The hemlock woolly adelgid (HWA), Adelges tsugae Annand, (Hemiptera: Adelgidae) is an invasive species in the eastern United States and Canada. This scale-like insect feeds on the xylem ray parenchyma cells of eastern hemlock, Tsuga canadensis (L.) Carrière, (Pinales: Pinaceae) and Carolina hemlock, Tsuga caroliniana Englem., resulting in multiple physical and physiological symptoms, all of which commonly leads to tree mortality within 4 to 10 years. Currently, HWA has spread throughout most of the eastern hemlock range and all of the range of Carolina hemlock and threatens the forest ecosystems they serve. In its introduced range, there are no natural enemies specialized to feed on HWA and suppress HWA populations below damaging levels. In the urban environment and at some public locations, the use of chemicals, such as imidacloprid, have been used to temporarily protect hemlocks, however; this does not provide a permanent solution. In the 1990's a classical biological control program for HWA was created with the goal to suppress HWA populations below damaging levels to reduce tree mortality, and to reduce the negative effects associated with the loss of either hemlock species. Presently, four predators have become the main focus of this program: Laricobius nigrinus Fender, Laricobius osakensis Shiyake and Montgomery (Coleoptera: Derodontidae), Leucotaraxis argenticollis (Zetterstedt), and Leucotaraxis piniperda (Malloch) (Diptera: Chamaemyiidae). Throughout the eastern US, La. nigrinus has established populations and expanded its distribution beyond where it has been released. Although it has successfully established and continues to spread, its effect on eastern hemlock health has been unknown. A 1-year study, carried out in 2021, revealed that La. nigrinus predation on the sistens generation not only reduced the sistens generation, but also had a similar net photosynthetic rate to the negative control on 1-2 year old shoot growth. In addition, significantly more shoots were produced for treatment branches with the highest La. nigrinus density. This also led to a significantly reduced aestivating sistens density compared to the no predator treatment. However, predation was insufficient to reduce HWA populations below the suggested damaging threshold of eastern hemlock (< 4 HWA/cm), suggesting that additional predators would be needed. Recently, the western strain of Le. argenticollis had been released in the eastern US. However, its phenology in the eastern US was unknown. A 2-year study, from 2021-2023, revealed that Le. argenticollis eggs and larvae were present when both generations of HWA adults with eggs and HWA nymphs were present. In addition, Le. argenticollis was capable of completing development in NY and VA, showing promise that this species could establish populations in the eastern US. Timing the adult fly releases into cages at the time when sistens were producing eggs resulted in Le. argenticollis phenology to synchronize well with HWA phenology at both locations, thus providing additional support for its release in the eastern US. Another 2-year study conducted in 2021-2022, at five locations (MD, NC, PA, TN, and VA), sought to determine how the predation of La. nigrinus and Le. piniperda, alone and together, would impact the sistens and progrediens generations, as well as how their predation would affect the aestivating sistens density and new shoot growth production. From this study, La. osakensis was found as the dominant Laricobius species present at the TN site, and similarly to La. nigrinus, reduced intact ovisac density in. This provided a small preview in La. osakensis capabilities and how this predator may be impacting HWA populations at locations where it has established. In order to assess Le. piniperda's impact on the progrediens generation, mesh cages were applied to treatment branches at each site. The study found that Le. piniperda could reproduce at all field sites and that the percentage of disturbed progrediens was significantly higher for treatments with Le. piniperda present at all field sites, except at PA in 2022. However, the affect of both of these predators on the aestivating sistens density and eastern hemlock health was inconclusive. In addition, the percentage of new shoot growth was highly variable among treatments at all field sites, and there was no treatment effect on the mean length of new shoot growth produced at all field sites. The results of this study suggests that in order to fully understand the affects of these predators on HWA populations and on hemlock tree health, longer term studies should be conducted. However, this study does provide some insight on the capabilities of these predators in the short term and supports the continued release and redistribution of these predators. / Doctor of Philosophy / The hemlock woolly adelgid (HWA), is an invasive species in the eastern United States and in Canada. This small scale-like insect was introduced likely on nursery stock material from southern Japan and was first discovered in the eastern US in 1951 in Richmond, Virginia. Two generations of HWA occur each year known as the sistens and progrediens; both reproduce without males. This insect feeds on eastern hemlock and Carolina hemlock, and since its discovery has spread through most of their range. While this insect feeds, it weakens the tree, eventually causing tree death. In eastern North America, eastern hemlock is one of the major tree species in the forest and provides a unique habitat for various mammals, birds, fish, and invertebrate species. To protect this habitat, multiple management strategies have been developed. The two main management strategies that are in use are the use of pesticides and the release of predators of HWA. Pesticides have been useful in the urban environment and in parks and forests, but they only last for a short time and can lead to negative affects to the environment. Since the 1990's, the release of insect predators originally found in the native range of HWA, has occurred with the goal of predator establishment, to lower HWA populations, and to lower hemlock tree death in the eastern US. Four small insect predators have been the main focus of this program: Laricobius nigrinus, Laricobius osakensis, Leucotaraxis argenticollis, and Leucotaraxis piniperda. So far La. nigrinus and La. osakensis have populations at multiple locations throughout the eastern US, and a low number of sites have been recently found for Le. argenticollis and Le. piniperda. Past studies have indicated that La. nigrinus has a significant impact to the sistens generation of HWA, but its affect to eastern hemlock health has been unclear. A 1-yr study conducted in VA in 2021, released La. nigrinus adults into mesh cages, with two different population sizes of beetles, to determine if La. nigrinus predation on HWA populations lead to an indirect effect on eastern hemlock health at the branch level. Results confirmed that La. nigrinus had a significant impact to the sistens generation of HWA. Photosynthesis of branches with La. nigrinus, in June, was similar to the photosynthesis of the negative control branches, which started with a lower number of sistens present on the branch at the beginning of the study. Suggesting that branches with La. nigrinus may be healthier compared to branches without La. nigrinus. Laricobius nigrinus predation also significantly reduced the number of aestivating sistens on new shoot growth, but not below damaging levels (< 4 HWA/cm). This suggested that additional predators may be needed to target the progrediens generation of HWA so that HWA populations do not return to damaging levels after La. nigrinus predation. To learn more about the potential for Le. argenticollis to survive and reproduce in the eastern US, a 2-yr study was conducted in 2021 – 2023, to determine when different life stages of Le. argenticollis could occur in NY and VA. Results revealed that Le. argenticollis was able to reproduce in the mesh cages and complete its life cycle. In addition, Le. argenticollis eggs and larvae were present at the time when HWA adults were producing eggs and when HWA nymphs were present for both generations of HWA. These results provide insight on Le. argenticollis life cycle and shows that Le. argenticollis has a good chance in surviving throughout the eastern US. A 2-yr study at five locations in the eastern US (MD, NC, PA, TN, and VA), tried to combine two predators, La. nigrinus and Le. piniperda, to determine if their combined impact would be sufficient to reduce HWA populations and affect eastern hemlock tree health. Three assessments were conducted each year to determine the impacts of La. nigrinus and Le. piniperda on HWA populations and on eastern hemlock shoot growth production as a representation of tree health. Results confirmed that La. nigrinus significantly reduced the sistens generation at all locations, except at TN. At the TN site, La. osakensis was found to be the main predator of the sistens generation and had a similar affect on the sistens generation as La. nigrinus. Leucotaraxis piniperda was able to reproduce at all locations and also had a significant affect on progrediens ovisacs at four of the locations in 2021 (MD, NC, PA, and VA) and at three of the locations in 2022 (MD, NC, and VA). However, the results on the number of aestivating sistens on new shoot growth and eastern hemlock health were inconclusive. Therefore, it is unclear as to how the presence of both of these predators' affect eastern hemlock health. A long term study may be needed in order to understand the relationship between these predators and their indirect affect to eastern hemlock health.

