Invasive species often encounter climatic conditions that differ significantly from those of their native range. These environmental shifts may trigger phenotypic responses, resulting through some combination of adaptation and plasticity, that enable the invader to persist under novel thermal regimes. In this dissertation, I examine phenotypic changes in a tropical lizard that has successful invaded a cooler temperate climate, specifically examining traits that may promote survival and reproduction in their new range. First, I examined physiological traits, as I predicted greater cold tolerance would be necessary to survival in the invasive range. I found that invasive populations tolerated lower temperatures, exhibited greater maximum sprint speeds, and had higher metabolic rates than native populations. Next, I examined how life-history traits may change in the invasive range in order to facilitate reproduction under shorter breeding and growing seasons. I found that compared to native females, invasive females had shorter interlaying intervals and produced eggs that hatched more quickly. Once I quantified changes physiological and life-history traits that may have aided in successful establishment, I executed a common garden study to determine whether changes were the result of adaptation or plasticity. I found that differences in critical thermal minimum, metabolic rate, interlaying interval, and incubation period were maintained in lab-reared offspring, while measures of sprint speed converged. My results provide evidence that life history and physiology can evolve rapidly during invasion. These findings are useful to understanding contemporary evolution, and also provide valuable insight on how species respond to environmental shifts, both during invasions and as a result of climate change. / Doctor of Philosophy / When species invade a new area, they often face different climates that make can make survival and reproduction challenging. In response, species may alter traits in order to adjust to new temperatures and conditions. In this dissertation, I examine trait changes in a tropical lizard that has successfully invaded a cooler temperate climate, specifically examining traits that may help them to survive and reproduce in their new range. First, I examined physiological traits, as I predicted greater cold tolerance would be necessary to survival in the invasive range. I found that invasive populations tolerated lower temperatures, could sprint faster, and had higher metabolism than native populations. Next, I examined how reproductive traits may change in the invasive range in order to facilitate reproduction under shorter breeding and growing seasons. I found that compared to native females, invasive females had less time between egg lays and produced eggs that hatched more quickly. Once I assessed how traits may have changed in the new range, I determined whether changes resulted from evolution or not. I found that differences in low temperature tolerance, metabolic rate, the time between egg lays, and incubation period were the result of evolution, while sprint speed did not seem to be the result of evolution. My results provide evidence that traits can evolve rapidly during invasion, allowing invasive species to persist and spread in new areas.

Peatlands are a diverse type of wetland ecosystem, characterized by high levels of soil organic matter, that provide a wide array of ecosystem services including water storage and filtration, carbon sequestration, and unique habitats. Draining peatlands degrades their resilience to future disturbances, notably including high intensity, soil-consuming fires. Peat soil fires are unique in that they can smolder vertically through the soil column, with consequences ranging from large carbon emissions to altered hydrology and dramatic shifts in vegetation communities. In this work we had two complementary objectives to understand both the drivers and impacts of smoldering fires at the Great Dismal Swamp (VA and NC, USA). First, we developed and verified a new method to model peat burn depths with readily available water level and peat hydraulic property data. Our findings suggest that drainage weakens both short- and long-term controls on peat burn depths by reducing soil moisture and by decreasing peat water holding capacity. To address the impacts of smoldering fires, we quantified the abundance of the noxious Phragmites australis in a large fire scar and the extent to which altered hydrology influenced its occurrence. We did so by leveraging satellite imagery, random forest models, LiDAR data, and water table observations. Our results suggest that P. australis is aided by a hydrologic regime generated, in part, from the combined effects of drainage and deep smoldering fires. Our conclusions from these two studies contribute to the scientific understanding of smoldering peat fires and can inform management efforts. / Master of Science / Peatlands are a diverse type of wetland ecosystem that have characteristically thick levels of organic-rich soil, known as peat. Peatlands are home to a variety of unique plants and animals, store large amounts of carbon, and provide water storage functions. Peatlands were historically drained to enable development and conversion to other land usages, which had many unintended consequences like increasing their risk to wildfires that consume soil organic matter. An intense peat fire can smolder down through the peat, with impacts ranging from large releases of carbon to changes in water levels and vegetation communities. In this work we had two objectives aimed at understanding the drivers and impacts of smoldering peat fires in the Great Dismal Swamp (GDS) (VA and NC, USA). First, we developed and verified a new method of modeling how deep peat fires burn by using readily available water level and soil property data. Our findings suggest that drainage weakens both the short- and long-term controls on peat fire burn depths by reducing soil moisture and by limiting the ability of peats to hold water. We also studied how water levels in a post-peat consuming fire environment influence the amount of the weedy Phragmites australis. We did so by using satellite imagery, elevation data, and water table observations. Results from this investigation suggest that the combined effects of drainage and deep smoldering fires help to create ideal conditions for P. australis invasion and establishment. Our findings from these two studies add to the scientific understanding of smoldering peat fires and may inform land management decisions.

