Foundation species loss and biodiversity of the herbaceous layer in new England forests

Abstract

Eastern hemlock (Tsuga canadensis) is a foundation species in eastern North American forests. Because eastern hemlock is a foundation species, it often is assumed that the diversity of associated species is high. However, the herbaceous layer of eastern hemlock stands generally is sparse, species-poor, and lacks unique species or floristic assemblages. The rapidly spreading, nonnative hemlock woolly adelgid (Adelges tusgae) is causing widespread death of eastern hemlock. Loss of individual hemlock trees or whole stands rapidly leads to increases in species richness and cover of shrubs, herbs, graminoids, ferns, and fern-allies. Naively, one could conclude that the loss of eastern hemlock has a net positive effect on biodiversity. What is lost besides hemlock, however, is landscape-scale variability in the structure and composition of the herbaceous layer. In the Harvard Forest Hemlock Removal Experiment, removal of hemlock by either girdling (simulating adelgid infestation) or logging led to a proliferation of early-successional and disturbance-dependent understory species. In other declining hemlock stands, nonnative plant species expand and homogenize the flora. While local richness increases in former eastern hemlock stands, between-site and regional species diversity will be further diminished as this iconic foundation species of eastern North America succumbs to hemlock woolly adelgid.