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The non-native hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae), has caused a significant decline of eastern hemlock, Tsuga canadensis L. (Pinales: Pinaceae), and Carolina hemlock, Tsuga caroliniana Engelmann (Pinales: Pinaceae), in eastern North America. Biological control of HWA has focused on the use of 2 Laricobius spp. (Coleoptera: Derodontidae), natural predators of HWA, which require arboreal and subterranean life phases to complete their development. In its subterranean phase, Laricobius spp. are subject to abiotic factors including soil compaction or soil-applied insecticides used to protect hemlock from HWA. This study used 3D X-ray microcomputed tomography (micro-CT) to identify the depth at which Laricobius spp. burrows during its subterranean lifecycle, characterize pupal chamber volume, and determine whether soil compaction had a significant effect on these variables. The mean burrowing depth in the soil of individuals was 27.0 mm ± 14.8 (SD) and 11.4 mm ± 11.8 (SD) at compaction levels of 0.36 and 0.54 g/ cm3, respectively. The mean pupal chamber volume was 11.15 mm3 ± 2.8 (SD) and 7.65 mm3 ± 3.5 (SD) in soil compacted at 0.36 and 0.54 g/cm3, respectively. These data show that soil compaction influences burrowing depth and pupal chamber size for Laricobius spp.This information will help us better identify the effect of soil-applied insecticide residues on estivating Laricobius spp. and soil-applied insecticide residues in the field. Additionally, these results demonstrate the utility of 3D micro-CT in assessing subterranean insect activity in future studies.

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

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

The non-native hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae), has caused a significant decline of eastern hemlock, Tsuga canadensis L. (Pinales: Pinaceae), and Carolina hemlock, Tsuga caroliniana Engelmann (Pinales: Pinaceae), in eastern North America. Biological control of HWA has focused on the use of 2 Laricobius spp. (Coleoptera: Derodontidae), natural predators of HWA, which require arboreal and subterranean life phases to complete their development. In its subterranean phase, Laricobius spp. are subject to abiotic factors including soil compaction or soil-applied insecticides used to protect hemlock from HWA. This study used 3D X-ray microcomputed tomography (micro-CT) to identify the depth at which Laricobius spp. burrows during its subterranean lifecycle, characterize pupal chamber volume, and determine whether soil compaction had a significant effect on these variables. The mean burrowing depth in the soil of individuals was 27.0 mm ± 14.8 (SD) and 11.4 mm ± 11.8 (SD) at compaction levels of 0.36 and 0.54 g/ cm3, respectively. The mean pupal chamber volume was 11.15 mm3 ± 2.8 (SD) and 7.65 mm3 ± 3.5 (SD) in soil compacted at 0.36 and 0.54 g/cm3, respectively. These data show that soil compaction influences burrowing depth and pupal chamber size for Laricobius spp.This information will help us better identify the effect of soil-applied insecticide residues on estivating Laricobius spp. and soil-applied insecticide residues in the field. Additionally, these results demonstrate the utility of 3D micro-CT in assessing subterranean insect activity in future studies.

Laricobius nigrinus Fender (Coleoptera: Derodontidae), a predatory beetle native to western North America, has been released since 2003 for management of hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae), a non-native pest killing hemlocks in eastern North America. Over 420,000 L. nigrinus have been released in the eastern USA from field and lab-reared sources, 14,000 of which were deployed in the Commonwealth of Virginia, USA. In order to determine the establishment rates of L. nigrinus in Virginia, surveys were conducted in 2017 and 2018 at all release sites within this state. During the study, stand-level HWA densities were estimated, and hemlock tree health and predator–prey ratios were quantified. The identification of Laricobius spp. recoveries were made using microsatellite analysis. During the period of the study, L. nigrinus were found to have established at 82% of Virginia release sites and were the primary species recovered (80%). Both Laricobius rubidus (18%) and hybrids (2%) were also recovered. Stand-level HWA densities varied greatly over sites and years but showed a general decline in year two of the study. Establishment at such a high percentage of release sites suggests that the climate in Virginia is suitable for the predator and the insect is adaptable to the wide variety of site conditions where hemlocks typically grow.

Laricobius nigrinus Fender (Coleoptera: Derodontidae), a predatory beetle native to western North America, has been released since 2003 for management of hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae), a non-native pest killing hemlocks in eastern North America. Over 420,000 L. nigrinus have been released in the eastern USA from field and lab-reared sources, 14,000 of which were deployed in the Commonwealth of Virginia, USA. In order to determine the establishment rates of L. nigrinus in Virginia, surveys were conducted in 2017 and 2018 at all release sites within this state. During the study, stand-level HWA densities were estimated, and hemlock tree health and predator–prey ratios were quantified. The identification of Laricobius spp. recoveries were made using microsatellite analysis. During the period of the study, L. nigrinus were found to have established at 82% of Virginia release sites and were the primary species recovered (80%). Both Laricobius rubidus (18%) and hybrids (2%) were also recovered. Stand-level HWA densities varied greatly over sites and years but showed a general decline in year two of the study. Establishment at such a high percentage of release sites suggests that the climate in Virginia is suitable for the predator and the insect is adaptable to the wide variety of site conditions where hemlocks typically grow.

Hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae), has devastated eastern hemlock (Tsuga canadensis [L.] Carriere) in a major portion of its native range in eastern North America. Population dynamics of HWA in the absence of predators have been studied for decades. After many years and much effort directed towards rearing and releasing biological control agents to manage HWA, one of these agents, Laricobius nigrinus Fender (Coleoptera: Derodontidae), is now successfully established at significant densities at sites from the southern to the mid-Atlantic states of the eastern U.S. However, high densities of HWA still persist at many locations throughout the region and spread of HWA and associated damage to hemlock continues. Population models for HWA have suggested that even upwards of 90% predation on eggs laid by the overwintering sistens generation will have minimal effect in reducing the population densities of HWA, if HWA are at high density. In this study, we tested the ability of L. nigrinus to reduce HWA densities, and experimentally tested these model predictions to better understand what impact, if any, L. nigrinus has on HWA densities. By using predator exclusion cages at field sites with well-established populations of L. nigrinus, we were able to record HWA densities, fecundity, overwintering mortality, and predation by L. nigrinus, as well as the proportion of branch tips producing new growth on study trees. Using our field-collected data, we refitted the model in ways that allowed us to predict what population densities we could expect for the following summertime progrediens generation given previous HWA density and levels of L. nigrinus. In both years, we found that despite high rates (greater than 80% ovisac predation) of predation by L. nigrinus on uncaged branches compared to caged branches, there were no significant differences in subsequent densities of the HWA spring generation between caged and uncaged treatments, as pr

Hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae), is an invasive pest causing significant mortality to eastern and Carolina hemlock in eastern North America. Since 2003, management of HWA has included targeted release of the HWA predator Laricobius nigrinus Fender (Coleoptera: Derodontidae), native to western North America. Establishment of L. nigrinus at release sites is well documented, but investigations of its impact on HWA populations have been limited. A four-year (2014-2018), two-phase study using predator exclusion cages to assess the impact of L. nigrinus on HWA was conducted at nine previous release sites in the eastern United States. Significantly more HWA sistens ovisacs were disturbed on no-cage and open-cage branches than on caged branches where predators were excluded. Mean disturbance levels on cage, no-cage and open-cage branches was 8, 38, and 27 percent, respectively. Seven of nine sites had a mean HWA ovisac disturbance greater than 50% for at least one year. Winter temperatures were also a significant factor in overall mortality of the sistens generation with a mean of 46% on study branches. Six of nine sites had a mean overall mortality (winter mortality and predation) greater than 80% for at least one year. Larvae of Laricobius spp. were recovered at all sites during this study. Sequencing of the COI gene from recoveries in Phase One (2015 and 2016) indicated that 88% were L. nigrinus and 12% were L. rubidus LeConte. Microsatellite analysis performed during Phase Two (2017 and 2018) indicated that approximately 97% of larval recoveries were L. nigrinus, 2% were hybrids of L. nigrinus and L. rubidus, and 1% were L. rubidus. Results of this study suggest that L. nigrinus can significantly impact the HWA sistens generation ovisacs and continued investment in the use of this species as a biological control is recommended.

Hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae), has devastated eastern hemlock (Tsuga canadensis [L.] Carriere) in a major portion of its native range in eastern North America. Population dynamics of HWA in the absence of predators have been studied for decades. After many years and much effort directed towards rearing and releasing biological control agents to manage HWA, one of these agents, Laricobius nigrinus Fender (Coleoptera: Derodontidae), is now successfully established at significant densities at sites from the southern to the mid-Atlantic states of the eastern U.S. However, high densities of HWA still persist at many locations throughout the region and spread of HWA and associated damage to hemlock continues. Population models for HWA have suggested that even upwards of 90% predation on eggs laid by the overwintering sistens generation will have minimal effect in reducing the population densities of HWA, if HWA are at high density. In this study, we tested the ability of L. nigrinus to reduce HWA densities, and experimentally tested these model predictions to better understand what impact, if any, L. nigrinus has on HWA densities. By using predator exclusion cages at field sites with well-established populations of L. nigrinus, we were able to record HWA densities, fecundity, overwintering mortality, and predation by L. nigrinus, as well as the proportion of branch tips producing new growth on study trees. Using our field-collected data, we refitted the model in ways that allowed us to predict what population densities we could expect for the following summertime progrediens generation given previous HWA density and levels of L. nigrinus. In both years, we found that despite high rates (greater than 80% ovisac predation) of predation by L. nigrinus on uncaged branches compared to caged branches, there were no significant differences in subsequent densities of the HWA spring generation between caged and uncaged treatments, as pr

Hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae), is an invasive pest causing significant mortality to eastern and Carolina hemlock in eastern North America. Since 2003, management of HWA has included targeted release of the HWA predator Laricobius nigrinus Fender (Coleoptera: Derodontidae), native to western North America. Establishment of L. nigrinus at release sites is well documented, but investigations of its impact on HWA populations have been limited. A four-year (2014-2018), two-phase study using predator exclusion cages to assess the impact of L. nigrinus on HWA was conducted at nine previous release sites in the eastern United States. Significantly more HWA sistens ovisacs were disturbed on no-cage and open-cage branches than on caged branches where predators were excluded. Mean disturbance levels on cage, no-cage and open-cage branches was 8, 38, and 27 percent, respectively. Seven of nine sites had a mean HWA ovisac disturbance greater than 50% for at least one year. Winter temperatures were also a significant factor in overall mortality of the sistens generation with a mean of 46% on study branches. Six of nine sites had a mean overall mortality (winter mortality and predation) greater than 80% for at least one year. Larvae of Laricobius spp. were recovered at all sites during this study. Sequencing of the COI gene from recoveries in Phase One (2015 and 2016) indicated that 88% were L. nigrinus and 12% were L. rubidus LeConte. Microsatellite analysis performed during Phase Two (2017 and 2018) indicated that approximately 97% of larval recoveries were L. nigrinus, 2% were hybrids of L. nigrinus and L. rubidus, and 1% were L. rubidus. Results of this study suggest that L. nigrinus can significantly impact the HWA sistens generation ovisacs and continued investment in the use of this species as a biological control is recommended.

