Woolly Adelgid Research Articles

Variable colonization by the hemlock woolly adelgid suggests infestation is associated with hemlock host species

Studies increasingly indicate host resistance likely influences hemlock woolly adelgid (HWA) populations. We describe coexisting HWA populations in four hemlock species, three from HWA’s native range (Tsuga chinensis, T. dumosa, and T. sieboldii) and one from its introduced range: T. canadensis. HWA populations were greater in T. canadensis and lesser or absent in T. chinensis and T. dumosa. HWA stylet insertion was observed the least in T. chinensis and T. dumosa, suggesting a physical and/or chemical feeding deterrent. While T. chinensis and T. dumosa consistently and distinctly differed from T. canadensis, T. sieboldii grouped with T. canadensis or T. chinensis and T. dumosa in different experiments, suggesting mechanisms influencing HWA populations differ by host species. HWA were found more often on tops of T. chinensis and T. dumosa branches but equally on tops and bottoms of T. canadensis branches, corresponding with locations of trichomes (which T. sieboldii lacks). Evidence from the literature about dissimilarities in hemlock hosts (e.g., HWA interactions and vulnerability, feeding damage response, cuticle thickness, terpenoid chemistry, arthropod and microbial communities) and native and introduced HWA populations (e.g., losses of sexual reproduction and primary spruce host) suggests a limit to applying information from native systems to management in introduced systems.

Combined Effects of Hemlock Woolly Adelgid (Adelges tsugae) Infestation and Treatment with Imidacloprid on Eastern Hemlock (Tsuga canadensis) Leaf Radiometry

Abstract The neonicotinoid systemic insecticide imidacloprid has proven to be an effective treatment for the prevention and control of hemlock woolly adelgid (Adelges tsugae Annand) in southeastern populations of eastern hemlock (Tsuga canadensis L.). Recent studies have demonstrated that imidacloprid and A. tsugae both stimulate salicylic acid-dependent physiological responses in plant tissues responsible for plant defenses against pathogens, timing of developmental outcomes including flowering and leaf senescence, and resilience to abiotic stressors. We evaluated the interacting effects of A. tsugae presence/absence and treatment with imidacloprid on leaf optical properties indicative of photosynthetic potential, photosynthetic efficiency, and tissue senescence. Our results indicated that A. tsugae changes lower canopy leaf optical properties indicative of reduced photosynthetic potential/efficiency and accelerated senescence in mature leaves. Imidacloprid was associated with declines in photosynthetic potential and showed a largely similar, though less pronounced, effect on leaf spectral properties to that of A. tsugae.

Hemlock woolly adelgid niche models from the invasive eastern North American range with projections to native ranges and future climates

Abstract: The hemlock woolly adelgid (Adelges tsugae Annand - HWA) is invasive in eastern North America where it causes extensive mortality to hemlock communities. The future of these communities under projected climate change is an issue of landscape ecological interest and speculation. We employed the MaxEnt algorithm with the random subset feature selection algorithm (RSFSA) in creating HWA niche models. Final models were ensembles of 12 statistically best models with six predictors each. Out of 119 climatic, topographic, and soil variables, 42 were used in at least one final model. Soil features, followed by climate and topographic features, were most common in selected models. The three most important variables among all models were November potential evapotranspiration, slope, and percent Ochrepts soil. The potential distributions of HWA within eastern North America were projected under historical and four future climate scenarios for 2050 and 2070 under low and high CO2 emissions. The mean of the minimum values for the minimum temperature of the coldest month from the 12 MaxEnt model projections in eastern North America was -15.8°C. This value was close to -15°C, the extreme minimum temperature found for both HWA occurrence points and previously reported HWA cold temperature limits. These results indicate that HWA may be close to equilibrium distribution in eastern North America under current climate. We also reverse-casted the eastern North American MaxEnt model back onto the HWA native ranges in eastern Asia and western North America. The projections match best with native ranges in Asian islands, such as Japan, and the Cascade Mountains in western North America. Statistically significant HWA range shifts of 221-468 km northwards and 110-164 km eastwards were projected by the 12 models for 2050-2070. The 2070 high CO2 emission scenario models projects HWA suitability throughout most of the northern range of eastern hemlock.

