Orientador: Gustavo Quevedo Romero / Texto em português e inglês / Tese (doutorado) - Universidade Estadual de Campinas, Instituto de Biologia / Made available in DSpace on 2018-08-24T16:18:04Z (GMT). No. of bitstreams: 1 Omena_PaulaMunhoz_D.pdf: 3562543 bytes, checksum: 5d44970e0b040040755da9a84ed62a42 (MD5) Previous issue date: 2014 / Resumo: Com a intensificação das transformações dos sistemas naturais pela atividade humana, o entendimento dos processos que afetam as comunidades e o funcionamento dos ecossistemas tornou-se um tema central para a ecologia contemporânea. As interações entre as espécies, bem como as interações entre as espécies e os componentes abióticos do meio ambiente, afetam tanto os padões de diversidade biológica como funções ecossistêmicas. Neste trabalho, buscou-se compreender o papel relativo das interações predador-presa e dos componentes abióticos sobre as propriedades das comunidades e funcionamento dos ecossistemas. No primeiro capítulo, investiguei o papel do tamanho do habitat sobre os efeitos de predadores terrestres na estrutura de comunidades e funcionamento de ecossistemas adjacentes. O tamanho do habitat modera cascatas tróficas dentro de ecossistemas, deste modo, esperei que efeitos similares do tamanho do habitat poderiam afetar cascatas tróficas que ocorrem através dos limites dos ecossistemas. No segundo capítulo, explorei predições relacionadas a variação no formato de pirâmides tróficas de biomassa ao longo de gradientes ambientais (tamanho do habitat, concentração de detritos e produtividade). Além disso, investiguei a contribuição relativa dos efeitos de consumo direto e do risco de predação nas interações predador-presa como mecanismos estruturadores de pirâmides tróficas de biomassa. No terceiro capítulo, tive como objetivo investigar os efeitos de dois extremos de um gradiente ambiental e os efeitos de predadores sobre os componentes da diversidade de presas detritívoras (i.e., diversidade funcional, filogenética e taxonômica). Além disso, explorei o papel relativo do ambiente, dos predadores e dos componentes da diversidade sobre o funcionamento ecossistêmico (i.e., decomposição e fluxo de nitrogênio). Os resultados encontrados nos três capítulos demonstram que fatores abióticos são cruciais na determinação das propriedades das comunidades, interações predador-presa e, consequentemente, no funcionamento ecossistêmico. O papel relativo dos efeitos cascatas de predadores é altamente dependente das condições ambientais que medeiam as interações entre predadores e presas / Abstract: In the face of the increasing transformation of environmental conditions by human activity, understanding the processes that affect communities and ecosystem functioning has become fundamental goals in ecology. The interactions between coexisting species and, their interactions with the abiotic components of environment, affect the patterns of biological diversity and functions of ecosystems. In this study, I sought to understand the relative role of predator-prey interactions and of abiotic factors on the communities¿ properties and ecosystem functioning. In the first chapter, I investigated the role of habitat size in mediating the effects of terrestrial predators on the structure of communities and the functioning of adjacent ecosystems. Habitat size mediated trophic cascades within ecosystems; therefore, I expected that similar effects of habitat size affect cross-ecosystem trophic cascades. In the second chapter, I explored predictions related to the variation of trophic pyramids of biomass across environmental gradients (i.e., habitat size, detritus concentration and productivity). Furthermore, I investigated the relative contribution of consumptive and non-consumptive effects of predators on the shape of trophic pyramids of biomass. In the third chapter, I sought to investigate the effects of two extremes of an environmental gradient and the effects of predators on the components of detritivores diversity (i.e., functional, phylogenetic and taxonomic). Moreover, I explored the relative role of environment, predators and detritivore diversity components on the functioning of ecosystems (i.e., detritus processing and nitrogen flux). The results from this study demonstrated that abiotic factors are crucial determinants of community properties, predator-prey interactions and ecosystem functioning. The relative role of the predators cascading effects are strongly dependent on the environmental conditions which mediate the interactions between predators and prey / Doutorado / Ecologia / Doutora em Ecologia

