Logging response alters trajectories of reorganization after loss of a foundation tree species.

Forest insect outbreaks cause large changes in ecosystem structure, composition, and function. Humans often respond to insect outbreaks by conducting salvage logging, which can amplify the immediate effects, but it is unclear whether logging will result in lasting differences in forest structure and dynamics when compared with forests affected only by insect outbreaks. We used 15 years of data from an experimental removal of Tsuga canadensis (L.) Carr. (Eastern hemlock), a foundation tree species within eastern North American forests, and contrasted the rate, magnitude, and persistence of response trajectories between girdling (emulating mortality from insect outbreak) and timber harvest treatments. Girdling and logging were equally likely to lead to large changes in forest structure and dynamics, but logging resulted in faster rates of change. Understory light increases and community composition changes were larger and more rapid in the logged plots. Tree seedling and understory vegetation abundance increased more in the girdled plots; this likely occurred because seedlings grew rapidly into the sapling- and tree-size classes after logging and quickly shaded out plants on the forest floor. Downed deadwood pools increased more after logging but standing deadwood pools increased dramatically after girdling. Understory light levels remained elevated for a longer time after girdling. Perhaps because the window of opportunity for understory species to establish was longer in the girdled plots, total species richness increased more in the girdled than logged plots. Despite the potential for greater diversity in the girdled plots, Betula lenta L. (black birch) was the most abundant tree species recruited into the sapling- and tree-size classes in both the girdled and logged plots and is poised to dominate the new forest canopy. The largest difference between the girdling and logging treatments-deadwood structure and quantity-will persist and continue to bolster aboveground carbon storage and structural and habitat diversity in the girdled plots. Human responses to insect outbreaks hasten forest reorganization and remove structural resources that may further alter forest response to ongoing climate stress and future disturbances.

N and P constrain C in ecosystems under climate change: Role of nutrient redistribution, accumulation, and stoichiometry.

We use the Multiple Element Limitation (MEL) model to examine responses of 12 ecosystems to elevated carbon dioxide (CO2 ), warming, and 20% decreases or increases in precipitation. Ecosystems respond synergistically to elevated CO2 , warming, and decreased precipitation combined because higher water-use efficiency with elevated CO2 and higher fertility with warming compensate for responses to drought. Response to elevated CO2 , warming, and increased precipitation combined is additive. We analyze changes in ecosystem carbon (C) based on four nitrogen (N) and four phosphorus (P) attribution factors: (1) changes in total ecosystem N and P, (2) changes in N and P distribution between vegetation and soil, (3) changes in vegetation C:N and C:P ratios, and (4) changes in soil C:N and C:P ratios. In the combined CO2 and climate change simulations, all ecosystems gain C. The contributions of these four attribution factors to changes in ecosystem C storage varies among ecosystems because of differences in the initial distributions of N and P between vegetation and soil and the openness of the ecosystem N and P cycles. The net transfer of N and P from soil to vegetation dominates the C response of forests. For tundra and grasslands, the C gain is also associated with increased soil C:N and C:P. In ecosystems with symbiotic N fixation, C gains resulted from N accumulation. Because of differences in N versus P cycle openness and the distribution of organic matter between vegetation and soil, changes in the N and P attribution factors do not always parallel one another. Differences among ecosystems in C-nutrient interactions and the amount of woody biomass interact to shape ecosystem C sequestration under simulated global change. We suggest that future studies quantify the openness of the N and P cycles and changes in the distribution of C, N, and P among ecosystem components, which currently limit understanding of nutrient effects on C sequestration and responses to elevated CO2 and climate change.

Broadening the ecological mindset.

Over the past three decades, the Harvard Forest Summer Research Program in Ecology (HF-SRPE) has been at the forefront of expanding the ecological tent for minoritized or otherwise marginalized students. By broadening the definition of ecology to include fields such as data science, software engineering, and remote sensing, we attract a broader range of students, including those who may not prioritize field experiences or who may feel unsafe working in rural or urban field sites. We also work towards a more resilient society in which minoritized or marginalized students can work safely, in part by building teams of students and mentors. Teams collaborate on projects that require a diversity of approaches and create opportunities for students and mentors alike to support one another and share leadership. Finally, HF-SRPE promotes an expanded view of what it means to become an ecologist. We value and support diverse career paths for ecologists to work in all parts of society, to diversify the face of ecology, and to bring different perspectives together to ensure innovations in environmental problem solving for our planet.

