Keystone species are widely distributed across aquatic and terrestrial ecosystems and are fundamental in preserving the structure, diversity, and stability of an ecological community due to its disproportionately large impact on its community relative to its biomass. As biodiversity of ecosystems becomes more threatened with urbanization and habitat destruction, it is imperative to understand a keystone species' role in maintaining ecosystem function. One of the ways to do so is by examining their significance and connection to the ecosystem food web. Within North American freshwater ecosystems is the pebble nest-building minnow, the bluehead chub (Nocomis leptocephalus; "chub"). Chubs provide spawning habitat for not only themselves, but for other minnows, collectively called "nest associates". In this work, I observe the predatory and potential mutualistic interactions between chubs, nest associates, and their predators. In Chapter 1, I observe spawning nests to identify the predators of adult chubs, nest associates, and embryos. I further investigate how nest visibility covariates including minnow activity, minnow abundance, nest size (area), and nest growth affect predator encounter rate to spawning nests. I found a total of 23 diverse taxa to prey on the adult minnows and minnow embryos on chub spawning nests, 14 predators of which had not been reported in literature. One of these predators was the common snapping turtle (Chelydra serpentina; "turtle"). Additionally, I found that activity, abundance, nest size (area), and nest growth had a significant effect on predator encounter rate, attracting predators to seek spawning nests for prey. In Chapter 2, I investigate the effect of ambient temperature on turtle epizoic coverage during the spawning season and provide preliminary evidence of a potential cleaning symbiotic mutualism between the turtle and minnows. I found that epizoic coverage decreases during the duration of a minnow spawning season after an initial increase with early summer warming, and my results also present unique and shared bacterial communities across three sources, the ambient environment, gut contents of minnows, and turtles. The results additionally revealed there to be bacterial communities unique between minnows and turtles that were not identified in the ambient environment. Overall, this study is first to systematically document predators of chub spawning nests and first to provide preliminary evidence of a cleaning symbiotic mutualism between a freshwater turtle and minnow species (or freshwater turtles and fish in general), which, thus far, has not been explored in freshwater ecosystems. This work demonstrates that chub spawning nests are a crucial entity of the freshwater food web structure across Nocomis' distribution range and reveals that chub spawning nests create an interconnection between a diversity of fauna in a freshwater ecosystem. / Master of Science / Ecological communities often include species that are essential in ensuring the overall stability and biodiversity of an ecosystem. These species, otherwise called keystone species, play a crucial role in facilitating interconnections within the ecosystem's food web. The bluehead chub (Nocomis leptocephalus; "chub") is an example of a keystone minnow found in North American freshwater streams. This minnow engages in a complex, distinguished act when it reproduces, making mounded, pebble nests using only its mouth. Chubs are not the only minnow species interested in this engineering complexity. Various minnow species called "nest associates" reproduce on the nests as well, providing an appearance of a mutualism: all species involved benefit from the interaction. While this interaction has been observed, there is limited research identifying predators of chub nests and if there are potential mutualisms with any of these predators. In this work, I identify predatory and mutualistic interactions between chubs, nest associates, and their predators. In Chapter 1, I identify predators of chub nests and observe variables that attract these predators to the nests. In Chapter 2, I explore a potential, mutualistic interaction between these minnows and an identified predator from this research, the common snapping turtle (Chelydra serpentina; "turtle"), whereby minnows feed on potentially harmful growth of algae and bacteria on turtles, while turtles benefit from the cleaning. For Chapter 1, my results revealed that a chub nest is a hotspot for predator diversity, showing 23 diverse taxa as predators, in which 14 of the identified taxa are novel for ecological literature. Additionally, variables that were observed to attract predators to chub nests were minnow activity, minnow abundance, nest size (area), and nest growth. Results for Chapter 2 demonstrated that there are unique bacterial communities between turtles and minnows that are not found in the stream environment, therefore providing preliminary evidence of mutualistic interaction between the coexisting species. Overall, this study is the first to systematically document predators of chub nests. This study is also first to investigate a mutualistic interaction between minnows and turtles in a freshwater ecosystem, an area that has not been previously explored, unlike similar interactions in marine ecosystems. Cohesively, the keystone species, the chub, and their reproductive nests, are important for the aquatic food web structure and the interconnectedness to their overall ecosystem function. This research further stewards scientific knowledge about how important Nocomis are to natural freshwater ecosystems.

