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The hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae), is an invasive pest causing significant ecological and economic damage to certain hemlock tree (Tsuga (Endlicher) Carrière, Pinales:Pinaceae) species. In response to this invasive threat, biological control strategies have been implemented, introducing natural predators such as Laricobius nigrinus Fender (Coleoptera: Derodontidae) and, more recently, Laricobius osakensis Montgomery and Shiyake (Coleoptera: Derodontidae), as specialist predators against A. tsugae. However, the genetic and morphological similarities between L. osakensis and both L. nigrinus and the native beetle, Laricobius rubidus LeConte (Coleoptera: Derodontidae), pose challenges in their identification. Effective monitoring of released predators is integral to evaluating the success of biological control measures. Environmental DNA (eDNA) holds potential for various detection applications, including species monitoring. In this study, we developed specific primers and probes targeting the mitochondrial cytochrome oxidase 1 gene sequences, achieving high specificity despite their 95% sequence similarity. With an optimal annealing temperature of 60 °C, our tools effectively differentiated L. osakensis from the other 2 beetles and demonstrated eDNA detection sensitivity down to 2 copies/µl. This research underscores the potential of precise molecular tools for advancing biological control and biodiversity assessment against invasive threats like A. tsugae. [ABSTRACT FROM AUTHOR]

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In hemlock stands within eastern US forests, classical biological control has been one of the main strategies used to manage the hemlock woolly adelgid, Adelges tsugae Annand. Specialist predator species may offer a management solution to help regulate A. tsugae populations. In the Pacific Northwest, a suite of specialist predators has been a focus of research and includes 2 species of silver fly, Leucotaraxis argenticollis (Zetterstedt) and Leucotaraxis piniperda (Malloch) (Diptera: Chamaemyiidae). Leucotaraxis spp. phenology has been documented in the Pacific Northwest, but the phenology of either western Leucotaraxis species is unknown in the eastern United States.This study sought to document the phenology of Le. argenticollis in NY in 2021 and in VA in 2021 and 2022. Nylon mesh cages were applied over eastern hemlock branches infested with A. tsugae to contain Le. argenticollis adults. Biweekly and monthly branch samples were taken in 2021 and 2022, documenting all life stages of A. tsugae and of Le. argenticollis that were observed. In 2021 and 2022, Le. argenticollis adults and eggs were present during the oviposition stage of the 2 generations of A. tsugae. In addition, Le. argenticollis larvae were present when A. tsugae ovisacs had eggs and while A. tsugae nymphs of both generations were present.These observations indicate that Le. argenticollis phenology is well synchronized with A. tsugae in the eastern United States. [ABSTRACT FROM AUTHOR]

Pre-emptive silvicultural treatments can enhance host-tree vigor as a component of an integrated management approach to mitigate invasive species' impacts. Hemlock woolly adelgid (HWA, Adelges tsugae Annand, 1928) threatens the health and existence of eastern hemlock-dominated forests. We propose that increasing canopy light exposure via silvicultural thinning can reduce hemlock's vulnerability to HWA. Three stands with high hemlock densities on the Allegheny National Forest were selected for thinning. There was no HWA infestation and half of each stand was thinned. Circular plots were established surrounding 96 hemlock "subject trees" targeted for crown release on 3–4 sides. Similar criteria were used to select 90 hemlocks in untreated plots. We visited subject trees pre-thinning and 5 and 10 years post thinning and measured the following: diameter, total height, live crown ratio, and crown area. Temporal changes in diameter were not significant between treatments, but trees in thinned plots averaged 5 cm larger after 10 years. Live crown ratio was maintained in thinned stands and decreased after year 5 in controls. Crown area increased in thinned plots and decreased in controls. Residual basal area significantly influenced growth but varied between treatments and among stands. [ABSTRACT FROM AUTHOR]

Hemlock wooly adelgid (Adelgis tsugae Annand) (HWA) has invaded much of eastern hemlock's (Tsugae canadensis L. Carrière) native range. Arborists and forest managers have successfully handled this pest using either contact or systemic pesticides. One of these pesticides, imidacloprid, has often been applied using a soil or trunk injection. Although imidacloprid has been labeled as a basal bark spray to control HWA, minimal information regarding its efficacy is available. This study compared bark treatments to soil treatment of imidacloprid at high and low application rates, as well as the use of a bark adjuvant. The results showed that basal bark treatments were as effective as soil treatments. Hence, basal bark treatments of imidacloprid can be an effective method for control of HWAs in eastern hemlock trees. A bark adjuvant may not be necessary, as it was not found to influence the amount of imidacloprid in the tissues. [ABSTRACT FROM AUTHOR]

Copyright of Forestry Chronicle is the property of Canadian Institute of Forestry and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

Sympatric host-associated genetic differentiation is a prominent pattern that could lead to speciation. In insects, there are numerous examples of host-associated differentiation among herbivores that prefer different plants, and parasitoids that prefer different hosts, but few examples for specialist predators. We developed new microsatellite loci for two species of silver fly, Leucotaraxis argenticollis (Zetterstedt) and L. piniperda (Malloch) (Diptera: Chamaemyiidae), being evaluated as biological control agents for the hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae), in eastern North America where it is a nonnative pest. We obtained DNA from specimens of both fly species feeding on native A. tsugae in western North America, as well as on other western and eastern adelgid species. We performed population genetic analyses using the new loci and DNA barcode sequences. Our results confirmed east–west allopatric divergence and uncovered nested genetic differentiation associated with different adelgid prey species and their host plants in western North America for both species of silver flies. For both species, there is also evidence for a longer history of diversification in the west, with ancestral specialization of feeding on pine adelgids, which was retained after range expansion to the east. More recently, divergence to feeding on new adelgid prey species occurred in the west. Our findings are consistent with the hypothesis that host-alternating life cycles in Adelgidae may provide temporary escape from specialist predators. We discuss the implications for biological control efficacy and potential for lineage hybridization as western flies are released in the east to control A. tsugae. [ABSTRACT FROM AUTHOR]