Invasive species are a growing issue that will compound under climate change. Rising temperatures, fluctuating precipitation and new transportation pathways will create new opportunities for invasive establishment. A direct and impactful consequence of climate change is the removal of climatic barriers to invasive survival. Species distribution modeling (SDM) for invasives must include an evaluation of future establishment potential so that managers can prioritize regions forecasted as high risk under climate change. Climatic SDMs effectively support pan-invasion risk assessments by forecasting potential invaded areas globally where climatic barriers have shifted the potential for establishment. Rarely is regional-scale climate variation considered in invasive SDMs, despite its relevance for pests that establish outside their native regional climate. Here, I apply a climatic pest risk framework to the Spotted Lanternfly grape pest (Lycorma delicatula, SLF). I assess how climate change shifts the establishment potential of SLF across important viticultural regions worldwide. I contrast an ensemble of three regional-scale SDMs to a global-scale SDM, which provided multiple predictions on how future regional climate variation might shift national SLF risk levels, impacting the global wine market. I found that the global suitable area for SLF will increase under climate change, with range expansion outpacing contractions by about 1.1 million km2. Expansions will primarily occur at present northern range edges in Europe, North America, and East Asia, and contractions will occur across the southern hemisphere. Next, 307 global viticultural regions (29% of 1,063 total sampled) will decrease in risk for SLF establishment and only 532 (50% of 1,063) will remain at any risk under climate change. Loss in SLF establishment risk under climate change followed a latitudinal gradient in the northern hemisphere. Meanwhile, only 85 known SLF populations (11% of 769 rarefied sample) will destabilize under climate change. Populations within the US and South Korean invaded ranges will remain stable with respect to climate. Our regional-scale ensemble emphasized the importance of mean winter temperature as a constraint on SLF establishment, with activity dropping sharply at -3°C. This method for regional-scale ensemble modeling should be utilized in similar invasive or climate change SDM applications to make more refined SDM predictions and to reduce uncertainty. Viticulturalists can and should use our provided tools and model framework to understand the risk of SLF establishment at their locality as climate change removes barriers to this pest’s establishment globally. / Biology

1.) Sentinel traps were used to survey for parasitoids of frugivorous drosophilids in Virginia fruit cropping systems, and determine if parasitoids were attacking invasive flies Drosophila suzukii (Matsumura) and Zaprionus indianus Gupta (Diptera: Drosophilidae) in the field. Two parasitoids of frugivorous drosophilids, Leptopilina boulardi (Barbotin, Carton, and Kelner-Pillault) (Hymenoptera: Figitidae) and Pachycrepoideus vindemiae (Rondani) (Hymenoptera: Pteromalidae), were reared, but only one P. vindemiae was reared from D. suzukii, and no parasitoids were reared from Z. indianus. Most parasitoids were reared from alternate host Drosophila melanogaster Meigen and other wild drosophilids. 2.) The ability of these parasitoids to attack D. melanogaster, D. suzukii and Z. indianus under controlled conditions was tested. Larval parasitoid L. boulardi did not develop on D. suzukii or Z. indianus, just D. melanogaster. Pupal parasitoid P. vindemiae successfully developed on all three fly species, but also increased pupal fly mortality. 3.) Olfactometry was used to ascertain if L. boulardi and P. vindemiae are selective about the type of fruit their hosts feed in. Results showed that among cherry, raspberry, blueberry, grape, and banana, L. boulardi preferred raspberry and banana to cherry, and preferred grape least, but no fruit was most preferred. Insufficient data were obtained for P. vindemiae. We conclude that parasitoids of Virginia are unlikely to provide effective biological control for D. suzukii or Z. indianus, and classical biological control should be investigated as a pest management option. Olfactometry results indicate tritrophic selectivity by Drosophila parasitoids, suggesting multiple parasitoids could be required for effective biological control. / Master of Science in Life Sciences / The spotted wing drosophila (henceforth SWD) is a globally invasive vinegar fly originating from southeast Asia, and is economically damaging to producers of small fruit, such as berries, cherries, and grapes. The African fig fly (henceforth AFF), a relative of SWD, is another recently invasive species to North America (originating from Africa) that often occurs simultaneously with SWD infestations, but its economic threat to North America is still unclear. With the economic threat posed by SWD, and potential threat posed by AFF, it is important to understand their relationships with other organisms in their environment, especially natural enemies, such as parasitoid wasps. Such information is integral for researchers to develop effective control methods, and will help determine if natural enemies can be used to our advantage as biological control agents. Biological control also helps to limit the use of chemical insecticides, mitigating the development of insecticide resistance in the pests. This project employed unique field trapping methods and laboratory bioassays to investigate the relationships of SWD and AFF with parasitoid wasps in affected fruit cropping systems in southwestern Virginia. We discovered that parasitoids of vinegar flies are present in Virginia fruit cropping systems, but they do not help to control populations of SWD and AFF. The parasitoids that are present prefer to attack other fly species, are unable to attack SWD and AFF, or do not attack in high enough numbers to have an impact on SWD or AFF populations. Biological control success is more likely to come from parasitoid species that have co-evolved with SWD and AFF in their native ranges.

