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Simple Summary: Understanding where insect pests inhabit and how they interact with their environment is important for creating sampling and management plans. This study focused on the hemlock woolly adelgid, a pest that affects hemlock trees, to find the best way to sample these pests and understand their distribution in hemlock stands in the USA. We randomly selected 24 branches from each of 46 infested hemlock trees and counted the egg sacs of hemlock woolly adelgid on each branch to determine the best sampling unit. We also divided three hemlock forest areas into grids and counted the egg sacs in each grid cell to study the spatial distribution pattern of hemlock woolly adelgid. This study found that hemlock woolly adelgids tended to aggregate at the tips of branches and that a 50 cm branch from the lower canopy was the best sampling unit. The hemlock woolly adelgid also showed spatial aggregation in the hemlock stands, and its abundance was correlated with tree height and tree diameter at breast height. These findings help in better sampling and managing hemlock woolly adelgid. Understanding the spatial patterns of insect pests and their associations with their environments is crucial for developing effective sampling and management plans. This study was conducted to identify optimal sampling units for the hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae) and to characterize its spatial distribution patterns in hemlock (Tsuga canadensis (L.) Carrière) stands in West Virginia, USA. To determine the optimal sampling unit, we randomly selected 24 branches from each of 46 A. tsugae-infested hemlock trees. The locations and number of A. tsugae ovisacs on each branch were recorded and the coefficient of variation was used to choose the optimal sampling units. To determine the spatial patterns of A. tsugae, each of the three 1-ha hemlock stands was divided into 100 grids, and ovisac counts were taken from the center of each grid. Semivariograms and spatial analysis by distance indices (SADIE) were used to analyze the spatial patterns of A. tsugae. In addition, various environmental and biological factors were measured to explore their spatial associations with A. tsugae. The results of this study revealed that the A. tsugae ovisacs exhibited spatial aggregation within branches, predominantly at the tips, and a 50 cm branch approximately 3 m above the ground was the optimal sampling unit. The spatial aggregation of A. tsugae in the hemlock stands was evident, and positive spatial associations were found between A. tsugae populations and factors including the aspect, tree diameter at breast height, and tree height. These findings offer valuable insights for the sampling and management of A. tsugae. [ABSTRACT FROM AUTHOR]

Widespread mortality of eastern hemlock (Tsuga canadensis [L.] Carr.) has been occurring due to the introduction of hemlock woolly adelgid (Adelges tsugae Annand) (HWA), threatening millions of hectares of hemlock-dominated forests in the eastern United States. HWA feeds at the base of needles and removes stored carbohydrates, which can impact leaf-level physiology, contributing to the decline of the tree. However, these physiological mechanisms in HWA-infested hemlocks are still not clearly understood. We investigated hemlock leaf physiology year-round at three forested sites with various degrees of infestation. At each site, half the trees were treated with imidacloprid (Merit® 2 F, Bayer, Kansas City, MO, USA) while the rest were left untreated. Imidacloprid is widely used to control HWA but can itself have phytotoxic effects. After one growing season, there was an increase in photosynthetic rates (7.5%, p = 0.0163) and stomatal conductance (7.1%, p = 0.0163) across sites in the trees treated with imidacloprid. After two years, the imidacloprid treatment also increased bud break from 22.5% to 88.7% at Fishburn (the most severely impacted site) and from 22.7% to 58.9% at Mountain Lake (the least impacted site), and slightly increased chlorophyll fluorescence for treated trees at Fishburn. Chemical treatment also slightly increased water use efficiency at Mountain Lake. These results suggest that HWA is causing tree mortality largely through a reduction in leaf area caused by decreasing bud break and also by a slight, but significant, reduction in leaf-level photosynthesis and stomatal conductance. [ABSTRACT FROM AUTHOR]

The hemlock woolly adelgid (HWA), Adelges tsugae, is an invasive pest of Tsuga spp. in eastern North America. Scymnus coniferarum is a predaceous beetle that was collected from HWA in the western United States. Limited knowledge of this insect in its native habitat led to studies to evaluate its potential for biological control of HWA. Seasonal abundance was sampled at six sites in Tacoma, WA, twice monthly, for one year on different host trees of potential adelgid prey. Tree species included Pseudotsuga menziesii, Pinus contorta, Pinus monticola, and Tsuga heterophylla. Scymnus coniferarum adults were found on all conifer species, except P. menziesii. Each conifer species supported a different adelgid species, suggesting S. coniferarum feeds on multiple adelgid species. More S. coniferarum were found on pine than hemlock. DNA barcoding of S. coniferarum found two distinct clusters that differed by 6% divergence. Beetles in each cluster were co-habiting the same conifer species, and they could not be distinguished morphologically. Further taxonomic studies are needed to understand the significance of DNA barcode sequence divergence. Because S. coniferarum was more frequently associated with pine adelgids than HWA, and because of remaining taxonomic uncertainty, S. coniferarum may not be suitable for HWA biological control. [ABSTRACT FROM AUTHOR]

