<p><a>Natural enemy ecology strives to integrate the fields of disease ecology and community ecology to forge a broader understanding of how pathogens and predators structure communities. To advance this field, we need a greater emphasis on: 1) quantifying pathogen-mediated effects on community structure and comparing these effects to those observed with predators and 2) determining the interactive effects of combined natural enemies on communities. I conducted a mesocosm experiment designed to assess the individual and combined effects of predators (dragonfly larvae and adult water bugs) and a pathogen (ranavirus) on a larval amphibian community. Additionally, I conducted laboratory experiments to assess whether ranavirus exposure increases the vulnerability of tadpoles to predation. In my laboratory experiments, I found that virus exposure increased predation rates with dragonflies, but not water bugs. For tadpoles in the dragonfly treatments, the probability of survival for virus-exposed tadpoles was 66-77% lower compared to unexposed tadpoles. This data suggests that predators may selectively remove infected individuals from the population, which can enhance the magnitude of the healthy herds effect. I found that the risk level of the predators largely explained effects on the community. For instance, high-risk dragonflies reduce overall survival to 30% whereas low-risk water bugs only reduced survival to 67%. Additionally, I found that virus reduce survival to 62%, which was comparable to effect of the low-risk predator. Interestingly, all three natural enemies influenced community structure (i.e. species relative abundance) in unique ways. These results demonstrate that pathogens can have effects similar to predators on communities, and that natural enemy identity is important when considering impacts on community structure. When predators were combined with the virus, I found that mortality was relatively unchanged from the predator-only treatments suggesting less than additive effects of combined natural enemies. This result was driven by the healthy herds effect; the presence of dragonflies reduced overall infection prevalence in the community to 7% compared to 30% in the virus-only treatment. This effect was observed in the water bug treatments, to a lesser degree, suggesting that predator risk or efficiency contributes the magnitude of the effect. Collectively, my work demonstrates the importance of examining the individual and combined effects of natural enemies on ecological communities.</a></p>

<p>Since the pre-industrial age, the Earth has been warming at unparalleled rates, and this warming is changing climate and weather, creating a more extreme global hydrological cycle. In this dissertation, I explore how these changes to the hydrological cycle may act ecosystem and community level responses of terrestrial plants in the Midwestern United States. In this region, it is projected that mean annual precipitation (MAP) will increase, but precipitation will become more variable across and within seasons. Ecosystem structure and function are vulnerable to changes in hydrologic patterns, including changes in biogeochemical cycles, plant productivity, and plant community structure and function. In this dissertation, I explore how changes in precipitation will alter these processes using two field experiments, and I suggest potential hypotheses that could explain drought-induced community change.</p><p><br></p><p>In chapter 1, I explore how alterations to seasonal precipitation in the winter and summer act ecosystem and community processes in a temperate deciduous forest. Biogeochemical processes and plant communities are sensitive to changes in abiotic conditions, and these conditions will alter forest succession, particularly juvenile woody plant species. Using a fully factorial experiment, I manipulated winter snowfall and summer precipitation to create wet, dry, and control (ambient conditions) treatments and investigated how changes in seasonal precipitation would act mineralization rates, woody plant recruitment, and understory composition. I found that the effects of winter and summer precipitation on these processes acted independently of one another in this system, and the system was resistant to changes in mineralization rates and understory composition. Woody plant recruitment may be more sensitive to altered precipitation, as recruitment of at least one of the four species planted, Lindera benzoin, was impacted by changes in seasonal precipitation. Snow removal treatments reduced germination and increased summer precipitation decreased the relative growth rate of this species. In the short term, slight changes to woody plant recruitment may have little impact on long-term forest succession, but as these changes persist over longer periods of time, they could alter the direction of succession, which could lead to changes in the understory community composition and nutrient cycling.</p> <p><br></p><p>The second and third chapters explore the effects that drought intensification will have on terrestrial plant communities. Numerous studies have investigated the effects of individual droughts on ecosystem and community responses, but the effects that both the timing and duration of drought have on these responses remain largely unknown. To explore this gap in the literature, I conducted a eld experiment using rainout shelters to reduce growing season precipitation, creating dry periods that varied in length and timing. Drought can impact productivity and diversity in this system, and the timing in which the drought occurs influences these effects. Surprisingly, I found that the length of drought did not affect productivity or community composition.</p> <p><br></p><p>The final chapter introduces the Community Response to Extreme Drought framework CRED), which addresses the potential temporal progression of mechanisms and plant-plant interactions that may lead to community changes during and after a drought. The mechanisms for the temporal evolution of community-level drought responses are not fully understood, but plant-plant interactions, both competitive (-) and facilitative (+), are increasingly being recognized as important drivers of community compositional changes. The CRED framework provides hypotheses for the roles that plant-plant interactions have on drought-induced community change. CRED addresses how system-specific variables and the intensity of drought may influence the strength of plant-plant interactions over time, and ultimately the systems resistance and resilience to drought. </p><p><br></p><p>The results from this dissertation work have revealed that more research needs to be done to fully understand how changes in precipitation regimes and patterns will affect terrestrial ecosystems and plant communities. A better understanding of how ecosystems and communities respond to drought timing and length can help improve climate models and restoration strategies.</p>

