The timing of biological events in autumn, or autumn phenology, is an important factor in many ecosystem processes. Leaf senescence terminates the growing season, fruiting is important for seed dispersal and frugivorous wildlife, bird migration concludes the breeding season and is a high-mortality event, and insect diapause ends the active season for insects. Climate change and the spread of invasive species have the potential to shift autumn events and ecological processes. However, autumn has been neglected in the phenology literature, and there are many gaps in our knowledge of basic phenological patterns in this season, as well as how they are affected by anthropogenic changes. To address these gaps, I first synthesized the literature on how climate change affects autumn phenology. I found that shifts in autumn phenology can alter reproductive capacities, exacerbate invasions, increase disease transmission rates, reshuffle enemy-prey dynamics, and alter interactions between species. With a focus on autumn interactions between birds and fleshy-fruited plants, my colleagues and I then observed patterns of fruit phenology, using herbarium specimens of 55 species collected across New England, and over 400 species in the living collections of 5 international botanical gardens. Last, I monitored fleshy fruit phenology and abundance at Manomet, a migratory stopover site in coastal Massachusetts, and compared those patterns to seeds identified from landbird fecal samples collected across the autumn season. I found that the sequence of fruiting is moderately consistent from year to year and place to place, and has a significant phylogenetic signal. In wild plants, invasive species fruit, on average, nearly one month later than native species. Considering many landbirds are migrating through New England later over time and in warm years, this suggests birds are increasingly likely to encounter invasive fruits during late-autumn migration. However, bird diets do not reflect the increased availability of invasive fruits in late-autumn; rather, birds show a preference for native fleshy fruits throughout the autumn season. These findings add to our knowledge of how climate change and species invasions affect autumn synchrony, and highlight the importance of native, rather than invasive, fruits as a food source for migratory landbirds. / 2019-11-07T00:00:00Z

Environmental degradation is accelerating worldwide, yet environmental conservation remains limited by funding. Tackling this limitation requires not only absolute increases in funding, but improved prioritisation of actions. On a global scale island ecosystems are of high priority, with invasive species one of their most significant threats. In this thesis I investigate prioritisation of invasive grazing species control, incorporating ecological, economic, and social concerns, on the island of Bonaire, Caribbean Netherlands. To enable the trade-off of potential grazer control options for their ecological impacts I modelled the relationship between of grazer density and vegetation, and watershed vegetation and the coral reef. I found negative relationships for goat and pig grazing with grass presence, and for donkey grazing with ground cover. Coral cover below 10m showed a positive relationship to ground cover, and, surprisingly, a negative relationship to tree biomass. Because conservation action is most likely to be sustainable when connected to funding, I conducted choice experiments with SCUBA divers, which estimated a positive willingness to pay for reef health improvements achieved using terrestrial grazer control. Through communication with local policy makers and practitioners I identified three options for grazer control, eradication, population reduction, or fencing, and estimated costs and social acceptability for each option. Though the ecological models predicted eradication to have the highest impacts on the terrestrial and marine ecosystem, lower costs and higher social acceptability identified fencing as the most suitable option for grazer control on Bonaire in the short term, with the potential to be funded through a fee on SCUBA divers. Through linking ecological, economic, and social considerations within a real world conservation context I illustrate the importance of looking beyond only ecological improvements when prioritising conservation action. This research is directly applicable to policy and practise on Bonaire.