Made available in DSpace on 2015-03-04T18:51:19Z (GMT). No. of bitstreams: 1 TeseLeonardoGama.pdf: 673646 bytes, checksum: b1f31519bf1d9bd07a3dde2a7574423b (MD5) Previous issue date: 2010-04-01 / Coordenacao de Aperfeicoamento de Pessoal de Nivel Superior / A food web comprises exchanges of matter and energy that occur among species and between biotic and abiotic environment. Given that abiotic components form the basal resources, the approach of this work consists of evaluating the effects of nutrients input in strategic models that describe food web and chain dynamics. Its focus lies on the determination of the nature of equilibrium populations as well as on their dynamics for different functional responses. Strategic models that describe the behavior of interactive populations under nutrient inputs are an important basis for outlining general phenomena that occur in community dynamics. / Uma rede trófica reúne as trocas de matéria e energia que ocorrem entre as espécies e entre o meio biótico e abiótico. Visto que os componentes abióticos formam a fonte de recursos basais, a abordagem deste trabalho consiste na avaliação dos efeitos da entrada de nutrientes alóctones em modelos estratégicos que descrevem a dinâmica de redes e cadeias tróficas, concentrando-se na determinação das características das populações de equilíbrio e das dinâmicas das espécies com diferentes respostas funcionais. Modelos estratégicos que contêm informações acerca do comportamento de populações interativas frente à entrada de nutrientes são uma base importante no delineamento de fenômenos gerais que podem ocorrer dentro da dinâmica de comunidades.

Les systèmes écologiques sont des systèmes complexes qui ne peuvent pas être d´écrits par un unique modèle mathématique. De nombreux modèles peuvent être construits pour un même système, selon les internets du modélisateur et ses choix dans la construction du modèle. Quel est l’impact de ces choix dans la construction du modèle sur les prédictions de la dynamique des systèmes écologiques et les informations qu’elles fournissent sur la résilience de ces systèmes est la question générale qui guide le travail présente dans cette thèse. Cette thèses focalise sur un choix entre formulations de modèle basées sur des mécanismes biologiques et qui décrivent les données empiriques avec la même efficacité. Ces modèles sont proches l’un de l’autre, donc on s’attendrait `a ce que leurs prédictions soient similaires. Cependant, nous montrons avec un exemple générique de modèle prédateur-proie que des formulations similaires du processus de prédation peuvent prédire des dynamiques qualitativement différentes en terme de: (i) nombre et type d'états stables, et (ii) réponse et résilience du système face à une perturbation extérieure. Ces différences dans les prédictions du modèle sont expliquées par une analyse mathématique détaillée du modèle prédateur-proie. Ensuite, ce modèle est étendu à des réseaux trophiques compos´es de dizaines d’espèces. La complexité de ces réseaux (nombre d'espèces et d’interactions) explique leur persistance, alors que leur dynamique temporelle est fortement affectée par la fonction utilisée pour modéliser la prédation. Des méthodes sont ´également proposées pour quantifier la sensibilités d’un modèle. Finalement, nous montrons que si un minimum de détails biologiques sont pris en compte, des modèles prédateurs-proies sont moins sensibles `a la formulation de la prédation. Ceci nous donne des pistes pour gérer les incertitudes dans la construction d’un modèle, qui sont intrinsèques à la complexité des systèmes naturels. / Ecological systems are complex systems which cannot be described by a single mathematical model. Multiple modelsof a same system can be built, depending on modeller’s interests and on its choices during model construction. Howfar these choices in model construction can affect the predicted dynamics of ecological systems and the informationthey provide on their resilience? is the general question that leads the research presented in this thesis. This thesisfocuses on a choice between model formulations that are based on biological mechanisms and describe empiricaldata with the same accuracy. These models are close to each other, so they are expected to predict similar systemdynamics. However, we show through a generic example of predator-prey model that similar formulations of thepredation process can predict qualitatively different system dynamics in term of: (i) number and type of stablestates, and (ii) system response to external disturbance and its potential for recovery. These differences in modelpredictions are explained by a detailed mathematical analysis of the predator-prey model. Next, this model isextended to complex food webs made of tens of species. The complexity of these networks (number of species andinteractions) drives their persistence, whereas their temporal dynamics is strongly affected by the function used tomodel predation. Methods to quantify model sensitivity are also proposed. Finally, we show that if a minimumlevel of biological details is included, predator-prey models are less sensitive to predation formulation. This providea clue to deal with uncertainties in model construction, which are intrinsic to the complexity of natural systems.