Here I provide the first report on Geukensia granosissima patterns of abundance along a tidal gradient within a mixed mangrove stand located in Tampa Bay, Florida, USA. Specifically, I examined 1) the relationship between G. granosissima size and density with mangrove root type (e.g. prop root, pneumatophore), and of density within the intertidal zones; and 2) the possible role of predation in shaping the lower zonation patterns displayed. Transect surveys located along the lower and upper population limit boundaries were conducted every two months over a ten-month period. Variables measured include size distribution, density of mussels, above ground mangrove prop and pneumatophore roots. To evaluate potential predator influence on mussel distribution, predator exclusion experiments were conducted in March and June 2016, using mock pneumatophore platforms at both high and low tidal elevations. Surveys indicated that over all dates mean mussel densities and percent cover were higher along the lower limit tidal elevation [mean (± SD) = 1280.3 ± 665.9 m-2 and 20.6 ± 3.78% respectively], versus that in the higher limit tidal elevation [102.4 ± 50.7 m-2 and 0.52 ± 0.17%]. Survivorship of mussels in the predator exclusion platforms placed at higher position within lower edge of the mussel tidal distribution was approximately 100% on both experimental dates. During March 2016, mussel survivorship was lower when predators had access to mussels with the greatest loss of mussels in the lower (26% survivorship) vs. higher (66.5%) tidal elevations after 48h. Similarly, when predators had access to mussels in June experiments, after just 24 h mussel loss was greater at lower (1% survivorship) vs. higher (80% survivorship) tidal elevations, possibly reflecting differences in predator densities, identities, and/or functional responses. Overall trends in the patterns of distribution of this population of G. granosissima suggest that the lower tidal boundary of the mussel is shaped by predation but that predation plays a lesser role in the high tidal areas.

Seascape ecology provides a useful framework from which to understand the processes governing spatial variability in ecological patterns. Seascape context, or the composition and pattern of habitat surrounding a focal patch, has the potential to impact resource availability, predator-prey interactions, and connectivity with other habitats. For my dissertation research, I combined a variety of approaches to examine how habitat quality for fishes is influenced by a diverse range of seascape factors in sub-tropical, back-reef ecosystems. In the first part of my dissertation, I examined how seascape context can affect reef fish communities on an experimental array of artificial reefs created in various seascape contexts in Abaco, Bahamas. I found that the amount of seagrass at large spatial scales was an important predictor of community assembly on these reefs. Additionally, seascape context had differing effects on various aspects of habitat quality for the most common reef species, White grunt Haemulon plumierii. The amount of seagrass at large spatial scales had positive effects on fish abundance and secondary production, but not on metrics of condition and growth. The second part of my dissertation focused on how foraging conditions for fish varied across a linear seascape gradient in the Loxahatchee River estuary in Florida, USA. Gray snapper, Lutjanus griseus, traded food quality for quantity along this estuarine gradient, maintaining similar growth rates and condition among sites. Additional work focused on identifying major energy flow pathways to two consumers in oyster-reef food webs in the Loxahatchee. Algal and microphytobenthos resource pools supported most of the production to these consumers, and body size for one of the consumers mediated food web linkages with surrounding mangrove habitats. All of these studies examined a different facet of the importance of seascape context in governing ecological processes occurring in focal habitats and underscore the role of connectivity among habitats in back-reef systems. The results suggest that management approaches consider the surrounding seascape when prioritizing areas for conservation or attempting to understand the impacts of seascape change on focal habitat patches. For this reason, spatially-based management approaches are recommended to most effectively manage back-reef systems.