Abstract Environmental DNA (eDNA) analysis can be a powerful tool for the early detection of invasive organisms. However, research on terrestrial eDNA detection from foliage surfaces has been limited. In this study, we developed methods to capture and detect eDNA using qPCR from an invasive forest pest, hemlock woolly adelgid (Adelges tsugae), and three of its biological control predators Leucotaraxis piniperda, Leucotaraxis argenticollis, and Laricobius nigrinus. We designed four highly efficient qPCR assays with a low limit of detection (1–10 copies/reaction). The assay targeting A. tsugae was species‐specific. The assays targeting Le. piniperda, and Le. argenticollis were biotype‐specific in addition to being species‐specific demonstrating applications of eDNA analysis beyond species‐level detection. The La. nigrinus assay also detected DNA from closely related and hybridizing Laricobius rubidus. The eDNA methods were evaluated against traditional detection methods. We collected foliage samples from three strata (bottom, middle, and top) of eastern hemlock trees to detect the presence of A. tsugae. The detection of the biological control predators was evaluated using western hemlock foliage samples collected from the predators' native range in western Washington. The eDNA methods had significantly higher positive detection rates (2.8–4.5 times) than conventional methods of all target species. The strata of sampling were not significant in determining the presence of A. tsugae infestation. The eDNA concentration positively correlated with the observed density for all species. This study demonstrates the efficacy of eDNA analysis as a more sensitive tool for early detection of A. tsugae and to track the establishment of its biological control predators.

Hemlock woolly adelgid (HWA), Adelges tsugae Annand, threatens hemlock forests throughout eastern North America. Management efforts focus on early detection of HWA to ensure rapid management responses to control and stop the spread of this pest. This study's goal was to identify an affordable, efficient trap to aid with airborne environmental DNA (eDNA) sampling approaches as an early monitoring tool for HWA. We initially compared HWA detection success between a standard sticky trap, commonly used for HWA monitoring, and trap designs potentially compatible with eDNA protocols (i.e. passive trap, funnel trap, and motorized trap). Passive, funnel, and motorized traps' estimated capture success probabilities compared to sticky traps were 0.87, 0.8, and 0.4, respectively. A secondary evaluation of a modified version of the motorized trap further assessed trap performance and determined the number of traps needed in a set area to efficiently detect HWA. By modifying the original motorized trap design, its estimated capture success probability increased to 0.67 compared to a sticky trap. Overall, the cumulative capture success over the 16-week sampling period for the motorized trap was 94% and 99% for the sticky trap. The number of traps did impact capture success, and trap elevation and distance to infested hemlocks influenced the number of adelgids captured per trap. As eDNA-based monitoring approaches continue to become incorporated into invasive species surveying, further refinement with these types of traps can be useful as an additional tool in the manager's toolbox. [ABSTRACT FROM AUTHOR]

Hemlock woolly adelgid (Adelges tsugae) is the most important pest of hemlocks in the eastern United States, where it completes three generations a year. We investigated the impact of temperature (8, 12, 16, and 20°C) on the estivation and postaestivation stages of the A. tsugae sistens generation. Temperature significantly impacted development and survival of this generation. The highest mortality occurred at the coolest temperature (8°C). Adelges tsugae developed rapidly as the temperature increased and optimum temperatures for development ranged between 17 and 22°C for the different instars. The estimated lower temperature threshold was 0°C for second instar nymphs and 3 –5°C for the other instars and the preoviposition period. Estivating first-instar sistentes resumed development (as evidenced by segments becoming visible) after 40–100 d at the constant temperatures (fastest at 16°C) then required only 105 degree-days (DD) for 50% of the individuals to molt. Subsequent instars developed rapidly (another 470 DD total to reach adult), and oviposition began at ∼623 DD from the time the first instars resumed development. This study provides valuable data required to develop an annual phenology model for A. tsugae which will assist in timing monitoring and control treatments. Graphical Abstract [ABSTRACT FROM AUTHOR]

Overstory basal area, ericaceous shrub cover (Kalmia latifolia L. and Rhododendron maximum L.), and fuels (i.e., woody fuel loads and depths and O Horizon thickness) were assessed within Great Smoky Mountains National Park, USA, in 2003 − 2004. Due to recent wildfire activity within the southern Appalachian Mountain region (including Great Smoky Mountains National Park), the potential spread and expansion of ericaceous shrubs, and the impacts of the hemlock woolly adelgid (Adelges tsugae Annand) on eastern hemlock (Tsuga canadensis (L.) Carrière), these same ecosystem components were again assessed in 2019. Elevation and moisture regime (xeric, intermediate, and mesic) were included in this assessment as potential influential factors. An evaluation of repeated measurements from 40 plots suggested that O Horizon thickness did not change significantly over the 16-year period, but increased as elevation increased, and moisture regime (xeric O Horizon thickness > mesic O Horizon thickness) was a significant, related factor. The sum of 1-, 10-, and 100-h fuel loads (fuels less < 7.6 cm diameter) increased, whereas woody fuel depth decreased over the 16-year period. No significant changes in 1000-h fuel loads (> 7.6 cm diameter), total woody fuel loads, ericaceous shrub cover, total basal area, or live T. canadensis basal area were observed. Live T. canadensis basal area decreased with increasing elevation. Dead, standing T. canadensis basal area increased from 2003–2019, and that increase was most pronounced as elevation increased on xeric and intermediate sites. Overall, we found that: 1. hypothesized increases in total woody fuel loads and ericaceous shrub cover were not present; and 2. elevation and moisture regime were most related to observed changes in vegetation and fuel condition. [ABSTRACT FROM AUTHOR]