The predatory beetle, Laricobius nigrinus Fender (Coleoptera: Derodontidae), has been released in the eastern U.S. since 2003 for the management of hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae). The establishment of L. nigrinus at release sites in Virginia was assessed in 2017 and 2018. Sampling was performed in both years to determine presence or absence L. nigrinus. Stand level HWA densities, tree health, predator-prey ratios, and Laricobius spp. identifications were also assessed at each site. Laricobius nigrinus established at 82% of sites and was the primary species recovered. HWA densities appeared to decline, and tree health appeared to improve in year two. Predator-prey ratios were lower than those indicated in the native range, however much is left to be understood about the dynamics of this system in its introduced range. A separate two-year study to assess the impact of L. nigrinus on HWA was initiated in 2014 (Phase One) at nine sites in the eastern U.S. Significant predation of HWA sistens ovisacs was demonstrated during this period, therefore, it was continued from 2016-2018 (Phase Two) to provide longer-term evaluations. Predator exclusion cages were used to monitor predator and prey populations. In Phase Two of the study, mean ovisac disturbance rates on no-cage branches were significantly greater than caged branches and were as high as 80%, suggesting that L. nigrinus can have a significant impact on the sistens generation. Microsatellite analysis of Laricobius larvae indicated that L. nigrinus was the primary species recovered at study sites.

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.

The predatory beetle, Laricobius nigrinus Fender (Coleoptera: Derodontidae), has been released in the eastern U.S. since 2003 for the management of hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae). The establishment of L. nigrinus at release sites in Virginia was assessed in 2017 and 2018. Sampling was performed in both years to determine presence or absence L. nigrinus. Stand level HWA densities, tree health, predator-prey ratios, and Laricobius spp. identifications were also assessed at each site. Laricobius nigrinus established at 82% of sites and was the primary species recovered. HWA densities appeared to decline, and tree health appeared to improve in year two. Predator-prey ratios were lower than those indicated in the native range, however much is left to be understood about the dynamics of this system in its introduced range. A separate two-year study to assess the impact of L. nigrinus on HWA was initiated in 2014 (Phase One) at nine sites in the eastern U.S. Significant predation of HWA sistens ovisacs was demonstrated during this period, therefore, it was continued from 2016-2018 (Phase Two) to provide longer-term evaluations. Predator exclusion cages were used to monitor predator and prey populations. In Phase Two of the study, mean ovisac disturbance rates on no-cage branches were significantly greater than caged branches and were as high as 80%, suggesting that L. nigrinus can have a significant impact on the sistens generation. Microsatellite analysis of Laricobius larvae indicated that L. nigrinus was the primary species recovered at study sites.

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.

The hemlock woolly adelgid, Adelges tsugae Annand, is an invasive insect that threatens hemlock species in eastern North America. Several species from the genus Laricobius are predators of A. tsugae in its native areas of Asia and the western United States. Two Laricobius species have been released as biological control agents: Laricobius nigrinus Fender, and Laricobius osakensis Montgomery and Shiyake. Laricobius rubidus LeConte is an adelgid predator native to the Eastern United States, where it can feed and complete development on A. tsugae opportunistically. Laboratory assays were conducted to assess the cold hardiness of these three Laricobius species, including two distinct populations of L. osakensis, by measuring the supercooling points of each species from November 2016 through March 2017. This information may be useful for choosing the best-suited biological control agent for a particular region to control A. tsugae. There was a significant difference between the overall mean supercooling point of L. rubidus compared to the other Laricobius spp. There were also significant differences of supercooling points between L. rubidus and both strains of L. osakensis in January, and significant differences between L. rubidus and all other strains in February. L. rubidus appear better adapted to cold extremes in the eastern U.S. than imported Laricobius spp.

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.

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.

The hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae), is an invasive pest from Japan that is causing significant hemlock mortality in the eastern U.S. The most promising control method is biological control. Beetles in the genus Laricobius are adelgid specialists. In 2005, Laricobius osakensis, was discovered in Japan and in 2010 was approved for release in the eastern United States and there have now been more than 60 releases. In 2014, the polar vortex resulted in significant HWA mortality in the eastern U.S., depleting the food source for L. osakensis, which resulted in low field recoveries of them. In the 2015-2016 field season, there were 14 larvae recovered from the field, and the 2016-2017 field season yielded 90 larvae. A significant correlation was found between Laricobius beetles recovered and HWA density, between HWA density and plant hardiness zone, and a negative correlation between Laricobius beetles recovered and average tree health. Understanding more about the supercooling point of L. osakensis gave insight to its ability to survive subfreezing temperatures as occurred in 2014. Comparison of the supercooling point between the northern and southern populations of L. osakensis, and to the other released HWA predator, L. nigrinus, will allow us to determine which biological control agent is best suited for release in the coldest regions. The overall mean supercooling points of northern L. osakensis was -13.52 oC, southern L. osakensis was -13.42 oC, and L. nigrinus was -13.57 oC. There were no significant differences between species or populations.