Marine Protected Areas (MPAs) are a management tool used to protect and sustain many ecologically and economically important fish species from overexploitation by recreational and commercial fishing. Lingcod (Ophiodon elongatus) and some of its prey species, such as rockfish (Sebastes spp.), are species that are protected from fishing in some California MPAs. Lingcod is an apex predator that consumes a variety of fish and invertebrate species. In this study, I sought to assess the effect of an MPA on the abundance, size and diet of Lingcod. I hypothesized that Lingcod in a no-take MPA would be more abundant and larger than Lingcod in an adjacent reference site (REF) that was open to fishing. Furthermore, I hypothesized that diet would differ between Lingcod in caught the MPA and Lingcod in the REF. I collected Lingcod from the Point Buchon State Marine Reserve (MPA) and an adjacent REF site that was open to fishing. I measured, weighed, sexed, and collected stomach contents from Lingcod using the gastric lavage (stomach pumping) technique. Then, I identified prey items from Lingcod stomach contents down to the lowest taxonomic level possible and quantified diet composition by percent by occurrence, percent by number, and percent by mass. Lingcod in the MPA consumed more fish prey items than Lingcod in the REF site. Lingcod in the REF consumed more cephalopod prey items than Lingcod in the MPA. I analyzed the four most common prey items (rockfish, anchovies, flatfish, and octopus) for nutritional content. My data suggest that Lingcod increased in size and abundance in a no-take MPA because they do not suffer from fishing mortality. However, a more nutritious diet could also contribute to a biologically significant advantage for Lingcod in the MPA. To address this would require further research focused on calculating the net energy (gross energy extracted from the prey item minus the energetic costs of handling and digesting the prey item) obtained by Lingcod from consuming different fish and cephalopod prey items. MPAs can be an effective management tool for protecting fish stocks, although, it is important to understand the interspecific interactions between predator and prey species to adaptively mange MPAs and the species that reside within them.

Numerous studies have focused on the drivers of diversity and stability of communities, especially under global change. However, multi-dimensionality of ecosystems due to biotic components (e.g predation, competition and adaptive dynamics) and abiotic factors (e.g. disturbances, resource dynamics and their distinct attributes) cause context-dependent outcomes and challenge the predictions. There are still controversies around complex community dynamics under varying regimes, however, finding mechanistical explanations will illuminate the fate of multispecies assemblages. Using model microbial communities, consisting of bacterial prey and protist predator, combined with simulation modelling and advanced statistics, this thesis investigated the impact of imposed disturbances (i.e. increased dilution rates that simulate density-independent mortality as press or pulse disturbances) (i) on transient recovery dynamics of a simple microbial food web, and (ii) on bacterial abundance, diversity and community structure in the absence or presence of a protist predator. In addition, this thesis questioned the impacts of species interactions and rapid trait shifts, as a response to predation and competition, on the community dynamics and stability. Our results revealed that the predator suffered more from disturbances over longer time periods. Reduced predation pressure caused a transient phase of prey release during and even after disturbances. Recovery time depended on the strength and duration of disturbances, however, coupling to an alternative resource increased the chance of fast recovery and stabilized the communities. In multi-species prey communities, bacterial abundance, diversity, and community composition were more affected by predation than by the disturbances and resource dynamics. Predator abundance, on the other hand, was strongly affected by the type of disturbance imposed. Importantly, community attributes had differential sensitivities, as reflected by their different response and recovery dynamics. Prey community dynamics varied more temporally andwere less stable under predation stress, while prey diversity increased significantly. Predation rapidly induced anti-predation traits, which altered population dynamics of both prey and predator. More importantly, predator and the resistant prey, in turn, elevated the number of direct cause-effect relationships between the community members. Our findings are not limited to the studied system and can be used to understand the dynamic response and recovery potential of many natural predator-prey or host-pathogen systems. They can be used as a base for future studies to illuminate the debates on the future communities.:Summary Zusammenfassung 1 Scope and Outline 2 General Introduction 2.1 Context dependency of community dynamics 2.2 Ecological disturbances 2.2.1 Transient dynamics and stability 2.2.2 Catastrophic shifts 2.3 Species interactions and evolutionary dynamics under environmental