In a changing climate, it is increasingly important to be able to model environmental effects on food webs, and to do that, one must have appropriate dynamic models. I present a shallow lake ecosystem model where producers, grazers, carnivores, piscivores, and detritivores are coupled through resource (light, nutrients and detritus) fluxes between the benthic and pelagic habitats and through carnivore life history events (ontogenetic habitat and diet shifts). The two habitats each contain primary producers, grazers, carnivores and detritivores. Within the habitats, there is strong top-down regulation, but across habitat boundaries, bottom-up interactions drive production. In the absence of piscivores, stage-structured carnivores cause intriguing patters of alternative stable states. Notably, the model predicts a lesser dependence on benthic production with detritus presence. Model predictions are largely in agreement with empirical studies. The results have implications for management of freshwater, and for the interpretation of previous models.

<p>The abundance of predators and their impact on ecosystem dynamics is a vividly discussed topic in current ecology. In my studies, incorporating field observations, field experiments and theoretical modeling, I explored the importance of predators and predation in a low arctic tundra ecosystem in northern Norway. This involved observing the abundance and spatial activity of predators (raptors and small mustelids); manipulating the abundance of predators (spiders and birds) in an arthropod community; and exploring the theoretical consequences of intraguild predation on the coexistence among predators.</p><p>The results show that predation is important both in the arthropod assemblage and, depending on the productivity of the community, in the vertebrate assemblage. In arthropod communities predators are at least as abundant as their prey, whereas in the vertebrate part of ecosystem, predators are substantially less abundant than their prey. Still, in both cases predators had strong impact on their prey, influencing the abundance of prey and the species composition of prey assemblages. The impact of predation cascaded to the plant community both in the reticulate and complex arthropod food web and in the linear food chain-like vertebrate community. In the vertebrate-based community we could even observe the long time scale effect on plant community composition.</p><p>Within the predator community, exploitation competition and intraguild predation were the structuring forces. As the arthropod communities consist of predators with different sizes, intraguild predation is an energetically important interaction for top predators. As a consequence, they reduce the abundance of intermediate predators and the impact of intermediate predators on other prey groups. Moreover, being supported by intermediate predators, top predators can have stronger impact themselves on other prey groups.</p><p>In vertebrate communities, intraguild predation seems to be unimportant as energetic link, instead it manifests as an extreme version of interference competition. Therefore intraguild predation reduces the likelihood of coexistence, as it is due limited prey diversity and intense exploitative competition already precarious in the low arctic tundra.</p><p>In conclusion, predators have strong impact on their prey, especially in the more productive parts of the low arctic tundra. This applies even to the food webs with complex and reticulate structure, and these effects carry through the community both in the short time scale of population growth and on the long time scale of population generations.</p>

The abundance of predators and their impact on ecosystem dynamics is a vividly discussed topic in current ecology. In my studies, incorporating field observations, field experiments and theoretical modeling, I explored the importance of predators and predation in a low arctic tundra ecosystem in northern Norway. This involved observing the abundance and spatial activity of predators (raptors and small mustelids); manipulating the abundance of predators (spiders and birds) in an arthropod community; and exploring the theoretical consequences of intraguild predation on the coexistence among predators. The results show that predation is important both in the arthropod assemblage and, depending on the productivity of the community, in the vertebrate assemblage. In arthropod communities predators are at least as abundant as their prey, whereas in the vertebrate part of ecosystem, predators are substantially less abundant than their prey. Still, in both cases predators had strong impact on their prey, influencing the abundance of prey and the species composition of prey assemblages. The impact of predation cascaded to the plant community both in the reticulate and complex arthropod food web and in the linear food chain-like vertebrate community. In the vertebrate-based community we could even observe the long time scale effect on plant community composition. Within the predator community, exploitation competition and intraguild predation were the structuring forces. As the arthropod communities consist of predators with different sizes, intraguild predation is an energetically important interaction for top predators. As a consequence, they reduce the abundance of intermediate predators and the impact of intermediate predators on other prey groups. Moreover, being supported by intermediate predators, top predators can have stronger impact themselves on other prey groups. In vertebrate communities, intraguild predation seems to be unimportant as energetic link, instead it manifests as an extreme version of interference competition. Therefore intraguild predation reduces the likelihood of coexistence, as it is due limited prey diversity and intense exploitative competition already precarious in the low arctic tundra. In conclusion, predators have strong impact on their prey, especially in the more productive parts of the low arctic tundra. This applies even to the food webs with complex and reticulate structure, and these effects carry through the community both in the short time scale of population growth and on the long time scale of population generations.