The prevalence of hypoxia in aquatic systems has increased in recent decades and climate change is expected to worsen the extent and severity of hypoxic phenomena worldwide. Moreover, aquatic hypoxia has produced adverse ecological consequences and stimulated research interest within the Laurentian Great Lakes. The physiological stress imposed by hypoxia reduces habitat quality for most aquatic biota and causes changes in patterns of resource use and food web dynamics. We conducted a review of the primary literature to identify trends in prevalence of Great Lakes hypoxia research and broadly classify the unique hypoxic conditions afflicting Great Lakes ecosystems. We found that the majority of research effort toward Great Lakes hypoxia is focused on the Lake Erie central basin. Our review further revealed that this does not characterize the breadth of hypoxic phenomena that occur throughout the Great Lakes region. We then utilized a long-term monitoring dataset provided by the U.S. Environmental Protection Agency Great Lakes National Program Office to quantify the impact of Lake Erie central basin hypoxia on habitat quality of several fish species. We found that bioenergetics-based growth-rate potential models have a potential application as the framework for the development of biological endpoints that measure the effects of hypoxia on aquatic biota. Finally, we utilized stable isotope analysis to look for broad spatial and temporal trends in resource utilization within distinct regions of the Lake Erie central basin, with hypoxia and large-scale hydrodynamic patterns serving as potential driving patterns for spatial differences.

<p>Amur honeysuckle (<i>Lonicera maackii</i>) is a shrub endemic to East Asia which has become invasive and nearly ubiquitous to the forest ecosystems of eastern North America. Through its extended growing season, competitive ability, and potential allelopathy, Amur honeysuckle alters native herbaceous-layer plant communities and inhibits the regeneration of native tree species. As such, it is representative of a range of invasive shrub species imported from East Asia. My thesis contributes to questions fundamental to the understanding this and other invasive shrubs: 1) How do species become invasive? 2) How can the invasions of the species be effectively controlled? 3) How does the ecosystem respond to treatment of the invasive species?</p> <p>First, I examined the role of self-compatibility within Amur honeysuckle. I compared the berry production, seed production, and germination rates between closed-pollinated and open-pollinated flowering branches of Amur honeysuckle individuals across multiple types of invasions (heavy, light, and sprouting). I found that Amur honeysuckle not only possesses the ability to self-pollinate, but that it can produce viable self-pollinated seed sets. This ability may help explain how Amur honeysuckle able to invade isolated forest patches far removed from the main invasion.</p> <p>Second, I evaluated the effectiveness of a novel herbicide adjuvant in reducing the amount of herbicide needed in the ‘cut-stump’ method of controlling Amur honeysuckle. Combining various concentrations of the most common herbicide in the world, glyphosate, with concentrations of cellulases derived from fungi, 2XL, I examined whether the cellulases improved the effectiveness of glyphosate, potentially by increasing glyphosate movement into the vascular tissue of Amur honeysuckle through the degradation of cell walls. While 2XL was not an effective adjuvant, glyphosate concentrations of less than half the recommended dosages were equally effective as higher concentrations in preventing sprouting of treated stumps. The ineffectiveness of 2XL may imply a need for protein-mitigated diffusion of glyphosate across cell walls and into the vascular tissue of the plant, which would be inhibited by the breakdown of cell walls.</p> Finally, I tested how deeper intensities of mulching-head treatments affected the sprouting response of Amur honeysuckle and the response of the herbaceous-layer plant community after treatment. I found a negative relationship between the volume of sprouting Amur honeysuckle and increasing depth of mulching-head treatment. Additionally, increasing mulching-head intensities were correlated with increased herbaceous layer diversity and conservation value as represented by Floristic Quality Index. Mulching-head treatments are a promising tool in controlling heavy invasions of non-native shrubs.