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 calculated for eggs, larvae, pre-pupae, pupae, and egg to adult were 54.6, 161.3, 196.1, 212.8, and 666.7, respectively. The laboratory derived degree-day model for predicting egg hatch was validated with field data collected in British Columbia. Observed median egg hatch at three of four sites over two years was within 15% of the predicted DD value (54.6). / Ph. D.

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. / Master of Science in Life Sciences / Hemlock woolly adelgid (HWA), an invasive insect native to Asia and western North America, is a significant threat to two native hemlock species in the eastern U.S. Since 2003, a predatory beetle, Laricobius nigrinus, has been released for management of HWA. In biological control programs such as this one, it is important to evaluate the ability of predators to establish and impact prey in areas where they are introduced, as this helps guide future management decisions. As such, a study was conducted to examine the ability of L. nigrinus to establish after being released at 26 locations in Virginia. In those investigations we found that L. nigrinus established at 82% of the sites. Although promising, longer-term studies are needed to understand if the predation of HWA by L. nigrinus helps improve the health of our native hemlocks. A separate two-phase study was carried out from 2014-2016 and 2016-2018 evaluating the impact of L. nigrinus on HWA at nine release sites in the eastern U.S. In Phase Two reported here, cages were used to exclude L. nigrinus on some HWA infested branches while on others, no cage was applied to allow free access to HWA. Comparisons between branches were made to determine the level of predation by L. nigrinus. These studies showed a significant impact by L. nigrinus on the winter generation of HWA with as many as 80% of those insects being attacked on study branches which indicates that this species has potential as an effective predator.