Based on a field study and several mesocosm experiments, I evaluated the use of pigments as chemotaxonomical biomarkers for phytoplankton community composition in the Baltic Sea and I examined effects of inorganic nutrients on the dynamics of carotenoids and thiamine (vitamin B1) at the phytoplankton–copepod level in marine pelagic food webs. My results show that HPLC pigment analysis combined with CHEMTAX data processing was an accurate alternative to microscopic analysis of Baltic Sea phytoplankton. Experimental supply of N, P and Si affected copepod growth and biochemical status via changes in biomass and composition of their phytoplankton diet. Net population growth rates were generally higher when phytoflagellates dominated (low Si:N ratio) and lower when diatoms dominated (high Si:N ratio). Copepod body concentrations of astaxanthin decreased with fertilization. Correlations with reduced under-water irradiance were consistent with the photo-protective function of this antioxidant. Thiamine concentrations in phytoplankton also decreased with fertilization. In copepods, low Si:N ratios resulted in higher thiamine concentrations than high Si:N ratios. Thiamine concentration and degree of phosphorylation were useful as indicators of thiamine shortage both in phytoplankton and copepods. The concentrations of thiamine and astaxanthin in the copepod communities were positively correlated. As copepods constitute a major link between pelagic primary producers and higher trophic levels, fertilization effects may be responsible for astaxanthin and thiamine deficiencies in salmon suffering from the M74 syndrome, which appeared concurrently with large-scale eutrophication in the Baltic Sea. As both thiamine and astaxanthin are deficient in M74-affected salmon, there is a need for physiological and molecular investigations of possible interactions between the two compounds in living cells.

The purpose of this study was to identify if terrestrial arthropod predators on Baltic Sea shores vary in their use of marine versus terrestrial food items, and to construct a bottom-up food web for Baltic Sea shores. The inflow of marine nutrients in the area consists mainly of marine algal detritus and emerging aquatic insects (e.g. phantom midges, Chironomidae). Diets of coastal arthropods were examined using carbon and nitrogen stable isotope analysis, and a two source mixing model was used to examine proportions of marine carbon to diets. The results suggest that spiders are the terrestrial predators mainly utilizing nutrients and energy of marine origin on Baltic Sea shores, while insect predators such as beetles and hemipterans mainly utilize nutrients and energy derived from terrestrial sources, possibly due to differences in hunting behaviour. That spiders are the predators that benefit the most from the marine inflow suggest that eventual effects of marine subsidies for the coastal ecosystem as a whole are likely mediated by spiders.

Species diversity may simultaneously be a cause and a consequence of variability in population, community and ecosystem properties. Ecology has traditionally focused on elucidating the causes of biodiversity. However, in the last decade and a half ecologists have asked the opposite question: What are the consequences of species diversity? The majority of these studies elucidated the effects of species diversity within single trophic levels. Incorporating trophic complexity is the next step in this research program. In this dissertation I investigated the causes of resource species diversity, as well as the impacts that resource diversity has on rates of consumption and the stability of population, community and ecosystem properties over time in planktonic food webs. The high diversity of phytoplankton found in nature appears to defy the competitive exclusion principle, and elucidating the mechanisms which maintain this diversity continues to be a challenge. In general, variability in limiting factors is required to maintain non-neutral species diversity, but this variability can be generated by forces outside of the competitive community (i.e. exogenous), or may be the outcome of competitive interactions themselves (i.e. endogenous). Using microcosm experiments, I showed that endogenously generated variability in limiting factors was more effective at maintaining phytoplankton species diversity over the long-term, although the strength of this effect depended on the composition of the phytoplankton community. Existing resource diversity has been proposed to generally weaken consumer-resource interaction strengths and limit consumer control of resource biomass. This is because more diverse resource communities are more likely to contain inedible, unpalatable, toxic or non-nutritious species. However, when resource communities contain multiple palatable species, diversity may also accelerate consumption. Using grazing experiments with multiple zooplankton consumer species, I found that the mechanism, direction and magnitude of modulation of consumption depended on the feeding selectivity of the consumer and the composition of the resource community. By altering consumer-resource interaction strengths in the short-term, resource species diversity may impact the stability of consumer-resource dynamics in the long-term. In separate microcosm experiments, I investigated the influence of resource species diversity, community composition and consumer feeding selectivity on population, community, and ecosystem properties over time. Diversity had positive effects on phytoplankton population biomass, resource community biomass, the rate of photosynthesis, the standing stock of particulate nutrients, and the generalist consumer’s population density. It also stabilized resource community biomass and the stocks of particulate nutrients over time. Unexpectedly, diversity did not stabilize either of the consumer populations, regardless of feeding selectivity. This suggests that effects of diversity on resource community properties do not impact consumer dynamics linearly. Resource community composition was generally more important than resource species diversity in determining food web properties. The importance of community composition in determining both the causes and consequences of resource diversity in these experiments points to the importance of species’ traits and the outcomes of their interactions. I suggest that the use of complex adaptive systems theory and trait-based approaches in the future will allow a consideration of the feedbacks between the causes and consequences of species diversity in food webs. / Graduate

