Understory Vegetation in Old-Growth and Second-Growth Tsuga Canadensis Forests in Western Massachusetts

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.Organismic and Evolutionary BiologyOther Research Uni

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

Effects of Hemlock Woolly Adelgid and Elongate Hemlock Scale on Eastern Hemlock Growth and Foliar Chemistry

In the eastern United States, two invasive specialist insects share a native host plant, Eastern hemlock, Tsuga canadensis. In recent years, much research has focused on the impacts of the hemlock woolly adelgid (Adelges tsugae) because of the detrimental effects it has on hemlock growth and survival. In contrast, the invasive elongate hemlock scale (Fiorinia externa) is thought to have only minor impacts on hemlock. We infested hemlock saplings with each insect and compared them with control (i.e., neither insect herbivore) saplings to assess how early infestations impact Eastern hemlock health (measured using new branch growth, foliar %N, and C:N ratio). Our study showed that, at equal densities, the two insects differed in their effect on Eastern hemlock. F. externa did not impact plant growth or foliar chemistry over the course of the 2-yr experiment. A. tsugae significantly reduced plant growth and caused a reduction of %N in the first year of the experiment. By the end of the experiment, A. tsugae trees had the same %N in their foliage as control and F. externa trees but drastically reduced growth patterns. The most likely explanation for this result is the greater growth in control and F. externa saplings during the second year resulted in the dilution of available foliar N over a larger amount of newly produced plant tissue. For early infestations of both insects, our study suggests that management plans should focus on the more detrimental A. tsugae

The Past, Present, and Future of the Hemlock Woolly Adelgid (\u3cem\u3eAdelges tsugae\u3c/em\u3e) and Its Ecological Interactions with Eastern Hemlock (\u3cem\u3eTsuga canadensis\u3c/em\u3e) Forests

The nonnative hemlock woolly adelgid is steadily killing eastern hemlock trees in many parts of eastern North America. We summarize impacts of the adelgid on these forest foundation species; review previous models and analyses of adelgid spread dynamics; and examine how previous forecasts of adelgid spread and ecosystem dynamics compare with current conditions. The adelgid has reset successional sequences, homogenized biological diversity at landscape scales, altered hydrological dynamics, and changed forest stands from carbon sinks into carbon sources. A new model better predicts spread of the adelgid in the south and west of the range of hemlock, but still under-predicts its spread in the north and east. Whether these underpredictions result from inadequately modeling accelerating climate change or accounting for people inadvertently moving the adelgid into new locales needs further study. Ecosystem models of adelgid-driven hemlock dynamics have consistently forecast that forest carbon stocks will be little affected by the shift from hemlock to early-successional mixed hardwood stands, but these forecasts have assumed that the intermediate stages will remain carbon sinks. New forecasting models of adelgid-driven hemlock decline should account for observed abrupt changes in carbon flux and ongoing and accelerating human-driven land-use and climatic changes

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

Organismic and Evolutionary Biolog

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

Asymmetric Biotic Interactions and Abiotic Niche Differences Revealed by a Dynamic Joint Species Distribution Model

A species’ distribution and abundance are determined by abiotic conditions and biotic interactions with other species in the community. Most species distribution models correlate the occurrence of a single species with environmental variables only, and leave out biotic interactions. To test the importance of biotic interactions on occurrence and abundance, we compared a multivariate spatiotemporal model of the joint abundance of two invasive insects that share a host plant, hemlock woolly adelgid (HWA; Adelges tsugae) and elongate hemlock scale (EHS; Fiorina externa), to independent models that do not account for dependence among co‐occurring species. The joint model revealed that HWA responded more strongly to abiotic conditions than EHS. Additionally, HWA appeared to predispose stands to subsequent increase of EHS, but HWA abundance was not strongly dependent on EHS abundance. This study demonstrates how incorporating spatial and temporal dependence into a species distribution model can reveal the dependence of a species’ abundance on other species in the community. Accounting for dependence among co‐occurring species with a joint distribution model can also improve estimation of the abiotic niche for species affected by interspecific interactions

Species Diversity Associated with Foundation Species in Temperate and Tropical Forests

Foundation species define and structure ecological communities but are difficult to identify before they are declining. Yet, their defining role in ecosystems suggests they should be a high priority for protection and management while they are still common and abundant. We used comparative analyses of six large forest dynamics plots spanning a temperate-to-tropical gradient in the Western Hemisphere to identify statistical “fingerprints” of potential foundation species based on their size-frequency and abundance-diameter distributions, and their spatial association with five measures of diversity of associated woody plant species. Potential foundation species are outliers from the common “reverse-J” size-frequency distribution, and have negative effects on alpha diversity and positive effects on beta diversity at most spatial lags and directions. Potential foundation species also are more likely in temperate forests, but foundational species groups may occur in tropical forests. As foundation species (or species groups) decline, associated landscape-scale (beta) diversity is likely to decline along with them. Preservation of this component of biodiversity may be the most important consequence of protecting foundation species while they are still common

Peak radial growth of diffuse-porous species occurs during periods of lower water availability than for ring-porous and coniferous trees

Climate models project warmer summer temperatures will increase the frequency and heat severity of droughts in temperate forests of Eastern North America. Hotter droughts are increasingly documented to affect tree growth and forest dynamics, with critical impacts on tree mortality, carbon sequestration and timber provision. The growing acknowledgement of the dominant role of drought timing on tree vulnerability to water deficit raises the issue of our limited understanding of radial growth phenology for most temperate tree species. Here, we use well-replicated dendrometer band data sampled frequently during the growing season to assess the growth phenology of 610 trees from 15 temperate species over 6 years. Patterns of diameter growth follow a typical logistic shape, with growth rates reaching a maximum in June, and then decreasing until process termination. On average, we find that diffuse-porous species take 16-18 days less than other wood-structure types to put on 50% of their annual diameter growth. However, their peak growth rate occurs almost a full month later than ring-porous and conifer species (ca. 24 +/- 4 days; mean +/- 95% credible interval). Unlike other species, the growth phenology of diffuse-porous species in our dataset is highly correlated with their spring foliar phenology. We also find that the later window of growth in diffuse-porous species, coinciding with peak evapotranspiration and lower water availability, exposes them to a higher water deficit of 88 +/- 19 mm (mean +/- SE) during their peak growth than ring-porous and coniferous species (15 +/- 35 mm and 30 +/- 30 mm, respectively). Given the high climatic sensitivity of wood formation, our findings highlight the importance of wood porosity as one predictor of species climatic sensitivity to the projected intensification of the drought regime in the coming decades.Peer reviewe