Woolly Adelgid Research Articles

Assessment of the potential for hybridisation between Laricobius nigrinus (Coleoptera: Derodontidae) and Laricobius osakensis, predators of the hemlock woolly adelgid (Hemiptera: Adelgidae)

In 2003, Laricobius nigrinus Fender was introduced into the eastern United States as a biological control agent of the hemlock woolly adelgid (Adelges tsugae Annand). Following its release, it was discovered that L. nigrinus was hybridising and producing viable progeny with Laricobius rubidus LeConte, a species native to eastern North America. Recently, Laricobius osakensis Montgomery and Shiyake was imported from Japan into the USA as a potential biological control agent of hemlock woolly adelgid. Hybridisation between L. nigrinus and L. rubidus led to interest in the outcome of interactions between L. osakensis and the other two Laricobius spp. The purpose of this study was to determine if L. osakensis could mate with L. nigrinus, if they could produce hybrid progeny, and whether mating interferes with reproductive output. Laricobius spp. were observed mating directly following emergence and found to be capable of producing sterile eggs in the absence of a mating event. Laboratory and confined field studies found no evidence that L. osakensis and L. nigrinus could produce hybrid progeny and the interaction between the two species did not result in a lower reproduction associated with interspecific mating attempts. Interbreeding should therefore not have an impact on biological control using these species. Fecundity experiments showed that L. osakensis produced eggs earlier in the season and at a higher rate than L. nigrinus, suggesting that L. osakensis may have the potential to be an even more successful biological control agent than L. nigrinus.

Where’s woolly? An integrative use of remote sensing to improve predictions of the spatial distribution of an invasive forest pest the Hemlock Woolly Adelgid

Non-native pests and pathogens present serious challenges to the management of forested ecosystems around the world. Early detection of pest and pathogen invasions may allow timely control and prevention methods to be implemented. Species distribution models (SDMs) and remote sensing (RS) methods have both been used effectively to determine locations of pest and pathogen damage. However, previous work integrating these two methods has rarely used RS metrics that have biological meaning. We use RS difference indices that show changes in forest cover from defoliation in order to map Hemlock Woolly Adelgid (HWA), Adelges tsugae, locations using MaxEnt in the Delaware Water Gap National Recreation Area (DWGNRA). Brightness, greenness, wetness, and Normalized Difference Vegetation Index (NDVI) were calculated from Landsat Thematic Mapper (TM) images for December 1982 and 2010. A difference for each index was created by subtracting the 1982 value from the 2010 value. We compared two models, one using difference indices and the other using 2010 indices along with other ancillary data layers, to determine if the more complicated but more biologically relevant difference indices were necessary for improved model performance. Variables with low importance were removed from both models, leaving NDVI, Wetness, soil, and elevation in the two final models. The difference model had an improvement in accuracy of three percent, across a number of threshold values. Despite this small difference in accuracy, however, the infected area predicted by the difference model (5.1% of total area) was approximately ½ of that predicted by the single year model (9.6% of total area). These results suggest that using remote sensing difference indices improves model accuracy slightly in terms of errors of omission, but also decreases predicted area of forest infestation by about 50%, suggesting that errors of commission would be substantially reduced using the difference approach. This method can provide forest managers more accurate information on the best locations to sample and treat.

Linking remote sensing and various site factors for predicting the spatial distribution of eastern hemlock occurrence and relative basal area in Maine, USA

Introduced invasive pests are perhaps the most important and persistent catalyst for changes in forest composition. Infestation and outbreak of the hemlock woolly adelgid (Adelges tsugae; HWA) along the eastern coast of the USA, has led to widespread loss of hemlock (Tsuga canadensis (L.) Carr.), and a shift in tree species composition toward hardwood stands.Developing an understanding of the geographic distribution of individual species can inform conservation practices that seek to maintain functional capabilities of ecosystems. Modeling is necessary for understanding changes in forest composition, and subsequent changes in biodiversity, and one that can be implemented at the species level. By integrating the use of remote sensing, modeling, and Geographic Information Systems (GIS) coupled with expert knowledge in forest ecology and disturbance, we can advance the methodologies currently available in the literature on predictive modeling.This paper describes an approach to modeling the spatial distribution of the less common but foundational tree species eastern hemlock throughout the state of Maine (∼84,000km2) at a high resolution. There are currently no published accuracy assessments on predictive models for high resolution continuous distribution of eastern hemlock relative basal area that span the geographic extent covered by our model, which is at the northern limit of the species’ range. A two stage mapping approach was used where presence/absence was predicted with an overall accuracy of 85% and the continuous distribution (percent basal area) was predicted with an accuracy of 84%. Overall, these findings are quite good despite high variability in the training dataset and the general minor component that eastern hemlock represents in the primary forest types in Maine.Eastern hemlock occurs along the southern half of the state stretching the east-west span with little to no occurrence in the northern regions. Several environmental and site characteristics, particularly average yearly maximum and minimum temperatures, were found to be positively correlated with hemlock occurrence. Eastern hemlock dominated stands appeared predominantly in the southwest corner of the state where HWA monitoring efforts can be focused. Given the importance of climate variables in predicting eastern hemlock, forecasts of future range shifts should be possible using data generated from climate scenarios.