change 2.3.1 Species interactions and coexistence 2.4 Eco-evolutionary dynamics 2.5 Community assembly mechanisms 2.6 Dealing with complexities 2.6.1 Microbial model systems as a tool in ecology 2.6.2 Correlation, causation and the future of predictions 2.7 Aims of this study 3 Community Dynamics under Disturbances 3.1 Transient recovery dynamics of a predator-prey system 4 Interactions of Community Drivers 4.1 Interactions between predation and disturbances shape prey communities 5 Species Interactions and Evolutionary Dynamics Shaping Communities 5.1 Summary 5.2 Introduction 5.2.1 Predator-Prey Dynamics and Community Stability 5.2.2 Causal inferences 5.3 Aim of the study 5.4 Methods 5.4.1 Organisms 5.4.2 Microcosm experiments and estimation of species abundances 5.4.3 Statistical analysis 5.5 Results 5.5.1 Community dynamics 5.5.2 Dynamics of prey diversity and community stability 5.5.3 Causal links between the species dynamics 5.6 Discussion 5.7 Synopsis 6 General Discussion 6.1 Communities under disturbances: Predator{ prey dynamics 6.2 Temporal species dynamics and community assembly Synthesis and Outlook 7.1 Increasing complexity of species interactions 7.2 Going further from causal links 7.3 Metacommunities References 8 Appendix 8.1 Declaration of the authorship 8.2 Author contributions of published articles 8.3 List of publications and conference contributions 8.4 Acknowledgments 8.5 Supplementary material for Chapter 3 8.6 Supplementary material for Chapter 4 8.7 Supplementary material for Chapter 5

>Magister Scientiae - MSc / In principle, an enrichment of resources in predator-prey systems show prompts destabilisation of a framework, accordingly, falling trophic communication, a phenomenon known to as the \Paradox of Enrichment" [54]. After it was rst genius postured by Rosenzweig [48], various resulting examines, including recently those of Mougi-Nishimura [43] as well as that of Bohannan-Lenski [8], were completed on this problem over numerous decades. Nonetheless, there has been a universal none acceptance of the \paradox" word within an ecological eld due to diverse interpretations [51]. In this dissertation, some theoretical exploratory works are being surveyed in line with giving a concise outline proposed responses to the paradox. Consequently, a quantity of di usion-driven models in mathematical ecology are evaluated and analysed. Accordingly, piloting the way for the spatial structured pattern (we denote it by SSP) formation in nonlinear systems of partial di erential equations [36, 40]. The central point of attention is on enrichment consequences which results toward a paradoxical state. For this purpose, evaluating a number of compartmental models in ecology similar to those of [48] will be of great assistance. Such displays have greater in uence in pattern formations due to diversity in meta-population. Studying the outcomes of initiating an enrichment into [9] of Braverman's model, with a nutrient density (denoted by n) and bacteria compactness (denoted by b) respectively, suits the purpose. The main objective behind being able to transform [9]'s system (2.16) into a new model as a result of enrichment. Accordingly, n has a logistic- type growth with linear di usion, while b poses a Holling Type II and nonlinear di usion r2 nb2 [9, 40]. Five fundamental questions are imposed in order to address and guide the study in accordance with the following sequence: (a) What will be the outcomes of introducing enrichment into [9]'s model? (b) How will such a process in (i) be done in order to change the system (2.16)'s stability state [50]? (c) Whether the paradox does exist in a particular situation or not [51]? Lastly, (d) If an absurdity in (d) does exist, is it reversible [8, 16, 54]? Based on the problem statement above, the investigation will include various matlab simulations. Therefore, being able to give analysis on a local asymptotic stability state when a small perturbation has been introduced [40]. It is for this reason that a bifurcation relevance comes into e ect [58]. There are principal de nitions that are undertaken as the research evolves around them. A study of quantitative response is presented in predator-prey systems in order to establish its stability properties. Due to tradeo s, there is a great likelihood that the growth rate, attack abilities and defense capacities of species have to be examined in line with reviewing parameters which favor stability conditions. Accordingly, an investigation must also re ect chances that leads to extinction or coexistence [7]. Nature is much more complex than scienti c models and laboratories [39]. Therefore, di erent mechanisms have to be integrated in order to establish stability even when a system has been under enrichment [51]. As a result, SSP system is modeled by way of reaction-di usion di erential equations simulated both spatially and temporally. The outcomes of such a system will be best suitable for real-world life situations which control similar behaviors in the future. Comparable models are used in the main compilation phase of dissertation and truly re ected in the literature. The SSP model can be regarded as between (2018-2011), with a stability control study which is of an original.