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. / Master of Science in Life Sciences / <i>Laricobius nigrinus</i> is a beetle predator of hemlock woolly adelgid (HWA), an aphid like insect covered by wool. HWA is a serious pest of both eastern and Carolina hemlock, two important tree species found in Appalachia which provide species diversity and habitat to numerous animal and plant species. Damage to hemlock trees include death, dieback, and disease. We began releasing <i>L. nigrinus</i> in 2003 in the eastern United States to manage HWA. There are now over 900 documented releases of <i>L. nigrinus</i> from Maine to Georgia. Nine field sites were set up to assess the impact <i>L. nigrinus</i> is having on HWA populations throughout the geographic range of its releases. At many of these sites we found that <i>L. nigrinus</i> adults and larvae are significantly reducing HWA populations. A large proportion of the predators of HWA recovered at these sites are <i>L. nigrinus</i>, which shows that this species is primarily responsible for the observed predation. The second study conducted assessed for establishment of <i>L. nigrinus</i> at release sites around Virginia. <i>Laricobius nigrinus</i> was found at four of the fourteen sites sampled. Continued sampling is necessary to get a more accurate assessment of establishment since many of the sites were recovering from low HWA populations resulting from extreme cold temperatures in 2014 and 2015.

Hemlock woolly adelgid, Adelges tsugae, native to Asia and the Pacific Northwest of North America (Pacific Northwest), has devastated eastern hemlock (Tsuga canadensis) in a major portion of its range in the eastern U.S. After many years and much effort directed towards rearing and releasing biological control agents to manage HWA, one of these agents, Laricobius nigrinus, native to the Pacific Northwest, is now well-established in sites from the southern to the mid-Atlantic states of the eastern U.S. However, there have yet to be studies of its efficacy in lowering A. tsugae densities, and there has been no noticeable drop in A. tsugae densities. Population models for A. tsugae have suggested that even upwards of 90% predation on eggs laid by the overwintering generation will have minimal effect in reducing the population densities of A. tsugae, if A. tsugae are at high density, due to compensatory density-dependent survival in the progrediens generation. Additionally, no studies showing insect predators are indeed what regulate A. tsugae in its native range exist. We established predator exclusion experiments, and recorded A. tsugae densities, mortality factors, and fecundity for multiple generations in both the native and invaded ranges. In the invaded range, we studied A. tsugae populations in sites with well-established populations of L. nigrinus to test its efficacy in reducing A. tsugae and tested model predictions of minimal difference in A. tsugae densities between treatments with and without predators. In the Pacific Northwest we tested the effect of insect predators and tree species, western (Tsuga heterophylla) and eastern hemlock, on populations of A. tsugae. In the invaded range we found that L. nigrinus predation was significantly higher in unbagged branches, however, model predictions were validated, and there was no effect of predation by treatment on the A. tsugae summer generation. In our plots in the Pacific Northwest we found that tree effects were not significant, but that summer-active predators were significantly lowering levels A. tsugae densities on unbagged branches. Our study demonstrates the importance of summer-active predators in reducing A. tsugae and suggest that summer- and winter-active predators are needed to suppress A. tsugae to innocuous densities.

Competitive interactions among two specialist predators, <i>Laricobius nigrinus</i> Fender (Coleoptera: Derodontidae) and <i>Sasajiscymnus tsugae</i> Sasaji and McClure (Coleoptera: Coccinellidae), and a generalist predator, <i>Harmonia axyridis</i> Pallas (Coleoptera: Coccinellidae), of hemlock woolly adelgid were evaluated using laboratory, field and video studies. The two specialist predators are part of a biological control program for <i>A. tsugae</i>, and the potential for competition among these species and previously established generalist predators is unknown. In laboratory studies of predator groups in Petri dish assays, the only significant negative effects from competition occurred among conspecifics, resulting in reduced net egg production by <i>L. nigrinus</i> and <i>H. axyridis</i> and reduced feeding by <i>H. axyridis</i>. In contrast, heterospecific combinations showed non-interference. In longer duration field studies of predator groups, held in branch enclosures, predator survival and feeding were not significantly affected by additional predators. Net reproduction was again significantly reduced by conspecifics, while heterospecifics showed non-interference for all predator responses. All predators reduced the number of <i>A. tsugae</i> nymphs of the next generation relative to no-predator controls; however, <i>L. nigrinus</i> had much greater impact overall due to the large number of progeny produced. Video studies revealed that predator behavior varied qualitatively and quantitatively by species, and did not appear to be coordinated temporally or spatially. All species exhibited continuous activity patterns that were punctuated by longer periods of rest. The specialist predators were more selective of feeding and oviposition sites, and rested at more concealed locations than <i>H. axyridis</i>. Conspecifics significantly altered the time allocated to specific behaviors for <i>L. nigrinus</i> and <i>H. axyridis</i>, resulting in reduced predator effectiveness due to increased searching and decreased feeding and oviposition. All predator groups maintained a high degree of spatial separation relative to assay size, suggesting that chemical or tactile cues may be used to regulate their distributions. Overall, these studies suggest that the three predator species will be compatible in this system. Management implications include using multiple-predator species combinations over single-species for biological control of <i>A. tsugae</i> and implementing low-density releases to reduce the potential negative effects of intraspecific competition. / Ph. D.

