Biological invasions have been observed throughout the world for centuries, often with major consequences to biodiversity and food security. Tying invasion to species identity and associated traits has led to numerous hypotheses on why, and where, some species are invasive. In recent decades, attention to intraspecific variation among invaders has produced questions about their adaptation to climate, land use, and environmental change. I examined the intraspecific variation of invasive Johnsongrass's (Sorghum halepense (L.) Pers.) seedling stress response, propagule cold tolerance, and large-scale niche dynamics for correlation with populations' climatic and ecotypic (i.e., agricultural vs. non-agricultural) origin. Overall, I found a greater number of home climate effects than ecotypic effects on various traits. Non-agricultural seed from cold climates and agricultural seed from warm climates germinated more and faster, while non-agricultural seedlings showed uniform chlorophyll production regardless of home soil carbon origin, unlike their agricultural counterparts. Neither seedling stress response nor propagule cold tolerance interacted with ecotype identity; however, drought stress varied with population origins' aridity and soil fertility, and seed from warm/humid and cold/dry climates was most germinable. Comparison of seed and rhizome cold tolerance also suggested that the latter is a conserved trait that may be limiting S. halepense poleward range expansion. This physiological limit, an unchanged cold temperature niche boundary between continents and ecotypes, and a narrowed niche following transition to non-agricultural lands all imply low likelihood of spread based on climatic niche shift. Instead, evidence points to range expansion driven primarily by climate change and highlights agriculture's role in facilitating invasibility. This tandem approach to climate and land use as drivers of intraspecific variation is transferable to other taxa and can help refine our conception of and response to invasion in the Anthropocene. / Doctor of Philosophy / Exotic invasive species are a global problem, threatening biodiversity and biosecurity now and in the future. In the last several decades, ecologists have studied many individual invaders and their traits to understand what drives their spread. More recently, abundant differences in traits between populations within an invasive species have raised questions about humans' role in facilitating invasion through climate change, land use, and other disturbances. I studied the invasive Johnsongrass's (Sorghum halepense (L.) Pers.) response to drought, nutrient limitation, and freezing to detect differences between populations based on their climate and ecotype (agricultural vs. non-agricultural) origin. I also tracked differences in the climates the species occupied across the globe and North America and projected its future distribution under climate change. Overall, I found a greater number of home climate effects than ecotypic effects on various traits. Non-agricultural seed from cold climates and agricultural seed from warm climates germinated the most, while non-agricultural seedlings performed consistently regardless of soil carbon origin, unlike their agricultural counterparts. In addition, drought stress varied with population origins' rainfall and soil fertility, and seed germination favored warm/humid and cold/dry origin. Rhizome (underground stem) cold tolerance appears to be a trait that limits S. halepense poleward range expansion. Along with no change in the coldest climates occupied worldwide and no spread to new climates with transition to non-agricultural lands, this implies that Johnsongrass is unlikely to expand its range without external forces. Instead future range expansion will likely be driven by climate change. This coupled approach to climate and land use affecting invasion is transferable to other species and can help refine both our concepts and response strategies.

This dissertation consists of three essays, of which, two involve assessing the value of tree cover in urban communities and the other evaluates cooperative management of an invasive species by urban communities. The first chapter summarizes the three topics and briefly describes the motivation, methods applied and main conclusions in each study. Chapter 2 presents a meta-analysis of hedonic property value studies on the value of tree cover. A meta-regression was performed using implicit value estimates for tree cover at property-level from various locations in the U.S. along with auxiliary data of county-level tree cover to investigate the relationship between tree cover and implicit-price estimates for residential properties. The study found that on average 35 percent and 40 percent tree cover respectively at property-level and county-level optimize the benefits to the property owners in urban areas. These results provide insights to forward-looking communities to adapt their tree planting and protection efforts to lessen climate-induced impacts. Chapter 3 applies a first-stage Hedonic property price model to estimate preference for tree cover in urban communities using single-family house sales data from multiple property markets across the U.S. The study analyses how home owners' preference for tree cover vary across the landscape and across cities. Further, it identifies what factors affect these variations via the general inferences obtained from an internal meta-analysis. The study confirms the heterogeneity of preferences as affected by the differences in the abundance of tree cover in study locations, regional differences and household characteristics. These findings add to the hedonic literature and provide useful information for future urban planning. Chapter 4 focuses on cooperative management of invasive species in landscapes with mixed land ownerships. This study analyzes the effect of the land ownership on the management efforts between an infested municipality and an uninfested municipality when a transferable payment scheme is involved in the cooperative agreement. A dynamic optimization problem was set up to evaluate the case of Emerald ash borer (EAB) control in multiple jurisdictions in the Twin Cities, Minnesota. The results suggest that when the infested municipality has more public lands and when the transfer payments are efficiently used to implement greater control, the municipalities are more likely to commit to bargaining, and smaller transfer payments paid over a longer span of time are sufficient for optimal control of the spread of invasive species across the municipalities. The last chapter concludes the three studies and discusses the insights for future research. / Ph. D.

