| 1. |
- Uszko, Wojciech, 1985-
(författare)
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Effects of warming and nutrient enrichment on feeding behavior, population stability and persistence of consumers and their resources
- 2016
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Consumer-resource interactions are the basic building blocks of every food web. In spite of being a central research theme of longstanding interest in ecology, the mechanisms governing the stability and persistence of consumer-resource interactions are still not entirely understood. In particular, theoretical predictions on consumer-resource stability along gradients of temperature and nutrient enrichment diverge widely and are sometimes in conflict with empirical results. In this thesis I address these issues from the angle of the functional response, which describes a consumer’s feeding rate as a function of resource density. Specifically, I explore mechanistic, nutrient-based consumer-resource interaction models with respect to the influence of feeding behavior (the shape of the functional response), environmental temperature, nutrient enrichment, and resource quality on consumer-resource stability and persistence. In order to parameterize these models I performed extensive laboratory experiments with pairs of freshwater pelagic algae and grazers of the genus Daphnia, which are widespread, ecologically important model organisms.I found a sigmoidal type III functional response in every studied Daphnia-algae species pair. The exact form of its shape is described by an exponent b which is determined by fitting functional response models to the experimental data. A high value of b can stabilize consumer-resource systems under the otherwise destabilizing influence of nutrient enrichment, as predicted by a novel stability criterion relating b to the consumer’s prey handling time, food conversion efficiency and mortality. Estimated parameter values and, consequently, stability predictions are sensitive to the method of parameter estimation, and I propose a new estimation procedure that minimizes parameter uncertainty. Because many consumers’ feeding rates depend on temperature, warming is expected to strongly affect food web stability. In functional response experiments over a broad temperature gradient, I found that the attack rate coefficient and the maximum ingestion rate of Daphnia are hump-shaped functions of temperature. Moreover, the functional response exponent increases with warming towards stronger type III responses. Plugging these findings into a nutrient-based consumer-resource model, I found that predator persistence is a U-shaped function of temperature in nutrient enrichment-temperature space. Enrichment easily turns the system unstable when the consumer has a type II response, whereas a type III response opens up a large region of stability at intermediate, for the consumer optimal, temperatures. These findings reconcile seemingly conflicting results of earlier studies of temperature effects on consumer-resource dynamics, which can be mapped as special cases onto the enrichment-temperature space. I finally demonstrate the utility of three key model ingredients - temperature dependence of rate parameters, a mechanistic description of the dynamics of algal resources, and a type III functional response in Daphnia - by successfully implementing them in the description and explanation of phytoplankton-Daphnia dynamics in a mesocosm experiment exploring effects of warming on the spring succession of the plankton.
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| 2. |
- Wickman, Jonas, 1985-
(författare)
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Evolution of Ecological Communities in Spatially Heterogeneous Environments
- 2019
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Evolutionarily stable communities are the endpoints of evolution, and ecological communities whose traits are under selection will eventually settle into them. Hence, the properties of such communities are of particular interest, as they can persist over long evolutionary time scales. The notion of an evolutionarily stable strategy - an evolved strategy that cannot be beat by any other once established - has now been part of theoretical ecology for almost 50 years, and the theory for evolutionarily stable strategies and communities, and how they are reached has become increasingly versatile. However, for environments where conditions vary in space, so-called heterogeneous environments, efficient analytical and numerical tools for studying evolutionarily stable communities and how they come about have been lacking. Hence, many questions regarding how evolutionarily stable diversity is generated and maintained when ecological and evolutionary forces vary in space remain unexplored. In particular, how spatially averaged selection and selective forces derived from spatial variability can act together to either promote or inhibit evolutionarily stable diversity is not well understood. In this thesis, I use a two-pronged approach towards answering such questions by developing the necessary analytical and numerical tools for assembling and analyzing evolutionarily stable communities in heterogeneous environments, and by then employing these tools to study communities of resource competitors and food webs. Specifically, I derive expressions for directional and stabilizing/disruptive selection when the spatially heterogeneous ecological dynamics of a community are described by reaction-diffusion equations. These expressions allow us to understand selection across an environment in terms of local selection pressures, and also enable efficient numerical implementations of evolutionary community assembly procedures that lead to evolutionarily stable communities. Applied to the communities of resource competitors and food webs I find that the selective forces derived from spatially averaged selection and those derived from spatial variability can act both in concert or in opposition. If these forces act in opposition and if the spatial variability of local selection is high, a high diversity of organisms can form even when spatially averaged selection is stabilizing. In contrast, if spatially averaged selection is disruptive, it can prevent more diverse communities from forming by creating few globally unbeatable strategies. However, these forces can also act disruptively in concert to create more diverse communities. Together, these results demonstrate a surprising variety of qualitatively different outcomes when evolutionarily stable communities are assembled in heterogeneous environments.
