@article{KathBoitGuilletal.2018, author = {Kath, Nadja J. and Boit, Alice and Guill, Christian and Gaedke, Ursula}, title = {Accounting for activity respiration results in realistic trophic transfer efficiencies in allometric trophic network (ATN) models}, series = {Theoretical ecology}, volume = {11}, journal = {Theoretical ecology}, number = {4}, publisher = {Springer}, address = {Heidelberg}, issn = {1874-1738}, doi = {10.1007/s12080-018-0378-z}, pages = {453 -- 463}, year = {2018}, abstract = {Allometric trophic network (ATN) models offer high flexibility and scalability while minimizing the number of parameters and have been successfully applied to investigate complex food web dynamics and their influence on food web diversity and stability. However, the realism of ATN model energetics has never been assessed in detail, despite their critical influence on dynamic biomass and production patterns. Here, we compare the energetics of the currently established original ATN model, considering only biomass-dependent basal respiration, to an extended ATN model version, considering both basal and assimilation-dependent activity respiration. The latter is crucial in particular for unicellular and invertebrate organisms which dominate the metabolism of pelagic and soil food webs. Based on metabolic scaling laws, we show that the extended ATN version reflects the energy transfer through a chain of four trophic levels of unicellular and invertebrate organisms more realistically than the original ATN version. Depending on the strength of top-down control, the original ATN model yields trophic transfer efficiencies up to 71\% at either the third or the fourth trophic level, which considerably exceeds any realistic values. In contrast, the extended ATN version yields realistic trophic transfer efficiencies 30\% at all trophic levels, in accordance with both physiological considerations and empirical evidence from pelagic systems. Our results imply that accounting for activity respiration is essential for consistently implementing the metabolic theory of ecology in ATN models and for improving their quantitative predictions, which makes them more powerful tools for investigating the dynamics of complex natural communities.}, language = {en} } @article{ColomaGaedkeSivonenetal.2019, author = {Coloma, Sebastian and Gaedke, Ursula and Sivonen, Kaarina and Hiltunen, Teppo}, title = {Frequency of virus-resistant hosts determines experimental community dynamics}, series = {Ecology : a publication of the Ecological Society of America}, volume = {100}, journal = {Ecology : a publication of the Ecological Society of America}, number = {1}, publisher = {Wiley}, address = {Hoboken}, issn = {0012-9658}, doi = {10.1002/ecy.2554}, pages = {10}, year = {2019}, abstract = {Parasites, such as bacterial viruses (phages), can have large effects on host populations both at the ecological and evolutionary levels. In the case of cyanobacteria, phages can reduce primary production and infected hosts release intracellular nutrients influencing planktonic food web structure, community dynamics, and biogeochemical cycles. Cyanophages may be of great importance in aquatic food webs during large cyanobacterial blooms unless the host population becomes resistant to phage infection. The consequences on plankton community dynamics of the evolution of phage resistance in bloom forming cyanobacterial populations are still poorly studied. Here, we examined the effect of different frequencies of a phage-resistant genotype within a filamentous nitrogen-fixing Nodularia spumigena population on an experimental plankton community. Three Nodularia populations with different initial frequencies (0\%, 5\%, and 50\%) of phage-resistant genotypes were inoculated in separate treatments with the phage 2AV2, the green alga Chlorella vulgaris, and the rotifer Brachionus plicatilis, which formed the experimental plankton community subjected to either nitrogen-limited or nitrogen-rich conditions. We found that the frequency of the phage-resistant Nodularia genotype determined experimental community dynamics. Cyanobacterial populations with a high frequency (50\%) of the phage-resistant genotype dominated the cultures despite the presence of phages, retaining most of the intracellular nitrogen in the plankton community. In contrast, populations with low frequencies (0\% and 5\%) of the phage-resistant genotype were lysed and reduced to extinction by the phage, transferring the intracellular nitrogen held by Nodularia to Chlorella and rotifers, and allowing Chlorella to dominate the communities and rotifers to survive. This study shows that even though phages represent minuscule biomass, they can have key effects on community composition and eco-evolutionary feedbacks in plankton communities.