David A. Orwig

Loss of foundation species: consequences for the structure and dynamics of forested ecosystems

In many forested ecosystems, the architecture and functional ecology of certain tree species define forest structure and their species-specific traits control ecosystem dynamics. Such foundation tree species are declining throughout the world due to introductions and outbreaks of pests and pathogens, selective removal of individual taxa, and over-harvesting. Through a series of case studies, we show that the loss of foundation tree species changes the local environment on which a variety of other species depend; how this disrupts fundamental ecosystem processes, including rates of decomposition, nutrient fluxes, carbon sequestration, and energy flow; and dramatically alters the dynamics of associated aquatic ecosystems. Forests in which dynamics are controlled by one or a few foundation species appear to be dominated by a small number of strong interactions and may be highly susceptible to alternating between stable states following even small perturbations. The ongoing decline of many foundation species provides a set of important, albeit unfortunate, opportunities to develop the research tools, models, and metrics needed to identify foundation species, anticipate the cascade of immediate, short- and long-term changes in ecosystem structure and function that will follow from their loss, and provide options for remedial conservation and management.

Forest response to the introduced hemlock woolly adelgid in southern New England, USA.

ORWIG, D. A. AND D. R. FOSTER (Harvard University, Harvard Forest, Petersham, Massachusetts 01366). Forest response to the introduced hemlock woolly adelgid in southern New England, USA. J. Torrey Bot. Soc. 125:60-73. 1998.-Hemlock woolly adelgid (HWA), Adelges tsugae, an introduced aphid-like insect from Asia, is expanding its range across the northeastern United States through the range of Tsuga canadensis (eastern hemlock) and can severely reduce or eliminate this important late-successional species. As part of a study investigating standand landscape-level forest dynamics resulting from HWA infestation, we examined initial community response of eight T. canadensis stands in south-central Connecticut. Our major objectives were to assess mortality patterns in T. canadensis, evaluate subsequent changes in stand microenvironment, and relate these and stand composition to initial patterns of regeneration, understory response, and community reorganization. Tsuga canadensis damage varied broadly across the study area ranging from near zero to greater than 95% mortality. All size and age classes sampled were attacked by HWA, although smaller trees exhibited higher mortality rates than larger trees. All remaining T. canadensis sampled in seven of the eight stands were infested with HWA and over 90% suffered at least 50% foliar loss. Substantial accumulations of downed woody debris have developed in stands with severe HWA damage. Canopy gaps created by HWA damage significantly increased the amount of light reaching the forest floor and resulted in rapid understory vegetation responses. Prolific Betula lenta (black birch) establishment occurred in stands with moderate to severe T. canadensis mortality. In addition, opportunistic herbaceous species (Erechtites hieracifolia, Phytolacca americana) and exotic species (Ailanthus altissima, Microstegium vimineum) have recently invaded these stands. Due to mortality from HWA, T. canadensis seedlings were scarce in sampled stands, suggesting that advance regeneration and seedbanks will not be important mechanisms for T. canadensis reestablishment. Tsuga canadensis cannot sprout following defoliation and has no apparent resistance to HWA. Therefore, dramatic reductions in T. canadensis across broad geographical areas appear imminent if HWA dispersal continues unimpeded and no effective natural enemies of HWA are found.

Variation in radial growth responses to drought among species, site, and canopy strata