Winter‐season extreme minimum temperatures may play a major role in limiting population growth and spread of the hemlock woolly adelgid (HWA) (Adelges tsugae Annand) (Hemiptera: Adelgidae), an invasive sap‐feeding insect that has caused extensive mortality of hemlock trees (Tsuga spp.) in many eastern United States (US) forests. This atypical insect feeds throughout the winter but populations can sustain high mortality when winter‐season extreme minimum temperatures drop below −20 to −30°C. Detection of HWA in Michigan during 2015 motivated interest in HWA winter survival in the US Great Lakes region. Here, we used the parameter‐elevation regressions on independent slopes model (PRISM) gridded daily minimum temperature dataset to construct a 1981–2018 climatology of extreme minimum temperatures in the Great Lakes region, the first such effort for this region. Metrics examined include absolute and mean annual extreme minimum temperatures (defined as the lowest daily minimum temperature during the study period and the study‐period mean lowest daily minimum temperature during each calendar year, respectively), and the frequency of daily minimum temperatures below −20 and −30°C. Minimum temperature patterns we identified support the following two hypotheses: first, proximity to water, surface elevation and latitude are the principal controls for extreme minimum temperatures in the Great Lakes region; second, the modifying influence of the relatively warm Lake Michigan serves to protect locations within about 10–25 km of the lakeshore from severe and potentially lethal temperatures for HWA. Analysis of projected minimum temperatures at the end of the 21st century (2080–2099) reveals a range of HWA distribution expansion scenarios. Although the original motivation for this study arose from interest in potential HWA mortality, a climatological study of extreme minimum temperatures has potentially broad relevance to, for example, human health and safety and forest ecology. [ABSTRACT FROM AUTHOR]

Hemlock woolly adelgid (HWA; Adelges tsugae Annand (Hemiptera: Adelgidae)) is the cause of widespread mortality of Carolina and eastern hemlock (Tsuga caroliniana Engelmann and T. canadensis (L.) Carrière) throughout the eastern United States (U.S.). Since its arrival in the northeastern U.S. HWA has steadily invaded and established throughout eastern hemlock stands. However, in 2018, anecdotal evidence suggested a sharp, widespread HWA decline in the northeastern U.S. following above-average summer and autumn rainfall. To quantify this decline in HWA density and investigate its cause, we surveyed HWA density in hemlock stands from northern Massachusetts to southern Connecticut and analyzed HWA density and summer mortality in Pennsylvania. As native fungal entomopathogens are known to infect HWA in the northeastern U.S. and rainfall facilitates propagation and spread of fungi, we hypothesized high rainfall facilitates fungal infection of aestivating nymphs, leading to a decline in HWA density. We tested this hypothesis by applying a rain-simulation treatment to hemlock branches with existing HWA infestations in western MA. Our results indicate a regional-scale decline and subsequent rebound in HWA density that correlates with 2018 rainfall at each site. Experimental rain treatments resulted in higher proportions of aestivating nymphs with signs of mortality compared to controls. In conjunction with no evidence of increased mortality from extreme winter or summer temperatures, our results demonstrate an indirect relationship between high rainfall and regional HWA decline. This knowledge may lead to better prediction of HWA population dynamics. [ABSTRACT FROM AUTHOR]

Efficient separation of insects from plant material for quantification and collection is an important component of entomological research. This paper reports on a novel, easily replicable container designed to efficiently collect two different biological control agents dispersing from hemlock (Tsuga spp.) foliage infested with the invasive hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae). The container utilizes a simplified Berlese-style funnel design to collect Laricobius spp. (Coleoptera: Derodontidae) larvae dropping from the foliage into a removable bottom jar, a central jar to house the foliage sample, and a removable top jar to collect adult silver flies (Leucopis spp. Diptera: Chamaemyiidae) emerging from puparia on the twigs. The efficacy of two designs (with and without a funnel leading to the top collection jar) was evaluated using western hemlock [ Tsuga heterophylla (Raf.) Sarg.] foliage naturally colonized with HWA and the two predator genera. All Laricobius larvae were effectively collected in the bottom jar, and the addition of an inverted funnel leading to the top collection jar increased the proportion of Leucopis flies reaching the target jar from 60% to 94%. This 'Lari-Leuco' container is presented as a research and motoring tool to benefit the integrated pest management program for HWA in eastern North America and for potential use in simultaneously separating ascending and descending life stages in other insect-plant or predator-prey systems. [ABSTRACT FROM AUTHOR]

Imidacloprid, a neonicotinoid pesticide, is used to prevent the spread of the hemlock woolly adelgid, currently affecting Eastern Hemlock trees across North America. When the pesticide is sprayed directly onto soil around infested trees (soil drenching), it can run off into aquatic systems, with potential negative effects on biota. Simultaneously, climate change may lead to faster pool drying, which acts as an additional stressor for sensitive species such as amphibians. We evaluated the sublethal effects of imidacloprid (10 ppb), and interaction with shorter hydroperiods on the larval behavior, growth, and survival of a model organism, the wood frog (Rana sylvatica). We performed 3 behavioral experiments evaluating swimming speed, time spent swimming, and distance the larvae swam. We found that larvae raised in 10 ppb imidacloprid or shorter hydroperiod did not differ in their swimming time, distance, and speed from nonexposed larvae. Naïve larvae exposed for 20 min to 10‐ to 500‐ppb concentrations also showed similar performance to nonexposed larvae. However, when we applied a stimulus halfway through each experiment, we found that larvae exposed to 10 ppb imidacloprid (short and long term) swam shorter distances and spent less time swimming, suggesting that imidacloprid exposure may slow reaction time, potentially increasing the risk of predation. To minimize impacts on pool‐breeding amphibians, imidacloprid application to combat the invasive hemlock woolly adelgid should use trunk injection and avoid soil drenching. Environ Toxicol Chem 2021;40:1840–1849. © 2021 SETAC [ABSTRACT FROM AUTHOR]

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.