Orientador: Denise de Cerqueira Rossa-Feres / Resumo: As interações entre espécies influenciam a estrutura de comunidades, a dinâmica das populações, a morfologia, a fisiologia e o comportamento das espécies. A mortalidade é um efeito direto promovido pelo predador, mas também há outros efeitos indiretos decorrentes da presença do predador, como alterações no comportamento da presa. Nos ambientes aquáticos a presa pode detectar o risco de predação por meio de sinais visuais, mecânicos e químicos. Ao detectar o predador, a presa pode usar estratégias, como formação de cardume, mudança no padrão de uso de habitat e diminuição da atividade natatória para reduzir a predação. Apesar de diminuir a taxa de predação, as estratégias antipredatórias tem custos para as presas e configuram um trade-off. Os comportamentos que aumentam a chance de escapar do predador prejudicam a atividade e o forrageamento dos girinos, diminuindo seu crescimento e suas chances de sobrevivência. Os aspectos morfológicos e fisiológicos são importantes na ocupação da coluna d’água por girinos, mas, será que a predação também é uma pressão que determina o modo como os girinos ocupam a coluna d’água? Entender como a predação influencia o comportamento dos girinos pode elucidar aspectos da dinâmica populacional e estruturação de comunidades, bem como dos mecanismos que regulam o padrão evolutivo dos anuros. Nesse contexto, esta dissertação, apresentada na forma de manuscrito, apresenta um estudo no qual foi testado experimentalmente como um predador aquático (náia... (Resumo completo, clicar acesso eletrônico abaixo) / Abstract: Interactions between species influence community structure, population dynamics, morphology, physiology and species’ behavior. Mortality is a direct effect caused by the interaction with a predator, but even the simple presence of predators can lead to other indirect effects such as alterations on prey’s behavior. In aquatic environments, prey may detect predation risk by visual, mechanical, and chemical cues. When detecting a predator, the prey can use strategies to escape predation such as shoal formation, changes on habitat use pattern, and reduction on swimming activity. Despite decreasing predation rates, the antipredatory strategies have costs for the prey and set up a trade-off. For example, the same behavior that raises the prey’s chance to escape from a predator, decreases its foraging activity, leading thus to a reduction in the prey’s growth and survivorship. Among tadpoles, morphological and physiological aspects are important to determine water column occupation. But could predation also be an important pressure that determines the way tadpoles occupy the water column? Understanding how predation influences tadpoles’ behavior may elucidate populational dynamics and community structuring aspects, as well as the mechanisms that regulate anuran evolutionary patterns. In this sense, this dissertation presents a study, in a manuscript form, that tested experimentally the influence of an aquatic predator (Odonata water nymph; Micrathyria sp.) on the use of space, activ... (Complete abstract click electronic access below) / Mestre

Daily rhythms occur in numerous physiological and behavioral processes across an immense diversity of taxa, but there remain few cases in which mechanistic links between rhythms of trait expression and organismal fitness have been established. We construct a dynamic optimization model to determine whether risk allocation provides an adaptive explanation for the daily foraging rhythm observed in many species using the orb-weaving spider Cyclosa turbinata as a case study. Our model predicts that female C. turbinata should generally start foraging at lower levels of energy reserves (i.e., should be less bold) during midday when predators are most abundant. We also find that individuals’ foraging efficacy determines whether daily rates of encounters with predators or prey more strongly influences boldness under high risk. The qualitative model predictions are robust to variation in our parameter estimates and likely apply to a wide range of taxa. The predictions are also consistent with observed patterns of foraging behavior under both laboratory and field conditions. We discuss the implications of our study for understanding the evolution of daily rhythms and the importance of model predictions for interpreting empirical studies and generating additional hypotheses regarding behavioral evolution.

