Quantifying deer and turkey leaf litter disturbances in the eastern deciduous forest: have nontrophic effects of consumers been overlooked?

Vertebrate herbivores and omnivores modify forest regeneration not only via direct and indirect trophic pathways, but also via nontrophic pathways such as litter and soil disturbances that may favor the regeneration of some species but not others. Deer are overabundant throughout vast portions of the eastern deciduous forest, and turkeys, once nearly extirpated, are now far more common; their foraging habits disturb litter over large areas, though this has rarely been evaluated. We quantified the size and frequency of litter disturbances created by both deer and turkeys. In addition, we tested the hypothesis that excluding vertebrates via fences would cause a reduction in the abundance and size of litter disturbances. We tested this hypothesis inside and outside six exclosures and adjacent control plots in an old-growth deciduous forest in Pennsylvania. Bare soil patches were ∼60% smaller and between 50% and 90% less abundant inside exclosures. The mean size of turkey litter disturbances was large (mean > 30 m²) and significantly greater than deer disturbances (p = 0.002), though turkey disturbances were less frequent. Our findings should apply broadly and are some of the first to demonstrate the extent of turkey disturbances. In addition, we demonstrate that exclosure studies reduce physical disturbances as well as browsing, both of which may synergistically act to cause changes in forest communities. We also caution that many deer fences exclude other vertebrates such as turkeys, which are important herbivores, seed predators, and major agents of disturbance. Consequently, we argue that studies that use fences to exclude deer should explicitly consider nontrophic indirect effects, particularly leaf disturbances and the potential impact of other large consumers as well (e.g., turkeys).