Survivors, Not Invaders, Control Forest Development Following Simulated Hurricane

Wind disturbance profoundly shapes temperate forests but few studies have evaluated patterns and mechanisms of long-term forest dynamics following major windthrows. In 1990, we initiated a large hurricane simulation experiment in a 0.8-ha manipulation (pulldown) and 0.6-ha control area of a maturing Quercus rubra–Acer rubrum forest in New England. We toppled 276 trees in the pulldown, using a winch and cable, in the northwesterly direction of natural treefall from major hurricanes. Eighty percent of canopy trees and two-thirds of all trees ≥5 cm dbh (diameter at breast height) suffered direct and indirect damage. We used 20 years of measurements to evaluate the trajectory and mechanisms of forest response after intense disturbance. Based on the patch size and disturbance magnitude, we expected pioneer tree and understory species to drive succession. The first decade of analyses emphasized tree seedling establishment and sprouting by damaged trees as the dominant mechanisms of forest recovery in this extensive damaged area. However, despite 80% canopy damage and 8000-m2 patch size, surviving overstory and advance regeneration controlled longer-term forest development. Residual oaks make up 42% of stand basal area after 20 years. The new cohort of trees, dominated by black birch advance regeneration, contributes 30% of stand basal area. There were shifts in understory vegetation composition and cover, but few species were gained or lost after 20 years. Stand productivity rebounded quickly (litterfall recovered to pre-disturbance levels in six years), but we predict that basal area in the pulldown will lag behind the control (which gained 6 m2/ha over 20 years) for decades to come. This controlled experiment showed that although the scale and intensity of damage were great, abundant advance regeneration, understory vegetation, and damaged trees remained, allowing the forest to resist changes in ecosystem processes and invasion by new species.Organismic and Evolutionary Biolog

The Relative Contributions of Seed Bank, Seed Rain, and Understory Vegetation Dynamics to the Reorganization of Tsuga Canadensis Forests After Loss due to Logging or Simulated Attack by Adelges tsugae

Profound changes are occurring in forests as native insects, nonnative insects, or pathogens irrupt on foundation tree species; comprehensive models of vegetation responses are needed to predict future forest composition. We experimentally simulated hemlock woolly adelgid (Adelges tsugae Annand) infestation (by girdling trees) and preemptive logging of eastern hemlock (Tsuga canadensis (L.) Carrière) and compared vegetation dynamics in replicate 90 m × 90 m treatment plots and intact hemlock stands from 2004 to 2010. Using Chao–Sørensen abundance-based similarity indices, we assessed compositional similarities of trees, shrubs, forbs, and graminoids among the seed bank, seed rain, and standing vegetation over time and among treatments. Post-treatment seed rain, similar among treatments, closely reflected canopy tree composition. Species richness of the seed bank was similar in 2004 and 2010. Standing vegetation in the hemlock controls remained dissimilar from the seed bank, reflecting suppressed germination. Recruits from the seed rain and seed bank dominated standing vegetation in the logged treatment, whereas regeneration of vegetation from the seed bank and seed rain was slowed due to shading by dying hemlocks in the girdled treatment. Our approach uniquely integrates multiple regeneration components through time and provides a method for predicting forest dynamics following loss of foundation tree species.Organismic and Evolutionary BiologyOther Research Uni

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

Experimentally Testing the Role of Foundation Species in Forests: The Harvard Forest Hemlock Removal Experiment

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.Organismic and Evolutionary BiologyOther Research Uni

Comparing Tree‐Ring and Permanent Plot Estimates of Aboveground Net Primary Production in Three Eastern U.S. Forests

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

Comparing Tree-Ring And Permanent Plot Estimates Of Aboveground Net Primary Production In Three Eastern U.S. Forests

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

Building a foundation: land-use history and dendrochronology reveal temporal dynamics of a Tsuga canadensis (Pinaceae) forest

Organismic and Evolutionary Biolog

Regional Invasive Species & Climate Change Management Challenge: Forest Pest Risk is Heating Up

Insect pests and pathogens, and climate change, each threaten forest health. But what happens when the two are combined? Climate change brings pests to new areas, makes pests more damaging, reduces trees’ defenses to pests, and can alter how forests recover after pest disturbance. Strategies for managing the combined impacts of forest pests and climate change include preventing new pest introductions, resisting pest spread by treating individual trees and diversifying forest stands, promoting more resilient forests that can rebound from pests, and helping forests transition to a state better adapted to our future climate

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

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.Organismic and Evolutionary Biolog