The red imported fire ant, Solenopsis invicta, is an abundant and aggressive component of early-successional communities in the southeastern United States. After disturbance, it rapidly invades new habitats, and once there, it has strong competitive and predatory effects on the existing arthropod community. In upland coastal plain pine forests at the Savannah River Site in South Carolina, I conducted two studies of fire ant ecology. In my first study (chapter 1), I investigated the way in which fire ants colonize early-successional road and powerline cuts through forests, and I tested whether some of these linear habitats provided better fire ant habitat than others. I found that fire ant mound density (#/ha) was similar in narrow dirt roads and in wider roads with the same intermediate level of mowing disturbance, and that density was lower in wide powerline cuts where the vegetation is only removed every five years. Furthermore, mound density was greatest near the edges of cleared roads and powerline cuts and was also greater on the northern sides of roads and powerline cuts where there was less shading from the adjacent forest. Results from this study suggest that allowing increased shading from adjacent forest vegetation, especially along northern roadside edges, would limit the suitability of road and powerline cuts as fire ant habitat, thereby slowing invasion. In my second study (chapter 2), I examined the impact of fire ants on arthropod and plant species in early-successional forest gaps. In a tritrophic system, I tested whether the top-down effect of insect herbivore consumption by fire ants was strong enough to cascade through two trophic levels and improve plant growth and fitness. I compared this potential effect to that of other arthropod predators in the community. I found that fire ants controlled the level of tissue damage to plant leaves by herbivores, but that the damage was not severe enough to influence plant growth or fitness. Fire ants had stronger negative interactions with insect herbivores than other predators in the community, and for this reason, fire ants can be considered keystone predators. This project provides further evidence that fire ants successfully invade even small patches of early-successional habitat that exist within larger matrices of uninhabitable, late-successional forest, and that once there, they have a dramatic effect on the arthropod community. Restricting the amount of early-successional habitat within southeastern forests, either as permanent road and powerline cuts or as temporary gaps, would limit the potential for strong and detrimental effects by the invasive fire ant. / Master of Science

Species vary phenotypically and genetically across their environmental range limits, and this variation can influence ecological processes. Ecologically meaningful intraspecific variation might be particularly important in the context of agricultural weeds and exotic invaders, because intraspecific variation in these species might allow them to rapidly adapt to their unusually dynamic and variable environments. In a greenhouse study, we explored intraspecific variation in the size, rhizome production, and competitive ability of the global invader, Johnsongrass (Sorghum halepense), representing populations from agricultural and non-agricultural habitats across its introduced North American range. We also used these populations to explore the relationship between phenotypic variation and genomic endoreduplication responses to the common stresses herbicides, competition, and clipping. Endoreduplication occurs when plants increase their genome size by increasing their nuclear chromosome number, with some evidence showing correlations with stress response. We found that Johnsongrass plants from agricultural habitats were larger than plants from non-agricultural habitats, but there was no difference between habitats in either rhizome production or competitive ability. Two of the five herbicides we tested, primisulfuron and imazethapyr, had the strongest suppressive effects on Johnsongrass, and also stimulated the greatest rates of endoreduplication. Furthermore, agricultural populations showed higher levels of endoreduplication. We found no overall effect of competition on endoreduplication, although endoreduplication was higher for non-agricultural populations than agricultural populations. When competing with corn, but not with conspecifics, Johnsongrass roots increased endoreduplication by 13%. Clipping induced substantial endoreduplication, but there was no difference between agricultural and non-agricultural populations. Our results suggest that endoreduplication may play a role in some, but not all, stress responses in Johnsongrass. Furthermore, our results indicate that Johnsongrass has adapted in some phenotypic and genomic ways to agricultural habitats in North America. Such adaptation may play a role in this species' success as both an agricultural weed and an exotic invader. / Master of Science in Life Sciences

Invasive species are reported as one of the top current and future concerns for the health and functioning of native ecosystems. In response, identification of invasive plant impacts are one of the top most studied aspects of plant invasions. Yet we still know very little about invasive plant impacts, and many land managers remove invasive plants not because of known negative impacts, but because of the general negative connotation associated with invasions. Here, I develop and utilize integrative methods to more holistically measure the ecological impacts of invasive plants. I develop a meta-analysis of current invasive plant impact literature, I integrate independent ecosystem metrics into holistic measures of total ecological impact, I examine the potential for legacy and temporal effects in newly established and recently managed invasions, and I measure ecological impact of co-invaded ecosystems. Through these studies, I find that magnitude, and not direction, of impact better represents actual ecosystem changes when evaluating invasions holistically. I also find invasive plant management may not only fail to remove long term legacy effects, but may in some cases further negatively impact the plant community. Finally, I find that co-invaded systems are not intrinsically worse off than single invaded systems. Collectively, these studies help to better our understanding of the impacts of invasive plants and their implications for management, and show that simple attempts at eradication may not always be desirable. / Ph. D.