Lack of forest tree seedling recruitment and enhanced tree and shrub growth characterizes post-Tsuga canadensis mortality forests in the southern Appalachians

The loss of Tsuga canadensis from invasion by hemlock woolly adelgid (Adelges tsugae, HWA) has altered ecosystem structure and function in forests across the eastern United States. In southern Appalachian forests, T. canadensis co-occurred with hardwood species and an evergreen Rhododendron maximum shrub layer in riparian and cove positions. In this region, HWA infestation was detected in 2003, with mortality reaching 97% by 2014. In this study we examined responses of light, soil moisture, tree seedling density, and overstory and understory vegetation growth from 2004 to 2014 following HWA infestation and mortality of T. canadensis. We hypothesized that seedling recruitment and vegetation growth would continue to increase over time as observed with initial trends reported through 2009, and that species that associate with ectomycorrhizal (ECM) fungi may benefit more than those that do not due to the pulse of organic material in these stands from dead T. canadensis trees. Light transmission measured at 1 m above the ground increased from 2006 to 2009, but gradually decreased from 2015 to 2017. Basal area of overstory non-Tsuga trees increased only marginally, and there was no recruitment of tree seedlings to the overstory size class, even though seedling density of deciduous species increased initially. Increased basal area and stem density of R. maximum may explain the light and seedling responses, as this species can inhibit tree seedling recruitment by limiting light and nutrients. Overstory species with the highest basal area increment (BAI) in the post-T. canadensis stands were Pinus rigida, Betula lenta and Quercus coccinea, which all associate with ECM fungi. However, not all ECM tree species grew significantly more following T. canadensis mortality compared to pre-mortality growth rates—only those ECM species that had high growth rates prior to mortality did. After a decade, growth of both overstory trees and R. maximum has not compensated for the loss of T. canadensis. Active management of R. maximum, which may involve the removal of the evergreen shrub and soil organic layer, may be required to allow for diverse tree seedling recruitment; and subsequently, restore riparian forest structure, diversity, and function.

Hemlock Woolly Adelgid (Hemiptera: Adelgidae) Management in Forest, Landscape, and Nursery Production.

Hemlock woolly adelgid, Adelges tsugae (Annand) (Hemiptera: Adelgidae), has caused significant damage to both eastern [Tsuga canadensis (L.) Carrière] and Carolina hemlock (Tsuga caroliniana Englemann) (Pinales: Pinaceae) since it was first reported in the eastern United States. This adelgid is particularly damaging to these hemlock species due to a lack of co-evolved plant defenses and natural enemies able to suppress hemlock woolly adelgid populations. Management of hemlock woolly adelgid relies heavily on insecticides to prevent death of vulnerable trees. Biological control programs have released natural enemies of hemlock woolly adelgid to aid in control at the landscape level. Quarantine restrictions on hemlock are in place in some regions of the United States and Canada. These quarantines impact sales and shipment of hemlock trees from nurseries as well as other hemlock products. A review of insect biology, description of life stages, damage, management options, and quarantine restrictions for hemlock woolly adelgid is presented.

Do Terrestrial Salamanders Indicate Ecosystem Changes in New England Forests?

Long-term ecological research (LTER) and monitoring programs accrue invaluable ecological data that inform policy and improve decisions that enable adaptation to and mitigation of environmental changes. There is great interest in identifying ecological indicators that can be monitored easily and effectively to yield reliable data about environmental changes in forested ecosystems. However, the selection, use, and validity of ecological indicators to monitor in LTER programs remain challenging tasks for ecologists and conservation biologists. Across the eastern United States of America, the foundation tree species eastern hemlock (Tsuga canadensis (L.) Carrière) is declining and dying from irruptions of a non-native insect, the hemlock woolly adelgid (Adelges tsugae Annand). We use data from the Harvard Forest LTER site’s Hemlock Removal Experiment together with information from other eastern US LTER sites to show that plethodontid salamanders can be reliable indicators of ongoing ecological changes in forested ecosystems in the eastern USA. These salamanders are abundant, they have a history of demographic stability, are both predators and prey, and can be sampled and monitored simply and cost-effectively. At the Harvard Forest LTER, red-backed salamanders (Plethodon cinereus Green) were strong indicators of intact forests dominated by eastern hemlock (Tsuga canadensis); their high site fidelity and habitat specificity yielded an indicator value (robust Dufrêne and Legendre’s “IndVal”) for this species of 0.99. Eastern red-spotted newts (Notopthalmus viridescens viridescens Rafinesque) were better indicators of nearby stands made up of young, mixed hardwood species, such as those which replace hemlock stands following adelgid infestation. At the Hubbard Brook and Coweeta LTER sites, plethodontid salamanders were associated with intact riparian habitats, which may also be dominated by eastern hemlock. We conclude that plethodontid salamanders satisfy most criteria for reliable ecological indicators of environmental changes in eastern US forests.