Contrasting effects of two exotic invasive hemipterans on whole‐plant resource allocation in a declining conifer

AbstractInvasive herbivores can cause widespread dieback of naïve native hosts in the invaded range. Some consume leaves, some bore through wood, whereas others, such as piercing‐sucking insects, alter plant resource allocation through changes to source‐sink dynamics and depletion of long‐term stores. Invasive sap‐sucking herbivores that target cells critical to resource transport and storage may have particularly large effects. Herbivory by two exotic hemipterans, hemlock woolly adelgid (HWA), Adelges tsugae Annand (Adelgidae), and elongate hemlock scale (EHS), Fiorinia externa Ferris (Diaspididae), have very different effects on eastern hemlock, Tsuga canadensis (L.) Carrière (Pinaceae). Although these insects differ in both timing and feeding site on their hemlock host, the reasons for their differential effects are poorly understood. Here, using potted seedlings in a common garden, we examined the effects of these two herbivores on resource uptake and allocation immediately after an initial attack. We labeled the plants with a single pulse of 13CO2 and a supply of 15NH415NO3 every third day to obtain a whole‐plant perspective on resource uptake and allocation. After 10 weeks of controlled infestation, plants were measured and divided into tissue types (needles, branches, main stem, and roots). In each tissue we quantified biomass, 13C, 15N, total carbon (C), nitrogen (N), protein, and starch pools. Hemlock woolly adelgid feeding decreased new needle biomass by 34%, increased 13C allocation to roots and main stems by 130%, and increased 15N allocation to old foliage by 18%. Hemlock woolly adelgid infestation also resulted in increased starch storage in old branches over new needles, and marginally increased protein content plant‐wide. Elongate hemlock scale infestation resulted in a different growth pattern, with a 27% increase in biomass allocation to the main stem. Elongate hemlock scale also caused a 23% increase in N allocation to roots and main stem. Increases in resource allocation to main stem and belowground may indicate herbivore‐induced changes to storage patterns, or compensatory increased fine root growth to facilitate nutrient and water uptake. These resource allocation effects likely underlie the rapid and dramatic decline of hemlock in response to HWA feeding, and the considerably milder effects of EHS feeding.

Behavioral Responses ofLaricobiusspp. and Hybrids (Coleoptera: Derodontidae) to Hemlock Woolly Adelgid and Adelgid Host Tree Odors in an Olfactometer

The predatory species Laricobius nigrinus (Fender) and Laricobius osakensis (Shiyake and Montgomery) (Coleoptera: Derodontidae) have been released for biological control of hemlock woolly adelgid (Adelges tsugae; Hemiptera: Adelgidae) in eastern North America. L. osakensis is native to Japan, whereas L. nigrinus is endemic to the Pacific Northwest of the United States and Canada. After release, L. nigrinus was found to hybridize with the native eastern species, Laricobius rubidus (LeConte). The purpose of this study is to observe prey location behaviors of these three Laricobius species and L. nigrinus × L. rubidus (Ln × Lr) hybrids. Olfactometer bioassays were used to test response to host odors of adelgid-infested eastern hemlock, uninfested eastern hemlock, and uninfested eastern white pine. Predators reacted in the olfactometer more quickly when adelgid-infested foliage was included as a choice. L. nigrinus preferred infested eastern hemlock over uninfested eastern white pine, and L. rubidus preferred uninfested eastern white pine over uninfested eastern hemlock. Laricobius hybrids did not show a preference for foliage types known to be primary adelgid hosts (eastern hemlock and eastern white pine). Unequal preference by species of Laricobius for host trees of different adelgid prey could therefore be maintaining Laricobius species barriers despite hybridization. L. osakensis for this study were reared in the laboratory, whereas other species in this study were collected from the field, yet still were attracted to infested and uninfested eastern hemlock. This species also responded most quickly in the olfactometer, which is encouraging for successful biological control with this species.