Landscape-scale maps of tree species densities are important tools for managing ecosystems threatened by forest pests. Eastern hemlock dominates riparian forests throughout its range. As a conifer in a deciduous landscape, hemlock plays an ecohydrological role, especially when other species are dormant. The nonnative, hemlock woolly adelgid has caused widespread hemlock decline and mortality. We used two existing basal area raster layers first to identify Chesapeake Bay subwatersheds with ≥6 percent hemlock basal area and second to quantify hemlock basal area densities within fixed-width riparian buffers of 50 m, 100 m, 250 m, and 500 m. Hemlock densities were higher in riparian zones compared with entire subwatersheds. In five subwatersheds, 50 m and 100 m zones had higher percentages of pixels with ≥25 percent hemlock basal area. We produced maps identifying hemlock riparian densities in the Pine Creek Watershed, which managers can use to prioritize sites for supplemental conifer planting under anticipated hemlock decline. [ABSTRACT FROM AUTHOR]

Eastern hemlock (Tsuga canadensis [L.] Carrière) is an important component of the riparian ecosystem. Due to the widespread establishment of hemlock woolly adelgid (Adelges tsugae Annand)(HWA) across the range of eastern hemlock, woodland trees may be infested for extended periods (years), resulting in their decline. Imidacloprid, a systemic neonicotinoid insecticide, may be used as a strategy in forested settings to manage HWA while more long-term solutions become established, such as biological controls. Symptoms of prolonged infestation include extensive dieback and thinned canopies. In this study, trees with a diameter at breast height (DBH) of 24.7 ± 2.7 SD cm in poor condition were treated with imidacloprid. Trees were treated once by trunk-injection (IMA-jet) or by soil drench in the Greenbrier area of the Great Smoky Mountains National Park, Gatlinburg, TN, USA. Changes in tree growth and HWA density were measured for 3 consecutive years. Imidaclopridtreated trees recovered, whereas the untreated trees declined. Imidacloprid treatments resulted in significantly higher 3-year mean percent growth (65.6% to 71.7% of tips) compared to the untreated controls (10.5% of tips). HWA density 3-year means in the imidacloprid-treated trees (0.10 to 1.09 per cm) likewise were statistically different to the untreated trees (2.72 per cm). The extended activity of imidacloprid-treated hemlock was attributed to storage in the symplast (xylem ray parenchyma) and to perennial needle retention. This study demonstrates that trunk-injection with IMA-jet is effective against HWA and comparable with soil drench to protect trees in the long term (≥ 4 years). [ABSTRACT FROM AUTHOR]

Neonicotinoids are one of the most widely used classes of insecticides in the world. The neonicotinoid imidacloprid is commonly applied to hemlock (Tsuga spp.) stands in eastern North America to reduce tree mortality from infestations of the invasive hemlock woolly adelgid (HWA; Adelges tsugae). While laboratory and mesocosm studies have determined that imidacloprid can bioaccumulate in anurans and cause sublethal effects, no field studies have investigated whether salamanders or insects in streams adjacent to HWA treatments bioaccumulate imidacloprid or if sublethal effects are detectable in wild salamanders. We assessed relationships between imidacloprid exposure and stream salamander health in West Virginia, USA, using concentration of the stress hormone corticosterone and body condition indices (BCI) as response variables. Of 107 Desmognathus salamanders from 11 sites tested for bioaccumulation, we detected imidacloprid in 47 salamanders. Of 15 benthic macroinvertebrate samples tested, we detected imidacloprid, imidacloprid-urea, and imidacloprid-olefin in 15, 13, and 1 sample, respectively. Based on 115 Desmognathus salamanders sampled at 11 sites for stress hormone responses, corticosterone concentration increased with imidacloprid concentration in stream water. For 802 salamanders sampled at 48 sites, BCI decreased as concentration of imidacloprid in stream water increased, but explanatory power was low. Our study suggests that chronic leaching of imidacloprid from treated hemlock stands into adjacent streams has the potential to negatively affect aquatic organisms and may provide a route of exposure to higher trophic levels.