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.

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.

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.

The insecticide imidacloprid, has been found to be highly effective in suppressing hemlock woolly adelgid, Adelges tsugae. Laricobius nigrinus is a predatory beetle released as a biological control of adelgids in the eastern U.S. This project was designed to develop a pest management strategy that utilizes both tactics concurrently within the same site. It will assess the efficiency of this strategy in reducing HWA populations and improving the health of hemlock forests. The project was started in 2010 and data were collected annually through 2016. The project spanned three sites in three different states (KY, WV, and TN). Results show that tree health has generally declined across all sites for each year. HWA population index values are highly variable and are more strongly influenced by the occurrence of low winter temperatures than by treatment effect. Cross-correlation analysis of tree health and HWA population, revealed characteristics of their temporal relationship. In two of the three sites, tree health lagged up to three years behind changes in HWA population, and HWA populations lagged approximately one year behind changes in tree health. L. nigrinus did not establish at any site as of 2016. The lack of sustained recovery of the beetle may be attributable to the occurrence of extremely cold temperatures during the winters of 2014 and 2015 which produced subsequent crashes in the HWA population at two of the three sites. In TN, the L. nigrinus population may have never established due to a decline in the HWA population shortly after release.

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

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

The insecticide imidacloprid, has been found to be highly effective in suppressing hemlock woolly adelgid, Adelges tsugae. Laricobius nigrinus is a predatory beetle released as a biological control of adelgids in the eastern U.S. This project was designed to develop a pest management strategy that utilizes both tactics concurrently within the same site. It will assess the efficiency of this strategy in reducing HWA populations and improving the health of hemlock forests. The project was started in 2010 and data were collected annually through 2016. The project spanned three sites in three different states (KY, WV, and TN). Results show that tree health has generally declined across all sites for each year. HWA population index values are highly variable and are more strongly influenced by the occurrence of low winter temperatures than by treatment effect. Cross-correlation analysis of tree health and HWA population, revealed characteristics of their temporal relationship. In two of the three sites, tree health lagged up to three years behind changes in HWA population, and HWA populations lagged approximately one year behind changes in tree health. L. nigrinus did not establish at any site as of 2016. The lack of sustained recovery of the beetle may be attributable to the occurrence of extremely cold temperatures during the winters of 2014 and 2015 which produced subsequent crashes in the HWA population at two of the three sites. In TN, the L. nigrinus population may have never established due to a decline in the HWA population shortly after release.

Management of hemlock woolly adelgid (HWA), Adelges tsugae (Hemiptera: Adelgidae), is currently being implemented through several different methods including the release of host-specific predators such as Laricobius nigrinus Fender (Coleoptera: Derodontidae). Releases of this predator began in 2003. In 2014, an assessment of the efficacy of L. nigrinus at release sites from New Jersey to Georgia. Field sites were chosen based on the presence of moderate to high densities of HWA, that L. nigrinus was released at least four years prior to the start of the study, and that the predator was determined to be established at that site. Three treatments were set up at each of the sites: no cage, closed exclusion cage, and open cage. Three assessments were taken during key points throughout the season in order to monitor both HWA and L. nigrinus populations. Larval predator were recovered from most of the sites in year one and in higher numbers in year two. Many sites at which L. nigrinus were recovered showed high predation rates of HWA in uncaged samples ranging from 1.5 to 47.3% in year one and 0 to 66% in year 2. A survey for L. nigrinus establishment at previous release sites in VA was conducted. These sites date back as far as 2003 and as recently as 2015. Beat sheeting and branch clippings were conducted to recover Laricobius spp. adults and larvae, respectively. The recovered insects were then identified to species through genetic analysis. A mix of the introduced L. nigrinus and the native L. rubidus LeConte (Coleoptera: Derodontidae) were recovered at some sites, and only L. rubidus were recovered from others. Higher numbers of Laricobius were recovered in spring of year two. Overall, tree health in Virginia decreased from spring 2015 to 2016.

The hemlock woolly adelgid (HWA), Adelges tsugae Annand, is an invasive pest of eastern hemlock, Tsuga canadensis (L.) Carriere and Carolina hemlock Tsuga caroliniana Englem. in the eastern United States. A newly reported beetle predator for HWA, Scymnus (Pullus) coniferarum Crotch (Coleoptera: Cocinellidae) preys on the pest in the western United States, and was approved for release in the eastern United States for the control of HWA. This research investigated the viability of S. coniferarum as a biological control agent of A. tsugae in the eastern United States, as well as the ecological dynamics between S. coniferarum and host prey species in its native range of western North America. In objective one, S. coniferarum predation, reproductive potential, and survival were evaluated in field-cages on adelgid infested T. canadensis in southwestern Virginia. Adult S. coniferarum fed on both generations and all life stages of A. tsugae at rates comparable to other adelgid-specific predators, and survived for extended periods of time in the field. In objective two, host-range tests for S. coniferarum were conducted in a series of no-choice and paired-choice feeding, oviposition, and development studies. Scymnus coniferarum adults fed on all adelgid species, and completed development on HWA and Adelges piceae Ratz. Scymmnus coniferarum oviposition was extremely low. In the final objective, Douglas-fir, Pseudotusga menziesii Mirb., Shore pine, Pinus contorta Dougl., western white pine, Pinus monticola Dougl., and western hemlock, Tsuga heterophylla (Raf.) Sarg. host tree species were sampled in Tacoma, Washington to investigate the life history of S. coniferarum and associated adelgid prey species in the western United States. Scymnus coniferarum adults were found on both pine species, Douglas fir, and western hemlock, and seemed to move between host tree species seasonally. Each host tree supports a different adelgid species, and a limited diet of strictly HWA in host-ra