Conifer responses to a stylet‐feeding invasive herbivore and induction with methyl jasmonate: impact on the expression of induced defences and a native folivore

Abstract Trees attacked by multiple herbivores need to defend themselves against dynamic biotic challenges; appropriate responses to one stressor can elicit hormonal responses that are antagonistic to another. Hemlock (Tsuga canadensis) infestation by hemlock woolly adelgid (HWA; Adelges tsugae) results in the accumulation of the defensive hormone salicylic acid. We explored the potential for HWA infestation to interfere with anti‐folivore‐induced defence signalling and its implications for a native folivore (hemlock looper; Lambdina fiscellaria). Hemlocks were infested with HWA and/or sprayed with methyl jasmonate; foliar defences were analyzed and foliage quality for looper larvae was assessed. Both treatments activated foliar defensive traits, including a HWA‐mediated increase in peroxidase activity and an accumulation of cell wall‐bound phenolics and lignin, as well as a methyl jasmonate‐mediated increase in lipoxygenase activity. The two treatments had an additive effect on other defensive traits and both treatments negatively affected looper performance. These results suggest that salicylic acid and jasmonic acid are not strictly antagonistic in conifers and that both have a role in anti‐folivore defence signalling. The present study illustrates the need for a better understanding of hormone signalling, cross‐talk and induced responses in conifers.

Spring bird migration as a dispersal mechanism for the hemlock woolly adelgid

In eastern North America, the invasive hemlock woolly adelgid (Adelges tsugae Annand), has expanded northward at a pace that exceeds predictions from mechanistic models, suggesting successful long-distance dispersal despite the only viable dispersive phase being a flightless nymph, or “crawler.” We hypothesize that migrating birds may contribute to long-distance dispersal of crawlers by passively transporting them in their plumage during northward migration. We collected hemlock woolly adelgid crawlers from the plumage of wild birds in Connecticut hemlock forests in spring and summer 2016–2017 and evaluated the factors that influence crawler loads on wild birds. Of 456 birds examined, 40 individuals of 22 species carried adelgid crawlers. Crawler loads varied strongly over time, showing a mid-spring peak that mirrored the phenological pattern in crawler abundance. However, crawler load was not affected by either local crawler abundance at capture sites or the degree of bird species association with hemlock forests. To test whether dispersed crawlers could start new invasions, we experimentally simulated avian-assisted dispersal of adelgids onto uninfested nursery hemlocks. Although rare, crawlers placed on birds did settle successfully on experimental branches during the adelgid’s summer generation. Our study confirms that birds carry hemlock woolly adelgid crawlers most often during the period of peak spring songbird migration, and that crawlers can move off bird plumage to settle on hemlock foliage. Bird-mediated, long-distance dispersal of crawlers likely has played a key role in hemlock woolly adelgid spread, and with warming temperatures, this mechanism may continue to be important for future range expansion.

Quantifying spatio-temporal variation of invasion spread.

