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Tsuga canadensis (L.) Carr. (eastern hemlock), a long lived and shade tolerant coniferous tree species native to eastern North America, is currently threatened by the hemlock woolly adelgid (Adelges tsugae Annand, HWA). This exotic, invasive insect poses a serious threat to T. canadensis stands throughout their native range. The loss of this unique tree species is often coupled with numerous ecological consequences. HWA-induced mortality has exceeded 95% in some forest types, and will likely continue to decimate T. canadensis populations in the coming years. Shortly after HWA was found in Great Smoky Mountains National Park (GSMNP), long-term monitoring plots were established across a gradient of T. canadensis associated forest communities to monitor HWA impacts and stand decline. Our objectives were to (1) determine how T. canadensis populations have changed in conjunction with HWA infestation, and (2) document how HWA has affected vegetative community dynamics following five to six years of infestation in GSMNP. We assessed T. canadensis associated overstory communities and determined that crown density, top dieback, and T. canadensis stem density has significantly declined over the observational period. Furthermore, we found that midstory T. canadensis trees were experiencing greater rates of decline than those of the overstory. Even with these losses, overstory and midstory community composition did not significantly change from 2003 to 2008/09. Many studies have documented rapid pulses of understory vegetative productivity in conjunction with T. canadensis defoliation, which often allows increased amounts of solar radiation to reach the forest floor. Although we observed significant T. canadensis mortality at most plots sampled, understory species composition did not significantly change following HWA infestation. We attribute this lack of species response to the overwhelming abundance of Rhododendron maximum in the mid/understory, which restricts increased light from reaching the forest floor despite the formation of small overstory gaps. However, we found that understory composition significantly varied among ecological groupings, and differences were largely associated with soil properties and elevation. Overall, results of this project suggest that T. canadensis may largely disappear from southern Appalachian forests resulting from infestations of HWA. Although community composition of T. canadensis associated stands has not yet been altered, changes are likely to occur as duration of infestation progresses.

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Thesis (Ph. D.)--Ohio State University, 2007. / Title from first page of PDF file. Includes bibliographical references (p. 157-170).

Thesis (M.S.)--West Virginia University, 2007. / Title from document title page. Document formatted into pages; contains vii, 55 p. : ill., maps. Includes abstract. Includes bibliographical references (p. 47-50).

An invasive insect, hemlock woolly adelgid (HWA), has initiated widespread hemlock decline and mortality in the Great Smoky Mountains National Park (GSMNP). Riparian hemlock mortality impacts on vegetative and aquatic systems of first-order, headwater streams were evaluated. Reference sites for this study were representative of the best available conditions within the GSMNP, with initial stages of HWA presence. Impacted sites were defined as areas with over 90 percent hemlock mortality. Impacted streams had decreased canopy coverage and increased light availability. Residual red maple, yellow birch, and sweet birch capitalized on the loss of hemlock, with increases in relative basal area and species importance values. Rosebay rhododendron responded with increased density and height at impacted sites, thereby preventing woody regeneration. Since long-term regeneration and post-mortality canopy recruitment are limited, alterations of vegetative composition and structure in the stream riparian zone are expected. A seasonal assessment (Sept. 2009 – March 2010) of aquatic impacts revealed increased diurnal variation in stream temperature and exhibited cooler temperatures during the colder months than reference streams. Impacted streams exhibited greater amounts and larger size classes of large woody debris (LWD). Higher concentrations for several nutrients, including silicon, sodium, potassium, magnesium, chlorine, and copper were detected in impacted streams than detected in reference streams. Impacted streams were characterized by higher pH and increased acid neutralization capacity, while reference streams exhibited nitrate concentrations three times higher than impacted stream concentrations. Reference streams were experiencing the initial stage of HWA-induced defoliations, increasing stream nitrate concentrations, while impacted sites had levels suggesting nitrate concentrations have returned to pre-infestation levels. A seasonal assessment of macroinvertebrates found species diversity, abundance, and taxa richness were not affected by hemlock mortality. Impacted streams had a lower density of Chironomidae and Ephemeroptera when compared to reference streams, while Pleuroceridae snails were virtually eliminated at impacted stream sites. Reference streams were dominated by the collector/filter functional feeding group (FFG), while impacted streams were dominated by the scraper FFG. Hemlock mortality induced by the presence of HWA has resulted in short-term impacts to vegetative and aquatic dynamics in stream riparian areas of the Great Smoky Mountain National Park.

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Environmental stressors such as nutrient deficiency and insect infestation can significantly impact tree health. Despite much research on the ecological effect on forests in the northeastern United States due to calcium depletion and hemlock woolly adelgid infestation, little is known regarding the physiological mechanisms altered by these stress factors. I tested the hypothesis that calcium depletion, associated with sugar maple decline, compromises water transport processes as a result of calcium-related reductions in cell growth and stabilization. A survey of forest-grown sugar maples from a long-term replicated calcium-manipulation study showed no significant impact of calcium deficiency on wood density, stem hydraulic conductivity (Ks), or vulnerability to cavitation (VC). In vitro removal of xylem-bound calcium showed no impact on VC or air seeding thresholds (Pt). Results suggest that sugar maple decline is not caused by compromises in xylem function due to calcium deficiency. I also tested the hypothesis that hemlock woolly adelgid (Adelges tsugae Annand) (HWA) infestations impact water transport processes and nutrient partitioning in eastern hemlock trees. HWA infestation resulted in higher Ks due to an increase in average tracheid lumen area associated with the proliferation of false rings. HWA-infested trees exhibited higher rates of net photosynthesis and significant changes in foliar nutrient partitioning. These results are the first to demonstrate increases in Ks and alterations in foliar cation levels in response to HWA infestation. In two additional studies, I investigated methods for evaluating the structure and function of xylem networks. Using sequential sectioning of aerial roots of epiphytic aroids, I directly quantified the topographic relation of vessels in a single organ with measurements of vessel length, diameter, vessel end overlap length, and vessel stelar orientation. In a separate study, I explored the relationship between vessel length and measurements of Pt. In establishing guidelines for estimating whole-stem cavitation with the use of single vessel air injection, I demonstrate that calculations of Pt are influenced by stem length measured and removal of native emboli prior to testing. Improvements in tools to quantify xylem structure and function will enhance our ability to understand the responses of forest trees to environmental stress.