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

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

The hemlock woolly adelgid, Adelges tsugae Annand is an invasive insect that frequently causes hemlock (Tsuga spp.) mortality in the eastern United States. Studies have shown that once healthy hemlocks become infested by the adelgid, nutrients are depleted from the tree, leading to both tree decline and a reduction of the adelgid population. Since A. tsugae is dependent on hemlock for nutrients, feeding on trees in poor health may affect the ability of the insect to obtain necessary nutrients and may consequently affect their physiological and population health. Trees were categorized as lightly or moderately impacted by A. tsugae based on quantitative and qualitative tree health measurements. Population health of A. tsugae on each tree was determined by measuring insect density and peak mean fecundity; A. tsugae physiological health was determined by measuring insect biomass, total carbon, carbohydrate, total nitrogen, and amino nitrogen levels. Adelges tsugae from moderately impacted trees exhibited significantly greater fecundity than from lightly impacted trees. However, A. tsugae from lightly impacted hemlocks contained significantly greater levels of carbohydrates, total nitrogen, and amino nitrogen. While the results of the physiological analysis generally support our hypothesis that A. tsugae on lightly impacted trees are healthier than those on moderately impacted trees, this was not reflected in the population health measurements. Adelges tsugae egg health in response to tree health should be verified. This study provides the first examination of A. tsugae physiological health in relation to standard A. tsugae population health measures on hemlocks of different health levels.

The hemlock woolly adelgid, Adelges tsugae Annand is an invasive insect that frequently causes hemlock (Tsuga spp.) mortality in the eastern United States. Studies have shown that once healthy hemlocks become infested by the adelgid, nutrients are depleted from the tree, leading to both tree decline and a reduction of the adelgid population. Since A. tsugae is dependent on hemlock for nutrients, feeding on trees in poor health may affect the ability of the insect to obtain necessary nutrients and may consequently affect their physiological and population health. Trees were categorized as lightly or moderately impacted by A. tsugae based on quantitative and qualitative tree health measurements. Population health of A. tsugae on each tree was determined by measuring insect density and peak mean fecundity; A. tsugae physiological health was determined by measuring insect biomass, total carbon, carbohydrate, total nitrogen, and amino nitrogen levels. Adelges tsugae from moderately impacted trees exhibited significantly greater fecundity than from lightly impacted trees. However, A. tsugae from lightly impacted hemlocks contained significantly greater levels of carbohydrates, total nitrogen, and amino nitrogen. While the results of the physiological analysis generally support our hypothesis that A. tsugae on lightly impacted trees are healthier than those on moderately impacted trees, this was not reflected in the population health measurements. Adelges tsugae egg health in response to tree health should be verified. This study provides the first examination of A. tsugae physiological health in relation to standard A. tsugae population health measures on hemlocks of different health levels.

The hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Sternorrhyncha: Adelgidae) is a small invasive insect that frequently causes hemlock (Tsugae spp.) mortality in the eastern United States. Studies have shown that once healthy hemlocks become infested by the adelgid, nutrients are depleted from the tree, leading to both tree decline and a reduction of the adelgid population. Since A. tsugae is dependent on hemlock for nutrients, feeding on trees in poor health may affect the insect's ability to obtain necessary nutrients and consequently affect their population and physiological health. A cluster analysis, based on quantitative and qualitative tree health measurements, grouped sample trees into categories of lightly and moderately impacted trees. The A. tsugae population health on each tree was determined by measuring insect density, survival from aestivation, and peak fecundity. A. tsugae physiological health was determined similarly by measuring insect biomass, total carbon, carbohydrate, total nitrogen, and amino nitrogen. A. tsugae from moderately impacted trees exhibited significantly greater fecundity; however, A. tsugae from lightly impacted hemlocks contained significantly greater levels of carbohydrates, total nitrogen, and amino nitrogen. All A. tsugae physiological parameters increased significantly over time as the insects matured and reproduced regardless of tree health classification. While the results of the physiological analysis generally support our hypothesis that A. tsugae on lightly impacted trees are healthier than those on moderately impacted trees, this was not reflected in the population fitness measurements of the insects. Further examination of A. tsugae egg health may elucidate this apparent contradiction.

Hemlock woolly adelgid (Adelges tsugae Annand) is causing defoliation and mortality of eastern hemlock (Tsuga canadensis (L.) Carrière) in the eastern United States. The objectives of this study were to quantify changes in tree-ring width and wood anatomy for trees that survived adelgid infestation, and to contrast dendroclimatic relationships across a latitudinal gradient. Six sites spanning the current range of hemlock woolly adelgid (HWA) infestation were selected. At each site, 23 infested eastern hemlocks were cored and two trees were felled at the Virginia site and thin-sectioned using a sliding microtome for analysis of wood anatomy. Tree cores were cross-dated and ring widths were measured. For each site, t-tests were used to determine if there was a difference in radial growth pre- and post-HWA arrival. To compare differences in pre- and post-HWA cell properties, t-tests were used. For dendroclimatic analysis, Pearson correlation coefficients were calculated between radial growth and monthly climate variables. Three sites showed significant suppression in radial growth after HWA arrival and latewood produced post- HWA arrival had significantly smaller cells with reduced cell-wall thickness than latewood produced before HWA arrival. This indicates that HWA can reduce a tree's photosynthate production. The relationship between hemlock growth and climate also varied with latitude and site, with trees growing further south or on shallower soils being more sensitive to moisture levels. This sensitivity to drought can also partially explain the variation in hemlock response to adelgid feeding, as trees affected by moisture stress tend to be more sensitive to insect attack.