<p dir="ltr">North American beaver are ecosystem and habitat altering mammals with a wide distribution in North America. Beaver are associated with bodies of fresh water while utilizing riparian habitat by foraging on woody and herbaceous, terrestrial and aquatic vegetation, as well as creating lodges, dams, dens, and scent mounds. Chain O’Lakes State Park was chosen as a study area for surveying beaver activity and woody species communities. The aim of the study was to better understand the relationship between beaver, and specifically, the woody vegetation in their foraging and home range. There is a large swath of habitat that beaver can utilize and alter within Chain O’Lakes State Park including the lakes, streams, and forests that attract visitors. The factors that have been found to influence beaver utilization in Chain O’Lakes State Park (and North America at large) are woody species community composition, abundance of heavily preferred woody species, as well as a combination of both an absence of predators and an abundance of aquatic vegetation. Based on the distribution of heavily, occasionally, and rarely preferred woody species across the understory (new individuals) and overstory (mature individuals) stratum an inference can be made that most of the riparian habitat around the lakes will continue to and/or shift into a higher proportion of species that beaver heavily select. A minority of the habitat shall shift away from those species and towards rarely and/or occasionally selected species. I will provide USDA APHIS Wildlife Services that play a wildlife management role in Chain O’Lakes State Park with the survey information detailing how beaver and their structures interact with the ecosystem, hypothesized methods for preserving the woody species that are targeted by beaver foraging within the park, and supplementary information that may aid in maintaining the riparian habitats for the benefit of park’s biodiversity and wildlife persistence while continuing to provide an enriched experience within Northwest Indiana with the potential to inform the greater collection of parks and maintained land at large.</p>

<p dir="ltr">The lichen-feeding tiger moth tribe Lithosiini (Erebidae: Arctiinae) represent the largest radiation of invertebrate lichenivory. Caterpillars feed on lichen and as they feed, also sequester lichen polyphenolics, a behavior unique to these insects. The role of these compounds is believed to defend lithosiines against predators as larvae have been found to be protected against predators such as ants and moths to predators such as birds and bats. Experimental testing with controlled diets is necessary to fully make this connection, however little is known about host specifics for lithosiines. Furthermore, although lithosiines are monophyletic, the lack of a fully resolved phylogeny hampers investigation into many of the shallower level relationships, e.g. those among genera and species, within the group.</p><p dir="ltr">I addressed these knowledge gaps using the subtribe Cisthenina. Members of this group have been used to investigate predator-prey interactions and been included in morphological and molecular studies. Thus, while the group still needs attention, there is an ample amount of legacy loci data available for its members. I used these data to investigate the evolutionary relationships at the genus level, but to increase resolution in my analyses I additionally sampled taxa throughout the group with a recently developed anchored hybrid enrichment (AHE) probe set. I combined it with the legacy loci to both increase taxon sampling and resolution. I confirmed that trees made strictly from the legacy loci were unsuccessful and resulted in poorly supported relationships that made little sense. The addition of the AHE data greatly helped resolve relationships, however, there remained areas that were poorly supported and they appear to be genera with only a few loci. Thus, there is still room for improvement, but this offers a way for moving forward in lithosiine research, particularly to involve others who may have limited funding, equipment, and/or personnel and may only be able to afford legacy loci in diverse collaborations.</p><p dir="ltr">As the AHE probe set worked well with genus-level relationships I further attempted to use it in species delimitation of the notorious <i>Hypoprepia fucosa</i>-<i>miniata </i>species complex. Members of this group are varying shades of yellows, oranges and reds and have a convoluted taxonomic history. I gathered and organized over 4,000 specimens and using the AHE probe set found support for five distinct species. Interestingly, I used other morphological characters such as genitalia, but found no differences between species and a large amount of intraspecific variation. This suggests other courtship behaviors may be present and external morphology, i.e., color patterns, remain the best way to identify species. As part of this I am describing a new species and raising one from subspecies and as species are now readily distinguishable, they can be used for further investigations into lithosiines.</p><p dir="ltr">I used a member of this complex, <i>H</i>. <i>fucosa</i> to then evaluate the use of species distribution models (SDMs) to better understand their niche and how it relates to plausible lichen hosts. I evaluated 17 lichen species from two lichen genera, <i>Physcia </i>(13 species) and <i>Myelochro</i><i>a </i>(4 species)<i>. </i>These genera were selected based on previous feeding assays and the metabolites found in them have also been found in <i>H</i>. <i>fucosa </i>further suggesting caterpillars may feed on them. SDMs typically only use environmental factors to define and predict species niches. I compared the niches described by traditional SDMs to assess how similar they were, but I also investigated the use of lichens as biotic factors in the models. I assessed the influence each lichen had on the moth’s distribution found the niche of every lichen to be significantly different than that of the moth and their inclusion in SDMs of <i>H</i>. <i>fucosa </i>to improve model performance. This suggests <i>H</i>. <i>fucosa </i>caterpillars to be polyphagous, but to have some connection with these lichens. Further investigation with live specimens is needed, but these results support this as an effective way to describe lithosiine niches to better understand lichen feeding.</p>

