Regeneration dynamics in remnant Tsuga canadensis stands in the northern Lake States: Potential direct and indirect effects of herbivory

We examined the effects of white-tailed deer (Odocoileus virginianus) herbivory and microsite limitation on Tsuga canadensis regeneration in 39 randomly selected remnant T. canadensis stands in Michigan's Upper Peninsula. Deer of the region migrate to and congregate in T. canadensis stands in winter resulting in strong seasonal habitat use patterns. In each study stand, we quantified vegetation, microsite availability, and deer use (via pellet counts). While some stands contained high densities of T. canadensis regeneration (stems < 4.0 cm dbh), we found complete T. canadensis regeneration failures in 6 out of 39 stands. Additionally, 17 and 22 stands respectively, had complete failures in the small and large sapling categories. General linear models suggested that deer use was the primary limiting factor in the small sapling size class, even at relatively low levels of deer use. T. canadensis seedling density was positively associated with the availability of high-decay coarse woody debris and negatively associated with basal area of Acer saccharum in the overstory. This latter association may be due, at least in part, to negative effects of broadleaf litter on T. canadensis establishment and a general trend toward increasing Acer abundance in the regeneration layer. Our results suggest that differential tolerance to browsing (Tsuga vs. Acer) in conjunction with reduced germination substrate availability may set up a scenario where successful T. canadensis establishments is more limited by legacy and indirect than direct effects given contemporary levels of deer use.