Assessment of Imidacloprid and Its Metabolites in Foliage of Eastern Hemlock Multiple Years Following Treatment for Hemlock Woolly Adelgid, Adelges tsugae (Hemiptera: Adelgidae), in Forested Conditions.

Widespread decline and mortality of eastern hemlock, Tsuga canadensis (L.) Carrière, have been caused by hemlock woolly adelgid, Adelges tsugae (Annand) (HWA) (Hemiptera: Adelgidae). The current study is a retrospective analysis conducted in collaboration with Great Smoky Mountains National Park (GRSM) to determine longevity of imidacloprid and its insecticidal metabolites (imidacloprid olefin, 5-hydroxy, and dihydroxy) in GRSM's HWA integrated pest management (IPM) program. Foliage samples were collected from three canopy strata of hemlocks that were given imidacloprid basal drench treatments 4-7 yr prior to sampling. Foliage was analyzed to assess concentrations in parts per billion (ppb) of imidacloprid and its metabolites. Imidacloprid and its olefin metabolite were present in most, 95 and 65%, respectively, branchlets 4-7 yr post-treatment, but the 5-hydroxy and dihydroxy metabolites were present in only 1.3 and 11.7%, respectively, of the branchlets. Imidacloprid and olefin concentrations significantly decreased between 4 and 7 yr post-treatment. Concentrations of both imidacloprid and olefin were below the LC50 for HWA 5-7 yr post-treatment. Knowledge of the longevity of imidacloprid treatments and its metabolite olefin can help maximize the use of imidacloprid in HWA IPM programs.

Benthic Collector and Grazer Communities Are Threatened by Hemlock Woolly Adelgid-Induced Eastern Hemlock Loss

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.

Large wood in central Appalachian headwater streams: controls on and potential changes to wood loads from infestation of hemlock woolly adelgid

AbstractLarge wood (LW) is an important component of forested headwater streams. The character of LW loads reflects a balance between adjacent valley processes that deliver LW to the channel (herein recruitment processes) and stream channel processes that either retain or transport LW through the reach (herein retention processes). In the central Appalachian Mountains, USA, LW characteristics in headwater streams located in eastern hemlocks (Tsuga candensis) forests are expected to change because of infestation of hemlock woolly adelgid (Adelges tsugae, HWA), an exotic, invasive insect. We examined LW characteristics in 24 headwater streams ranging from un‐infested to severe infestation, as determined by hemlock canopy health. The objectives of this work were to: (i) quantify wood loads; (ii) assess the relative importance of valley recruitment and in‐stream retention mechanisms in controlling reach‐scale wood loads; and (iii) assess if there was a detectable influence of HWA on LW loads. We hypothesized that LW loads would be similar to other forested streams in eastern USA and dominated by recruitment processes. In addition, higher LW loads would correspond with advanced HWA infestation. Mean wood frequency was 38 pieces/100 m ± 17 (standard deviation); mean wood volume was 3.69 m3/100 m ± 2.76. In general, LW load characteristics were influenced by both recruitment and retention parameters; jam (accumulations ≥ 3 pieces) characteristics were dominated by retention parameters. Results suggest that adjacent stand basal area influences LW loads and once LW is recruited to the channel, streams lack sufficient hydraulic driving forces, despite having lower resistance structures, to transport LW out of the reach. Sites in moderate decline had higher proportions of short (1–2 m and 1–4 m) and very long (>10 m) LW with higher frequency of jams that were low in volume. We present a hypothesized conceptual model of expected changes to LW loads associated with HWA infestation and hemlock mortality. Copyright © 2015 John Wiley & Sons, Ltd.

Comparison of Suspended Branch and Direct Infestation Techniques for Artificially Infesting Hemlock Seedlings with the Hemlock Woolly Adelgid for Resistance Screening

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.