Trophic interactions within aquatic ecosystems are complex, with many different pathways facilitating transfer of energy and nutrients among trophic levels and many different mechanisms that influence energy and nutrient transfer. This is illustrated in the “top down” and “bottom up” regulatory effects on aquatic food webs, through which primary producer biomass and, therefore, herbivore and carnivore densities, are influenced by both nutrient availability (bottom up) and densities of consumers at higher trophic levels (top down). In an aquatic food web, planktivore presence can directly alter zooplankton density via consumption, while indirectly shaping phytoplankton biomass via reduced herbivore abundance and the release of nutrients due to excretion, egestion, and decomposition. Novel species introduced into an established food web may have important consequences. An invasive species may impact an invaded food web through competition, predation, alteration of nutrient cycling, or, potentially, through facilitation of native species or other invasives. For example, an invasive planktivore may shift zooplankton density or community composition, thereby facilitating phytoplankton blooms. Such a planktivore may also compete with and, potentially, replace native species. Moreover, an invasive species that reaches high densities within its invaded range may serve as an important nutrient sink as it consumes a high biomass of native species or a nutrient source via excretion or decomposition. Two such invasive species with the capacity to dramatically alter native food web dynamics are bighead (Hypophthalmichthys nobilis) and silver carp (H. molitrix; collectively, bigheaded carp). Bigheaded carp are large-bodied, planktivorous fishes that were introduced into the United States in the 1970s and have since spread throughout much of the Mississippi River and its tributaries. These species currently threaten the Great Lakes, where they may constitute a threat to native planktivores such as gizzard shad (Dorosoma cepedianum) and commercially important species such as walleye (Sander vitreus), although there remains a great deal of uncertainty surrounding their potential ecosystem impacts. Consumption of both zooplankton and phytoplankton has been observed in bigheaded carp, although their impact on primary producer biomass is not well understood. Although field observations suggest that condition and abundance of native planktivores, including gizzard shad and bigmouth buffalo (Ictiobus cyprinellus), as well as zooplankton density, have declined following the bigheaded carp invasion, there is little direct, experimental evidence of bigheaded carp food web impacts. Therefore, I sought to examine the effects of bigheaded carp on native ecosystems through a series of mesocosm experiments at the Southern Illinois University pond facility. My primary objectives were to 1) observe potential competition between bigheaded carp and the native gizzard shad, 2) evaluate effects of bigheaded carp predation on zooplankton and phytoplankton communities, 3) assess impacts of bigheaded carp decomposition on nitrogen and phosphorus availability, and 4) measure the rate at which bigheaded carp excrete nitrogen and phosphorus. In order to elucidate the impacts of bigheaded carp on gizzard shad growth and survival, zooplankton and phytoplankton densities, and nitrogen and phosphorus availability in the pelagic and benthic pools and to determine whether gizzard shad experience a diet shift in response to bigheaded carp presence, I performed two mesocosm experiments with three treatments: gizzard shad only, gizzard shad, bigheaded carp, and fishless control (Chapter 1). I predicted that bigheaded carp would reduce zooplankton densities but that gizzard shad, which are both detritivorous and planktivorous, would be unaffected due to their ability to use detritus as an alternative food source. Additionally, both predator release via zooplankton consumption and increased nutrient availability from bigheaded carp excretion would stimulate phytoplankton. I found that gizzard shad survival was reduced by bigheaded carp presence but that surviving gizzard shad did not experience a decline in growth in the bigheaded carp plus gizzard shad treatments. This may have been due to the ability of gizzard shad to consume detritus, as foreguts of sampled gizzard shad in Experiment 2 contained mostly detritus. Moreover, phytoplankton density declined in the presence of silver carp in Experiment 2, suggesting silver carp herbivory. In addition, nitrogen and phosphorus availability in either the pelagic or benthic pools did not appear to be impacted by bigheaded carp presence. After demonstrating experimentally the overall negative impact of bigheaded planktivory on native food webs, I focused my remaining two chapters on the effects of silver carp on nutrient availability. In Chapter 2, I outline a decomposition experiment testing for potential changes in pelagic and benthic nitrogen and phosphorus availability and, in turn, phytoplankton, zooplankton, and macroinvertebrate densities in response to silver carp decomposition. Although silver carp die offs have been reported throughout the Midwest, little is known about the magnitude of those die offs and their consequences for the ecosystem. In this study, silver carp decomposition did not appear to alter nutrient availability or densities of phytoplankton or invertebrates. However, in comparison to northern streams in which salmon spawning and decomposition provide an important nutrient subsidy, the mesocosms used in this study have relatively higher background nutrient concentrations. Thus, silver carp decomposition, at least at the densities studied, may have little importance to