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 lower on branches with predators than those without predators. This impact was demonstrated on both the winter and spring generation of HWA. The feasibility of caged field releases of L. nigrinus was determined in a third field study. An estimated 10, 000 L. nigrinus eggs were liberated in field cages in spring 2003. The density of adelgids in the subsequent (spring) generation of the adelgids was significantly lower on branches with larval activity than those with no predators. Despite extensive sampling, no F1 adults were observed, however F2 adults were recovered in the fall of 2004, 20 months after release. The L. nigrinus life stages incurring high mortality during rearing were identified and factors affecting survival in the feeding and non-feeding life stages were investigated. Studies on the survival and feeding of adults, length of ovipositional period, density per cage, and survival of larvae were conducted. The effect of type of pupation medium, moisture level, disturbance, soil sterilization, temperature, and photoperiod on survival of the non-feeding stages and time of emergence from aestivation was examined. The most noteworthy finding, regarding rearing, is that aestivation can be extended by storing adults at high temperatures and long daylength throughout the summer and decreasing the temperature and daylength in the fall. Based on the results of these studies, procedures for rearing L. nigrinus have been developed and it is currently being reared at two other institutions. To date, over 8, 000 adults produced at Virginia Tech have been released in 6 states and both F1 and F2 adults have been recovered from multiple locations. / Ph. D.

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 full efficacy of L. nigrinus predation was likely underestimated because this study assessed experimental release sites. Interactive roles of biotic and abiotic pressures on A. tsugae should continue to be evaluated at L. nigrinus introduction sites. / Ph. D.

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 function. In addition, we did not observe a significant behavioral response to host-volatiles. It seems unlikely that this species uses olfaction in long-range host location. We identified 10 monoterpenes that were consistently expressed in the hemlock volatile profile and were unable to isolate volatile emissions from HWA. There is an increased monoterpene release rate from HWA-infested hemlock foliage as compared to uninfested foliage apparently driven indirectly by HWA through a reduction in new growth at branch tips. In addition there was a slight but statistically significant change in the percent composition of the individual compounds. We see potential in developing a more efficient sampling procedure for L. nigrinus through utilization of olfactory cues. More biological assays must be conducted to determine whether an attractive blend exists and electrophysiological assays are required to isolate physiologically active compounds. However, our data suggest that P. tsugae is not likely to be reliant on olfaction in long-range host location. / Master of Science

The biology and host-specificity of <i>Laricobius kangdingensis</i> sp. n. (Coleoptera: Derodontidae), a new predator of hemlock woolly adelgid (HWA), <i>Adelges tsugae</i> Annand (Homoptera: Adelgidae) were investigated and compared with that of a related predator of HWA, <i>L. nigrinus</i> 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. <i>Laricobius kangdingensis</i> 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 <i>L. nigrinus</i> 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 <i>L. nigrinus</i>. <i>Laricobius kangdingensis</i> sp. n. individuals completed development only at 12 and 15ºC; however, these adults did not emerge from aestivation after pupation. <i>Laricobius kangdingensis</i> 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, <i>Adelges abietis</i> (L.), and pine bark adelgid, <i>Pineus strobi</i> (Hartig)], the pine needle scale, <i>Chionaspis pinifoliae</i> (Fitch), and the woolly alder aphid, <i>Paraprociphilus tessellatus</i> (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, <i>A. laricis</i> Vallot; the balsam woolly adelgid, <i>A. piceae</i> (Ratzeburg); pine bark adelgid; and pine needle scale, but not on woolly alder aphid or elongate hemlock scale, <i>Fiorinia externa</i> Ferris. Although no larvae completed development in any of the bioassays, 6 out of 15 larvae (40%) in the HWA treatment reached the pre-pupal stage. One larva out of 6 reached the pre-pupal stage on pine bark adelgid. / Master of Science