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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]

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]

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]

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]

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. [ABSTRACT FROM AUTHOR]

Seven species of hemlock (Tsuga spp.)--four from North America and three from Asia--were evaluated in replicated plots in Katonah, New York, United States (USDA Plant Hardiness Zone 6b) as potential replacements for eastern hemlock (Tsuga canadensis), which is gradually being extirpated from landscapes in the eastern United States. by the invasive hemlock woolly adelgid (Adelges tsugae). Trends reported in an earlier study (Weston and Harper 2009) continued but were exaggerated after an additional three years of observation. For example, Chinese hemlock (T. chinensis) continued to show the greatest potential as a replacement for T. canadensis as mortality was very low, overall plant health was exceptional, and tolerance to A. tsugae was robust. Early indicators suggest that T. chinensis may also be readily propagated from hardwood cuttings under appropriate greenhouse conditions. These characteristics suggest that T. chinensis may indeed become a viable replacement for T. canadensis, and a valuable addition to landscapes in the eastern U.S. [ABSTRACT FROM AUTHOR]

The hemlock woolly adelgid ( Adelges tsugae Annand) is an invasive species causing high mortality of eastern hemlock ( Tsuga canadensis (L) Carriere) in the forests of eastern North America. Recent findings revealed that sapling eastern hemlocks artificially infested with hemlock woolly adelgid in a plantation setting responded to the insect with an array of induced resin volatile changes. Here we determine if eastern hemlocks growing beneath a forest canopy respond to hemlock woolly adelgid infestation with the same patterns of constitutive and inducible volatile resin production as those plantation specimens. We inoculated previously uninfested branches of mature and immature hemlocks in a central New England forest with hemlock woolly adelgid. We then sampled twig tissue of infested and uninfested trees in late spring, early summer, and mid-autumn, after known intervals of adelgid activity when an induced response might be expected. We identified and quantified resin volatiles by gas chromatography-mass spectrometry. Constitutive levels of total monoterpenoids, sesquiterpenoids, and combined resin volatiles were all several-fold more abundant in forest trees than those previously measured in a plantation setting, pointing to further study of the influence of site factors on hemlock volatile production. Hemlock woolly adelgid infestation induced an array of changes in eastern hemlock's volatile profile, including many-fold increases in benzyl alcohol and methyl salicylate accumulation. Despite differences in constitutive concentrations of volatiles between the two sites, our findings verify that hemlock woolly adelgid elicits patterns of resin volatile induction in forest-grown eastern hemlocks quite similar to those previously observed in plantation grown trees. [ABSTRACT FROM AUTHOR]

Eastern North American forests have effectively lost two major tree species (American chestnut and American elm) in the last 100 years and two more, eastern and Carolina hemlock, will be functionally extinct over much of their ranges within a couple of decades. The loss of eastern hemlock is of particular concern because hemlock is: (1) a foundation species; (2) one of the longest-lived tree species over much of temperate eastern North America; and (3) sensitive to climatic variation and ecosystem disturbance, making it an ideal species for the reconstruction of environmental history. Unlike American chestnut, we have a small window of opportunity to salvage environmental histories from hemlock before they are lost. In this progress report, we review the extensive body of science derived from this paleoenvironmental archive and urge scientists from eastern North America to sample and archive old-growth hemlock while living and dead material remain. Here we describe a community-based approach to salvaging paleoenvironmental archives that could serve as a model for collections from other foundation species currently threatened by exotic forests pests and pathogens (e.g. whitebark pine, ash). The approach supports Schlesinger’s (2010) call for ‘translational ecology’ by building connections between scientists, students, environmental NGOs, and land managers focused on old-growth forests. [ABSTRACT FROM AUTHOR]

The article focuses on the explicit stochastic model which combines population and dispersal processes developed to investigate the invasion of hemlock woolly adelgid (HWA) in North America. It states that the model has predicted heterogenous risk of HWA invasion through time and across space. It says that differences in observed and modeled dynamics can be attributed to inability to capture direction and timing of long-distance dispersal event.

Abstract: Molecular diagnostics based on DNA barcodes can be powerful identification tools in the absence of distinctive morphological characters for distinguishing between closely related species. A specific example is distinguishing the endemic species Laricobius rubidus from Laricobius nigrinus, a biological control agent of hemlock woolly adelgid introduced into the eastern United States. This is especially important because their larvae are morphologically similar and are often collected together in stands where pines and hemlock grow together. Diagnostic nucleotide differences to distinguish species were determined using 157 L. nigrinus and 205 L. rubidus cytochrome oxidase I (COI) barcode DNA sequences. Two polymerase chain reaction (PCR) assays were developed: PCR followed by restriction length polymorphism (RFLP) and real-time PCR (qPCR) based on hydrolysis probes. The qPCR assay had limited success when applied to L. nigrinus originating from inland vs. coastal populations in the western United States. PCR–RFLP was successful, regardless of sample origin. These two species-specific assays provide a choice of diagnostic tools to best match the available lab equipment and management objectives for those using L. nigrinus as a biological control agent. [Copyright &y& Elsevier]