Submitted by FABIANA DA SILVA FRANÇA (fabiana21franca@gmail.com) on 2018-02-23T17:33:04Z No. of bitstreams: 1 Arquivo Total.pdf: 2718770 bytes, checksum: 9378d34888dfcacdd356cf58ac71eaa8 (MD5) / Made available in DSpace on 2018-02-23T17:33:04Z (GMT). No. of bitstreams: 1 Arquivo Total.pdf: 2718770 bytes, checksum: 9378d34888dfcacdd356cf58ac71eaa8 (MD5) Previous issue date: 2017-03-22 / The Rufford Foundation / Understanding the ecological role of species within an ecosystem depends largely on knowledge of its trophic relations. Such relations and the position of species within a foodweb are a central step in understanding the dynamics of the communities and the impacts that each species has on the trophic network compartments. The knowledge on the diet of the South American freshwater stingrays has increased considerably over the last years,but still, little is know about their ecological role. Thereby, the aim of this research is to evaluate the importance of the stingray Potamotrygon magdalenae in the structure of a freshwater food web. The feeding habits and the trophic ecology of this species were evaluated using stomach content analysis in combination with stable isotopes of carbon and nitrogen from samples collected in the middle Magdalena river basin, Colombia, and its ecological importance from topological analyzes of trophic networks. Potamotrygon magdalenae showed a specialized diet, feeding primarily on insects. No significant differences of δ13C and δ15N were observed in the muscle of the species between the hydrological periods of high and low waters. The species isotopic niche was intermediate. In terms of ecological importance, it played an intermediary role in the dispersion of indirect effects through the system, due to its intermediate values of centrality and topological importance. These results provide a starting point for further ecological studies about the South American freshwater stingray species, in order to assess their role in the ecosystem. On the other hand, theycomplement the existing studies of Potamotrygon magdalenae. / Compreender o papel ecológico das espécies no ecossistema depende amplamente do conhecimento de suas relações tróficas. Conhecer tais relações e a posição das espécies em umarede é um passo crucial para entender a dinâmica das comunidades e os impactos quecada espécie tem sobre os demais compartimentos. Oconhecimento sobre adieta das espécies de raias de água doce de América do Sul tem aumentado consideravelmente ao longo dos últimos anos, porém, é muito pouco o que se sabe sobre o papel ecológico das espécies nos seus respectivos ecossistemas. Assim, oobjetivo deste trabalho foi avaliar a importânciaecológica da raia Potamotrygon magdalenae na estrutura da rede trófica de um ecossistema de águas continentais. Foram avaliados os hábitos alimentares e a ecologia trófica da espécie através da análise de conteúdos estomacais e isótopos estáveis de carbono e nitrogênio a partir de amostras coletadas na bacia do médio rio Magdalena, Colômbia; e sua importância ecológica a partir de análises topológicas de redes tróficas. A espécie foi considerada um predador especialista de nível trófico intermediário, com preferência pelos insetos.Não foram observadas diferenças significativas de δ13C e δ15N na espécie entre os períodos hidrológicos de cheia e de águas baixas. O seu nicho isotópico foi intermediário. Em relação à importância ecológica, a espécie desempenhou um papel intermediário na dispersão de efeitos indiretos através do sistema, devido aos seus valores intermediários de centralidade e importância topológica. Estes resultadosconstituem um ponto de partida para novos estudos ecológicos das espécies de raias de água doce da América do Sul, que visem avaliar seu papel nos ecossistemas, e complementam os estudos existentes de Potamotrygon magdalenae.