Benefits of invasion prevention: Effect of time lags, spread rates, and damage persistence

Quantifying economic damages caused by invasive species is crucial for cost-benefit analyses of biosecurity measures. Most studies focus on short-term damage estimates, but evaluating exclusion or prevention measures requires estimates of total anticipated damages from the time of establishment onward. The magnitude of such damages critically depends on the timing of damages relative to a species' arrival because costs are discounted back to the time of establishment. Using theoretical simulations, we illustrate how (ceteris paribus) total long-term damages, and hence the benefits of prevention efforts, are greater for species that a) have short lags between introduction and spread or between arrival at a location and initiation of damages, b) cause larger, short-lived damages (as opposed to smaller, persistent damages), and c) spread faster or earlier. We empirically estimate total long-term discounted impacts for three forest pests currently invading North America — gypsy moth (Lymantria dispar), hemlock woolly adelgid (Adelges tsugae), and emerald ash borer (Agrilus planipennis) — and discuss how damage persistence, lags between introduction and spread, and spread rates affect damages. Many temporal characteristics can be predicted for new invaders and should be considered in species risk analyses and economic evaluations of quarantine and eradication programs.

Changes in canopy structure and ant assemblages affect soil ecosystem variables as a foundation species declines

The decline of Tsuga canadensis (eastern hemlock)—a foundation tree species—due to infestation by Adelges tsugae (hemlock woolly adelgid) or its complete removal from a stand by salvage logging dramatically affects associated faunal assemblages. Among these assemblages, species composition (richness and abundance) of ants increases rapidly as T. canadensis is lost from the stands. Because ants live and forage at the litter‐soil interface, we hypothesized that environmental changes caused by hemlock loss (e.g., increased light and warmth at the forest floor, increased soil pH) and shifts in ant species composition would interact to alter soil ecosystem variables. In the Harvard Forest Hemlock Removal Experiment (HF‐HeRE), established in 2003, T. canadensis in large plots were killed in place or logged and removed to mimic adelgid infestation or salvage harvesting, respectively. In 2006, we built ant exclosure subplots within all of the canopy manipulation plots to examine direct and interactive effects of canopy change and ant assemblage composition on soil and litter variables. Throughout HF‐HeRE, T. canadensis was colonized by the adelgid in 2009, and the infested trees are now declining. The experimental removal of T. canadensis from the canopy was associated with an increase in the rate of cellulose decomposition by >50%, and exclosure of ants from subplots directly reduced their soil nitrate availability by 56%. Partial least squares path models revealed sequential interactive effects prior to adelgid infestation: canopy change (as a proxy for associated environmental changes) altered both decomposition and ant assemblage structure; changes in ant assemblage structure and decomposition rates altered nitrogen availability. The results illustrate that biotic changes directly associated with decline of T. canadensis can have cascading effects on ecosystem nutrient availability and cycling.

Long-Term Effects of Imidacloprid on Eastern Hemlock Canopy Arthropod Biodiversity in New England

Abstract The systemic insecticide imidacloprid is commonly used to protect trees against attack by the Adelges tsugae (Hemlock Woolly Adelgid [HWA]), an invasive pest that threatens Tsuga canadensis (Eastern Hemlock) and T. caroliniana (Carolina Hemlock) in eastern North America. Although there have been some studies documenting the short-term (1–3 years) impact of imidacloprid on non-target arthropods in hemlock systems, almost nothing is known about the impact over longer time scales. Here, using a set of trees which were experimentally treated 3 and 9 years prior to this study, we found that while the impact of imidacloprid on HWA may be approaching the limits of detection and efficacy on trees treated 9 years ago, there is still an intermittently detectable impact on HWA density. Similarly, 9 years after application there is a subtle but detectable increase in arthropod richness and a shift in canopy-arthropod community composition. Results from the 3-year treated trees were, however, ambiguous, but m...

Invasive insect effects on nitrogen cycling and host physiology are not tightly linked.

Invasive insects may dramatically alter resource cycling and productivity in forest ecosystems. Yet, although responses of individual trees should both reflect and affect ecosystem-scale responses, relationships between physiological- and ecosystem-scale responses to invasive insects have not been extensively studied. To address this issue, we examined changes in soil nitrogen (N) cycling, N uptake and allocation, and needle biochemistry and physiology in eastern hemlock (Tsuga canadensis (L) Carr) saplings, associated with infestation by the hemlock woolly adelgid (HWA) (Adelges tsugae Annand), an invasive insect causing widespread decline of eastern hemlock in the eastern USA. Compared with uninfested saplings, infested saplings had soils that exhibited faster nitrification rates, and more needle (15)N uptake, N and total protein concentrations. However, these variables did not clearly covary. Further, within infested saplings, needle N concentration did not vary with HWA density. Light-saturated net photosynthetic rates (Asat) declined by 42% as HWA density increased from 0 to 3 adelgids per needle, but did not vary with needle N concentration. Rather, Asat varied with stomatal conductance, which was highest at the lowest HWA density and accounted for 79% of the variation in Asat. Photosynthetic light response did not differ among HWA densities. Our results suggest that the effects of HWA infestation on soil N pools and fluxes, (15)N uptake, needle N and protein concentrations, and needle physiology may not be tightly coupled under at least some conditions. This pattern may reflect direct effects of the HWA on N uptake by host trees, as well as effects of other scale-dependent factors, such as tree hydrology, affected by HWA activity.