<p dir="ltr">Non-native biological invasions are one of the leading concerns for global biodiversity. The establishment of invasive species reduces local biodiversity, shifts species composition, changes successional trajectories, and alters ecosystem functions. This thesis examines two aspects of invasive plants: (1) the distribution and the most important climatic and anthropogenic drivers of invasive trees across the eastern United States, and (2) an evaluation of invasive plant removal and herbaceous recovery from a government cost-share program that provides financial support for invasive plant management by private landowners.</p><p><br></p><p dir="ltr">Our first study focused on identifying the distribution of invasive trees, and the factors associated with their distribution. This is essential to predicting spread and planning subsequent management. Using USDA Forest Inventory Analysis (FIA) data and random forest modeling, we examined the distribution, and variables associated with the distribution, of invasive tree species. Invasive trees were found in 10,511 out of 299,387 FIA plots. Invasive species basal area and density (trees per ha; TPH) were highest within the central and southern Appalachian Mountains, Michigan, the Northeast, and the southern Coastal Plain of the United States. A random forest model of invasive species basal area (R²= 0.47, RMSE = 0.47) and density (R²=0.46, RMSE=0.50) vs. environmental variables found that both invasive basal area and density were most strongly associated with human footprint, followed by various climatic variables. An equivalent model of native tree basal (R²=0.53, RMSE=9.25) and TPH (R²=0.47, RMSE=8.64) found that native tree basal area and density were most strongly associated with aridity followed by various climatic variables. As human footprint increased, invasive tree basal area and density increased. These results suggest that the distribution of invasive trees is reliant on human alterations to forests.</p><p><br></p><p dir="ltr">Our second study focused on Environmental Quality Incentives Program (EQIP), a federal cost-share program that has provided $25 billion of financial assistance to farmers and non-industrial private forest owners. Few studies have examined whether this program facilitates the recovery of the herbaceous layer while decreasing the dominance of invasive plant species. We surveyed the herbaceous layer of EQIP-treated and untreated (reference) forests across three physiographic regions of Indiana. Using non-metric multidimensional scaling (NMDS) ordination and linear mixed effects models, we evaluated the species composition, richness, diversity, evenness, floristic quality index, and herbaceous-layer cover of EQIP and reference sites. We also used linear mixed models to evaluate how EQIP site treatment affected the diversity of native plant species. Sites treated with EQIP contracts typically had significantly higher native species richness, Shannon’s diversity, and floristic quality than reference sites. There were significant separations in species composition between EQIP treated and reference forests state-wide and in the southern non-glaciated region of Indiana, although composition overlapped between EQIP and reference forests. Our study suggests that EQIP-funded treatments promote increased species richness and diversity. However, the persistent overlap in species composition we observed may signify biotic homogenization due to a long-shared history of anthropogenic disturbances between EQIP and reference sites. Therefore, active restoration of the herbaceous layer might be needed to allow a full recovery after invasive removal.</p>

