees may have large impacts on forest dynamics. In this paper, we determined the effects of dispersal, shrubs (Rhododendron maximum), and density-dependent mortality on seed and seedling distributions of Southern Appalachian trees. We quantified the spatial distribution of seed rain, seed bank densities, first-year seedlings, and older than first-year seedlings in five vegetation plots. We fit models to these data assuming effects of limited dispersal, R. maximum (an understory shrub), and (or) density-dependent mortality (as well as a null model with none of these effects) and used best-fitting models to indicate which processes affected a particular species. We found that all factors examined limit species distributions, and thus, affect seedling dynamics. Seedling densities are higher near parent trees long after dispersal occurs. This pattern is less frequently observed for animal-dispersed species than for wind-dispersed species, presumably due to secondary dispersal of seeds by animals. Seedling densities of five species are decreased beneath R. maximum. Shade tolerance does not explain which species are affected, suggesting that factors other than low light are responsible for increased seedling mortality under this shrub. Our results suggest that density-dependent mortality affects four species, decreasing seedling densities close to parent trees. Dispersal, density-dependent mortality, and R. maximum all act in ways that may promote or limit diversity, illustrating that multiple factors are likely to control species diversity.