Patterns of Herbivory and Resource Utilization of Tree Seedlings Along an Altitudinal Gradient

In montane areas, climate change can shift tree species distributions upslope over time which can affect forest ecosystem structure and functioning. Seedlings of low-elevation temperate broadleaf trees establishing beyond their ranges at high elevations need to overcome both herbivore pressure and soil nutrient limitations. To assess the influence of these two stressors, we quantified leaf and stem herbivory, soil and foliar nutrient status, and two-year survival of experimentally grown seedlings of two hardwood species, sugar maple (Acer saccharum) and American beech (Fagus grandifolia) along elevation gradients in the Green Mountains of Vermont, USA. While insect foliar herbivory was reduced on maple seedlings growing beyond range boundaries, suggesting enemy escape, the opposite pattern was observed for beech. Mammalian browsing increased with elevation for both species. In general, foliar nutrient concentrations and resource utilization (that is, the relationship between soil nutrient availability and foliar nutrient status) declined with elevation for both species (although more so for maple, especially calcium), while toxic foliar aluminum concentrations increased for maple. Survival decreased with elevation for both species, but especially for sugar maple, linked more to declining foliar nutrient status than herbivory at higher elevations. Thus, the effects of herbivory and nutrient utilization on seedling survival can be critical for shaping tree species range shifts and, ultimately, species composition and forest ecosystem functioning.