Invasions by two non-native insects alter regional forest species composition and successional trajectories

While invasions of individual non-native phytophagous insect species are known to affect growth and mortality of host trees, little is known about how multiple invasions combine to alter forest dynamics over large regions. In this study we integrate geographical data describing historical invasion spread of the hemlock woolly adelgid, Adelges tsugae, and beech scale, Cryptococcus fagisuga, with regional forest inventory data collected by the US Forest Service’s Forest Inventory and Analysis program to quantify the individual and combined impacts of these pest species. This analysis indicates that regional impacts of these insects on their hosts occur surprisingly slow but act to change regional forest succession pathways. Because beech and hemlock commonly co-occur in eastern North American forests, invasions by the two pest species are altering the current and future composition of large forest regions through their impacts on these two late-successional species. Such results demonstrate how forest insect invasions can profoundly modify forest dynamic processes, resulting in long-term changes in forest ecosystems.

Effects of Light and Water Availability on the Performance of Hemlock Woolly Adelgid (Hemiptera: Adelgidae)

Eastern hemlock (Tsuga canadensis (L.) Carriere) is a dominant shade-tolerant tree in northeastern United States that has been declining since the arrival of the hemlock woolly adelgid (Adelges tsugae Annand). Determining where A. tsugae settles under different abiotic conditions is important in understanding the insect's expansion. Resource availability such as light and water can affect herbivore selectivity and damage. We examined how A. tsugae settlement and survival were affected by differences in light intensity and water availability, and how adelgid affected tree performance growing in these different abiotic treatments. In a greenhouse at the University of Rhode Island, we conducted an experiment in which the factors light (full-sun, shaded), water (water-stressed, watered), and adelgid (infested, insect-free) were fully crossed for a total of eight treatments (20 two-year-old hemlock saplings per treatment). We measured photosynthesis, transpiration, water potential, relative water content, adelgid density, and survival throughout the experiment. Adelgid settlement was higher on the old-growth foliage of shaded and water-stressed trees, but their survival was not altered by foliage age or either abiotic factor. The trees responded more to the light treatments than the water treatments. Light treatments caused a difference in relative water content, photosynthetic rate, transpiration, and water potential; however, water availability did not alter these variables. Adelgid did not enhance the impact of these abiotic treatments. Further studies are needed to get a better understanding of how these abiotic factors impact adelgid densities and tree health, and to determine why adelgid settlement was higher in the shaded treatments.

Survival and Near Extinction of Hemlock Woolly Adelgid (Hemiptera: Adelgidae) During Summer Aestivation in a Hemlock Plantation.

Varying densities of the spring generation of the hemlock woolly adelgid were manipulated on 16 previously uninfested eastern hemlocks in an open-field plantation in Massachusetts. In contrast to experimentally created hemlock woolly adelgid populations in a forest, as reported previously, there was no evidence of density-dependent survival on a tree-wide basis in the plantation in the spring (progrediens) generation. There was, however, comparable density-dependent survival of settled crawlers and sexupara production when samples of the population were examined from branches with high density. Plantation hemlocks had 9.3 times more foliage and 10 times lower adelgid densities per cm than the forest hemlocks. These results show that density-dependent processes in the progrediens generation may only be evident when hemlock woolly adelgid density reaches a critical threshold. In the sistens generation that begins in midsummer, we counted a mean of 126 settled crawlers on marked branch on all 16 trees, but none of these adelgids survived the mid-summer aestivation phase, perhaps due to higher temperatures that were recorded in the plantation compared with a nearby hemlock forest, where 16% of the adelgids survived the aestivation phase. Whole tree counts of overwintering adelgids revealed that the adelgid populations had gone extinct on 13 out of the 16 trees. Mortality in the midsummer aestivation phase often exceeds overwintering mortality that has been widely thought to be the main factor that limits adelgid population growth and spread, particularly in northern states.

