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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. 

Aims: Climate change is expected to shift climatic envelopes of temperate tree species into boreal forests where unsuitable soils may limit range expansion. We studied several edaphic thresholds (mycorrhizae, soil chemistry) that can limit seedling establishment of two major temperate tree species, sugar maple (arbuscular mycorrhizal, AM) and American beech (ectomycorrhizal, EM). Methods: We integrate two field surveys of tree seedling density, mycorrhizal colonization, and soil chemistry in montane forests of the Adirondack and Green Mountains (Mtns) in the northeastern United States. We conducted correlation and linear breakpoint analyses to detect soil abiotic and biotic thresholds in seedling distributions across edaphic gradients. Results: In the Green Mtns, sugar maple seedling importance (an index of species relative density and frequency, IV) declined sharply with low pH (< 3.74 in mineral soil) and low mycorrhizal colonization (< 27.5% root length colonized). Sugar maple importance was highly correlated with multiple aspects of soil chemistry, while beech was somewhat sensitive to pH only; beech mycorrhizal colonization did not differ across elevation. Mycorrhizal colonization of sugar maple was positively correlated with soil pH and conspecific overstory basal area. In the Adirondacks, sugar maple importance, but not beech, plateaued above thresholds in soil calcium (~ 2 meq/100 g) and magnesium (~ 0.3 meq/100 g). Conclusions: The establishment of sugar maple, but not beech, was impeded by both biotic and abiotic soil components in montane conifer forests and by soil acidity in temperate deciduous forests. These differences in species sensitivity to edaphic thresholds will likely affect species success and future shifts in forest composition.