Abstract Radial growth responses to drought were examined in the tree-ring records of six species growing within two locations of differing land-use history and soil moisture characteristics, and in overstory and understory canopy positions in northern Virginia. Tree species experienced differential ring-width reductions during or immediately following four severe drought periods occurring from 1930 to 1965 and were influenced by climatic variables including annual and summer temperatures, annual precipitation, and annual Palmer Drought Severity Index. Relative growth comparisons averaged across species before and after drought years indicated that understory trees on dry-mesic sites grew 11% faster after drought compared to pre-drought rates while mesic site trees in both canopy positions grew approximately 4% slower. Superposed epoch analysis indicated that Liriodendron tulipifera growing on mesic sites experienced greater ring-width reductions associated with drought than co-occurring, more drought-tolerant Quercus alba and Q. velutina. On dry-mesic sites, L. tulipifera also experienced greatly reduced growth as a result of drought but exhibited significant growth increases following individual drought events. Quercus alba was the only species that exhibited a consistent, significant ring-width decrease associated with all droughts on dry-mesic sites. In contrast, Pinus virginiana was least impacted by drought on dry-mesic sites but was much more impacted by drought on mesic sites, indicating a drought×site interaction for this species. Overstory Carya glabra and Q. alba experienced larger growth decreases during drought on dry-mesic versus mesic sites. Understory tree growth reductions did not differ between site types but were often significantly larger than overstory responses of the same species on mesic sites. Following drought, most trees exhibited growth reductions lasting 2–3 years, although several species experienced reductions lasting up to 6 years. The results of this study suggest that tree rings represent an important long-term proxy for leaf-level ecophysiological measurements of growth responses to drought periods.

Dendroecological Analysis of Successional Dynamics for a Presettlement- Origin White-Pine-Mixed-Oak Forest in the Southern Appalachians, USA

1 Prior to European settlement, Quercus alba, Castanea dentata, Carya spp. and Pinus strobus dominated Ridge and Valley forests in southern West Virginia. Dendroecological techniques were used to study the disturbance history and successional dynamics of one of the few remaining valley-floor remnants of this primary forest type. 2 The uneven-aged forest is presently dominated by P. strobus, Acer rubrum, Q. alba, Q. rubra and Q. velutina. Quercus alba and P. strobus represented most of the oldest and largest trees in the forest, while Acer, Fagus and Tsuga were the youngest and smallest. Maximum age was 295 years for Q. alba and 231 years for P. strobus. 3 There was continuous recruitment of Q. alba trees from 1700 to 1900. The abundance of P. strobus increased between 1830 and 1900, with a peak in the 1870s associated with releases in radial growth. The subsequent peak recruitment of Q. rubra and Q. velutina from 1880 to 1900 is suggestive of possible facilitation of these species by P. strobus. Following the cessation of Pinus and Quercus recruitment in 1900, Acer, Fagus and Tsuga abundance increased, particularly A. rubrum. 4 Radial growth chronologies across all species and age classes exhibited a series of major or moderate releases at regular intervals (typically every 20-30 years). However, the asynchronous nature of these releases suggests that they were caused by a series of small-scale disturbances, such as fire and wind-throw, each of which had localized impacts within the stand. 5 We believe that Pinus and Quercus were maintained in this stand during the 18th and 19th centuries by periodic disturbance, in particular fire, which would have eliminated later successional species. The lack of Pinus and Quercus recruitment after 1900 and the subsequent increase in Acer, Fagus and Tsuga indicates the transitional nature of this forest in the absence of fire. 6 Coupling of tree-ring chronologies and species establishment dates greatly increased our understanding of the disturbance history and dynamics of this old-growth forest, and we believe this represents an important approach to the study of species lifehistory attributes and ecological history in general.

Hemlock woolly adelgid in New England forests : canopy impacts transforming ecosystem processes and landscapes

Exotic insect pests may strongly disrupt forest ecosystems and trigger major shifts in nutrient cycling, structure, and composition. We examined the relationship between these diverse effects for the hemlock woolly adelgid (HWA, Adelges tsugae Annand) in New England forests by studying its impacts on local canopy processes in stands differing in infestation levels and linking these impacts to shifts in canopy nutrient cycling and stand and landscape effects. HWA initiated major changes in canopy biomass and distribution. Whereas uninfested trees exhibit a significant decline in canopy biomass from the center to the periphery and a positive correlation between total needle litter and estimated biomass, infested trees have significantly less total canopy biomass, produce less new foliage, shed relatively more needles, and exhibit no correlation between litter and canopy biomass. Foliar N content of infested trees was 20%–40% higher than reference trees, with the strongest increase in young foliage supporting the highest densities of HWA. Foliar %C was unaffected by HWA or foliar age. Epiphytic microorganisms on hemlock needles exhibited little variation in abundance within canopies, but colony-forming units of bacteria, yeast, and filamentous fungi were 2–3 orders of magnitude more abundant on medium and heavily infested than uninfested trees. Throughfall chemistry, quantity, and spatial pattern were strongly altered by HWA. Throughfall exhibits a strong gradient beneath uninfested trees, decreasing in volumes from the canopy periphery to the trunk by more than 45%. The amount of throughfall beneath infested trees exhibits no spatial pattern, reaches 80%–90% of the bulk precipitation, and is characterized by significantly higher concentrations of nitrogen compounds, dissolved organic carbon, and cations. Across the southern New England landscape there is a strong south-to-north gradient of decreasing hemlock tree and sapling mortality and understory compositional change that corresponds to the duration of infestation. Regionally, black birch (Betula lenta L.) is profiting most from hemlock decline by significantly increasing in density and cover. These findings suggest that it is necessary to study the connections between fast/small-scale processes such as changes in nutrient cycling in tree canopies and slow/integrative processes like shifts in biogeochemieal cycling and compositional changes at forest stands and landscapes to better understand the effects of an exotic pest species like HWA on forest ecosystem structure and function.