Ecohydrological processes occurring at or near the Earth's surface are strongly influenced by Eastern hemlock [EH; Tsuga canadensis (L.) Carrière], a foundation tree species of eastern North American forests. EH populations are currently threatened by the invasive hemlock woolly adelgid (HWA; Adelges tsugae Annand). HWA HWA populations have been expanding rapidly throughout the EH's range. Catchment‐scale research examining the hydrological consequences of HWA infestation is lacking, and plot‐scale studies remain conflicted in their findings. Given the complex relationships between canopy interception, unsaturated and saturated groundwater storage, and root water uptake, it is not immediately clear how EH loss will affect the hydrologic cycle. We investigated the impact of EH mortality on stream discharge characteristics across a regional sample of catchments utilizing both simulation and statistical modelling approaches. We first examined the relationship between various catchment characteristics, including EH health, and three hydrological variables through regression analysis. We then employed a non‐parametric statistical test to evaluate differences in hydrologic regime trends between non‐infested and infested catchments. Finally, we calibrated a physically based hydrologic model and considered differences in optimal model parameter values and simulated overland runoff between non‐infested and infested catchments. HWA presence modified several ecohydrological characteristics and precipitation partitioning between groundwater flows and surface runoff, potentially driving higher stream flashiness and overland flow, lower baseflow contributions and catchment storage, shorter flow‐path lengths, and variable source area dilation at infested sites. Our results suggest that EH decline will augment flooding potential associated with the increasing frequency and intensity of Atlantic Basin tropical cyclone events. Further, our physically based simulation provides more determinate results than regression analysis, indicating that a purely statistical methodology, commonly utilized in studying the relationship between landcover characteristics and hydrologic regime, neglects dynamic physical ecohydrologic relationships. [ABSTRACT FROM AUTHOR]

We developed an approach using sticky trap arrays as an early detection tool for populations of first-instar nymphs of the hemlock woolly adelgid (Adelges tsugae Annand), a pest of hemlocks (Tsuga spp. [Pinaceae]) in North America. We considered the detection rate of at least one nymph from trapping arrays consisting of one to six sticky panels, where we varied both the surface area of each trap that we assessed and the length of the trapping duration. We also estimated the time needed to set up, service, and assess groups of traps and attempted to relate capture of nymphs on traps to incidence and abundance of A. tsugae in the canopy above the traps. Arrays consisting of two traps provided a detection rate of 75% when 87.5% of the surface area of each trap was assessed, a process that required 38 min per array. The probability of detecting nymphs on traps left in the field for 5–6 d was similar to that for traps left for 12 d. The number of nymphs trapped in an array predicted the probability of finding A. tsugae in the canopy but only when all six traps were fully assessed. To reliably detect incipient A. tsugae infestations, we recommend placing arrays of traps at 1 km intervals along the perimeter of a stand during peak activity of first-instar sistentes nymphs and servicing these arrays every 5–7 d. [ABSTRACT FROM AUTHOR]

Invasive species are increasingly present in our ecosystems and pose a threat to the health of forest ecosystems. Practitioners are tasked with locating these invasive species and finding ways to mitigate their spread and impacts, often through costly field surveys. Meanwhile, researchers are developing remote sensing products to detect the changes in vegetation health and structure that are caused by invasive species, which could aid in early detection and monitoring efforts. Although both groups are working towards similar goals and field data are essential for validating RS products, these groups often work independently. In this paper, we, a group of researchers and practitioners, discuss the challenges to bridging the gap between researchers and practitioners and summarize the literature on this topic. We also draw from our experiences collaborating with each other to advance detection, monitoring, and management of the Hemlock Woolly Adelgid (Adelges tsugae; HWA), an invasive forest pest in the eastern U.S. We conclude by (1) highlighting the synergies and symbiotic mutualism of researcher–practitioner collaborations and (2) providing a framework for facilitating researcher–practitioner collaborations that advance fundamental science while maximizing the capacity of RS technologies in monitoring and management of complex drivers of forest health decline such as invasive species.

• Hemlock is a foundation species that supports distinct forest ecosystems. • The invasive hemlock woolly adelgid (HWA) has decimated hemlock in the eastern US. • HWA continues to spread north and has begun causing hemlock mortality in Nova Scotia. • We provide a decision framework to inform HWA management plans in eastern Canada. • Current lack of HWA control highlights the importance of an ecosystem restoration plan. The invasive hemlock woolly adelgid (HWA) has decimated hemlock stands across much of the eastern United States, and presents a significant threat to all eastern hemlock in Canada across Ontario, Quebec, New Brunswick, Nova Scotia, and Prince Edward Island, especially since the recent detection of its widespread establishment in southwest Nova Scotia. The spread and rising infestation level and impacts of HWA in this region serve as a warning for forest managers across eastern Canada to develop appropriate management plans and priorities. The HWA decision framework presented here aims to prepare forest managers in eastern Canada for the decisions and challenges that they will face, from prevention, detection, and control, to hemlock ecosystem restoration and management program evaluation. We review the strategies and tactics that are currently available, that are being developed, and that show the most promise to date. Given the nature of HWA, the long-term outlook for eastern hemlock in Canada will likely feature HWA as a component of hemlock ecosystems across much of the region, necessitating a comprehensive, adaptive management program to mitigate its ecosystem consequences. [ABSTRACT FROM AUTHOR]

The article discusses the article ‘‘Media representation of hemlock woolly adelgid management risks: a case study of sciencecommunication and invasive species control,'' by Leppanen et al. in the September 18, 2018 issue and mentions limitations to media reporting of science. It also mentions the comments made by Salom et al. on this article in terms of scientific communications, methods used in the study, and biological control of insects.