Laricobius nigrinus is an insect predator introduced into the eastern United States for the biological control of hemlock woolly adelgid, Adelges tsugae. Laricobius nigrinus dispersal characteristics and predation on A. tsugae were evaluated by monitoring its immature life stages that were distinguished from larvae of the native, L. rubidus using a portion of the cytochrome oxidase subunit one gene. Conventional polymerase chain reaction (PCR) followed by restriction fragment length polymorphism produced DNA fragment sizes that were unique to each species. Real-time PCR used a TaqMan assay with two species-specific nucleotide sequence probes labeled with fluorescence molecule VIC or FAM that indicated the DNA template was L. nigrinus or L. rubidus, respectively. Laricobius nigrinus beetles released in the lower hemlock crown (< 7 m) dispersed to the upper crown (>15 m) for oviposition. Four years post-release, its rate of spread into the surrounding forest was 75 m/yr and the number of L. nigrinus detected on hemlock trees was positively correlated with the density of A. tsugae, regardless of proximity to the release area. Adelges tsugae survival rate was lower on release trees than on geographically separated control trees for three years at a L. nigrinus release site in each Pennsylvania and North Carolina. Yet, the density of A. tsugae remained greater on release trees than control trees. In the eastern United States, the average density of A. tsugae was 3.8 times greater than the upper 95% confidence interval in the native range of L. nigrinus and A. tsugae, as observed in Seattle, WA. This contributed to the average ratio of predator to prey remaining 4.6 times lower than the 95% confidence interval observed in WA. Five to seven years post-release, hemlock health of release and geographically separated control trees declined at similar rates, though the average A. tsugae infestation level was 35% greater on release trees at the time of L. nigrinus release. The

Laricobius nigrinus is an insect predator introduced into the eastern United States for the biological control of hemlock woolly adelgid, Adelges tsugae. Laricobius nigrinus dispersal characteristics and predation on A. tsugae were evaluated by monitoring its immature life stages that were distinguished from larvae of the native, L. rubidus using a portion of the cytochrome oxidase subunit one gene. Conventional polymerase chain reaction (PCR) followed by restriction fragment length polymorphism produced DNA fragment sizes that were unique to each species. Real-time PCR used a TaqMan assay with two species-specific nucleotide sequence probes labeled with fluorescence molecule VIC or FAM that indicated the DNA template was L. nigrinus or L. rubidus, respectively. Laricobius nigrinus beetles released in the lower hemlock crown (< 7 m) dispersed to the upper crown (>15 m) for oviposition. Four years post-release, its rate of spread into the surrounding forest was 75 m/yr and the number of L. nigrinus detected on hemlock trees was positively correlated with the density of A. tsugae, regardless of proximity to the release area. Adelges tsugae survival rate was lower on release trees than on geographically separated control trees for three years at a L. nigrinus release site in each Pennsylvania and North Carolina. Yet, the density of A. tsugae remained greater on release trees than control trees. In the eastern United States, the average density of A. tsugae was 3.8 times greater than the upper 95% confidence interval in the native range of L. nigrinus and A. tsugae, as observed in Seattle, WA. This contributed to the average ratio of predator to prey remaining 4.6 times lower than the 95% confidence interval observed in WA. Five to seven years post-release, hemlock health of release and geographically separated control trees declined at similar rates, though the average A. tsugae infestation level was 35% greater on release trees at the time of L. nigrinus release. The

Prey-mediated effects of imidacloprid were evaluated for Laricobius nigrinus Fender and Sasajiscymnus tsugae Sasaji and McClure after feeding on hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae). Two methods were evaluated for detecting imidacloprid in hemlock tissues: a commercially available enzyme linked immunoassay (ELISA) kit and a high performance thin-layer chromatography technique for detecting and quantifying imidacloprid residues in hemlock wood and needle tissues. ELISA is advantageous because of its cost, sensitivity, and ease of use. However, matrix effects in the form of false positives and overestimated imidacloprid concentrations were evident in hemlock wood and needle tissue extracts. Matrix effects could be reduced by dilution with water, effectively raising the lower detection range of the kit from 0.2 to 200 ppb. High performance thin-layer chromatography was accurate, quick, easy to use, and matrix effects were not evident. However, the technique is sensitive in the lower ppm range and tissue samples from field-treated hemlocks are often in the ppb range, making this technique less desirable than more sensitive analytical methods. Lethal and sublethal effects on both predators were evident after eastern hemlock branches infested with HWA were spiked with imidacloprid in the laboratory. HWA mortality increased with dosage and time, and its 30 d LC50 was determined to be 242 ppb. Both predator species exhibited reduced survivorship and fitness parameters after feeding on HWA from the treated branches. In a topical application bio-assay, 6 d imidacloprid LD50 values for L. nigrinus and S. tsugae were 2.43 and 1.82 µg/g, respectively. Imidacloprid and its major metabolites in hemlock tissues were analyzed by liquid chromatography-tandem mass spectrometry. Imidacloprid recovery from beetle cadavers was correlated with beetle mortality from feeding on treated hemlock branches. Olefin was the primary imidacloprid metabolite rec