Eastern (Tsuga canadensis) and Carolina hemlocks (T. caroliniana) of eastern North America have been attacked by the non-native hemlock woolly adelgid (Adelges tsugae Annand) (HWA) since the first half of the 20th century. Unlike most insects, HWA develops through one generation from fall to late winter, exposing this insect to the lethal effects of winter temperatures. The mortality inflicted by winter temperatures on HWA determines the surviving population density as well as its ability to spread to uninfested areas. With the ongoing changes in climate, knowledge of this species' ability to survive and spread in the future can help land managers prepare for its management. This study began during the winter of 2014 and ended in the spring of 2017. During this period, winter mortality of HWA was recorded at 100 sites from Maine to Georgia (n = 209). Changes in population density from the sistens to the succeeding progrediens generation were recorded at 24 sites (n = 35). Models were developed to predict HWA mortality using the lowest minimum temperature prior to the mortality assessment date, the number of days with mean temperature < 1 °C, and the mean daily temperature of the three days preceding that minimum. Models were also developed to predict population density changes from the overwintering sistens generation to the following progrediens generation. Future projections under climate change showed increases in winter survival and population growth rates over time. Especially towards the northeastern edge of T. canadensis' distribution as minimum temperatures are predicted to increase at a greater rate. This will result in an increase in density throughout its current distribution and expansion northward causing an increase in its impact on eastern Tsuga spp. [ABSTRACT FROM AUTHOR]

Hemlock woolly adelgid (HWA), Adelges tsugae Annand, an invasive insect native to the Pacific Northwest and Asia, is responsible for widespread health decline and mortality of native hemlocks (Tsuga spp.) in the eastern United States. Shading and fertilizer has been found to affect the survival and health of both HWA and hemlocks. These abiotic factors have been studied separately but not in combination. In this three year study, eastern hemlock trees (1-2 m tall) were treated with pruning, fertilizer, and shade to determine their effects on hemlock tree health and HWA survival and density. Shade cloths were erected over individual trees, granulated fertilizer was applied, and trees were pruned annually. The total number of HWA were counted during the sistens and progrediens adult stages on the low, mid, and high branches on the north, east, south, and west sides of each tree for three years. Survival of aestivating sistens was recorded in artificially, naturally, and unshaded hemlocks. The mean of percent tips alive, branches alive, and foliage density was used to calculate a hemlock health index (scale of 0-100). Shade cloth reduced solar radiation to the trees to levels similar to a naturally-forested hemlock canopy, but did not alter temperature. Trees exposed to shade alone and shade plus fertilizer maintained the greatest HWA density. On unshaded trees, branches on the west side of the tree had lower HWA densities and branches high on the tree had the lowest HWA densities. Pruning plus fertilizer and shading plus fertilizer reduced tree health. Shaded trees had reduced branchlet new growth length. Survival of summer aestivating sistens was nearly twice the survival under artificially- and naturally-shaded trees compared to unshaded trees. There was an inverse density-dependent survival response for aestivating HWA under artificially-shaded and unshaded trees but not naturally-shaded trees. Unshaded hemlock trees had lower HWA densities due to increased mortality of summer aestivating sistens. Unshaded trees had better health and longer new growth branchlets due to increased exposure to solar radiation and lower HWA densities. Silvicultural thinning of hemlocks in forest stands could increase direct sunlight reaching the trees and help decrease HWA densities and improve hemlock health. [ABSTRACT FROM AUTHOR]

Eastern hemlock (Tsuga canadensis) is a foundation species in eastern North American forests. Because eastern hemlock is a foundation species, it often is assumed that the diversity of associated species is high. However, the herbaceous layer of eastern hemlock stands generally is sparse, species-poor, and lacks unique species or floristic assemblages. The rapidly spreading, nonnative hemlock woolly adelgid (Adelges tusgae) is causing widespread death of eastern hemlock. Loss of individual hemlock trees or whole stands rapidly leads to increases in species richness and cover of shrubs, herbs, graminoids, ferns, and fern-allies. Naively, one could conclude that the loss of eastern hemlock has a net positive effect on biodiversity. What is lost besides hemlock, however, is landscape-scale variability in the structure and composition of the herbaceous layer. In the Harvard Forest Hemlock Removal Experiment, removal of hemlock by either girdling (simulating adelgid infestation) or logging led to a proliferation of early-successional and disturbance-dependent understory species. In other declining hemlock stands, nonnative plant species expand and homogenize the flora. While local richness increases in former eastern hemlock stands, between-site and regional species diversity will be further diminished as this iconic foundation species of eastern North America succumbs to hemlock woolly adelgid. [ABSTRACT FROM AUTHOR]