All species on Earth occupy limited geographic space. More than a century of observational, experimental, and theoretical work investigating the factors that drive species distributions have demonstrated the importance of the interactions between abiotic, biotic, and demographic factors in determining why species are found where they are. However, it is still unclear when and where these factors interact to set species range limits. Filling the existing knowledge gaps is imperative for the accurate predictions of how species will respond to global change, and particularly for invasive species, many of which are expected to benefit from global change. Here, I sought to investigate the mechanisms that enable, as well as limit, the range expansion of the globally invasive plant Sorghum halepense (L.) Pers. (Johnsongrass). I performed a series of field and laboratory experiments to study population and range dynamics throughout Johnsongrass's North American distribution, and test for the effects of climate, local habitat, and competition on multiple functional traits. I found Johnsongrass consistently demonstrated impressive performance across varying environments, often growing more than 3 m tall, producing hundreds of flowering culms within a single growing season, and maintaining positive population growth rates, even under intense competition with resident weeds. I also found evidence that seed germination has adapted to varying climates encountered during Johnsongrass's range expansion resulting in a shift in the germination temperature niche from warmer to cooler as Johnsongrass spread from warmer climates in the south to more temperate climates in higher latitudes. This shift in the germination temperature niche may have been an important contributing factor in the range expansion of Johnsongrass by enabling the optimization of seed germination in varying climates. On the other hand, results from a field study suggested a possible trade-off between flowering time and growth in populations originating from the range periphery (i.e., range boundary) which may be limiting, or slowing, continued range expansion of Johnsongrass. Together, the outcomes of this work contribute to our understanding of the factors involved in the distribution of species, which is a fundamental goal of Ecology, and essential to accurately predict how invasive species will respond to global change. / Doctor of Philosophy / Invasive species threaten our natural ecosystems, our agricultural systems, and even our infrastructure, and we spend billions of dollars each year attempting to control them and reduce their negative impacts. Climate change, habitat destruction, and other forms of global change, will benefit many of these species, magnifying their impacts and promoting their invasion into new territories. Because of the damaging effects of invasive species, and the costs to control them, it is imperative that we are able to predict how they will respond to global change so that we can improve plans to reduce their impact and spread. First, we need to understand the processes that promote their invasion across large swaths of land. Just as importantly, we must study the processes that prevent their invasion of certain areas. Here, I investigated some of the processes that have facilitated, as well as hampered, the spread of the invasive plant Johnsongrass. For this work, I used Johnsongrass plants originating from different habitats, including regions where Johnsongrass is highly invasive and those where Johnsongrass is very rare. I found Johnsongrass originating from regions where it is highly invasive were able to grow very large and produce thousands of seeds that were able to germinate under a range of conditions. These traits may have contributed to the invasion success of this species. However, I found a different pattern for plants that originated from regions where Johnsongrass is rare. These plants reached reproductive age earlier and grew smaller across all environmental conditions, potentially due to the less hospitable climates of these range edges. These findings allow us to project into future climate change scenarios, because it is likely that, as temperatures warm, invasive species will be able to invade new regions, where they will impact the work of conservationists, natural resource professionals, agricultural produces, and other land managers.

<p> Kudzu (Pueraria montana) is an invasive woody vine widespread throughout the  southeastern United States, with recent studies predicting that its habitat will expand northward.  New occurrences and recent studies using climatic parameters suggest that the Midwestern  region of the United States is at the greatest risk of kudzu invasion. As there have already been  25 reports of kudzu within the Great Lakes basin, and no previous landscape models exist for the  basin, I developed probability models from existing spatial data (land cover, hydrology, geology,  annual precipitation, elevation, aspect, and known kudzu locations) using generalized additive,  bioclimate envelope, and maximum entropy methods. I further expanded each model to include  the basin and a 2.25-degree buffer in order to include 193 reported kudzu sites. For each  predictive model, I determined the area under the curve (AUC) for a receiver operating  characteristic curve (ROC) comparing false positive and false negative rates. I performed field  surveys at eight known sites of kudzu presence in Michigan, Indiana, and Ohio. Each presence  site was paired with a control (known kudzu absence site). I collected environmental data  including canopy cover, volumetric soil moisture, soil pH, litter depth, midstory species diversity  and diameter at breast height (DBH), and overstory basal area. Each environmental measure was  compared between kudzu presence and control survey sites as well as between in-basin and out?of-basin survey sites using a two-way ANOVA. Maximum entropy models produced the highest  AUC in both the basin and buffer models during model development. These models showed that  urban and disturbed habitats resulted in the greatest probability of potential habitat for kudzu. I  found no statistically significant differences in environmental characteristics between kudzu  absent and presence sites or between in- and out-of-basin sites, suggesting kudzu might be  dispersal-limited rather than limited by environmental characteristics. Continuing existing  management and further monitoring of kudzu spread is likely necessary to limit further  introduction and to mitigate spread of kudzu within the Great Lakes region.  </p>