付記する学位プログラム名: 社会を駆動するプラットフォーム学卓越大学院プログラム / 京都大学 / 新制・課程博士 / 博士(農学) / 甲第25317号 / 農博第2583号 / 新制||農||1104(附属図書館) / 京都大学大学院農学研究科森林科学専攻 / (主査)教授 北島 薫, 教授 小野田 雄介, 教授 井鷺 裕司 / 学位規則第4条第1項該当 / Doctor of Agricultural Science / Kyoto University / DGAM

ABSTRACT Alliaria petiolata (garlic mustard) is an aggressive non-native and invasive forb that negatively impacts native arbuscular mycorrhizal communities and inhibits or prevents the growth of native plant species. Invasive species mitigation and management strategies that use native revegetation vary in success. This study focuses on which species naturally regenerate in areas where A. petiolata has been mitigated to help inform restoration efforts. Seedling emergence of species within the seed bank of four plot types (uninvaded, invaded, chemically treated, and mechanically treated) were observed two years post restoration efforts to determine which native species are likely to persist to seedlings following management. Species abundance and percent cover of all plant species were recorded to evaluate the presence of species in addition to plant health and physiological differences. Native species abundance was significantly reduced within herbicide treated plots as compared to untreated and invaded plots (p=0.02). Plots treated with mechanical removal had the greatest percent cover of native plants as compared to all other plot types but were dominated by colonizing species which is typical of a disturbed habitat. Mechanical removal also resulted in a greater abundance and stability among functional groups of native species, than those treated with herbicide. Forb species dominated coverage of plots over other functional groups when treated with herbicide. Alliaria petiolata invasion and management methods significantly impacted forb and graminoid species, as they had significantly lower abundance in plots treated with herbicide. The results demonstrate that the method of removal as well as the presence of A. petiolata affects emergence of plant species from the seedbank. The additional disturbance of mechanical removal may alter successional trajectories following invasion. Herbicide treatment resulted in the most similar species abundance as the uninvaded reference plots, which had the lowest seedling emergence and percent cover.

Doctor of Philosophy / Department of Biology / Keith B. Gido / Many native freshwater animals are imperiled as a result of habitat alteration, species introductions and climate-moderated changes in disturbance regimes. Native conservation and nonnative species management could benefit from greater understanding of critical factors promoting or inhibiting native and nonnative success in the absence of human-caused ecosystem change. The objectives of this dissertation were to (1) explain spatiotemporal patterns of native and nonnative success, (2) describe native and nonnative response to uncharacteristic wildfire disturbance, and (3) test the hypothesis that wildfire disturbance has differential effects on native and nonnative species. This research was conducted across six sites in three reaches (tributary, canyon, and valley) of the unfragmented and largely-unmodified upper Gila River Basin of southwestern New Mexico. Secondary production was measured to quantify success of native and nonnative fishes prior to wildfires during 2008-2011. Native fish production was greater than nonnatives across a range of environmental conditions, although nonnative fish, tadpole, and crayfish production could approach or exceed that of native macroinvertebrates and fishes in canyon habitats, a warmwater tributary, or in valley sites, respectively. The second objective was accomplished by measuring biomass changes of a warmwater native and nonnative community during 2010-2013 before and after consecutive, uncharacteristic wildfires. Several native insect and fish taxa decreased after both wildfires, whereas nonnative decreases were most pronounced for salmonids and more limited for other taxa. Finally, effects of uncharacteristic wildfires followed by extreme flooding on metapopulations of native and nonnative fishes were contrasted during 2008-2013. Wildfire and flood disturbances increased extinction probabilities of all native fishes while leaving many nonnative fishes unaffected. These findings revealed a swinging pendulum of native and nonnative success, wherein wildfire disturbance resulted in a pendulum swing in favor of nonnatives. Ensuring the pendulum swings back in favor of natives will be facilitated by management activities that decrease wildfire size and intensity and maintain inherent ecosystem resilience.