- The spread of invasive species can have far-reaching environmental and ecological consequences. Understanding invasion spread patterns and the underlying process driving invasions are key to predicting and managing invasions. - We combine a set of statistical methods in a novel way to characterize local spread properties and demonstrate their application using simulated and historical data on invasive insects. Our method uses a Gaussian process fit to the surface of waiting times to invasion in order to characterize the vector field of spread. - Using this method, we estimate with statistical uncertainties the speed and direction of spread at each location. Simulations from a stratified diffusion model verify the accuracy of our method. - We show how we may link local rates of spread to environmental covariates for two case studies: the spread of the gypsy moth ( Lymantria dispar), and hemlock woolly adelgid ( Adelges tsugae) in North America. We provide an R-package that automates the calculations for any spatially referenced waiting time data.

Establishment and Distribution of Laricobius spp. (Coleoptera: Derodontidae), a Predator of Hemlock Woolly Adelgid, Within the Urban Environment in Two Localities in Southwest Virginia

Abstract Hemlock woolly adelgid (HWA), Adelges tsugae Annand, is a highly destructive non-native pest lethal to eastern hemlock, Tsuga canadensis (L.) Carrière, and Carolina hemlock, T. caroliniana Engelmann. Fifty years following the first observation of HWA in eastern North America, a predatory beetle, Laricobius nigrinus Fender, was evaluated, approved, and released as a biological control agent. Efforts were made to mass rear L. nigrinus with the goal of redistributing the beetles on public lands, typically in forest ecosystems, as a secondary option to silvicultural and chemical controls. The majority of L. nigrinus releases has occurred on public lands. Herein, we report the observation of L. nigrinus within the urban environment outside of known release locations. Two towns, Blacksburg and Radford, were divided into 0.40 km2 grids. A total of 27 and 19 grids were randomly selected from each town, respectively. Hemlocks were present in 44 and 42% of the grids surveyed in Blacksburg and Radford, respectively. In Blacksburg and Radford, 86 and 100% of the grids with hemlocks were infested with HWA, and of those infested hemlocks, Laricobius spp. was present in 100 and 75% of grids, respectively. A total of 154 Laricobius spp. (98% L. nigrinus and 2% Laricobius rubidus) adults were collected between each town. While it is unclear the level of control L. nigrinus has on reducing HWA's impact, the establishment of this biological control agent in the urban environment is an additional level of predation, that would otherwise not be present, for homeowners with HWA-infested trees.

Circumventing regulatory safeguards: Laricobius spp. and biocontrol of the hemlock woolly adelgid

Abstract Biological control entails introducing non‐native natural enemies to control weeds and pests. Instances of poorly regulated biocontrol have had harmful consequences leading to significant efforts to improve risk assessment of agents before release. Laricobius naganoensis (Coleoptera: Derodontidae) was inadvertently imported to eastern North America with Laricobius osakensis, introduced from Japan to control the hemlock woolly adelgid Adelges tsugae (Hemiptera: Adelgidae). The United States Department of Agriculture (U.S.D.A.) subsequently permitted introduction of L. naganoensis as a biocontrol agent and contaminant, requiring no study or monitoring owing to difficulty of the effort required to distinguish it from L. osakensis. The Plant Protection Act of 2000 (7 U.S.C. § § 7701–7786) assigns U.S.D.A. responsibility for vetting and reducing risk from biocontrol agents while ensuring the process is transparent, accessible, and based on scientific evidence. Typically, release of a new agent involves a 30‐day period when the public may review and comment on an assessment of risk associated with the introduction, an Environmental Assessment (EA). No public review and comment period was allowed before L. naganoensis’ introduction, and the EA was not made public. Biocontrol practitioners and stakeholders agree that critical scientific evaluation of multiple lines of evidence is necessary to evaluate risk associated with new introductions. The importance of accurate agent identification and avoidance of contamination has long been recognised, and procedures to minimise risk are well known. While such processes might be difficult to implement, those challenges should not justify a release that circumvents regulatory safeguards.

The Past, Present, and Future of the Hemlock Woolly Adelgid (Adelges tsugae) and Its Ecological Interactions with Eastern Hemlock (Tsuga canadensis) 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.