Species that become naturalized in a new geographical range, subsequently multiply and spread, and persist to the detriment of resident communities, are known as alien invasive species. Two aspects of species invasion – spread and ecological impact – were examined using Lantana camara L. (henceforth lantana) as the study system, specifically in the context of a seasonally dry tropical forest ecosystem of the Mudumalai Wildlife Sanctuary and National Park. Lantana is a thicket-forming woody shrub of South and central American origin, which is now widespread across the tropics. The thesis is divided broadly into two parts -the first part examines the influences of environmental factors on the distribution and spread of lantana while the second part focuses on the effects of lantana on the distribution, survival and growth of native woody species. Much of the work presented in this thesis was conducted within a 50 ha permanent plot (the Mudumalai Forest Dynamics Plot, MFDP hereon) in Mudumalai, chiefly because the history of invasion by lantana has been recorded here since 1989. The influence of changing resources on lantana invasion was explored at two scales -small spatial but fine temporal scale in the MFDP and at the level of the landscape. Available data on an 18 year chronosequence of changes in the qualitative density of lantana from the MFDP and field studies between the years 2009 and 2010 were used to determine the environmental correlates of lantana spread in time and space. It was found that biotic factors such as the presence of the shrub Helicteres isora and abiotic factors such as proximity to drainages and the combination of fire and drought promoted the intensification of lantana invasion in time while proximity to streams, higher total annual rainfall and low fire frequency contributed to lantana invasion at the landscape level. The impacts of lantana on the seedlings of native woody species were assessed in 10m x 10m plots within the MFDP. An initial enumeration of 60 such plots revealed that animaldispersed, dry forest habitat preferring species were most affected by the presence of dense lantana. A follow-up study comprising of growth and survival measurements made on 1105 seedlings over two years (2008-2010) further confirmed that dry forest preferring species were most affected by the presence of dense lantana and that this response at the community level was most likely influenced by the most abundantly sampled species in this habitat preference guild – Catunaregam spinosa. In conclusion, while the environmental correlates of lantana most likely promoted its invasion, only certain guilds of native species seemed to be negatively affected by the presence of lantana.

Marine and freshwater aquatic invasive species are a huge problem in the United States and its territorial waters with myriad ecological and economic impacts. Current management methods have fallen short. One possible solution is to harvest them for human consumption. This should be done through small scale, carefully regulated, scientifically based programs that are part of a larger overall management strategy. Four case studies (northern snakehead, European green crab, Asian carp, Chinese mitten crab) assessing edibility are included.

Invasive species are an increasing problem worldwide, threatening indigenous communities and species. Many human-made environments promote introductions of alien species and one such habitat is road verges. The invasive plant Lupinus polyphyllus is benefiting from these habitats and is today widespread along road verges in many parts of Sweden. However, it has been shown to be a problematic species, as it supresses native plants. Many of these plants originate from semi-natural grasslands, but have found a refuge in road verges. The Swedish Transport Administration has tried to control L. polyphyllus, but it is unclear to what extent it is possible. To manage an invasive species, it is often preferable to prevent further extension by limiting its dispersal, which can be done by reducing seed production. To find out how to prevent L. polyphyllus from spreading, I investigated how resprouting capacity and seed production was affected by cutting of L. polyphyllus at different times throughout the season, and at different heights. Plants in all plots resprouted after being cut, but resprouted leaf stalks grew taller in the plants cut early at the flowering stage, compared to the later cutting treatments. These were also the only ones producing new flowers and fruits, perhaps due to stored resources in roots or the fact that they were cut before the summer solstice. Plants cut higher above the ground produced more flowers and fruits and produced taller leaf stalks and might have used photosynthesizing aboveground parts to acquire resources for regeneration. Fruits were produced but not ripe before the mid-summer cutting. These seeds were still able to germinate after maturing on cut plants. However, they were more mould-infected than seeds maturing on non-cut plants. Seeds from cut plants were also lighter. Thus, seeds produced by the mid-summer cutting might disperse and germinate but may perhaps suffer from lower survival due to more frequent mould infections. The late-cut plants had already produced mature seeds, and thus, were probably able to disperse. Also, the later cutting treatments produced more seeds over the entire season. This indicates that to hinder dispersal by limiting seed production, it is best to cut Lupinus polyphyllus as early as possible, in the flowering stage, when seeds are not produced and to cut them entirely to the ground.