Hemlock Woolly Adelgid ( Adelges tsugae) is spreading across forests in eastern North America, causing mortality of eastern hemlock ( Tsuga canadensis [L.] Carr.) and Carolina hemlock ( Tsuga caroliniana Engelm.). The loss of hemlock from riparian forests in Great Smoky Mountains National Park (GSMNP) may result in significant physical, chemical, and biological alterations to stream environments. To assess the influence of riparian hemlock stands on stream conditions and estimate possible impacts from hemlock loss in GSMNP, we paired hardwood- and hemlock-dominated streams to examine differences in water temperature, nitrate concentrations, pH, discharge, and available photosynthetic light. We used a Geographic Information System (GIS) to identify stream pairs that were similar in topography, geology, land use, and disturbance history in order to isolate forest type as a variable. Differences between hemlock- and hardwood-dominated streams could not be explained by dominant forest type alone as forest type yields no consistent signal on measured conditions of headwater streams in GSMNP. The variability in the results indicate that other landscape variables, such as the influence of understory Rhododendron species, may exert more control on stream conditions than canopy composition. The results of this study suggest that the replacement of hemlock overstory with hardwood species will have minimal impact on long-term stream conditions, however disturbance during the transition is likely to have significant impacts. Management of riparian forests undergoing hemlock decline should, therefore, focus on facilitating a faster transition to hardwood-dominated stands to minimize long-term effects on water quality. [ABSTRACT FROM AUTHOR]

In western North America, infestations of the hemlock woolly adelgid, Adelges tsugae Annand (Hemiptera: Adelgidae), are common on orchard, ornamental, and roadside western hemlock, Tsuga heterophylla (Raf.) Sargent. However, these infestations rarely cause T. heterophylla mortality. Host tolerance and presence of endemic predators may be contributing to the relatively low levels of injury to T. heterophylla caused by A. tsugae. Field surveys of the arthropod community associated with A. tsugae infestations on 116 T. heterophylla at 16 sites in Oregon and Washington were conducted every 4-6 wk from January 2005 through November 2006. Fourteen uninfested T. heterophylla were also surveyed across 5 of the 16 sites. Immature A. tsugae predators collected in the field were brought to the laboratory for rearing. Eight species of hymenopterous parasitoids were reared from pupae of predators of A. tsugae in the laboratory. Two Pachyneuron spp. (Pteromalidae) and a Melanips sp. (Figitidae) were reared from Leucopis spp. (Diptera: Chamaemyiidae) puparia. Syrphoctonus pallipes (Gravenhorst) (Ichneumonidae), Woldstectius flavolineatus (Gravenhorst) (Ichneumonidae), Syrphophagus sp. (Encyrtidae), and Pachyneuron albutius Walker were reared from Syrphidae (Diptera) puparia. A Helorus sp. (Heloridae) was reared from a Chrysoperla sp. (Neuroptera: Chrysopidae) cocoon. Laboratory rearing did not show any direct association between parasitoids and A. tsugae. In the field survey, a total of 509 adult parasitic Hymenoptera representing 19 families and at least 57 genera were collected from T. heterophylla. Nonparametric analysis of community structure showed Pachyneuron spp. were strongly correlated to abundance of their Leucopis spp. hosts and to A. tsugae population score in the field. The possible impact of parasitism on Leucopis spp., potential A. tsugae biological control candidates for the eastern United States, is discussed. [ABSTRACT FROM AUTHOR]

An exotic insect, the hemlock woolly adelgid, Adelges tsugae Annand (Hem., Adelgidae), is spreading through eastern North America, killing hemlock trees [ Tsuga canadensis (L.) Carrière], and thereby impoverishing ecosystems. Adelges tsugae, like many alien invasive insects, is difficult to monitor or sample in the forest. Monitoring of A. tsugae has been hampered by lack of information about its distribution within tree crowns. In order to assist future monitoring and biocontrol of A. tsugae, this study investigates the crown distribution of A. tsugae by sampling from the entire height of mature hemlock trees in a forest with an established infestation. In addition to A. tsugae, sampling includes scale insects, which are another group of important pests on hemlock trees. This study demonstrates the utility of a randomized branch sampling (RBS) plan for monitoring both invasive insects as well as native insects that are difficult to sample. Results from the RBS show that in trees with high populations of A. tsugae, branches from the lower crown have slightly higher densities of A. tsugae than upper crown branches. In trees with low A. tsugae populations, the upper crown may have higher A. tsugae densities than the lower crown. North pointing branches also have higher densities of A. tsugae than branches pointing in other cardinal directions. Future sampling efforts for A. tsugae can take advantage of higher densities in certain portions of the crown to increase accuracy. [ABSTRACT FROM AUTHOR]

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