Modeling Benefit–cost Analysis of Hemlock Wooly Adelgid Host Plant Resistance with Emphasis on Least-toxic Chemical Treatment Alternatives

Chinese hemlock (Tsuga chinensis) exhibits a high level of resistance to the exotic insect hemlock woolly adelgid (HWA; Adelges tsugae) relative to the native and widely planted eastern hemlock (Tsuga canadensis). Furthermore, both chinese and eastern hemlock exhibit similar autecologic and aesthetic characteristics in urban and suburban environments. This study provides a comparative 25-year economic benefit-cost analysis (BCA), tracking estimated establishment and insect control costs for the two tree species. Eastern hemlock survival requires insecticide treatments when growing within the range of HWA. Insect control scenarios used and evaluated in this study include annual horticultural oil spray, biannual horticultural oil spray, biennial imidacloprid soil drench, and no treatment. The chinese hemlock scenario did not include chemical insect control because of the species’ host plant resistance (HPR) to HWA. Benefits were estimated using the U.S. Department of Agriculture, Forest Service’s i-Tree tool, which estimates economic benefits for ecosystem services (expressed in dollars). Benefit–cost ratios (BCRs) were developed using the present value for 25-year benefit and cost streams at 2% and 4% discount rates. Payback periods were also estimated for all options that had a calculated BCR greater than one. The benefit–cost analyses for each insect control scenario were evaluated, compared, and assessed through the lens of market potential. The costs exceed the benefits for all of the eastern hemlock scenarios. The benefits exceed costs for the chinese hemlock scenario. Results suggest that chinese hemlock is a viable alternative to eastern hemlock in view of its HPR and reduced associated costs over time. If chinese hemlock becomes more widely planted, it is expected to produce greater BCRs relative to chemical control options as a result of the lack of required, ongoing insect control treatment costs.

Media representation of hemlock woolly adelgid management risks: a case study of science communication and invasive species control

Public support can aid invasive species control and thus reduce negative impacts. Scientific communication can help or hinder efforts when associated risk and uncertainty are exaggerated or neglected, creating fear or distrust. Review of 104 media articles about managing a devastating forest insect, the hemlock woolly adelgid (Adelges tsugae), revealed that uncertainties and value judgments associated with claims of success were usually obscured or omitted, legitimate disagreement among experts about efficacy was omitted, and risks to non-targets were usually unaddressed. Discussion of chemical control in 83 articles included mention of non-target effects (e.g., to water quality, pollinators, and other native taxa) in only 11 articles and specificity was mentioned in only five instances. Biocontrol non-target impacts were mentioned in 11 of 83 cases and agent specificity was not mentioned in 71 of those cases. Ironically, while the value of the native ecosystem is used to justify management, possible non-target impacts of management on native ecosystems are largely ignored, notably hybridization between an introduced non-native biocontrol agent (Laricobius nigrinus) and a native beetle (L. rubidus) and ongoing releases of two non-native beetles, L. osaskensis colonies contaminated with a poorly understood, newly described species (L. naganoensis). To increase the public’s ability to make informed decisions, benefits of management techniques should be discussed along with risks, scientific disagreement, and uncertainty.

Phenology and Synchrony of Scymnus coniferarum (Coleoptera: Coccinellidae) with Multiple Adelgid Species in the Puget Sound, WA, USA

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.

Herbaceous-layer diversity and tree seedling recruitment are enhanced following Rhododendron maximum shrub removal

