<div> <p>Understanding the dispersal ability of invasive insects provides useful insights for developing effective management strategies. Historically, methods for marking insects for dispersal studies have been expensive, time-consuming, and labor-intensive, especially for woodboring beetles. In addition, capturing or rearing insects requires human handling, which can alter behavior. Immunomarking is a well-established technique for studying the dispersal of insects, however, it has not been broadly applied to woodborers. This study evaluates the potential for using immunoproteins applied directly to woodborer-infested trees to mark emerging beetles. <a>Specifically, in the first experiment I sprayed varying concentrations of ovalbumin (egg white) solution directly onto logs infested with emerald ash borer (EAB, <i>Agrilus planipennis </i>Fairmaire) (Buprestidae: Agrilini) and ELISA was used to detect the presence of protein on emerged beetles. To test the persistence of the mark, I applied varying concentrations of albumin to freeze-killed beetles, mounted them on pins, and placed them in an exposed location outdoors. Adult EAB self-marked as they emerged from protein-treated trees, with higher protein concentrations persisting for longer on the cuticle when exposed to sun and rain. </a>This technique offers a convenient, inexpensive and durable means of marking woodborers and circumvents the need for human handling, allowing for more natural behavior and more realistic estimates of dispersal. Protein self-marking may find application in studies of woodborer dispersal within natural forest environments.<a></a></p> </div> <br>

<div>Native and invasive bark and ambrosia beetles threaten the health and productivity of natural and planted forests worldwide. Management of these pests relies on semiochemical-based tactics, but these are often ineffective at monitoring for incipient populations or decreasing pest populations. The role of fungal and non-host volatiles in colonization behavior remains unknown for many important bark and ambrosia beetle species, thereby hindering their control. In this dissertation, I tested the hypothesis that fungal and tree-associated volatiles influence the host colonization behavior of bark and ambrosia beetles that affect hardwood trees. This work describes the identification of novel fungal and host-associated semiochemicals that may aid in future management of these important pests.</div><div><br></div><div>In Chapter 1, I review the current literature describing the volatile chemical ecology of bark and ambrosia beetles that inhabit hardwood trees. A review of groups with numerous identified semiochemicals, as well as considerations for future research is included.</div><div><br></div><div>In Chapter 2, I test the hypothesis that host colonization by the peach bark beetle (Phloeotribus liminaris) is chemically mediated by compounds associated with infested hosts. I found that benzaldehyde mediates colonization by the peach bark beetle, and that that benzaldehyde lures are effective attractants in field-trapping studies.</div><div><br></div><div>In Chapter 3, I test the hypothesis that ambrosia beetle attraction to host stress compounds can be modified by symbiotic fungal volatiles. I found that for three species of invasive ambrosia beetles individual fungal volatiles act as repellents, with species-specific differences in response to different compounds.</div><div><br></div><div>In Chapter 4, I test the hypothesis that attraction of the walnut twig beetle (Pityophthorus juglandis) to its pheromone lure can be enhanced by symbiotic fungal volatiles. I found that symbiotic fungal volatiles consistently enhance attraction of the beetles to their fungus, while one symbiotic fungal volatile of ambrosia beetle species repelled the walnut twig beetle.</div><div><br></div><div>In Chapter 5, I summarize results from each of the chapters and discuss patterns observed in the response to fungal and host-associated volatiles among the focal bark and ambrosia beetle species. I also discuss future research needs and directions to continue development of the knowledge surrounding scolytine chemical ecology and management of these pest beetle species.</div>

<p>Due to overpopulation and resource-poor habitat structure, deer threaten the<a> future of oak and other browse-sensitive species in hardwood forests. </a>Appropriate tools must be used to ensure desirable, diverse, and ecologically stable regeneration of future forests and the sustainability of native plant communities. We performed two experiments and a review to examine the effectiveness of available methods for managing browse of hardwood seedlings and to discover how these interact with each other and other silvicultural methods. First, we examined how fencing interacts with controlled-release fertilization, seed source (genetically select and non-select), and site type (afforested and reforested sites) to enhance the regeneration of planted northern red oak (<i>Quercus rubra </i>L.), white oak (<i>Quercus alba</i>), black cherry (<i>Prunus serotina</i>), and black walnut (<i>Juglans nigra</i>) at five sites in Indiana. Fencing proved to be the greatest determinant of seedling growth, survival, and quality. Fertilizer enhanced the early growth of white oak and black cherry, though for black cherry this occurred only inside fences. Select seed sources grew better and showed greater quality; however, the survival of select seedlings was limited by deer browse in absence of fences. Trees at afforested sites had lower survival if left non-fenced. Secondly, we also investigated how fencing and invasive shrub removal affected natural regeneration, species richness, and ground-layer plant cover under closed-canopy forests. Honeysuckle (<i>Lonicera maackii</i>) removal had a variable effect depending on species and site. Positive effects were most common for shade-intolerant species, while negative effects occurred for a few shade-tolerant species at some sites. Deer fencing had a positive effect on cherry and hackberry seedling density, and a negative effect on elm seedling density. Honeysuckle and deer fencing interacted antagonistically in some instances. Fencing without honeysuckle removal resulted in lower elm abundance and herbaceous-layer cover. In the densest invasions, leaving honeysuckle intact resulted in a complete lack of recruitment into the sapling layer. Our experiment suggests that invasive shrub removal and fencing be done together. Finally, we synthesized the existing literature on browse management options for hardwood regeneration to evaluate their relative effectiveness. Fences, tree shelters, repellents, facilitation by neighboring plants, deer population control, timber harvest, and slash all had positive effects on height growth of regenerating seedlings under deer browse pressure. Fences were more effective at reducing browse than repellents, while fertilizers increased browse and had no effects on growth. </p>