Eastern hemlock (Tsuga canadensis (L.) is a foundation species in eastern North America where it is under threat from the highly invasive, exotic hemlock woolly adelgid (Adelges tsugae). Eastern hemlock is especially important in riparian areas of Central and Southern Appalachia, so we compared the spatial and temporal composition of benthic collector-gatherers, collector-filterers, and grazers in headwater streams with hemlock-dominated riparian vegetation to those with deciduous tree-dominated riparian vegetation to evaluate the extent to which adelgid-induced hemlock loss could influence composition and abundance of these two functional feeding groups. We found differences in benthic invertebrate abundance and family-level diversity based on riparian vegetation and sampling approach, and, often, riparian vegetation significantly interacted with location or season. Collector-gatherers and grazers were more abundant in eastern hemlock streams in the summer, when hemlock litter is readily available and deciduous litter is relatively sparse. Riparian eastern hemlock appears to exert considerable influence on benthic invertebrate functional feeding group composition in headwater stream communities, as expected with a foundation species. With the loss of eastern hemlock due to adelgid-induced mortality, we should expect to see alterations in spatial and temporal patterns of benthic invertebrate abundance and diversity, with potential consequences to both benthic and terrestrial ecosystem function. [ABSTRACT FROM AUTHOR]

The hemlock woolly adelgid (Adelges tsugae Annand) is an invasive forest pest in eastern North America that has caused significant decline and mortality in populations of eastern hemlock (Tsuga canadensis (L.) Carr.) and Carolina hemlock (T. caroliniana Engelm.). The breeding of adelgid-resistant genotypes for reforestation activities is still in the early development phases, and most resistance screening programs have depended on labor-intensive direct artificial infestation techniques for introducing adelgids to target seedlings. We investigated the timing and effectiveness of a potentially less labor-intense suspended branch infestation technique compared to two levels of a direct infestation method. Results indicated that peak crawler emergence from adelgid infested hemlock branches occurred within a 10 to 14 day period and that crawler emergence was higher from non-hydrated compared to hydrated branches. Greater infestation pressure was achieved when using progrediens crawlers compared to sistens crawlers. In 2013, when the infestation attempts were most successful, the suspended branch technique induced the same or higher adelgid densities on target seedlings as the direct infestation techniques. Assuming an initial investment in infrastructure, the suspended branch approach could be a more time and cost effective method for inducing adelgid infestations for resistance screening of large numbers of candidate trees. [ABSTRACT FROM AUTHOR]

Extensive mortality of eastern hemlock, Tsuga canadensis (L.) Carrière, resulting from infestation by hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae), has occurred throughout the eastern United States. Although imidacloprid treatment can reduce tree mortality, its effectiveness can be influenced by several factors including tree water stress. The relationship between water stress and infestation rates is unknown, and an understanding of these could greatly increase the efficiency of management for this invasive insect. The primary objective of this study was to assess water stress at three levels of hemlock woolly adelgid infestations. Water stress was measured monthly for 13 months in eastern hemlocks classified as <25%, 25%-75%, and >75% infested. The highest level of water stress was found in those trees with hemlock woolly adelgid infestation levels greater than 75%. The lowest level of water stress was found in those trees with less than 25% hemlock woolly adelgid infestation levels. Knowledge of these effects can contribute to development of more effective chemical management strategies. [ABSTRACT FROM AUTHOR]

Mycopathogens of economically important exotic invasive insects in forests of northeastern USA have been the subject of research at the Entomology Research Laboratory, University of Vermont, for the last 20 years. Elongate hemlock scale, European fruit lecanium, hemlock woolly adelgid and pear thrips were analyzed for the presence of mycopathogens, in order to consider the potential for managing these pests with biological control. Fungal cultures isolated from insects with signs of fungal infection were identified based on morphological characters and DNA profiling. Mycopathogens recovered from infected insects were subdivided into three groups, i.e., specialized entomopathogenic; facultative entomopathogens; ubiquitous opportunistic contaminants. Epizootics were caused by fungi in the specialized group with the exception of M. microspora, P. marquandii and I. farinosa. Inoculation of insects in laboratory and field conditions with B. bassiana, L. muscarium and Myriangium sp. caused insect mortality of 45 to 95%. Although pest populations in the field seemed severely compromised after treatment, the remnant populations re-established themselves after the winter. Although capable of inducing high mortality, a single localized aerial application of a soil-dwelling fungus does not maintain long-time suppression of pests. However, it can halt their range expansion and maintain populations below the economic threshold level without the use of expensive insecticides which have a negative impact on the environment. [ABSTRACT FROM AUTHOR]

Hemlock woolly adelgid ( Adelges tsugae Annand, HWA) remains the single greatest threat to the health and sustainability of hemlock in the eastern USA. The loss of hemlock trees leads to further negative impacts on the diversity and stability of ecosystems in the eastern part of North America. It is, therefore, urgent to develop effective control measures to reduce HWA populations and promote overall hemlock health. Currently available individual and integrated approaches should continue to be evaluated in the laboratory and in the field along with the development of other new and innovative methods.