Multi-year ecosystem response to hemlock woolly adelgid infestation in southern New England forests

The introduced hemlock woolly adelgid (HWA) (Adelges tsugae Annand) has generated widespread tree decline and substantial mortality of eastern hemlock (Tsuga canadensis (L.) Carriere) throughout th...

The ecology of energy and nutrient fluxes in hemlock forests invaded by hemlock woolly adelgid

The hemlock woolly adelgid (HWA, Adelges tsugae Annand) is currently causing a severe decline in vitality and survival of eastern hemlock in North American forests. We analyzed the effects of light HWA infestation on vertical energy and nutrient fluxes from the canopy to the forest floor. Canopy throughfall, litter lysimeters, and laboratory litter microcosms were used to examine the effects of HWA-affected and unaffected throughfall on litter type, leachate, and litter chemistry. Early in the season adelgid infestation caused higher dissolved organic carbon (DOC; +24.6%), dissolved organic nitrogen (DON; +28.5%), and K (+39.3%) fluxes and lower inorganic nitrogen fluxes (-39.8%) in throughfall and in adjacent litter solutions collected beneath infested compared to uninfested trees. Needle litter collected beneath uninfested hemlock had significantly lower N concentrations compared to needles collected beneath infested trees, while no difference in N concentrations was found in birch litter. Bacteria were significantly more abundant on hemlock and birch litter beneath infested trees, while yeasts and filamentous fungi showed no consistent response to HWA throughfall. Litter microcosms showed that less DOC was leaching from birch than from hemlock needles when exposed to HWA throughfall. Overall, NH4-N and DON leachate concentrations were higher from birch than from hemlock litter. Thus, HWA-affected throughfall leads to qualitative and quantitative differences in nitrogen export from the litter layer. The N concentration of hemlock litter did not change with time, but the N concentration in birch litter increased significantly during the course of the experiment, especially when HWA-affected throughfall was applied. We suggest a nonlinear conceptual model for the temporal and vertical transition of energy and nutrient fluxes relative to progressing HWA infestation from a pure hemlock to a birch/maple-dominated forest. Progressive needle loss and changes in needle chemistry are likely to produce a humped-shaped DOC curve, while N fluxes initially decrease as infestation continues but rise eventually with hemlock decline and immigration of hardwood species. These findings suggest that it is necessary to understand the biology and specific physiological/trophic effects of exotic pests on their hosts and associated ecosystem processes in order to decipher the temporal dynamics, direction of change, and functional consequences.