Thesis (PhD (Conservation Ecology and Entomology))--University of Stellenbosch, 2009. / ENGLISH ABSTRACT: Natural and semi-natural ecosystems and human communities worldwide are under siege from a growing number of destructive invasive alien species. Alien species are those whose presence in an area is due to intentional or accidental introduction as a result of human activities. Some alien species become invasive, and some cause tremendous destruction to the ecosystem and their stability, but we do not yet understand fully the many factors that determine the levels of invasiveness in alien species. However, management of alien plants requires a detailed understanding of the factors that make them invasive in their new habitat. The aim of this study was to explore in detail the processes and potential for invasion of Schinus molle (Peruvian pepper tree) into semiarid savanna in South Africa and to examine the potential for this species to invade further in these ecosystems, and in other South African biomes. In this thesis I explored the patterns and processes of invasion of S. molle in semiarid savanna using small-scale experiments to investigate physical and ecological barriers to invasion that prevent or accelerate the invasion of this species. I examined factors such as pollination; seed production; seed dispersal; seed predation and viability, all known to contribute to invasiveness. I highlighted the critical role of microsite conditions (temperature, humidity, water availability) in facilitating S. molle seedling establishment in semi-arid savanna and demonstrated that microsite type characteristics need to be considered for management and monitoring of the species in South Africa. I demonstrated the ability of S. molle to out-compete indigenous woody plants for light and other resources and also showed that disturbance of natural ecosystems was not a prerequisite for invasion, although human activities such as tree planting have played a major role in disseminating this species in South Africa. Predicting the future distribution of invasive species is very important for the management and conservation of natural ecosystems, and for the development of policy. For this reason, I also assessed the present and potential future spatial distribution of S. molle in South Africa by using bioclimatic models and a simulation-based spread model. I produced accurate profiles of environmental conditions (both biophysical and those related to human activities) that characterize the planted and naturalized ranges of this species in South Africa, by linking species determinants, potential habitat suitability and likely spread dynamics under different scenarios of management and climate change. All those components provided insights on the dynamics of invasions by fleshy-fruited woody alien plants in general, and on S. molle invasions in South Africa in particular. I developed a conceptual model that described S. molle population dynamics leading to an understanding of the processes leading to the invasive spread of this species in South Africa. This work also emphasized the need for policy review concerning the invasive status of S. molle in South Africa, and recommendations are made for future research. / AFRIKAANSE OPSOMMING: Die natuurlike and half-natuurlike ekosisteme sowel as menslike gemeenskappe wereldwyd word bedreig deur ‘n groeiende hoeveelheid indringerplantspesies. Indringerplantspesies (daardie spesies wie se teenwoordigheid toegeskryf kan word aan opsetlike of toevallige inbringing deur menslike toedoen) is ‘n bedreiging nie net vanweë die massiewe verwoesting van die ekosisteme en ekosisteemstabilitiet nie, maar ook omdat ons nog nie ten volle verstaan hoe hulle van skaars in hul natuurlike omgewing tot dominant in hul nuwe habitat gaan nie. Bestuur van indringer plante vereis ’n begrip van biologise en ekologiese faktore wat lei tot hulle indringing in die nuwe habitat. Die primêre doel van hierdie studie was om in detail uit te vind wat die prosessesse en potensiaal is vir die indringing van S. molle (die Peruviaanse peper boom) in droë savanna en om indringingspatrone in droë savanna met huidige en potensieële toekomstige patrone in ander Suid-Afrikaanse biome te vergelyk. In hierdie tesis ondersoek ek die patrone en prosesse wat verband hou met die indringing van S. molle in droë savanna deur gebruik te maak van kleinskaalse eksperimente om fisiese en ekologise hindenisse te ondersoek wat indringing van S. molle in Suid-Afrika voorkom of versnel. Dit sluit faktore in wat bekend is om by te dra tot indringing van plant spesies, soos bestuiwing, saadproduksie, saadpredasie en kiemkragtigheid. Ek het die kritieke rol beklemtoon van mikroomgewingskondisies (temperatuur, humiditeit, waterbeskikbaarheid) in die fasilitering van S. molle saailingvestiging in droë savanna en het gedemonstreer dat die tipe mikroomgewingskarakteristieke in ag geneem moet word by betuur en monitering van die verspreiding van S. molle in Suid-Afrika. Ek het die vermoë van S. molle om inheemse plante te uitkompeteer gedemonstreer, en het gewys dat versteuring van natuurlike ekosisteme nie ’n voorvereiste vir S. molle indringing was nie, hoewel menslike aktiwiteite soos boomaanplantings ’n groot rol speel deur by te dra tot indringing van hierdie spesie in Suid-Afrika. Voorspelling van toekomstige verspreiding van indringerspesies is baie belangrik vir die bestuur en bewaring van natuurlike ekosisteme, sowel as vir ontwikkeling van wetgewing. Daarom is die huidige en potensiele toekomstige ruimtelike verspreiding van S. molle in Suid-Afrika bereken deur inkorporering van bioklimaatsmodelle en simulering gebasseer op ’n verspreidingsmodel. Ek het derhalwe akkurate profiele van omgewingstoestande (beide fisiese en daardie wat verband hou met menslike aktiwiteite) wat die aangeplante en natuurlike omvang van die spesie in Suid-Afrika kenmerk geproduseer deur spesiedeterminante, potensieële geskiktheid van habitatte en moontlike verspeidingsdinamika onder verskillende bestuursscenarios en kimaatsverandering te koppel. Al hierdie komponente verskaf insig in die dinamika van die indringing van houtagtige plante met vlesige vrugte oor die algemeen en S. molle in besonder in Suid- Afrika. Hierdie werk beklemtoon ook die behoefte vir hersiening van beleidsrigtings wat betrekking het op die indringerstatus van S. molle in Suid-Afrika en maak aanbevelings vir toekomstige navorsing.