Forest ecosystems dominated by Tsuga canadensis are undergoing fundamental changes in function and composition from infestations by hemlock woolly adelgid (Adelges tsugae). We proposed that the first step to restoring southern Appalachian riparian forests following T. canadensis mortality would be eliminating the evergreen shrub, Rhododendron maximum. We hypothesized that removing R. maximum would increase light transmittance, soil moisture and temperature; and subsequently, enhance herbaceous-layer diversity and promote tree seedling recruitment and survival. We tested these hypotheses at two locations, (CWT, Coweeta Hydrologic Laboratory; WOC, White Oak Creek) in the Nantahala Mountain Range of western North Carolina, both with heavy T. canadensis mortality and a dense R. maximum subcanopy. The treatments were designed to remove only soil O-horizon (FF), remove only R. maximum (CR), remove R. maximum and soil O-horizon (CFFR), and untreated, reference (REF). We installed permanent plots across treatments and locations and measured light transmittance (Qi/Qo), soil water content (θ), herbaceous-layer cover and diversity (Shannon’s index (H′cover) and species richness), and tree seedling recruitment.As expected, cutting the R. maximum subcanopy (CR and CFFR) immediately increased Qi/Qo in the spring months across locations, and it was sustained through the first growing season. θ was generally high across plots, averaging 26% during the growing season, and didn’t vary over time. By the second growing season (2017) after treatments, herbaceous-layer cover and diversity increased on CR and CFFR. Herbaceous-layer cover was significantly related to Qi/Qo (r2 = 0.22, p < 0.001) and θ (r2 = 0.13, p = 0.009), while diversity was only related to Qi/Qo (H′cover, r2 = 0.14, p < 0.001; species richness, r2 = 0.21, p < 0.001). Tree seedling density was related to Qi/Qo (r2 = 0.10, p = 0.001) and θ (r2 = 0.26, p < 0.001). Tree seedling density was low before treatment (1.4 ± 0.3 seedlings m−2) and increased by 10-fold in CR and CFFR two growing seasons after treatment. In CR, species with the highest density ranked Betula spp. > Acer rubrum > Quercus coccinea > Liriodendron tulipifera > Q. rubra. In CFFR, tree seedling recruitment ranked Betula spp. > A. rubrum > L. tulipifera. These vegetation responses have important implications for potential recovery of riparian forests following T. canadensis mortality.

First Report on Establishment of Laricobius osakensis (Coleoptera: Derodontidae), a Biological Control Agent for Hemlock Woolly Adelgid, Adelges tsugae (Hemiptera: Adelgidae), in the Eastern U.S.

The hemlock woolly adelgid (HWA) is an invasive insect species native to Japan causing significant hemlock mortality in the eastern United States. Laricobius spp. have been targeted as biological control agents because they are adelgid specialists. Laricobius osakensis Montgomery and Shiyake is native to the same region of Japan from which the strain of HWA found in the eastern United States originated. Studies in Japan found that it is phenologically synchronous with HWA. Following approval to release L. osakensis from quarantine in 2010, approximately 32,000 were released at a total of 61 sites starting in 2012. In winter of 2014 and 2015, periods of extreme cold temperatures throughout the eastern USA, as well as the polar vortex, resulted in extensive mortality to HWA, which likely delayed the establishment of L. osakensis. The ability of the beetle to survive and establish in the eastern United States is reported here. In the first year of this study (2015–2016), limited numbers of L. osakensis were recovered, as HWA populations were still rebounding. In the second year (2016–2017), 147 L. osakensis were collected at 5 of 9 sites sampled, coinciding with rebounding HWA populations. Larval recovery was much greater than adult recovery throughout the study. HWA density was directly correlated with warmer plant hardiness zones and recovery of Laricobius beetles was significantly correlated with HWA density. Our results suggest that L. osakensis is successfully establishing at several of the sampled release sites and that the best predictor of its presence at a site is the HWA density.

Shifts in vegetation and avian community structure following the decline of a foundational forest species, the eastern hemlock

Eastern hemlock (Tsuga canadensis) is undergoing widespread decline throughout the eastern United States due to the introduction of the nonnative hemlock woolly adelgid (Adelges tsugae). In light of hemlock's unique significance for avian communities, we examined the long-term response of the breeding bird community to the decline and die-off of this foundational forest species. We conducted variable-radius point counts to survey bird communities in both hemlock and hardwood stands in 2000, during the early stages of adelgid infestation, and again in 2015 and 2016, following significant hemlock decline. We also measured the severity of hemlock decline and associated vegetation variables in the same hemlock stands in which avian surveys were completed. We used multispecies occupancy models to examine species-specific and species group responses to hemlock decline. Results showed that, across hemlock stands, hemlock basal area declined from a mean of 56% of the total basal area to 46%, and that hemlock decline was correlated with current vegetation structure, including an increased deciduous understory. Species richness of hemlock-associated birds declined by an average of ∼1 species per survey location. The Black-throated Green Warbler (Setophaga virens) was the most strongly affected, declining in occupancy by 30%. All other species groups responded positively, with the strongest responses by species associated with the shrub layer, forest edge, and mature deciduous habitat. The species composition in hemlock and hardwood stands became more similar over time as the unique species assemblages in hemlock stands gave way to the avian community of the surrounding hardwood landscape, highlighting a trend toward homogenization of the avian community. Where hemlock-associated bird species persisted, their presence was correlated with a combined effect of greater hemlock basal area and better hemlock condition. Our results demonstrate that the decline of hemlocks has restructured vegetation communities and their associated avian communities over time, with strong implications for forest-specialist birds.