Defoliation severity is positively related to soil solution nitrogen availability and negatively related to soil nitrogen concentrations following a multi-year invasive insect irruption.

Understanding connections between ecosystem nitrogen (N) cycling and invasive insect defoliation could facilitate the prediction of disturbance impacts across a range of spatial scales. In this study we investigated relationships between ecosystem N cycling and tree defoliation during a recent 2015-18 irruption of invasive gypsy moth caterpillars (Lymantria dispar), which can cause tree stress and sometimes mortality following multiple years of defoliation. Nitrogen is a critical nutrient that limits the growth of caterpillars and plants in temperate forests. In this study, we assessed the associations among N concentrations, soil solution N availability and defoliation intensity by L. dispar at the scale of individual trees and forest plots. We measured leaf and soil N concentrations and soil solution inorganic N availability among individual red oak trees (Quercus rubra) in Amherst, MA and across a network of forest plots in Central Massachusetts. We combined these field data with estimated defoliation severity derived from Landsat imagery to assess relationships between plot-scale defoliation and ecosystem N cycling. We found that trees in soil with lower N concentrations experienced more herbivory than trees in soil with higher N concentrations. Additionally, forest plots with lower N soil were correlated with more severe L. dispar defoliation, which matched the tree-level relationship. The amount of inorganic N in soil solution was strongly positively correlated with defoliation intensity and the number of sequential years of defoliation. These results suggested that higher ecosystem N pools might promote the resistance of oak trees to L. dispar defoliation and that defoliation severity across multiple years is associated with a linear increase in soil solution inorganic N.

Long-term structural and biomass dynamics of virgin Tsuga canadensis-Pinus strobus forests after hurricane disturbance.

The development of old-growth forests in northeastern North America has largely been within the context of gap-scale disturbances given the rarity of stand-replacing disturbances. Using the 10-ha old-growth Harvard Tract and its associated 90-year history of measurements, including detailed surveys in 1989 and 2009, we document the long-term structural and biomass development of an old-growth Tsuga canadensis-Pinus strobus forest in southern New Hampshire, USA following a stand-replacing hurricane in 1938. Measurements of aboveground biomass pools were integrated with data from second- and old-growth T. canadensis forests to evaluate long-term patterns in biomass development following this disturbance. Ecosystem structure across the Tract prior to the hurricane exhibited a high degree of spatial heterogeneity with the greatest levels of live tree basal area (70-129 m2 /ha) on upper west-facing slopes where P. strobus was dominant and intermixed with T. canadensis. Live-tree biomass estimates for these stratified mixtures ranged from 159 to 503 Mg/ha at the localized, plot scale (100 m2 ) and averaged 367 Mg/ha across these portions of the landscape approaching the upper bounds for eastern forests. Live-tree biomass 71 years after the hurricane is more uniform and lower in magnitude, with T. canadensis currently the dominant overstory tree species throughout much of the landscape. Despite only one living P. strobus stem in the 2009 plots (and fewer than five stems known across the entire 10-ha area), the detrital legacy of this species is pronounced with localized accumulations of coarse woody debris exceeding 237.7-404.2 m3 /ha where this species once dominated the canopy. These patterns underscore the great sizes P. strobus attained in pre-European landscapes and its great decay resistance relative to its forest associates. Total aboveground biomass pools in this 71-year-old forest (255 Mg/ha) are comparable to those in modern old-growth ecosystems in the region that also lack abundant white pine. Results highlight the importance of disturbance legacies in affecting forest structural conditions over extended periods following stand-replacing events and underscore that post-disturbance salvage logging can alter ecosystem development for decades. Moreover, the dominant role of old-growth P. strobus in live and detrital biomass pools before and after the hurricane, respectively, demonstrate the disproportionate influence this species likely had on carbon storage at localized scales prior to the widespread, selective harvesting of large P. strobus across the region in the 18th and 19th centuries.

Foundation species loss affects vegetation structure more than ecosystem function in a northeastern USA forest.

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.