<p>Urban areas are warmer than surrounding rural areas. During the cold of winter, warming increases surrounding host temperature and may improve the overwintering survival of marginally hardy insects like evergreen bagworms. Similarly, during the summer, it has the potential to increase the fecundity and abundance of sap feeding insect pests such as scale insects in ways that change the capacity of their natural enemies to regulate their populations. </p> <p>Although in parts of Indiana winters can be cold enough to kill bagworm eggs, they thrive in cities. I conducted field experiments to determine the extent to which impervious surface near an infestation could keep temperatures warm enough to affect bagworm survival during cold of winter. My results suggest that the percentage of live eggs inside overwintering pupae decreased as ambient temperature drops. This response was moderated by the presence of impervious surface around an infested plant. Eggs found in bagworms collected from host trees surrounded by more impervious surface had a higher chance of survival than those collected from trees with low levels of hardscape. However, impervious surface has its limit such that egg mortality was not buffered by impervious surfaces at temperatures at or below -21.67°C. Similarly, I also conducted field experiments with sap feeding insects on honeylocust trees, a commonly planted tree in cities. Hot sites had a mean daily temperature more than 1.5 °C warmer than cool sites and scale insects were more abundant and fecund on trees in the hottest part of Indianapolis compared to cooler areas. No differences were observed in rates of parasitism on the scale insect. However, I found strong density dependence relation between parasitoids and scales abundance at scale density at or below the levels present in cool sites. The top-down regulation was prevalent at or below a critical density of scale hosts. Conversely, bottom-up regulation was prevalent above this host density as pests benefit from bottom-up factors. This suggests that urban habitats helped the scales to escape biological control by resident natural enemies above critical density of scale hosts. </p> <p>My findings can be useful to landscape designers to design landscapes that are less prone to insect pests. My finding adds to a growing body of evidence that suggests that planting urban trees with lesser amount of impervious surface can help reducing the urban warming effect and increase the regulation from natural enemies. </p>

<div>Anthropogenically driven global change is disrupting ecosystems and habitats of many plant species, straining the ability of native species to survive and reproduce. The overarching goal of this research was to holistically work towards restoration of a threatened tree species by connecting research from different disciplines. In order to do so, the threatened butternut tree (<i>Juglans cinerea</i>) and its hybrids were used as a case study. Hybridization can incorporate stress tolerance in plants and could be a potential restoration tool. Evidence in some wild butternut populations indicates that naturalized hybrids of butternut with Japanese walnut (<i>Juglans ailantifolia</i>) may be more tolerant to butternut canker disease (BCD) than butternut, but this has not been formally tested. Thus, chapter 2 examined potential BCD tolerance within and between unadmixed and hybrid butternut inoculated with two BCD fungal isolates. Differences in canker growth were observed by fungal isolate, which could help to explain some differences in BCD severity found among butternut populations. Smaller and fewer cankers and greater genetic gains were detected in hybrid families, demonstrating that hybrids warrant further evaluation as a possible breeding tool for developing BCD-resistant butternut trees.</div><div>However, even with increased disease tolerance, hybrids must possess similar ecophysiological tolerances to their native progenitor to be an effective replacement. Butternut is extremely cold hardy, but Japanese walnuts are native to a warmer ecosystem, indicating potential disparities in extreme temperature tolerances between the two species and their hybrids. Thus, samples from mature trees were subjected to cold and heat treatments to compare relative extreme temperature tolerances within butternut and between butternut, Japanese walnut, and their hybrids. Within butternut, trees from colder areas exhibited less cold damage than those from warmer areas. Differences in heat damage among provenances occurred but did not follow a clear trend. Butternut exhibited greatest cold tolerance, Japanese walnut exhibited greatest heat tolerance, and hybrids were intermediate. Thus, the utility of hybrids for restoration could be limited at the extremes of the species’ distributions.</div><div>A second, but different type of freeze test was conducted for chapter 4 using seedlings to gain a more nuanced understanding of cold tolerance within butternut and between butternut and its hybrids. No survival or damage differences were detected in butternut provenances, although seedlings from the coldest provenances experienced more delayed budbreak at the two warmest treatments than those from warmer provenances. Interspecific differences were not observed in dieback but were detected in survival and budbreak. The hybrids had greater survival than butternut from warmer provenances at the lowest temperature treatment (-38 °C), but given that temperatures that low are extremely unlikely to occur in those provenances, it is not anticipated to give the hybrids an advantage if planted in those areas. However, the hybrids’ earlier budbreak could limit the success of restoration with these hybrids in the coldest extents of butternut’s range. </div><div>If hybrids, as well as genetically modified (GM) trees, are successfully developed for effective disease tolerance and to serve as an ecologically suitable replacement, success of restoration using hybrids will ultimately depend on those directly responsible for replanting efforts. A survey was administered to land managers in 46 organizations in Indiana to gauge perceptions of hybrid and GM trees, as well as current use of hybrid trees. Land managers had stronger concern for ecological, rather than economic, issues. Agreement was highest for using hybrid and GM trees for “conservation and restoration of at-risk species”, “timber production”, and “non-timber products (fruit, syrup, etc.)”. However, perceptions varied by characteristics, such as concern type, age, and the type of land they managed. Ecological concern and the type of land being managed most strongly predicted current hybrid use. Overall, results indicate the majority of land managers in Indiana would likely be agreeable to recommendations towards using hybrids. However, most nonetheless had strong ecological concerns about their suitability as a native replacement. It is important to note, though, that consistent with the results of previous studies, great variation was seen within the performance and characteristics of the butternut hybrids in chapters 2-4. Thus, it may be possible with careful selection and breeding to harness this variation to develop disease tolerant and ecologically similar hybrids acceptable to land managers.</div>