Decline of a foundation tree species due to invasive insects will trigger net release of soil organic carbon

AbstractEastern U.S. forests are witnessing an ecologically disruptive decline in one of the region's distinctive foundation tree species, the eastern hemlock (Tsuga canadenis). The exotic insect pests hemlock woolly adelgid (Adelges tsugae, HWA) and elongate hemlock scale (Fiorinia externa) have greatly altered many forest ecosystems previously dominated by this important evergreen conifer. The consequences for ecosystem processes are far reaching because hemlock is often replaced by deciduous tree species, such as black birch (Betula lenta), which have strongly divergent effects on forest floor microenvironments and nutrient cycling. We took advantage of an accidental experiment initiated by patch‐level timber harvesting ~30 yr ago to investigate how the removal of hemlock, and its replacement by deciduous trees, has affected leaf litter characteristics, soil organic layer mass, C:N content, and soil respiration rates. We also contrasted these areas to nearby forest plots where deciduous B. lenta has been dominant for almost a century. The inclusion of healthy, intact hemlock stands in the design, and the close proximity of plots, allowed for a powerful space‐for‐time approach to detect ecosystem changes that are likely to occur across the broader landscape with HWA‐induced hemlock loss in coming years. Three years of data collection from a series of plots in hemlock, young birch, and mature birch stands revealed dramatic differences in soil carbon pools and cycling. Between forests dominated by hemlock vs. mature birch, we saw a significant decrease in soil organic layer mass and in the C:N of the remaining organic material. Although hemlock and young birch stands showed no significant differences in soil respiration rates, mature birch stands had significantly higher soil respiration rates throughout the entire growing season, regardless of wet or dry years. Our results suggest that the carbon pool in the forest floor is likely to mobilize through greater decomposition with a 6.8× decline in soil organic layer C storage as hemlocks are replaced by deciduous trees, leading to a potential net release of ~4.5 tons C per hectare. We conclude that the ramifications of this change for carbon storage could be extensive, but may take decades to manifest.

Impact of Sirococcus shoot blight (Sirococcus tsugae) and other damaging agents on eastern hemlock (Tsuga canadensis) regeneration in Northeastern USA

In 2009, Sirococcus tsugae was first reported in Maine on eastern hemlock. Our goal was to quantify the impact of the shoot blight disease caused by this fungal pathogen of unknown origin on eastern hemlock regeneration. From 2013 to 2014, 59 long-term monitoring plots established by the US Forest Service (USFS) Forest Inventory and Analysis (FIA) program in New England and New York were surveyed to determine the impact of S. tsugae. Damage by hemlock woolly adelgid (Adelges tsugae), elongate hemlock scale (Fiorinia externa), white-tailed deer (Odocoileus virginianus), or other causes was also recorded. Disease incidence and severity (percentage of shoots blighted and percentage of crown defoliated) were assessed for 20 seedlings per plot. Sirococcus shoot blight symptoms were present in most plots (90%) and on most seedlings (72%). For the majority of seedlings, blight affected less than 10% of shoots, but the percentage of shoots blighted did range up to 75%. Similarly, needle loss was limited to less than 25% of the crown for most seedlings. Disease severity was positively correlated with overstory hemlock density. Using species-specific polymerase chain reaction (PCR) primers, Sirococcus tsugae was identified from samples collected in the majority of sites (68%) in New England and New York. In permanent plots at the Massabesic Experimental Forest in Maine, disease symptom severity increased from 16% blighted shoots in 2011 to 47% blighted shoots in 2013. Results confirm that Sirococcus shoot blight of eastern hemlock is more widespread in natural forests of northeastern USA than previously known and that symptoms can be severe (>75% blighted shoots) in some locations.