in-stream nutrient availability. Lastly, because I am interested in how bigheaded carp, particularly silver carp, alter nutrient dynamics in invaded food webs, it is necessary to calculate silver carp nitrogen and phosphorus excretion rates, as well as body nitrogen and phosphorus content (Chapter 3). Nutrient stoichiometry theory predicts a balance between the relative consumption of nutrients by an organism and the extent to which the organism retains nutrients in its tissues or excretes them. Thus, it is a useful tool in determining how an invasive species may alter nutrient availability via consumption and excretion. In Chapter 3, I describe the body and excretion N:P ratios for silver carp, which exhibit a lower body N:P ratio than excretion N:P, suggesting that these organisms may serve as a sink for phosphorus. Moreover, silver carp body excretion N:P ratios were higher than those reported for gizzard shad, suggesting that, in regions where silver carp may replace gizzard shad or lower gizzard shad population density via competition (Chapter 1), silver carp may alter nutrient cycling processes in aquatic ecosystems by shifting the overall available N:P ratio. Bigheaded carp may pose a significant threat to invaded ecosystems through their potential to compete with native species, reduce plankton densities, and alter nutrient availability. However, although bigheaded carp are expanding in range and approaching the Great Lakes, the full extent of their ecosystem impacts remain uncertain. Through my work on bigheaded carp food web impacts, particularly the influence of silver carp on native species and nutrient cycling processes, I have found that bigheaded carp have the capacity to negatively impact invaded ecosystems overall by reducing zooplankton, phytoplankton, and forage fish densities. Moreover, as bigheaded carp in particular continue to reach high densities as they expand in range, their capacity to alter relative nitrogen and phosphorus availabilities must be monitored to understand the extent of their influence. Due to their ability to disrupt top down and bottom up processes in freshwater ecosystems, bigheaded carp constitute a critical environmental issue in the Great Lakes area and throughout the Midwest and, thus, it is imperative to continue to experimentally assess how bigheaded carp interact with native species to the detriment or benefit of U.S. freshwater communities.

Invasões biológicas estão entre as principais causas da perda de biodiversidade no planeta. Ambientes isolados como as ilhas oceânicas e ambientes sujeitos a distúrbio antrópico são considerados mais propensos à invasão. Para as plantas, o sucesso na invasão pode ter relação com a superioridade na competição com as espécies nativas, que pode ocorrer através da produção de substâncias alelopáticas. Dentre as 100 principais espécies invasoras do planeta está a Leguminosa Leucaena leucocephala, que produz substâncias com potencial alelopático e está estabelecida em ilhas oceânicas tropicais em todo o mundo. No Brasil, a invasora foi introduzida na ilha de Fernando de Noronha, onde ocupa vastas áreas. Apesar da relevância desta ilha para a conservação da biodiversidade, não há informações essenciais para o manejo da invasora, como a situação da invasão e seus fatores determinantes. Neste trabalho, realizado em Fernando de Noronha, utilizamos experimentos para investigar a alelopatia como mecanismo associado à invasão e para avaliar o efeito de L. leucocephala sobre o estabelecimento de Erythrina velutina (Leguminosae), espécie nativa comum na ilha, mas frequentemente excluída das áreas invadidas por L. leucocephala. Não encontramos indícios de efeitos alelopáticos de L. leucocephala sobre a germinação de E. velutina, mas a exótica reduziu o crescimento e a sobrevivência da nativa. O efeito negativo é potencializado quando L. leucocephala está associada à Capparis flexuosa (Capparaceae), única espécie nativa frequentemente encontrada em áreas invadidas. Isoladamente, o efeito de C. flexuosa sobre E. velutina varia de positivo a neutro, evidenciando que o saldo das interações entre espécies nativas é alterado na presença de uma exótica. Adicionalmente, diagnosticamos a extensão atual da invasão e sua expansão nos últimos 20 anos, seus fatores determinantes e o impacto sobre a comunidade de plantas nativas em Fernando de Noronha. O diagnóstico da invasão mostrou que L. leucocephala está amplamente distribuída pela ilha, povoando densamente a maioria dos locais onde ocorre. A área ocupada pela espécie aumentou cerca de 40% nos últimos 20 anos, e não há restrições ambientais para o estabelecimento da exótica, embora ela seja favorecida pela atividade agropecuária. Em áreas invadidas o número de espécies nativas diminui quase pela metade e observamos uma tendência à homogeneização da comunidade. É provável que o alto grau de perturbação antrópica em Fernando de Noronha gere limitações à dispersão e modifique os ambientes tornando-os desfavoráveis para o estabelecimento de espécies nativas. Entretanto, há fortes evidências de que L. leucocephala causa mudanças ecológicas na ilha influenciando na perda espécies nativas. Considerando a importância biológica de Fernando de Noronha, ações de controle da expansão da exótica e restauração das áreas invadidas demonstram-se urgentes / Biological invasions are among the main causes of biodiversity loss on the planet. Isolated environments such as oceanic islands and disturbed environments are considered more prone to invasion. For plants, the invasion success may be related to advantages in competition with native species, which may occur through the production of allelopathic substances. Among the 100 most invasive species on the planet is the legume Leucaena