Scymnus camptodromus (Coleoptera: Coccinellidae) Larval Development and Predation of Hemlock Woolly Adelgid (Hemiptera: Adelgidae).

Development time and prey consumption of Scymnus (Neopullus) camptodromus Yu and Liu (Coleoptera: Coccinellidae) larvae by instar, strain, and temperature were evaluated. S. camptodromus, a specialist predator of hemlock woolly adelgid Adelges tsugae (Annand) (Hemiptera: Adelgidae), was brought to the United States from China as a potential biological control agent for A. tsugae. This beetle has been approved for removal from quarantine but has not yet been field released. We observed that temperature had significant effects on the predator's life history. The larvae tended to develop faster and consume more eggs of A. tsugae per day as rearing temperature increased. Mean egg consumption per day of A. tsugae was less at 15°C than at 20°C. However, as larvae took longer to develop at the lower temperature, the total number of eggs consumed per instar during larval development did not differ significantly between the two temperatures. The lower temperature threshold for predator larval development was estimated to be 5°C, which closely matches the developmental threshold of A. tsugae progrediens. Accumulated degree-days for 50% of the predator neonates to reach adulthood was estimated to be 424. Although temperature had a significant effect on larval development and predation, it did not impact survival, size, or sex ratio of the predator at 15 and 20°C. Furthermore, no remarkable distinctions were observed among different geographical populations of the predator.

Influence of Hemlock Woolly Adelgid Infestation Levels on Water Stress in Eastern Hemlocks within the Great Smoky Mountains National Park, U.S.A.

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.

Hemlock woolly adelgid alters fine root bacterial abundance and mycorrhizal associations in eastern hemlock

While the impact of aboveground herbivores on plant biomass and fitness has received considerable attention, there has been far less research on the corresponding belowground impacts. The belowground effects of aboveground feeding may be particularly noticeable for invasive and/or outbreaking herbivore species that reach high densities and can cause major damage and sometimes death. The hemlock woolly adelgid, Adelges tsugae, is an invasive pest on the eastern seaboard of the United States that feeds on a native shade-tolerant conifer, the eastern hemlock Tsuga canadensis. Trees rapidly decline and die following infestation, and the invasion of this insect has devastated hemlock populations from Georgia in the south to Maine in the north. Despite their substantial impact on tree health, we are unaware of any research into the adelgid’s effect on hemlock roots and the surrounding rhizosphere. We report the results of research assessing ectomycorrhizal root colonization, rhizosphere bacterial abundance, and root C:N ratios of infested and uninfested T. canadensis. We found that adelgid infestation decreased the percentage of root material colonized by ectomycorrhizal fungi by more than 67%. Rhizosphere bacterial abundance on fine roots was 25% lower on adelgid-infested versus uninfested trees, and roots of adelgid-infested trees contained significantly less carbon. Our results demonstrate that aboveground adelgid infestation can affect hemlock root composition and alter belowground interactions with ectomycorrhizal fungi and bacteria. This information demonstrates that above-belowground linkages can transmit the impact of herbivory far from the site of localized damage.

Field assessment of hybridization between Laricobius nigrinus and L. rubidus, predators of Adelgidae

Two adelgid predators, Laricobius nigrinus Fender and Laricobius rubidus LeConte, were recently discovered to produce hybrid progeny in the eastern United States. L. rubidus is native to eastern North America where it feeds on pine bark adelgid (Pineus strobi Hartig) and L. nigrinus is native to western North America and was introduced to the eastern United States in 2003 for biological control of hemlock woolly adelgid (Adelges tsugae Annand). Currently, L. nigrinus and L. rubidus form a mosaic of hybrid zones throughout the eastern United States. It is not known whether these zones will be maintained over time and whether hybridization will impact the efficacy of biological control or result in displacement of L. rubidus. Sampling from 2007 to 2012 on eastern hemlock (Tsuga canadensis Carrière) showed a clear increase in L. nigrinus, a decrease in L. rubidus, and a steady proportion of hybrids. Sampling from 2010 to 2012 on both eastern hemlock and white pine (Pinus strobus L.) at ten sites showed that L. nigrinus was more dominant on hemlock and L. rubidus was more dominant on white pine, which may demonstrate habitat preference and promote segregation between the two species. Site factors were tested for a relationship with the proportion of hybrids. The number of years L. nigrinus had been present at the site was the only factor that showed a relationship. The results suggest that L. nigrinus may displace L. rubidus on hemlock, but not on white pine at sites where they are both present.