The hemlock woolly adelgid (HWA) (Adelges tsugae [Annand]) (Hemiptera: Adelgidae) is a non-native lethal pest to eastern hemlock (Tsuga canadensis [L.] (Pinales: Pinaceae), and Carolina hemlock (Tsuga caroliniana [Engelmann]) (Pinales: Pinaceae). In the early 2000's, a robust biological control initiative using Laricobius spp. (Insecta: Coleoptera) was launched with the goal of decreasing HWA's impact to hemlock and the associated ecosystems on landscape level. Since then, two Laricobius agents, Laricobius nigrinus Fender (Coleoptera: Derodontidae) and Laricobius osakensis Montgomery and Shiyake (Coleoptera: Derodontidae) have been evaluated, approved, and released. The production and subsequent release of these agents has largely occurred through mass rearing efforts by Virginia Tech. A descriptive analysis of these rearing and release data over the past 16 years at Virginia Tech revealed that the mass production of these agents has been consistently limited by: 1) excessively high rates of subterranean mortality and 2) early subterranean emergence. Very little is known about the subterranean life cycle of these species in the field in terms of survivorship and timing of emergence. Additionally, the thermal limits and rates of development are unknown for L. osakensis and were thus further investigated. In 2019, surveys within the urban environment in two counties in southwest Virginia were conducted to determine if these agents have dispersed from their original release location. Laricobius nigrinus was documented to have established outside of release sites in 100 and 75% of the surveyed grids containing HWA infested hemlocks. The establishments of these species in easily accessible locations allowed for the use of field-caught larvae, in addition to laboratory-produced larvae, for experiments. Experiments were conducted in 2019-2020 to determine the overall subterranean survivorship in relation to site condition, differences in survivorship between field-caught vs. laboratory-reared, and the seasonal timing of emergence. We found that the average field subterranean survivorship (17.1 ± 0.4%) was significantly less than the historical production of these agents in the laboratory (37.5 ± 13.6%). We found that as soil pH and organic matter depth increased, survivorship decreased. While the majority of emergence occurred as expected in the fall, a surprising amount of emergence also occurred in spring and summer. The rate of development and the minimum threshold temperature for each life stage were determined through constant temperature experiments, and were used in the construction of a phenological forecast model. Laricobius osakensis minimum threshold temperature (3.6°) from egg to adult was similar to that of other Laricobius spp., but completed development at higher a constant temperature (22°C). Our results suggest L. osakensis can handle the same colder climates as L. nigrinus, but may be better adapted for warmer locations than L. nigrinus. Laboratory rearing, establishment, subterranean survivorship, and thermal requirements of Laricobius spp. (Coleoptera: Derodontidae), biological control agents released for Adelges tsugae (Hemiptera: Adelgidae) / Doctor of Philosophy / Eastern and Carolina hemlocks are long lived, shade tolerant evergreen tree species that support a unique ecological assemblage of plants and animals. However, following the accidental introduction of a highly invasive aphid-like insect pest first found in the 1950s, known as the hemlock woolly adelgid (HWA), significant mortality of hemlocks in the eastern U.S. was eventually observed. In order to mitigate this pest's impact and to prevent the further collapse of this unique ecological system, a robust management program was initiated. Today, the primary management strategy to reduce HWA's deleterious effects is through the use of chemical and biological treatments. Biological treatments in the form of insect biological control agents, have focused on the use of two closely related beetle species in the genus Laricobius spp. These species were imported from their respective native ranges, biologically evaluated, and approved for release by federal and state agencies. Since then, the beetles have been mass-produced by multiple governmental and academic agencies and released on infested landscape on mostly public land. Key aspects of these species' biology in terms of their population patterns, development, distribution capabilities, and potential supplementary diet remains unknown. Therefore, investigations into these aforementioned aspects were conducted from 2018-2021 in both laboratory and field experiments. In order to supply beetle larvae for field experiments and to better understand how well these predatory beetles were establishing across the landscape, we investigated heavily urbanized areas for their presence. We found that L. nigrinus was established outside of release locations, in 100 and 75% of the surveyed grids containing HWA in southwest Virginia. Laricobius spp. spend a significant amount of their lives in a presumed dormant state beneath the soil during the summer months. Observations over the past 17 yrs. of rearing Laricobius spp. revealed relatively high rates of subterranean mortality and subterranean early emergence, both of which are substantial mortality factors. Through field experiments, the survivorship of these beneficial beetles was found to be 17.1 ± 0.4% and their seasonal emergence was not limited to fall.Comparing the average survivorship from our field study (17.1 ± 0.4% ) to that of the past 17 yrs. of laboratory production (37.5 ± 13.6%), there was significantly less survivorship. The developmental and phenological biology of one of the two Laricobius spp., L. osakensis was determined and found to be similar to that of other beetles. However, L. osakensis was able to complete development at 22°C, the highest developmental temperature recorded for any Laricobius spp. The similar phenological biology of these two predators will allow for precise larval sampling as a metric for establishment. Results and conclusions from these data have provided further knowledge to scientists, land managers, and public stakeholders regarding the benefits of Laricobius spp.

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