This thesis focuses on the guppy, Poecilia reticulata, as an invasive species. Its non-native distribution, the biology behind its success and the reasons for its introduction are examined. A worldwide email survey revealed that the guppy is established in at least 73 countries outside of its native range and that mosquito control schemes and the release of unwanted aquarium fish are the two primary routes of introduction. Knowledge gaps were identified; primarily the scarcity of scientific evidence for negative impacts of guppy introductions and similarly for mosquito control efficacy. Replicated mesocosm experiments demonstrated that female guppies are capable of routinely establishing populations, and that these retain behavioural viability over several generations. The first mesocosm study suggested that founders with very different evolutionary histories were equally good at establishing populations. The second mesocosm study suggested that monandrous females were extremely successful at establishing behaviourally viable populations, with no decline in behavioural variation. The effectiveness of guppies as mosquito control agents was examined in two related foraging experiments. The first study found little evidence for the presence of ‘prey switching’ in guppies, questioning the validity of previous work advocating their introduction to stabilise prey populations. The second study revealed a preference for non-vector mosquito larvae in a two-prey system. However, both mosquito species were consumed equally readily when habitat complexity increased. The presence of conspecifics affected female foraging behaviour. The presence of males reduced the strength of prey preference in the first study, and the presence of conspecifics of either sex removed prey preference in the second. Both demonstrate that multi-prey systems have important implications for the efficacy of poeciliids in biological control. Despite severe demographic bottlenecks, their adaptability and ability to rapidly increase in numbers enable guppies to establish and persist when introduced. Such bottlenecks are typical of introduction scenarios, warning that particular caution should be exerted when introducing this species, or other livebearing fish, to natural water bodies.

Since their discovery in Yellowstone Lake in 1994, Lake Trout (Salvelinus namaycush) have been the object of an intensive gillnet suppression program due to their predation on native Yellowstone Cutthroat Trout (Oncorhynchus clarkii bouvieri). Managers are also interested in targeting early life stages to augment suppression. A benthic sled was used to sample for Lake Trout eggs at 24 locations, hypothesized to be spawning sites, that encompassed a range of depths, slopes, and substrate composition to determine the location and characteristics of spawning sites in Yellowstone Lake. Lake Trout eggs were collected at seven sites, five of which had not been previously confirmed as spawning sites. Habitat characterization at these sites indicate Lake Trout spawning in Yellowstone Lake is limited to areas with rocky substrate, but is not constrained to areas with interstitial spaces or contour breaks as is seen within the species' native range. Lake Trout fry were captured around Carrington Island, an additional spawning site in Yellowstone Lake, in 2014 and 2015. These fry were significantly larger at each developmental stage, consumed more food beginning at earlier stages, and were captured much later into the summer than fry captured at a spawning site in Lake Champlain. The lack of potential egg and fry predators in Yellowstone Lake could be driving these differences in spawning site selection and fry behavior. This information will allow managers to identify additional spawning locations for suppression and evaluate the impact their efforts might have on the Lake Trout population in Yellowstone Lake.