Tree Regeneration Structure Following Beech Bark Disease-Motivated Harvests: Factors Associated with Patterns and Management Implications

Abstract

In the northern Appalachian region of North America, mortality of mature American beech (Fagus grandifolia Ehrh.) via the introduced beech bark disease (BBD) can result in dense thickets of beech saplings that inhibit the regeneration of other species. It is unknown if similar structures characterize more recently infested managed forests in the Great Lakes region. If these dense beech sapling layers do exist, management would be aided by knowing which site/regional factors they are associated with and by identifying particular sapling structures that may threaten the sustainability of these forests under current management paradigms. To examine these patterns, we used a natural experiment with sample plots in 69 unevenly aged, selection silviculture-managed, maple (Acer spp.)-dominated northern hardwood stands. Our stands were dispersed across northern Michigan, USA and had undergone BBD-motivated partial harvests favoring beech removal (mean = 5.5 years before measurement). In each stand, we quantified tree regeneration structure in relation to winter deer use (fecal pellet count density), site quality (habitat type), geographic region (Eastern Upper Peninsula and Northern Lower Peninsula), and multiple measures of overstory stand density. We also examined the density effects of taller regeneration strata on subordinate strata. Across sites, the small sapling recruit class (i.e., >137 cm tall and <5 cm diameter at 137 cm tall) was dominated by beech and was often dense (44% of subplots > 2000 stems ha−1 and 16% of subplots > 5000 ha−1) but never exceeded the > 10,000 stems ha−1 reported in the northern Appalachian region. Beech sapling density was higher in the Northern Lower Peninsula, on lower quality sites, at lower postharvest overstory densities, and on sites with higher densities of preharvest overstory beech. In contrast to the beech-dominated small sapling recruit class, seedlings (i.e., <25 cm tall) were generally more species diverse than sapling strata and were dominated by maple species. Although generally dense, seedling density was negatively related to small sapling recruit density, suggesting that saplings may suppress the seedling stratum. The general pattern for the small sapling recruit layer of browsing-insensitive beech (and ironwood, Ostrya virginiana Mill. K. Koch) dominance and low representation of browsing-sensitive species (e.g., Acer spp.) circumstantially supports the notion that regeneration structure is heavily influenced by deer. However, current deer use was generally low in our stands, and relationships with tree regeneration structure were weak. Instead, regeneration structure is likely shaped by a combination of factors operating at long time scales (i.e., legacies of deer browsing pressure, selection silviculture (given beech and ironwood are shade tolerant), overstory composition, and site quality) and by those effects that are more proximal, such as postharvest overstory density. Minimum stocking criteria for species considered desirable for management (e.g., sugar maple and Acer saccharum Marshall) suggest many stands are inadequately stocked in the sapling recruit classes. Although future regeneration dynamics are unclear, current patterns suggest that many stands with high beech/ironwood small sapling recruit densities may require management intervention to overcome insufficient recruitment of species targeted for management.