Highlights • Following Tsuga canadensis mortality, increase in light transmission was transient. • Basal area of co-occurring trees increased only marginally. • Increased density and size of Rhododendron maximum. • No recruitment of sapling to overstory, despite initial increased seedling density. • Light and seedling responses may be linked to increased abundance of R. maximum. Abstract 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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Bacterial endosymbionts of sap‐sucking insects provide their host with a number of beneficial qualities, including the supply of nutrition, defense against parasitoids, and protection from heat stress. Damage to these bacterial associates can therefore have a negative impact on the fitness of their insect host. We evaluated observational and experimental factors regarding the nonnative hemlock woolly adelgid (Adelges tsugae Annand) (Hemiptera: Adelgidae) to help understand the roles of its three recently identified symbionts, including under heat stress conditions. The prevalence of A. tsugae's facultative symbiont (Serratia symbiotica) was examined at different spatial scales to determine how variable infection rates are for this symbiont. There was no significant difference found in infection rates between adelgids on a tree, within a plot, or within a state. However, significantly more adelgids in Georgia (95%) had S. symbiotica compared to those in New York (68%). Microsatellite genotyping of the adelgids found that this difference was most likely not the result of a second introduction of A. tsugae into eastern North America. Comparison of S. symbiotica proportions between first and fourth instars showed that symbiont absence did not affect the ability of A. tsugae to survive aestivation. Evaluations of symbiont densities within each adelgid found that when S. symbiotica was absent, the density of obligate symbionts was significantly higher. Exposure to heat stress (32.5 °C) was not consistently correlated with changes in symbiont densities over a 4‐d period. Overall, we have shown that symbiont prevalence and densities vary within the broad population of A. tsugae in eastern North America, with potentially significant effects upon the ecology of this important pest. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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 novelway 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. [ABSTRACT FROM AUTHOR]

• Sustainability of eastern hemlock is threatened by the invasive insect Adelges tsugae. • Canopy gaps improved crown health and radial growth of infested understory hemlocks. • Tree competition index and pest density were predictive of crown decline post-release. • Results suggest silvicultural release can complement biological control of A. tsugae. The hemlock woolly adelgid (HWA, Adelges tsugae Annand) is an invasive forest insect threatening the sustainability of eastern hemlock (Tsuga canadensis (L.) Carrière) in eastern North America. In this study, cutting small gaps in the forest canopy above understory eastern hemlocks was evaluated as a potential tool for use in the integrated pest management of HWA. Two gap sizes and two gap creation methods (felling vs. girdling trees) were assessed for effects on HWA density, hemlock crown health, hemlock growth, and regeneration of competing tree species, at three different latitude groups in the southern Appalachian Mountains. In all three latitude groups, the treatment that removed the most canopy competition (Large Fell) increased the production of new hemlock shoots, reduced the percentage of dead shoots, and increased basal area increment of target hemlocks relative to unreleased trees. Smaller gaps and those that removed fewer competing live trees (Girdle treatments) also improved hemlock health in the southernmost latitude group. Positive effects of canopy gaps on hemlock crown condition and growth occurred despite similar or higher HWA densities on gap-released vs. unreleased trees. In regression analysis, more than half the variability in crown transparency change and basal area growth post-treatment was explained by the competition index of live trees remaining, average HWA density index, and tree size (dominant tree height and hemlock dbh). In the southernmost latitude group, the regeneration density of deciduous species increased significantly in the three years following gap creation and were highest in the Large Fell treatment. Results support the hypothesis that the increased availability of sunlight (and possibly other limited resources) resulting from canopy gaps (radius approximately ¼ to ½ dominant tree height) improves the physiological tolerance of eastern hemlock to HWA infestation and could complement biological control by providing a more stable source of new shoots with adelgid prey for predatory insects. [ABSTRACT FROM AUTHOR]

Hemlock woolly adelgid (Adelges tsugae Annand, HWA) is an introduced invasive forest pest in eastern North America. Herbivory by this insect results in mortality to eastern hemlock (Tsuga canadensis L. Carr.) and Carolina hemlock (Tsuga caroliniana Engelm.). These species occur in landscapes where extreme topographic variation is common. The vegetation communities within these landscapes feature high diversity of tree species, including several other conifer species. Traditional forest inventory procedures and insect pest detection methods within these limited-access landscapes are impractical. However, further information is needed to evaluate the impacts of HWA-induced hemlock mortality. Accordingly, our goal was to develop a semi-automatic method for mapping patches of coniferous tree species that include the living hemlock component and tree mortality by the HWA using aerial images and LiDAR (light detection and ranging) to increase our understanding of the severity and pattern of hemlock decline. The study was conducted in the Linville River Gorge in the Southern Appalachians of western North Carolina, USA. The mapping task included a two-phase approach: decision-tree and support vector machine classifications. We found that about 2% of the forest canopy surface was covered by dead trees and 43% by coniferous tree species. A large portion of the forest canopy surface (over 55%) was covered by deciduous tree species. The resulting maps provide a means for evaluating the impact of HWA herbivory, since this insect was the only significant coniferous mortality agent present within the study site.

The southern Appalachian forests have been threatened by several large-scale disturbances, such as wildfire and infestation, which alter the forest ecosystem structures and functions. Hemlock Woolly Adelgid (Adelges tsugae Annand, HWA) is a non-native pest that causes widespread foliar damage and eventual mortality, resulting in irreversible tree decline in eastern (Tsuga canadensis) and Carolina (T. caroliniana) hemlocks throughout the eastern United States. It is important to monitor the extent and severity of these disturbances over space and time to better understand their implications in the biogeochemical cycles of forest landscapes. Using all available Landsat images, we investigate and compare the performance of Tasseled Cap Transformation (TCT)-based indices, Normalized Difference Vegetation Index (NDVI), Enhanced Vegetation Index (EVI), and Disturbance Index (DI) in capturing the spectral-temporal trajectory of both abrupt and gradual forest disturbances (e.g., fire and hemlock decline). For each Landsat pixel, the temporal trajectories of these indices were fitted into a time series model, separating the inter-annual disturbance patterns (low frequency) and seasonal phenology (high frequency) signals. We estimated the temporal dynamics of disturbances based on the residuals between the observed and predicted values of the model, investigated the performance of all the indices in capturing the hemlock decline intensity, and further validated the results with the number of individual dead hemlocks identified from high-resolution aerial images. Our results suggested that the overall performance of NDVI, followed by TCT wetness, was most accurate in detecting both the disturbance timing and hemlock decline intensity, explaining over 90% of the variability in the number of dead hemlocks. Despite the overall good performance of TCT wetness in characterizing the disturbance regime, our analysis showed that this index has some limitations in characterizing disturbances due to its recovery patterns following infestation.

