Environmental change drivers reduce sapling layer diversity in sugar maple-beech forests of aastern North America
A century of beech bark disease (BBD) in North America has transformed hardwood forests by reducing the canopy biomass of American beech ( Fagus grandifolia ), even as beech has come to dominate the sapling layer of many forests. We do not understand the extent to which environmental change drivers such as climate, acidic atmospheric deposition (and its legacy of acidified soils), and invasive disease (BBD) may have contributed to this transformation. We investigated how BBD effects and tree community composition varied along a well-documented soil acidity gradient in the northeastern United States. We surveyed overstory and sapling layer tree species composition, BBD effects, and soil chemistry on 30 watersheds in forests co-dominated by beech and sugar maple ( Acer saccharum ). We analyzed potential drivers of community composition, BBD, and beech sapling density using linear models and non-metric multidimensional scaling. Predictors accounted for soil chemistry, climate, overstory beech (importance value, IV), mortality, and BBD defect. Overall overstory species composition varied most along the acidity gradient, while beech and BBD severity varied along their own distinct environmental gradient. Species composition of the overstory and sapling layers diverged significantly, with the latter dominated by beech. Beech sapling density was positively related to the proportion of standing dead overstory beech and soil exchangeable aluminum, but was unrelated to the overall proportion of overstory beech or their BBD severity. The dominance of sapling layers by beech may have resulted from a gradual accumulation of canopy-opening events precipitated by BBD and sugar maple decline, the latter driven by stressors such as acidification and climate change.