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| 3. |
- Maier, Dominique Béatrice, 1982-
(författare)
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Combining limnology and paleolimnology : a refined understanding of environmental sediment signal formation in a varved lake
- 2017
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Paleoclimatic archives, such as lake sediments, extend our understanding of terrestrial and aquatic ecosystem dynamics in relation to climate variability beyond the period covered by instrumental data. In this context, annually laminated (i.e. varved) lake sediments are particularly valuable, as they offer high temporal resolution and undisturbed sediment. However, in order to extract reliable climate information from lake sediments, a careful calibration with the processes controlling the sediment formation is essential. This thesis combines limnological and paleolimnological data from a varved, boreal lake in northern Sweden (Nylandssjön, Nordingrå) collected over different time scales. The main aim of the thesis is to gain a more refined insight into which processes are reflected in the sedimentary diatom assemblage. More specifically, sequential sediment trap records were coupled with physical, chemical and biological lake monitoring and environmental data for comparison and validation with the varved sediment record. The main result of the thesis is that timing, succession and inter-annual variability of key limnological and environmental processes (e.g. ice-cover duration, lake over-turn or catchment run-off) are of major importance for the sedimentary diatom assemblage formation. Continuous monitoring of physico-chemical parameters over three consecutive years identified varying winter air temperature as a major factor influencing in-lake processes and hence the diatom record. Timing of lake over-turn and catchment run-off seemed to be the driver for monospecific diatom blooms, which are reflected in the annual sediment signal. The integrated annual diatom signal in the sediment was dominated by spring or autumn blooms, resulting either from a Cyclotella glomerata dominated spring bloom after relatively warm winter conditions, or a Asterionella formosa dominated autumn bloom after relatively cold winter conditions. The analysis of the diatom stratigraphy in the varved sediment over several decades corroborated the importance of climatic variables (late winter air temperature and NAO), even though the variables with the most predictive power for variance in the diatom data were associated with sediment composition (C, N and sedimentation rate) and pollution (Pb and Cu). Overall, the analysis of the drivers of inter-annual and decadal diatom assemblage fluctuations emphasizes the importance of winter air temperature, indicating that weather extremes may be disproportionately represented in annual sediment records in contrast to nutrient concentrations or sedimentation rate.
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| 4. |
- Thomsson, Gustaf, 1981-
(författare)
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Biomass patterns in boreal-subarctic lake food webs along gradients of light and nutrients
- 2015
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Licentiatavhandling (övrigt vetenskapligt/konstnärligt)abstract
- There is large natural variation in light and nutrient conditions across lakes. In the boreal-subarctic region most lakes are small, shallow and nutrient poor. In such lakes there is often sufficient light to support primary production at the lake bottom. An expectation for the future is that colored dissolved organic matter (cDOM) of terrestrial origin will increase in these lakes. cDOM depresses the underwater light climate but is often associated with elevated pelagic nutrient concentrations.A dynamical model of a coupled benthic-pelagic food web was explored for how lake ecosystems might respond to altered light and nutrient regimes. The model predicts that mobile carnivores (fish) control grazers and release primary producers from grazing pressure. Primary producers are therefore limited by their resources and cross-habitat interactions are dominated by spatially asymmetric competition for light and nutrients. At high light and low nutrient supply benthic algae out-compete pelagic algae for nutrients diffusing from the sediment, whereas pelagic algae shade out benthic algae at lower light and/or higher nutrient supply. Biomass patterns of benthic and pelagic consumers follow the patterns of primary production. In contrast, habitat coupling through carnivore movement has only a weak impact on biomass patterns in the model food web.Model predictions were compared with data from boreal-subarctic lakes covering a broad range of cDOM concentrations. In agreement with model expectations the following relationships with increasing light attenuation were observed: benthic primary and secondary production decreased, pelagic primary production showed a unimodal trend, and pelagic nutrient concentrations as well as the proportion of fish feeding in the pelagic habitat increased. As a consequence, both primary and fish production were negatively related to pelagic nutrient concentrations across lakes.In a comparative study of boreal-subarctic lakes covering a broad range of cDOM concentrations, a similar negative relationship was found between pelagic total nutrient concentrations and the biomass of epilithic algae. This was surprising, because epilithon cannot access nutrients from the sediment. Patterns in epilithon biomass were largely driven by nitrogen fixing cyanobacteria, which in turn were positively related to light supply. The data suggest that nitrogen fixing autotrophs may have a competitive advantage over other epilithic primary producers in low-cDOM, low-nutrient, high-light environments, and that patterns in epilithic biomass, nutrient sequestration and elemental stoichiometry depend upon which functional group is dominant in the epilithic biofilm.
