Person:

Orwig, David

Loss of foundation tree species rapidly alters ecological processes in forested ecosystems. Tsuga canadensis, an hypothesized foundation species of eastern North American forests, is declining throughout much of its range due to infestation by the nonnative insect Adelges tsugae and by removal through pre-emptive salvage logging. In replicate 0.81-ha plots, T. canadensis was cut and removed, or killed in place by girdling to simulate adelgid damage. Control plots included undisturbed hemlock and mid-successional hardwood stands that represent expected forest composition in 50–100 years. Vegetation richness, understory vegetation cover, soil carbon flux, and nitrogen cycling were measured for two years prior to, and five years following, application of experimental treatments. Litterfall and coarse woody debris (CWD), including snags, stumps, and fallen logs and branches, have been measured since treatments were applied. Overstory basal area was reduced 60%–70% in girdled and logged plots. Mean cover and richness did not change in hardwood or hemlock control plots but increased rapidly in girdled and logged plots. Following logging, litterfall immediately decreased then slowly increased, whereas in girdled plots, there was a short pulse of hemlock litterfall as trees died. CWD volume remained relatively constant throughout but was 3–4× higher in logged plots. Logging and girdling resulted in small, short-term changes in ecosystem dynamics due to rapid regrowth of vegetation but in general, interannual variability exceeded differences among treatments. Soil carbon flux in girdled plots showed the strongest response: 35% lower than controls after three years and slowly increasing thereafter. Ammonium availability increased immediately after logging and two years after girdling, due to increased light and soil temperatures and nutrient pulses from leaf-fall and reduced uptake following tree death. The results from this study illuminate ecological processes underlying patterns observed consistently in region-wide studies of adelgid-infested hemlock stands. Mechanisms of T. canadensis loss determine rates, magnitudes, and trajectories of ecological changes in hemlock forests. Logging causes abrupt, large changes in vegetation structure whereas girdling (and by inference, A. tsugae) causes sustained, smaller changes. Ecosystem processes depend more on vegetation cover per se than on species composition. We conclude that the loss of this late-successional foundation species will have long-lasting impacts on forest structure but subtle impacts on ecosystem function.

1. Problem statement– Foundation species define and structure ecological systems. In forests around the world, foundation tree species are declining due to overexploitation, pests and pathogens. Eastern hemlock (Tsuga canadensis), a foundation tree species in eastern North America, is threatened by an exotic insect, the hemlock woolly adelgid (Adelges tsugae). The loss of hemlock is hypothesized to result in dramatic changes in assemblages of associated species with cascading impacts on food webs and fluxes of energy and nutrients. We describe the setting, design and analytical framework of the Harvard Forest Hemlock Removal Experiment (HF-HeRE), a multi-hectare, long-term experiment that overcomes many of the major logistical and analytical challenges of studying system-wide consequences of foundation species loss. 2. Study design– HF-HeRE is a replicated and blocked Before-After-Control-Impact experiment that includes two hemlock removal treatments: girdling all hemlocks to simulate death by adelgid and logging all hemlocks >20 cm diameter and other merchantable trees to simulate pre-emptive salvage operations. These treatments are paired with two control treatments: hemlock controls that are beginning to be infested in 2010 by the adelgid and hardwood controls that represent future conditions of most hemlock stands in eastern North America. 3. Ongoing measurements and monitoring– Ongoing long-term measurements to quantify the magnitude and direction of forest ecosystem change as hemlock declines include: air and soil temperature, light availability, leaf area and canopy closure; changes in species composition and abundance of the soil seed-bank, understorey vegetation, and soil-dwelling invertebrates; dynamics of coarse woody debris; soil nitrogen availability and net nitrogen mineralization; and soil carbon flux. Short-term or one-time-only measurements include initial tree ages, hemlock-decomposing fungi, wood-boring beetles and throughfall chemistry. Additional within-plot, replicated experiments include effects of ants and litter-dwelling microarthoropods on ecosystem functioning, and responses of salamanders to canopy change. 4. Future directions and collaborations– HF-HeRE is part of an evolving network of retrospective studies, natural experiments, large manipulations and modelling efforts focused on identifying and understanding the role of single foundation species on ecological processes and dynamics. We invite colleagues from around the world who are interested in exploring complementary question.

We compared the understory communities (herbs, shrubs, and tree seedlings and saplings) of old-growth and second-growth eastern hemlock forests (Tsuga canadensis) in western Massachusetts, USA. Second-growth hemlock forests originated following clear-cut logging in the late 1800s and were 108–136 years old at the time of sampling. Old-growth hemlock forests contained total ground cover of herbaceous and shrub species that was approximately 4 times greater than in second-growth forests (4.02 ± 0.41%/m^2 versus 1.06 ± 0.47%/m^2) and supported greater overall species richness and diversity. In addition, seedling and sapling densities were greater in old-growth stands compared to second-growth stands and the composition of these layers was positively correlated with overstory species composition (Mantel tests, r > 0.26, P < 0.05) highlighting the strong positive neighborhood effects in these systems. Ordination of study site understory species composition identified a strong gradient in community composition from second-growth to old-growth stands. Vector overlays of environmental and forest structural variables indicated that these gradients were related to differences in overstory tree density, nitrogen availability, and coarse woody debris characteristics among hemlock stands. These relationships suggest that differences in resource availability (e.g., light, moisture, and nutrients) and microhabitat heterogeneity between old-growth and second-growth stands were likely driving these compositional patterns. Interestingly, several common forest understory plants, including Aralia nudicaulis, Dryopteris intermedia, and Viburnum alnifolium, were significant indicator species for old-growth hemlock stands, highlighting the lasting legacy of past land use on the reestablishment and growth of these common species within second-growth areas. The return of old-growth understory conditions to these second-growth areas will largely be dependent on disturbance and self-thinning mediated changes in overstory structure, resource availability, and microhabitat heterogeneity.

Old-growth forests are valuable sources of ecological, conservation, and management information, yet these ecosystems have received little study in New England, due in large part to their regional scarcity. To increase our understanding of the structures and processes common in these rare forests, we studied the abundance of downed coarse woody debris (CWD) and snags and live-tree size-class distributions in 16 old-growth hemlock forests in western Massachusetts. Old-growth stands were compared with eight adjacent secondgrowth hemlock forests to gain a better understanding of the structural differences between these two classes of forests resulting from contrasting histories. In addition, we used standlevel dendroecological reconstructions to investigate the linkages between disturbance history and old-growth forest structure using an information–theoretic model selection framework. Old-growth stands exhibit a much higher degree of structural complexity than secondgrowth forests. In particular, old-growth stands had larger overstory trees and greater volumes of downed coarse woody debris (135.2 vs. 33.2 m3 /ha) and snags (21.2 vs. 10.7 m3 /ha). Secondgrowth stands were characterized by either skewed unimodal or reverse-J shaped diameter distributions, while old-growth forests contained bell-shaped, skewed unimodal, rotated sigmoid, and reverse J-shaped distributions. The variation in structural attributes among oldgrowth stands, particularly the abundance of downed CWD, was closely related to disturbance history. In particular, old-growth stands experiencing moderate levels of canopy disturbance during the last century (1930s and 1980s) had greater accumulations of CWD, highlighting the importance of gap-scale disturbances in shaping the long-term development and structural characteristics of old-growth forests. These findings are important for the development of natural disturbance-based silvicultural systems that may be used to restore important forest characteristics lacking in New England second-growth stands by integrating structural legacies of disturbance (e.g., downed CWD) and resultant tree-size distribution patterns. This silvicultural approach would emulate the often episodic nature of CWD recruitment within old-growth forests.