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.

How, where, and why carbon (C) moves into and out of an ecosystem through time are long-standing questions in biogeochemistry. Here, we bring together hundreds of thousands of C-cycle observations at the Harvard Forest in central Massachusetts, USA, a mid-latitude landscape dominated by 80-120-yr-old closed-canopy forests. These data answered four questions: (1) where and how much C is presently stored in dominant forest types; (2) what are current rates of C accrual and loss; (3) what biotic and abiotic factors contribute to variability in these rates; and (4) how has climate change affected the forest's C cycle? Harvard Forest is an active C sink resulting from forest regrowth following land abandonment. Soil and tree biomass comprise nearly equal portions of existing C stocks. Net primary production (NPP) averaged 680-750 g CÁm À2 Áyr À1 ; belowground NPP contributed 38-47% of the total, but with large uncertainty. Mineral soil C measured in the same inventory plots in 1992 and 2013 was too heterogeneous to detect change in soil-C pools; however, radiocarbon data suggest a small but persistent sink of 10-30 g CÁm À2 Áyr À1. Net ecosystem production (NEP) in hardwood stands averaged~300 g CÁm À2 Áyr À1. NEP in hemlock-dominated forests averaged 450 g CÁm À2 Áyr À1 until infestation by the hemlock woolly adelgid turned these stands into a net C source. Since 2000, NPP has increased by 26%. For the period 1992-2015, NEP increased 93%. The increase in mean annual temperature and growing season length alone accounted for~30% of the increase in productivity. Interannual variations in GPP and NEP were also correlated with increases in red oak biomass, forest leaf area, and canopy-scale lightuse efficiency. Compared to long-term global change experiments at the Harvard Forest, the C sink in regrowing biomass equaled or exceeded C cycle modifications imposed by soil warming, N saturation, and hemlock removal. Results of this synthesis and comparison to simulation models suggest that forests across the region are likely to accrue C for decades to come but may be disrupted if the frequency or severity of biotic and abiotic disturbances increases.

Abstract.Forests account for a large portion of sequestered carbon, much of which is stored as wood in trees. The rate of carbon accumulation in aboveground plant material, or aboveground net primary productivity (aNPP), quantifies annual to decadal variations in forest carbon sequestration. Permanent plots are often used to estimate aNPP but are usually not annually resolved and take many years to develop a long data set. Tree rings are a unique and infrequently used source for measuring aNPP, and benefit from fine spatial (individual trees) and temporal (annual) resolution. Because of this precision, tree rings are complementary to permanent plots and the suite of tools used to study forest productivity. Here we evaluate whether annual estimates of aNPP developed from tree rings approximate estimates derived from colocated permanent plots. We studied a lowland evergreen (Howland, Maine), mixed deciduous (Harvard Forest, Massachusetts), and mixed mesophytic (Fernow, West Virginia) forest in the eastern United States. Permanent plots at the sites cover an area of 2-3 ha, and we use these areas as benchmarks indicative of the forest stand. We simulate random draws of permanent plot subsets to describe the distribution of aNPP estimates given a sampling area size equivalent to the tree-ring plots. Though mean tree-ring aNPP underestimates permanent plot aNPP slightly at Howland and Fernow and overestimates at Harvard Forest when compared with the entire permanent plot, it is within the 95% confidence interval of the random draws of equal-sized sampling area at all sites. To investigate whether tree-ring aNPP can be upscaled to the stand, we conducted a second random draw of permanent plot subsets simulating a twofold increase in sampling area. aNPP estimates from this distribution were not significantly different from results of the initial sampling area, though variance decreased as sampling area approaches stand area. Despite several concerns to consider when using tree rings to reconstruct aNPP (e.g., upscaling, allometric, and sampling uncertainties), the benefits are apparent, and we call for the continued application of tree rings in carbon cycle studies across a broader range of species diversity, productivity, and disturbance histories to fully develop this potential.