Prey-mediated effects of imidacloprid were evaluated for Laricobius nigrinus Fender and Sasajiscymnus tsugae Sasaji and McClure after feeding on hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae). Two methods were evaluated for detecting imidacloprid in hemlock tissues: a commercially available enzyme linked immunoassay (ELISA) kit and a high performance thin-layer chromatography technique for detecting and quantifying imidacloprid residues in hemlock wood and needle tissues. ELISA is advantageous because of its cost, sensitivity, and ease of use. However, matrix effects in the form of false positives and overestimated imidacloprid concentrations were evident in hemlock wood and needle tissue extracts. Matrix effects could be reduced by dilution with water, effectively raising the lower detection range of the kit from 0.2 to 200 ppb. High performance thin-layer chromatography was accurate, quick, easy to use, and matrix effects were not evident. However, the technique is sensitive in the lower ppm range and tissue samples from field-treated hemlocks are often in the ppb range, making this technique less desirable than more sensitive analytical methods. Lethal and sublethal effects on both predators were evident after eastern hemlock branches infested with HWA were spiked with imidacloprid in the laboratory. HWA mortality increased with dosage and time, and its 30 d LC50 was determined to be 242 ppb. Both predator species exhibited reduced survivorship and fitness parameters after feeding on HWA from the treated branches. In a topical application bio-assay, 6 d imidacloprid LD50 values for L. nigrinus and S. tsugae were 2.43 and 1.82 µg/g, respectively. Imidacloprid and its major metabolites in hemlock tissues were analyzed by liquid chromatography-tandem mass spectrometry. Imidacloprid recovery from beetle cadavers was correlated with beetle mortality from feeding on treated hemlock branches. Olefin was the primary imidacloprid metabolite rec

Different Laricobius nigrinus Fender release locations, numbers of predators, and timing of release were evaluated for biological control of the hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae). It established at 59% of the sites and location was the most important factor related with establishment and abundance, HWA density, and hemlock vigor index. Cold locations had poor establishment or low abundance, declines in HWA density, and increases in hemlock vigor over time. Paired release and control sites detected a predator impact on HWA density, but densities remained high and tree vigor declined. The phenology of L. nigrinus, L. rubidus LeConte, and HWA were studied at a field insectary and the species were highly synchronized. A cage exclusion study showed that HWA survival and density were lower and ovisac disturbance was higher when exposed to predation. To improve L. nigrinus monitoring, we compared beat sheets for adults or branch clipping for immatures, and the host searching behavior of L. nigrinus was studied to understand how it locates a tree and HWA. In the Appalachians, beat sheet sampling resulted in false negatives as larvae were collected by branch clipping. Adults orientated to a tree visually, fed when prey were present and flew when absent, and showed different search patterns on infested versus uninfested trees. In Seattle, both sampling methods detected L. nigrinus because the predator was common. Predator : prey ratios were high at heavily infested sites in Seattle and low in the eastern US, where is has been released recently. Partial life tables were constructed for HWA sistentes at four sites for 2 yr in Seattle. Unspecified causes of nymph and adult mortality were high and L. nigrinus was the dominant predator of ovisacs. Adult L. nigrinus abundance was positively related to HWA density and immature abundance was related to ovisac density, indicating an aggregation and numerical response to its prey. Laricobius nig

The biology and host-specificity of Laricobius kangdingensis sp. n. (Coleoptera: Derodontidae), a new predator of hemlock woolly adelgid (HWA), Adelges tsugae Annand (Homoptera: Adelgidae) were investigated and compared with that of a related predator of HWA, L. nigrinus Fender. Adults became active in June 2003; the males emerged first and in greater numbers than the females. The sex ratio of female to male was 1: 1.1. Laricobius kangdingensis sp. n. has four larval instars. The total number of eggs laid between 7 July 2003 and 7 August 2004 was 1327. Mean lifetime fecundity per female was 196.4 ± 53.4 eggs and 97.9 eggs for the P1 (parental gen.) and F1 generations, respectively. In comparison, lifetime fecundity for F1 L. nigrinus was 100.8 ± 89.6 eggs. Developmental rates were studied in fall 2003 and spring 2004. Combined with data gathered previously at 12 and 15ºC in early 2003, the 6, 9, and 18ºC data were used to determine lower temperature development thresholds for the egg (2.8ºC), larval (1.6ºC), and pre-pupal (5.8ºC) stages. The egg and larval thresholds are lower than those observed for L. nigrinus. Laricobius kangdingensis sp. n. individuals completed development only at 12 and 15ºC; however, these adults did not emerge from aestivation after pupation. Laricobius kangdingensis sp. n. appears to be host-specific. In paired-choice feeding tests, adults consumed significantly more HWA than two other adelgid species [eastern spruce gall adelgid, Adelges abietis (L.), and pine bark adelgid, Pineus strobi (Hartig)], the pine needle scale, Chionaspis pinifoliae (Fitch), and the woolly alder aphid, Paraprociphilus tessellatus (Fitch). In paired-choice oviposition tests, females laid more eggs on HWA-infested eastern hemlock than on any other test prey. In no-choice host suitability experiments larvae fed on HWA; eastern spruce gall adelgid; the larch adelgid, A. laricis

Hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae), is an introduced pest injurious to hemlock trees (Tsuga spp.) in the eastern United States. HWA currently infests hemlock in over 50% of its geographic range and has the potential to spread throughout its entire range. Since HWA populations in the eastern United States are not regulated effectively by natural enemies (Wallace and Hain 2000), classical biological control is the most promising option for controlling this pest in the forest setting. This work evaluates Laricobius nigrinus Fender (Coleoptera: Derodontidae), a predator associated with HWA in western North America, is being evaluated as a potential biological control agent for HWA (Cheah et al. 2004). Observations suggest that natural enemies may help keep HWA populations below injurious levels in the western United States (Mausel, pers. comm.). Laboratory studies have revealed that L. nigrinus feed selectively on HWA and field studies in British Columbia, Canada have shown that its lifecycle is highly synchronous with HWA phenology. This predator is being further evaluated for its suitability as a biological control agent. In order to be a viable candidate, L. nigrinus must survive under natural field conditions in the eastern United States, reduce the density of HWA, and it must be amenable to mass rearing. The survival, reproductive capability and predator efficiency of L. nigrinus were assessed over 3 years in the field. In addition, the environmental factors regulating processes in the life cycle to develop and improve rearing procedures for this insect were identified. In the first year, adults survived from February - May, laid up to 41 eggs/beetle, and consumed approximately 4.3 adelgids per day. In the second season, L. nigrinus adults survived from November - April, laid up to 38 eggs/beetle and consumed approximately 4.5 adelgids per beetle throughout the study. In both seasons, adelgid populations were significantly lo

Decline of the health of hemlocks in Delaware Water Gap National Recreation Area was noticeable in the southern areas of the park by 1992. The following year, a series of plots were established to monitor hemlock health and the abundance of hemlock woolly adelgid. This poster examines only the health rating of the hemlocks in the monitoring plots.

The hemlock woolly adelgid (HWA), Adelges tsugae Annand (Homoptera: Adelgidae), is forest pest introduced to eastern North America in the early 1950's. Although this pest occurs on both landscape and nursery stock as well as in natural stands of hemlock forest, pesticides are only practical and effective in urban settings. Ecological and economical considerations prevent utilization of chemical treatment in the forest setting, thus biological control is viewed as the most promising option for slowing the spread of HWA. It is essential for researchers to be able to accurately assess the population levels of biocontrol agents after release into the environment. No method currently exists for sampling HWA predators. This project was designed to determine whether two species of predators are able to utilize olfactory cues from eastern hemlock and/or HWA in host-finding. If predators use olfactory cues, we may develop an attractive sythnetic blend of compounds to draw them to a trap, thereby simplifying the sampling and improving its accuracy. To address this question we executed three experiments. The first involved examination of the antennae of the predators for the presence and abundance of olfactory sensilla. The second experiment was designed to detect a behavioral response by the predators following exposure to host volatile compounds. The final experiment involved identifying compounds emitted from eastern hemlock, and the affect of HWA-feeding on volatile emissions. Laricobius nigrinus Fender (Coleoptera: Derodontidae) antennae are densely populated with sensilla, several of which are potentially olfactory in function. In addition, we observed a behavioral response to olfactory cues which included altered flight behavior. However, the behavior was not clearly attraction. Pseudoscymnus tsugae Sasaji and McClure (Coleoptera: Coccinellidae) has few sensilla on a very short antennae and only one type of sensilla possesses wall pores suggestive of an olfactory functi

The biology of Laricobius nigrinus Fender (Coleoptera: Derodontidae) and its potential for control of hemlock woolly adelgid (HWA), Adelges tsugae Annand (Homoptera: Adelgidae) in the eastern United States were investigated. Laricobius nigrinus completed development on HWA in laboratory studies. There are four larval instars. Mean larval consumption was 225.9 and 252.3 HWA eggs at 12 and 18 °C, respectively. Post-aestivation activity period was 36.6 and 30.8 weeks for males and females at 13 °C, respectively. Mean lifetime fecundity was 100.8 eggs over a mean ovipositional period of 13.2 weeks. Laricobius nigrinus is host specific. In paired-choice and no-choice oviposition tests, L. nigrinus laid more eggs in HWA ovisacs than other test species. Laricobius nigrinus consumed more eggs of HWA than eggs of Adelges piceae (Ratzeburg) and Pineus strobi (Hartig) but not of Adelges abietis (L.). In larval development tests, L. nigrinus only completed development on HWA. A two-year field study conducted in British Columbia showed that: 1) the life cycle of HWA in British Columbia is similar to that previously reported in Virginia and Connecticut; 2) L. nigrinus adults undergo an aestival diapause that coincides with the diapause of the first instar HWA sistens. Adult activity beginning in autumn coincides with resumption of development of the sistens generation; 3) oviposition and subsequent larval development of L. nigrinus coincide with oviposition by HWA sistens adults. Temperature-dependent development was determined for egg, larval, pre-pupal and pupal stages of L. nigrinus at five constant temperatures (9, 12, 15, 18 and 21 °C). Development time was inversely proportional to temperature between 9 and 18 °C. Laricobius nigrinus did not complete development at 21 °C. Minimum developmental temperatures for eggs (6.5 °C), larvae (5.1 °C), pre-pupae (3.8 °C), and pupae (3.1 °C) were determined by extrapolation of linear regression equations to the x-intercept. Degree-days