The hemlock woolly adelgid (HWA; Adelges tsugae) is an invasive insect infestation that is spreading into the forests of the northeastern United States, driven by the warmer winter temperatures associated with climate change. The initial stages of this disturbance are difficult to detect with passive optical remote sensing, since the insect often causes its host species, eastern hemlock trees (Tsuga canadensis), to defoliate in the midstory and understory before showing impacts in the overstory. New active remote sensing technologies—such as the recently launched NASA Global Ecosystem Dynamics Investigation (GEDI) spaceborne lidar—can address this limitation by penetrating canopy gaps and recording lower canopy structural changes. This study explores new opportunities for monitoring the HWA infestation with airborne lidar scanning (ALS) and GEDI spaceborne lidar data. GEDI waveforms were simulated using airborne lidar datasets from an HWA-infested forest plot at the Harvard Forest ForestGEO site in central Massachusetts. Two airborne lidar instruments, the NASA G-LiHT and the NEON AOP, overflew the site in 2012 and 2016. GEDI waveforms were simulated from each airborne lidar dataset, and the change in waveform metrics from 2012 to 2016 was compared to field-derived hemlock mortality at the ForestGEO site. Hemlock plots were shown to be undergoing dynamic changes as a result of the HWA infestation, losing substantial plant area in the middle canopy, while still growing in the upper canopy. Changes in midstory plant area (PAI 11–12 m above ground) and overall canopy permeability (indicated by RH10) accounted for 60% of the variation in hemlock mortality in a logistic regression model. The robustness of these structure-condition relationships held even when simulated waveforms were treated as real GEDI data with added noise and sparse spatial coverage. These results show promise for future disturbance monitoring studies with ALS and GEDI lidar data.

The hemlock woolly adelgid (HWA) (Adelges tsugae) is causing widespread mortality of eastern hemlock (EH) (Tsuga canadensis [L.] Carrière) across its range. Unless ecological or cultural considerations are accounted for, few forest managers would find it economical to manage the HWA. However, new markets for EH may increase incentives to manage the HWA or at least bring more value to this low-grade tree species. Our research explores the feasibility of using EH as animal bedding, by comparing water absorption, microbial growth, and wood shaving production factors with eastern white pine (EWP) (Pinus strobus L.). Results showed that when dried to 10 percent and 30 percent moisture content, EH absorbed 281 percent and 176 percent of its weight in water, which was significantly lower than EWP, which absorbed 361 percent and 243 percent respectively. However, no difference was found between tree species when comparing the growth of pathogenic bacteria in bedding samples. Furthermore, no difference in volume of produced bedding was found when using a wood shaving machine. These findings suggest that EH is suitable for animal bedding, especially if the substantially lower stumpage value of the wood resource, as compared with EWP and other species within its range, compensates for the reduced moisture absorbing capability. [ABSTRACT FROM AUTHOR]

The ability to predict key phenological stages of insect pests is of prime importance in sampling and management programs. Hemlock woolly adelgid, Adelges tsugae (Annand) (Hemiptera: Adelgidae), is a non-native invasive pest of eastern and Carolina hemlock that is currently expanding its range westward through the central Appalachian region. We conducted intensive sampling of A. tsugae across four sites in West Virginia and western Maryland from March 2013 to May 2014, and developed a degree-day model (minimum temperature threshold of 4°C) to predict the temporal distribution of progrediens eggs, progrediens egg hatch, newly hatched sistens, and settled first-instar sistens. We then linked degree-day predictions to recent weather data collected from our study region from 2012 to 2016 to provide a range of calendar days over which these key phenological events occur. The initial occurrence of progrediens eggs and progrediens egg hatch is predicted to occur at 209 (mean date of 19 April) and 346 (mean date of 8 May) degree days, respectively. The initial occurrence of newly hatched sistens and settled first-instar sistens is predicted to occur at 852 (mean date of 19 June) and 1010 (mean date of 30 June) degree days, respectively. Because our study region represents a region in which A. tsugae continues to spread, this information should be useful in optimizing sampling programs and the timing of control tactics in the central Appalachian region. [ABSTRACT FROM AUTHOR]

Hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae), is an exotic pest of eastern hemlock, Tsuga canadensis (L.) Carrière (Pinales: Pinaceae), in the eastern United States. Two commonly used insecticides to manage adelgid are imidacloprid, a systemic neonicotinoid insecticide, and horticultural oil, a refined petroleum oil foliar spray. We have investigated the influence of imidacloprid and horticultural oil on spider abundance at different canopy strata in eastern hemlock. In total, 2,084 spiders representing 11 families were collected from the canopies of eastern hemlock. In beat-sheet and direct observation samples, the families Theridiidae, Araneidae, Salticidae, and Anyphaenidae were the most abundant. Significantly higher numbers of spiders were recorded on untreated control trees compared with trees treated with imidacloprid using soil drench and soil injection applications. Spider abundance in trees injected with imidacloprid and horticultural oil applications did not significantly differ from control trees. Spider abundance was significantly greater in the top and middle strata of the canopy than in the bottom stratum, where imidacloprid concentrations were the highest. Regression analysis showed that spider abundance was inversely associated with imidacloprid concentration. This research demonstrates that imidacloprid, when applied with selected methods, has the potential to result in reductions of spider densities at different strata. However, slight reductions in spider abundance may be an acceptable short-term ecological impact compared with the loss of an untreated hemlock and all the associated ecological benefits that it provides. Future studies should include investigations of long-term impact of imidacloprid on spiders associated with eastern hemlock. [ABSTRACT FROM AUTHOR]