leucocephala, which produces substances with putative allelopathic effects and is established on tropical oceanic islands worldwide. In Brazil, the invader was introduced on the island of Fernando de Noronha, where it occupies vast areas. Despite the relevance of this island for biodiversity conservation, important information for the management of the invasion, as its extension and determinants, do not exist. In this work we use experiments to investigate allelopathy as a mechanism associated with the invasion and to evaluate the effect of L. leucocephala on the establishment of Erythrina velutina, a native species common on the island, but often absent from invaded areas. We found no evidence of allelopathic effects of L. leucocephala in the germination of E. velutina, but the exotic reduced the growth and survival of the native. The negative effect is enhanced when L. leucocephala is associated with Capparis flexuosa, the only native species often found in heavily invaded areas. When alone, the effect of C. flexuosa on E. velutina varies from positive to neutral, indicating that the balance of interactions between native species is altered in the presence of an exotic. Additionally, we describe the current distribution of L. leucocephala and its expansion in the last 20 years in Fernando de Noronha. We also investigate the environmental and anthropic factors determining the invasion and the impact of L. leucocephala on the plant community. We found that L. leucocephala is widely distributed throughout the island, densely populating most places where it occurs. The area occupied by the species increased about 40% in the last 20 years, and there are no environmental restrictions for the establishment of exotic, although it is favored by farming. In invaded areas, the number of dominant native species decreased by almost half and we observed a tendency towards homogenization of the community. It is likely that the high degree of human disturbance in Fernando de Noronha poses dispersal limitations and modifies the environments making them unsuitable to the establishment of natives. However, there are strong evidences that L. leucocephala is driving ecological changes on the island that influence in native species loss. Considering the biological importance of Fernando de Noronha, actions to control the expansion of exotic and to restore the invaded areas are urgent

O uso de animais de montaria em áreas naturais tem gerado grande preocupação, pois podem gerar impactos ambientais. Um dos problemas que precisa ser investigado é a relação entre animais de montaria e a presença de plantas exóticas em áreas de alto valor ecológico. Assim, o presente trabalho teve como objetivo testar um protocolo de germinação para sementes de duas espécies de gramíneas, braquiária (Urochloa decumbens) e capim gordura (Melinis minutiflora) que passaram pelo trato digestório de cavalos. As sementes foram ofertadas aos animais junto com a ração com posterior coleta das fezes. No experimento foram realizados três tratamento. No tratamento \"braquiária\" foram ofertadas sementes de Urochloa decumbens, no tratamento \"capim gordura\" foram ofertadas sementes de Melinis minutiflora e o \"sem tratamento\" foi ofertada apenas a ração. Parte das sementes foram retiradas das fezes e submetidas a testes de germinação e o material restante foi utilizado para testar a capacidade germinativa das sementes diretamente nas fezes em um ambiente controlado. Na germinadora verificou-se que as sementes que passaram pelo trato digestório dos cavalos tiveram uma menor taxa de germinação comparadas com as sementes com tratamento de quebra de dormência. No tratamento feito em casa de vegetação as sementes que foram retiradas das fezes e que germinaram foram identificadas. Não houve diferença na taxa de germinação entre os cavalos, o que ocorreu entre os tratamentos. No tratamento braquiária predominou a germinação de Urochloa decumbens, no tratamento capim gordura a maior taxa de germinação de Melinis minutiflora e no sem tratamento predominou o nascimento de outras espécies. As sementes estudas tem potencial de germinar, crescer e frutificar, porém não se sabe se as espécies podem formar uma população capaz de colonizar novas áreas. / The use of riding animals in natural areas has caused great concern because their use cause environmental impacts. One of the problems that needs to be investigated is the relationship between the use of riding animals and the presence of exotic plants in natural areas. This study aimed to test a germination protocol for the of two species Urochloa decumbens and Melinis minutiflora that has passed through the digestive tract of the horse. The seeds were offered to horses and their feces were collected. Tree treatments were set, in \"braquiaria\" treatment seeds Urochloa decumbens were offered to the horses, in the \"capim gordura\" treatment seeds of Melinis minutiflora offered and the untreated were offered only horse food. Part of the seeds were removed from the feces and undergo germination test and the remain of the feces were used to test in a green house. The germination rate of the seed from de horse dung was lower than germination rate of the dormancy breaking treatment seeds. The seeds that germinate on the greenhouse were identified. There were no difference between the germination rate between treatments, but there was between species in braquiaria treatment predominated the germination of Urochloa decumbens in the grass fat treatment most Melinis minutiflora germination rate and the untreated predominated the germination of other species. Seeds from horse dung has the potential to germinate, grow and bear fruit, but it is not known if this species can form a population able to colonize new areas.