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| 5. |
- Vasconcelos, Francisco Rivera, 1984-
(författare)
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Effects of warming and browning on benthic and pelagic ecosystem components in shallow lakes
- 2017
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- The majority of lakes on Earth are shallow, unproductive and located at high latitudes. These lakes are experiencing big changes due to climate change, where two environmental drivers operate simultaneously, browning and warming. How they affect lake ecosystems is not well understood. Here, I addressed this issue by using a theoretical and an experimental approach. In particular, I generated model predictions and compared them with the results of a realistic large-scale experiment, where browning and warming were manipulated in a factorial design. In addition, model outcomes were compared with data from 12 unproductive lakes sampled along a gradient of browning. Another novelty of my thesis is that it integrates benthic and pelagic food web components in the model and experimental approaches. I found that browning affected the resources availability for benthic and pelagic producers in the model and in the experiment. With browning, benthic primary producers became increasingly light limited and declined, while pelagic producers became less nutrient limited and increased. Pelagic nutrient limitation was alleviated by two non-exclusive mechanisms. Browning directly enriched the water with nutrients, and browning indirectly increased the nutrient flowing from the sediment to the pelagic habitat via suppression of benthic producers. To tease apart these two mechanisms I applied structural equation modeling (SEM). The indirect evidence by SEM suggests that both mechanisms contributed equally to the pelagic nutrient concentration in the experiment. Interestingly, a model food web with only primary producers shows similar qualitative behavior as a food web with grazers and carnivores included. This happens because carnivorous fish exert strong top-down control in the more productive habitat, which relaxes grazing pressure on primary producers and increases resource limitation in the adjacent habitat. Biomass of benthic and pelagic consumers followed the same pattern as their resources. The lake data were largely congruent with model expectations and supported the findings of the experiment. Furthermore, the model also predicted a negative relationship between total phosphorus and both primary and fish production, which was observed across the 12 lakes. Warming effects were more complex. The model predicts that warming effects should depend on browning and are expected to be strongest in the more productive of the two (benthic and pelagic) habitats. For example, at low levels of browning the biomasses of benthic algae and fish are expected to decline with warming, which was observed in the experiment. In contrast, observed warming effects at high levels of browning deviated from model expectations. The mechanisms by which browning and warming interactively affect lake food webs are still poorly understood. This thesis offers a conceptual foundation for their further study through the integration of within- and between-habitat interactions.
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| 6. |
- Bergström, Ulf, 1971-
(författare)
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Spatial heterogeneity and biotic interactions : scaling from experiments to natural systems
- 2004
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Much of current ecological theory stems from experimental studies. These studies have often been conducted in closed systems, at spatial scales that are much smaller than the systems of interest. It is known that the outcome of these experiments may be seriously affected by artefacts associated with the caging procedures, as well as by the actual difference in spatial scale between experimental and target system. Yet, quantitative methods for estimating and removing artefacts of enclosure and for extrapolating experimental results to the scales of natural systems are largely lacking.The aim of this thesis was to confront some of the problems encountered when scaling from experiments to nature in studies on predator-prey systems, with focus on effects of changes in spatial heterogeneity. Specifically, I examined mechanisms that may cause consumption rate estimates to depend on the size of the experimental arena. I also studied methods for scaling up these process rate estimates to natural predator-prey systems. The studies were performed on invertebrate predator-prey systems found in the northern Baltic Sea. Initially, a descriptive study of small-scale distribution patterns was performed, in order to get background information on how the behaviour of the organisms was manifested in the spatial structure of the community. Experimental studies of two predator-prey systems exposed an artefact that may be widespread in experiments aiming at quantifying biotic interactions. It is caused by predator and prey aggregating along the walls of the experimental containers. This behaviour affects the encounter rate between predator and prey, thereby causing consumption rates to be scale-dependent. Opposing the common belief that larger arenas always produce less biased results, this scale effect may instead be reduced by decreasing arena size. An alternative method for estimating the magnitude of, and subsequently removing, the artefact caused by aggregation along the arena wall was presented.Once unbiased estimates of process functions have been derived, the next step is to scale up the functions to natural systems. This extrapolation entails a considerable increase in spatial heterogeneity, which may have important implications for the dynamics of the system. Moment approximation provides a method of taking the heterogeneity of natural populations into account in the extrapolation process. In the last study of the thesis, the concepts of moment approximation and how to estimate relevant heterogeneity were explained, and it was shown how the method may be used for adding space as a component to a dynamic predator-prey model. It was concluded that moment approximation provides a simple and useful technique for dealing with effects of spatial variation, and that a major benefit of the method is that it provides a way of visualising how heterogeneity affects ecological processes.