Eastern 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 6.89 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. [ABSTRACT FROM AUTHOR]

Nutritional bacterial symbionts enhance the diets of sap-feeding insects with amino acids and vitamins missing from their diets. In many lineages, an ancestral senior symbiont is joined by a younger junior symbiont. To date, an emergent pattern is that senior symbionts supply a majority of amino acids, and junior symbionts supply a minority. Similar to other hemipterans, adelgids harbor obligate symbionts, but have higher diversity of bacterial associates, suggesting a history of symbiont turnover. The metabolic roles of dual symbionts in adelgids and their contributions to the consortium are largely unexplored. Here, we investigate the symbionts of Adelges tsugae, the hemlock woolly adelgid (HWA), an invasive species introduced from Japan to the eastern United States, where it kills hemlock trees. The response of hemlocks to HWA feeding has aspects of a defensive reaction against pathogens, and some have speculated that symbionts may be involved. We sequenced the genomes of "Ca. Annandia adelgestsuga" and "Ca. Pseudomonas adelgestsugas" symbionts to detail their metabolic capabilities, infer ages of relationship, and search for effectors of plant defenses. We also tested the relationship of "Ca. Annandia" to symbionts of other insects. We find that both symbionts provide nutrients, but in more balanced proportions than dual symbionts of other hemipterans. The lesser contributions of the senior "Ca. Annandia" support our hypothesis for symbiont replacements in adelgids. Phylogenomic results were ambiguous regarding the position of "Ca. Annandia". We found no obvious effectors of plant defenses related to insect virulence, but hypothetical proteins in symbionts are unknown players. [ABSTRACT FROM AUTHOR]

Exploring genetic diversity within species of biological control agents can expose previously overlooked beneficial genotypes. This may be the case for two species of silver flies, Leucopis argenticollis and L. piniperda , predators of the hemlock woolly adelgid ( Adelges tsugae ) in the Pacific Northwest of North America. The same Leucopis species occur in eastern North America, where they feed on other adelgid species, but not on hemlock woolly adelgid, which became a pest in the region after its introduction from Japan earlier this century. We collected DNA sequence data from one mitochondrial and two nuclear genes for 606 Leucopis individuals to examine variation correlated with geographic origin and prey association. Specimens of L. argenticollis and L. piniperda collected from adelgids on hemlock and spruce in the West are phylogenetically divergent from conspecifics collected from adelgids on pine and spruce in the East. These results suggest that within each species, there are distinct lineages that specialize on different adelgid prey on different hosts in western versus eastern North America. The western lineages appear to be strong candidates for enhancing biological control of Japanese hemlock woolly adelgid in the eastern United States, given their widespread occurrence on western hemlock woolly adelgid in the Pacific Northwest, and the absence of these genotypes in the East. We also developed a PCR-RFLP assay based on the mitochondrial COI gene as an inexpensive and reliable way to distinguish the four genetic groups to document establishment and impact following release. [ABSTRACT FROM AUTHOR]

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

Hemlock woolly adelgid (Adelges tsugae Annand) (HWA) is an invasive insect that feeds upon the foliage of eastern hemlock (Tsuga canadensis (L.) Carrière) trees, leading to a decline in health and often mortality. The exact mechanism leading to the demise of eastern hemlocks remains uncertain because little is known about how HWA infestation directly alters the host's physiology. To evaluate the physiological responses of eastern hemlock during early infestation of HWA, we measured needle loss, xylem hydraulic conductivity, vulnerability to cavitation, tracheid anatomy, leaf-level gas exchange, leaf water potential and foliar cation and nutrient levels on HWA-infested and noninfested even-aged trees in an experimental garden. HWA infestation resulted in higher xylem hydraulic conductivity correlated with an increase in average tracheid lumen area and no difference in vulnerability to cavitation, indicating that needle loss associated with HWA infestation could not be attributed to reduced xylem transport capacity. HWA-infested trees exhibited higher rates of net photosynthesis and significant changes in foliar nutrient partitioning, but showed no differences in branch increment growth rates compared with noninfested trees. This study suggests that HWA-induced decline in the health of eastern hemlock trees is not initially caused by compromised water relations or needle loss. [ABSTRACT FROM AUTHOR]

Micromechanical properties that help mediate herbivore access may be particularly important when considering herbivorous insects that feed with piercing-sucking stylets. We used microindentation to quantify the micromechanical properties of hemlock, Tsuga spp., to quantify the hardness of the feeding site of the invasive hemlock woolly adelgid, Adelges tsugae. We measured hardness of the hemlock leaf cushion, the stylet insertion point of the adelgid, across four seasons in a 1 y period for four hemlock species growing in a common garden, including eastern, western, mountain, and northern Japanese hemlocks. Leaf cushion hardness was highest in the fall and winter and lowest in summer for all species. Northern Japanese hemlock had relatively greater hardness than the remaining species. Our data contributes an additional perspective to the existing framework within which greater susceptibility and subsequent mortality of eastern hemlocks is observed. The potential application of microindentation to understanding the nature and relevance of plant mechanical defenses in plant–herbivore interactions is also demonstrated and highlighted.