Biological invasions threaten biodiversity globally, and degraded ecosystems increase the potential for invaders to compete with threatened native populations. In natural systems, niche partitioning minimizes interspecific competition, but introduced species may alter expected outcomes by competing with ecologically similar species for scarce resources. Where food production is highly variable, coexistence of native and invasive competitors may depend on dietary niche flexibility. Territorial species under invasion face additional challenges to maintain economically defendable territories. From 2011-2016, we conducted removal and behavior experiments to determine effects of non-territorial introduced Abert’s squirrels (Sciurus aberti) on diet, space use, and territoriality of endangered Mount Graham red squirrels (MGRS; Tamiasciurus fremonti grahamensis) in their declining habitat in the Pinaleño Mountains, Arizona. We collected comparative data from Arizona sites of natural syntopy between Abert’s and Fremont’s squirrels (T. fremonti). Stable isotope analysis revealed similar dietary partitioning among populations. Experimental removals did not appear to affect MGRS diet. Space use by MGRS responded inconsistently to removals; territory sizes increased after the first removal, but did not change following the second removal. Territory sizes and body mass of MGRS were sensitive to conspecific population density and food production. Behavioral experiments showed MGRS were more aggressive than other Fremont’s squirrels (hereafter, red squirrels). Dietary flexibility of Abert’s squirrels may have facilitated coexistence with MGRS, possibly due to coevolved resource partitioning with red squirrels. However, aggressive territoriality toward Abert’s squirrels may incur fitness costs for MGRS especially during poor food production years. Climate change may reduce the advantage of ecological specialist species globally, and where introduced species are better-adapted to novel environmental conditions, native species may ultimately be replaced.

Grasslands in the western United States face severe environmental threats including those brought about by climate change, such as changes in precipitation regimes and altered fire cycles; land-use conversion and development; and the introduction, establishment, and spread of non-native species. Lehmann’s lovegrass (Eragrostis lehmanniana) was introduced to the southwestern United States in the early 1900s. Since its introduction, it has become the dominant grass in the mid-elevation grasslands of southern Arizona, including the Santa Rita Experimental Range (SRER), where it has displaced native grasses including Arizona cottontop, three awns, and gramas. Like all plants in terrestrial ecosystems, this grass harbors fungal symbionts that can be important for its establishment and persistence. This thesis focuses on fungal symbionts of Lehmann’s lovegrass and has two components. First, the diversity and distributions of endophytes in Lehmann’s lovegrass are evaluated in the context of biotic and abiotic factors in the SRER. Culturing from roots and shoots of Lehmann’s lovegrass at points beneath and outside the canopy of native mesquites, which are encroaching on grasslands over time, provides insight into how a single plant species can exhibit local variation in the composition of its symbionts. Second, the thesis is used as the basis for engagement of students in science, technology, engineering, and mathematics (STEM) through the development and implementation of classroom- and field activities centered on endophytes, which help high school students address core learning aims while also gaining real research experience. Engaging students in important questions relevant to their local environment can catalyze interest in science and help students cross the threshold into research. The contributions of such approaches with respect to learning not only fulfills key next-generation science standards and common core objectives, but provides students with a meaningful introduction to the excitement, importance, and accessibility of science.