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| 7. |
- Gudmundson, Sara, 1985-
(författare)
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Species Responses to Environmental Fluctuations : impacts of food web interactions and noise color
- 2017
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Species constantly experience changes in their environmental conditions owing to natural or human induces reasons. Understanding how species respond to these fluctuations are important for ecology, especially given the ongoing climate change. Empirical studies have shown that species respond differently to the same disturbance. However, our knowledge of what create these differences in the environmental response is limited and in most cases based on studies focusing on single species. In this thesis, I have taken a theoretical approach and used dynamical models to investigate how the population dynamics of species are affected by species-species interactions and environmental fluctuations. In the first paper (Paper I) I investigated how a species respond to environmental fluctuations when isolated or embedded in a food web. The study showed that species-species interactions had an effect in temporally positively autocorrelated environments (red noise) but not in uncorrelated environments (white noise). This was owing to species following their equilibrium densities in red environments which in turn enabled species-species interactions to come into play. Red environmental variables are more prominent in nature than white. Thus, these results show the importance of using a food web approach when analyzing species response to environmental fluctuations. The most commonly discussed effect of climate change is an elevated mean temperature. This shift is expected to affect the growth rate of many species. However, there is no robust theory of how we should expect species in food webs to respond to a rise in temperature. In the second paper (Paper II) I defined and studied the dynamic rate of food webs(DR) acting analogously to single species growth rate. I found that the higher DR the easier for species population densities to follow their equilibrium over time. Both DR and noise color changed the temporal relationship between the population and the environmental noise. Thus, it is of major importance to take the scale of time into consideration when investigating species response to environmental fluctuations. Another important factor which affect population dynamics is species spatial distributions. Dispersal between subpopulations enable individuals to rescue or prolong the time to extinction for the population seen as a whole. In the third paper (Paper III), I investigated how species in food webs respond to environments that varies both in time and space and compared the results with the one from single species. I found that single species were stabilized by an increased dispersal rate independent of the noise color. Species-species interactions had an effect for some of the species in these landscapes.At red asynchronous noise, one resource species in each food web had a local minimum in stability at low dispersal rate. Here, dispersal decoupled local population dynamics and prevented species from tracking their equilibriums. At high dispersal rates, all resource species and their single species counterparts were stabilized by dispersal as local patch dynamics lost its importance. Environmental noise together with the spatial dimension does seem to explain much of the stability properties of species on our planet. However, natural ecosystems are much more complex and species rich than the food web models I have used so far. Theoreticians have previously had a hard time describing stable complex systems that survive environmental fluctuations. Thus, in my fourth and last project (PaperIV) I investigated how species population dynamics are affected by environmental fluctuations when embedded in larger food webs. These systems were built by connecting food web modules with periodic boundary conditions (PBC). The PBC method has previously helped physicists to understand the nature of waves and particles by removing the edges in systems. I found that food web size does not have to have a negative effect on food web stability. I showed that by removing the destabilizing effect of edges it is possible to describe large stable food webs, more similar to natural ecosystems. Overall, the research presented here give new insights into species responses to environmental fluctuations. They especially highlight the importance of considering both species interactions and environmental noise color when studying population dynamics in a fluctuating environment. A food web approach is necessary when analyzing species population dynamics and planning for conservation actions, especially when studying the effects of climate change on biodiversity.
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