Nonnative forest insects and pathogens in the United States: Impacts and policy options

Abstract We review and synthesize information on invasions of nonnative forest insects and diseases in the United States, including their ecological and economic impacts, pathways of arrival, distribution within the United States, and policy options for reducing future invasions. Nonnative insects have accumulated in United States forests at a rate of ~2.5 per yr over the last 150 yr. Currently the two major pathways of introduction are importation of live plants and wood packing material such as pallets and crates. Introduced insects and diseases occur in forests and cities throughout the United States, and the problem is particularly severe in the Northeast and Upper Midwest. Nonnative forest pests are the only disturbance agent that has effectively eliminated entire tree species or genera from United States forests within decades. The resulting shift in forest structure and species composition alters ecosystem functions such as productivity, nutrient cycling, and wildlife habitat. In urban and suburban areas, loss of trees from streets, yards, and parks affects aesthetics, property values, shading, stormwater runoff, and human health. The economic damage from nonnative pests is not yet fully known, but is likely in the billions of dollars per year, with the majority of this economic burden borne by municipalities and residential property owners. Current policies for preventing introductions are having positive effects but are insufficient to reduce the influx of pests in the face of burgeoning global trade. Options are available to strengthen the defenses against pest arrival and establishment, including measures taken in the exporting country prior to shipment, measures to ensure clean shipments of plants and wood products, inspections at ports of entry, and post‐entry measures such as quarantines, surveillance, and eradication programs. Improved data collection procedures for inspections, greater data accessibility, and better reporting would support better evaluation of policy effectiveness. Lack of additional action places the nation, local municipalities, and property owners at high risk of further damaging and costly invasions. Adopting stronger policies to reduce establishments of new forest insects and diseases would shift the major costs of control to the source and alleviate the economic burden now borne by homeowners and municipalities.

A foundation tree at the precipice: Tsuga canadensis health after the arrival of Adelges tsugae in central New England

Hemlock (Tsuga canadensis) plays a unique role in Eastern forests, producing distinctive biogeochemical, habitat, and microclimatic conditions and yet has begun a potentially irreversible decline due to the invasive hemlock woolly adelgid (Adelges tsugae; HWA) that causes foliar damage, crown loss, and mortality of host trees. Understanding the regional, landscape, site, and stand factors influencing HWA spread and impact is critical for predicting future landscape dynamics and directing effective management. Using aerial photographs, we documented hemlock distribution throughout central Massachusetts and subsampled 123 stands to examine the spatial pattern of HWA and its impact on tree vigor and mortality since its arrival in 1989. In the study region, over 86,000 ha of hemlock forest were mapped in 5,127 stands. White pine (Pinus strobus), red oak (Quercus rubra), red maple (Acer rubrum), and black birch (Betula lenta) were common overstory associates. Hemlock abundance increased from south to north, commonly on western and northwestern slopes. Average stand size was 55 ha, overstory basal area ranged from 23 to 55 m2 ha−1 and overstory stem densities averaged 993 ha−1. By 2004, 40% of sampled stands were infested, but most stands remained in good health overall; only 8 stands contained high HWA densities and only two had lost >50% overstory hemlock. Out of fifteen stand and landscape predictor variables examined, only latitude and winter climate variables were related to HWA density. Cold temperatures appear to be slowing the spread and impact of HWA at its northern extent as HWA infestation intensity and hemlock mortality and vigor were significantly correlated with average minimum winter temperature. Contrary to predictions, there was no regional increase in hemlock harvesting. The results suggest that regional HWA-hemlock dynamics are currently being shaped more by climate than by a combination of landscape and social factors. The persistence and migration of HWA continues to pose a significant threat regionally, especially in the northern portion of the study area, where hemlock dominates many forests.

The Legacy of Episodic Climatic Events in Shaping Temperate, Broadleaf Forests

In humid, broadleaf-dominated forests where gap dynamics and partial canopy mortality appears to dominate the disturbance regime at local scales, paleoecological evidence shows alteration at regional-scales associated with climatic change. Yet, little evidence of these broad-scale events exists in extant forests. To evaluate the potential for the occurrence of large-scale disturbance, we used 76 tree-ring collections spanning ∼840 000 km2 and 5327 tree recruitment dates spanning ∼1.4 million km2 across the humid eastern United States. Rotated principal component analysis indicated a common growth pattern of a simultaneous reduction in competition in 22 populations across 61 000 km2. Growth-release analysis of these populations reveals an intense and coherent canopy disturbance from 1775 to 1780, peaking in 1776. The resulting time series of canopy disturbance is so poorly described by a Gaussian distribution that it can be described as “heavy tailed,” with most of the years from 1775 to 1780 comprising th...