Invasive species are becoming more common as human interactions within coastal waters and the aquarium trade continues to increase. The establishment of the invasive lionfish complex Pterois volitans and P. miles from the Indo-Pacific to the Western Atlantic, Gulf of Mexico, and the Caribbean Sea has had significant negative effects on reef fish biodiversity and economically important species. Their rapid colonization and success has been attributed to their biological and ecological life history traits as well as their absence of predation. Past research has highlighted these characteristics; however, there is a knowledge gap in lionfish parasitism. This research explored the enemy release hypothesis as a key success factor in rapid establishment in the invaded range on a biogeographical scale. The diversity of lionfish parasitism was compared among 15 geographically diverse sites within the invaded range, incorporating the time of introduction at each site. Eight new parasites are described for the first time in the invasive lionfish: (1) a Cymothoid isopod: Rocinela stignata, (2) four nematodes: Raphidascais sp., Contraceacum sp., Paracuria adunca and Hysterothylaceum sp., (3) one digenean: Tergestia sp., (4) two acanthacephalans: Serracentis sp. and Dollfusentis sp., and (5) two cestodes: Nybelinia sp. and Tentacularia sp. Lionfish from the east coast of Florida exhibited the highest abundance in parasite fauna while other invaded areas yielded low abundance and diversity. Comparisons between lionfish parasitism from the past native range studies and the invaded range suggest that vectors of time, life history traits, and trophic interactions structure the lionfish parasite community. Lionfish in the Western Atlantic and Caribbean were found to be host for generalists parasite species within the coastal ecosystem. Consequently, lionfish have relatively low parasite abundance, supporting the enemy release hypothesis and its direct relation to their invasion success.

The simultaneous increase in biological invasions and habitat alteration through the building of coastal infrastructure is playing an important role in reshaping the composition and functioning of nearshore marine ecosystems. This thesis examined patterns of marine invasions across anthropogenic and natural habitats and explored some of the processes that influence establishment and spread of invaders. The goals of this thesis were four-fold. First, I examined the habitat distribution of marine nonindigenous species (NIS) spanning several taxonomic groups and geographical regions. Second, I conducted systematic subtidal surveys in anthropogenic and natural habitats and investigated the distribution of nonindigenous ascidians on Southern Vancouver Island, British Columbia, Canada. Third, I tested methods for in-situ larval inoculations and utilized these techniques to manipulate propagule supply and assess post-settlement mortality of ascidians across habitat types. Fourth, I investigated the role of biotic resistance, through predation by native species, on the survival of ascidian colonies in anthropogenic and natural habitats. Results from this research showed that anthropogenic habitats are hubs for marine invasions and may provide beachheads for the infiltration of nearby natural sites. Specifically, a literature review of global scope showed that most NIS are associated with anthropogenic habitats, but this pattern varied by taxonomic group. Most algal and mobile invertebrate NIS were reported from natural habitats, while most sessile NIS were reported from artificial structures. Subtidal field surveys across both anthropogenic and natural habitats showed that nonindigenous ascidians were restricted largely to artificial structures on Southern Vancouver Island and that this pattern is consistent across their global introduced ranges. Field manipulations using the ascidian Botrylloides violaceus as a model organism, showed that post-settlement mortality is high and that large numbers of larvae or frequent introduction events may be needed for successful initial invasion and successful infiltration of natural habitats. Experiments also showed that predation by native species can limit the survival of B. violaceus in anthropogenic and natural habitats. This dissertation contributes knowledge about the patterns and processes associated with habitat invisibility; provides insight into factors affecting colonization; and supplies valuable information for predicting and managing invasions. / Graduate / 0329

The main objective of this research was to determine the seed bank longevity of Melinis repens at two Southern Florida sites. Seeds were divided among different exposure levels (shade versus sun) and depths (surface versus buried) and tested for baseline viability using 2,3,5-Triphenyl-tetrazolium chloride. Statistical analysis determined that at the pine rockland site there was a significant interaction between time, exposure, and depth. The initial mean viability at this site declined from 49.71% to 11.26% and 13.06% for sun/buried seeds and sun/surface seeds, respectively, by month 8. The mean viability of shade/surface seeds and shade/buried seeds declined to 24.56% and 22.06% after 8 months. There were no significant effects in the Florida scrub. In order for land managers to completely remove this species from a site, treatment with herbicide will need to continue for a minimum of one year to effectively kill all viable seeds in the seed bank.