This thesis focuses on the impact of spatially heterogeneous environments on the spatio-temporal behavior of planktonic systems. Specific emphasis placed is on the influence of spatial variations in the strength of random or chaotic movements (diffusion) of the organisms. Interaction between different species is described by ordinary differential equations. In order to describe movements in space, reaction–diffusion or advection–reaction–diffusion systems are studied. Examples are given for different approaches of diffusive motion as well as for the possible effects on the localized biological system. The results are discussed based on their biological and physical meanings. In doing so, different mechanisms are shown which are able to explain events of fast plankton growth near turbulent flows. In general, it is shown that local variation in the strength of vertical mixing can have global effects on the biological system, such as changing the stability of dynamical solutions and generating new spatiotemporal behavior. The thesis consists of five chapters. Three of them have been published in international peer-reviewed scientific journals. Chapter 1. Introduction: This chapter gives a general introduction to the history of plankton modeling and introduces basic ideas and concepts which are used in the following chapters. Chapter 2. Fokker-Planck law of diffusion: The influence of spatially in- homogeneous diffusion on several common ecological problems is analyzed. Dif- fusion is modeled with Fick’s law and the Fokker–Planck law of diffusion. A discussion is given about the differences between the two formalisms and when to use the one or the other. To do this, the discussion starts with a pure diffusion equation, then it turns to a reaction–diffusion system with one logistically growing component which invades the spatial domain. This chapter also provides a look at systems of two reacting components, namely a trimolecular oscillating chemical model system and an excitable predator–prey model. Contrary to Fickian diffusion, spatial inhomogeneities promote spatial and spatiotemporal pattern formation in the case of Fokker–Planck diffusion. A slightly modified version of this chapter has been published in the Journal of Mathematical Biology (Bengfort et al., 2016). Chapter 3. Plankton blooms and patchiness: Microscopic turbulent motions of water have been shown to influence the dynamics of microscopic species. Therefore, the number, stability, and excitability of stationary states in a predator– prey model of plankton species can change when the strength of turbulent motions varies. In a spatial system these microscopic turbulent motions are naturally of different strength and form a heterogeneous physical environment. Spatially neighboring plankton communities with different physical conditions can impact each other due to diffusive coupling. It is shown that local variations in the physical conditions can influence the global system in the form of propagating pulses of high population densities. For this, three local predator–prey models with different local responses to variation in the physical environment are considered. The degree of spatial heterogeneity can, depending on the model, promote or reduce the number of propagating pulses, which can be interpreted as patchy plankton distributions and recurrent blooms. This chapter has been published in the Journal Ecological Complexity (Bengfort et al., 2014). Chapter 4. Advection–reaction–diffusion model: Here, some of the models introduced in chapter 1 and 2 are modified to perform two dimensional spatial simulations including advection, reaction and diffusion. These models include assumptions about turbulent flows introduced in chapter 1. Chapter 5. Competition: Some plankton species, such as cyanobacteria, have an advantage in competition for light compared to other species because of their buoyancy. This advantage can be diminished by vertical mixing in the surround- ing water column. A non–spatial model, based on ordinary differential equations, which accounts for this effect is introduced. The main aim is to show that vertical mixing influences the outcome of competition between different species. Hystersis is possible for a certain range of parameters. Introducing a grazing predator, the system exhibits different dynamics depending on the strength of mixing. In a diffusively coupled horizontal spatial model, local vertical mixing can also have a global effect on the biological system, for instance, destabilization of a locally stable solution, or the generation of new spatiotemporal behavior. This chapter has been published in the Journal Ecological Modelling (Bengfort and Malchow, 2016).

Seasonal shifts in predation habits, from a generalist in the summer to a specialist in the winter, have been documented for the great horned owl (Bulbo virginialis) in the boreal forest. This shift occurs largely due to varying prey availability. There is little study of this switching behaviour in the current literature. Since season length is predicted to change under future climate scenarios, it is important to understand resulting effects on species dynamics. Previous work has been done on a two-species seasonal model for the great horned owl and its focal prey, the snowshoe hare (Lepus americanus). In this thesis, we extend the model by adding one of the hare's most important predators, the Canadian lynx (Lynx canadensis). We study the qualitative behaviour of this model as season length changes using tools and techniques from dynamical systems. Our main approach is to determine when the lynx and the owl may invade the system at low density and ask whether mutual invasion of the predators implies stable coexistence in the three-species model. We observe that, as summer length increases, mutual invasion is less likely, and we expect to see extinction of the lynx. However, in all cases where mutual invasion was satisfied, the three species stably coexist.