<p>Climate change has dramatically altered the ecological landscape of the eastern U.S., leading to shifts in phenological events and redistribution of tree species. However, shifts in phenology and species distributions have implications for the productivity of different populations and <a></a>the communities these species are a part of. Here, I utilized two studies to quantify the effects of climate change on forests of the eastern U.S. First, I used phenology observations at a common garden of 28 populations of northern red oak (<i>Quercus rubra</i>) across seven years to assess shifts in phenology in response to warming, identify population differences in sensitivity to warming, and correlate sensitivity to the productivity of the populations. Second, I utilized data from the USDA Forest Service’s Forest Inventory and Analysis Program to identify forest communities of the eastern U.S., assess shifts in their species compositions and spatial distributions, and determine which climate-related variables are most associated with changes at the community level. In the first study, I found that populations were shifting their spring phenology in response to warming, with the greatest sensitivity in populations from warmer, wetter climates. However, these populations with higher sensitivity did not have the highest productivity; rather, populations closer to the common garden with intermediate levels of sensitivity had the highest productivity. In the second study, I found that there were 12 regional forest communities of the eastern U.S., which varied in the amount their species composition shifted over the last three decades. Additionally, all 12 communities shifted their spatial distributions, but their shifts were not correlated with the distance and direction that climate change predicted them to shift. Finally, areas with the highest changes across all 12 communities were associated with warmer, wetter, lower temperature-variable climates generally in the southeastern U.S. Taken together, these studies provide insight into the ways in which forests are responding to climate change and have implications for the management and sustainability of forests in a continuously changing global environment.</p>

Pathogens and parasites are increasingly recognized as important components within host populations, communities, and ecosystems. Parasite contributions to ecosystem function most likely manifest as density-mediated impacts of parasites on their hosts, the direct contributions of parasite biomass to a system, and via parasite-induced changes in host behavior and physiology (trait-mediated impacts). Here, a framework was constructed that can be used to conceptualize parasite contributions to ecosystem function (Chapter 1). Then the influence of parasite attack on host movement was explored to further evince the mechanistic underpinnings of trait-mediated parasite impacts (Chapter 2). Additionally, mesocosms were created across a gradient of parasitism to examine how these mechanisms are likely to unfold at larger biological scales (Chapter 3). Lastly, a series of differential equations was created to model host-parasite-ecosystem interactions and generate theoretical predictions about how and when parasites are likely to influence ecosystem processes (Chapter 4). Parasites have many characteristics of ecosystem engineers, but their role has historically been ignored. These studies begin to explore the role that parasitism may have as one of the drivers of ecosystem processes.

<p>Mayflies are important macroinvertebrate members of steam communities and are highly sensitive to agricultural insecticides. In this dissertation I sought to explore whether Heptageniidae<i> </i>mayflies in Indiana have evolved greater tolerance to clothianidin, a neonicotinoid insecticide that has been widely applied over the predominant corn and soybean crops of the state for the last 20 years. Over the five chapters of this dissertation, I investigate different aspects of the effects of clothianidin on mayfly populations. First, I collected mayfly populations from streams around the state across a gradient of agricultural landscape to see if proximity to corn and soybean crops was associated with tolerance to the insecticide. I found significant variation in tolerance between the populations but only connected it to land use for one species and one insecticide. Second, I investigated the potential for increases in predation rates due to exposure to low concentrations of clothianidin by both a vertebrate and invertebrate predator. Sublethal exposure did increase the risk of predation by the invertebrate predator but not the vertebrate. Third, I collected four populations of mayflies and placed them in a common stream which is heavily impacted by agricultural runoff. The most tolerant of the four populations had the highest survival rates and was native to the polluted stream, suggesting that it was best adapted to the local conditions. Finally, the last chapter is a discussion of the factors influencing the evolution of contaminant tolerance, the implications for biomonitoring, and a series of recommendations for incorporating evolved tolerance into biomonitoring practices. Collectively, this work suggests that the lethal and sublethal effects of clothianidin exposure create conditions suitable for the evolution of tolerance in Heptageniid mayflies. While there are already differences in tolerance between populations, it seems likely that mayfly pesticide tolerance will continue to increase as pesticides continue to impact aquatic ecosystems. </p>