}, language = {en} } @article{RosenbaumRaatzWeithoffetal.2019, author = {Rosenbaum, Benjamin and Raatz, Michael and Weithoff, Guntram and Fussmann, Gregor F. and Gaedke, Ursula}, title = {Estimating parameters from multiple time series of population dynamics using bayesian inference}, series = {Frontiers in ecology and evolution}, volume = {6}, journal = {Frontiers in ecology and evolution}, publisher = {Frontiers Research Foundation}, address = {Lausanne}, issn = {2296-701X}, doi = {10.3389/fevo.2018.00234}, pages = {14}, year = {2019}, abstract = {Empirical time series of interacting entities, e.g., species abundances, are highly useful to study ecological mechanisms. Mathematical models are valuable tools to further elucidate those mechanisms and underlying processes. However, obtaining an agreement between model predictions and experimental observations remains a demanding task. As models always abstract from reality one parameter often summarizes several properties. Parameter measurements are performed in additional experiments independent of the ones delivering the time series. Transferring these parameter values to different settings may result in incorrect parametrizations. On top of that, the properties of organisms and thus the respective parameter values may vary considerably. These issues limit the use of a priori model parametrizations. In this study, we present a method suited for a direct estimation of model parameters and their variability from experimental time series data. We combine numerical simulations of a continuous-time dynamical population model with Bayesian inference, using a hierarchical framework that allows for variability of individual parameters. The method is applied to a comprehensive set of time series from a laboratory predator-prey system that features both steady states and cyclic population dynamics. Our model predictions are able to reproduce both steady states and cyclic dynamics of the data. Additionally to the direct estimates of the parameter values, the Bayesian approach also provides their uncertainties. We found that fitting cyclic population dynamics, which contain more information on the process rates than steady states, yields more precise parameter estimates. We detected significant variability among parameters of different time series and identified the variation in the maximum growth rate of the prey as a source for the transition from steady states to cyclic dynamics. By lending more flexibility to the model, our approach facilitates parametrizations and shows more easily which patterns in time series can be explained also by simple models. Applying Bayesian inference and dynamical population models in conjunction may help to quantify the profound variability in organismal properties in nature.}, language = {en} } @article{PerkinsPernaAdrianetal.2019, author = {Perkins, Daniel M. and Perna, Andrea and Adrian, Rita and Cermeno, Pedro and Gaedke, Ursula and Huete-Ortega, Maria and White, Ethan P. and Yvon-Durocher, Gabriel}, title = {Energetic equivalence underpins the size structure of tree and phytoplankton communities}, series = {Nature Communications}, volume = {10}, journal = {Nature Communications}, publisher = {Nature Publ. Group}, address = {London}, issn = {2041-1723}, doi = {10.1038/s41467-018-08039-3}, pages = {8}, year = {2019}, abstract = {The size structure of autotroph communities - the relative abundance of small vs. large individuals - shapes the functioning of ecosystems. Whether common mechanisms underpin the size structure of unicellular and multicellular autotrophs is, however, unknown. Using a global data compilation, we show that individual body masses in tree and phytoplankton communities follow power-law distributions and that the average exponents of these individual size distributions (ISD) differ. Phytoplankton communities are characterized by an average ISD exponent consistent with three-quarter-power scaling of metabolism with body mass and equivalence in energy use among mass classes. Tree communities deviate from this pattern in a manner consistent with equivalence in energy use among diameter size classes. Our findings suggest that whilst universal metabolic constraints ultimately underlie the emergent size structure of autotroph communities, divergent aspects of body size (volumetric vs. linear dimensions) shape the ecological outcome of metabolic scaling in forest vs. pelagic ecosystems.}, language = {en} } @article{RossbergGaedkeKratina2019, author = {Rossberg, Axel G. and Gaedke, Ursula and Kratina, Pavel}, title = {Dome patterns in pelagic size spectra reveal strong trophic cascades}, series = {Nature Communications}, volume = {10}, journal = {Nature Communications}, publisher = {Nature Publ. Group}, address = {London}, issn = {2041-1723}, doi = {10.1038/s41467-019-12289-0}, pages = {11}, year = {2019}, abstract = {In ecological communities, especially the pelagic zones of aquatic ecosystems, certain bodysize ranges are often over-represented compared to others. Community size spectra, the distributions of community biomass over the logarithmic body-mass axis, tend to exhibit regularly spaced local maxima, called "domes", separated by steep troughs. Contrasting established theory, we explain these dome patterns as manifestations of top-down trophic cascades along aquatic food chains. Compiling high quality size-spectrum data and comparing these with a size-spectrum model introduced in this study, we test this theory and develop a detailed picture of the mechanisms by which bottom-up and top-down effects interact to generate dome patterns. Results imply that strong top-down trophic cascades are common in freshwater communities, much more than hitherto demonstrated, and may arise in nutrient rich marine systems as well. Transferring insights from the general theory of nonlinear pattern formation to domes patterns, we provide new interpretations of past lake-manipulation experiments.}, language = {en} } @article{WojcikCeulemansGaedke2021, author = {Wojcik, Laurie Anne and Ceulemans, Ruben and Gaedke, Ursula}, title = {Functional diversity buffers the effects of a pulse perturbation on the dynamics of tritrophic food webs}, series = {Ecology and Evolution}, volume = {11}, journal = {Ecology and Evolution}, number = {22}, publisher = {John Wiley \& Sons, Inc.}, address = {Hoboken (New Jersey)}, issn = {2045-7758}, doi = {10.1002/ece3.8214}, pages = {15639 -- 15663}, year = {2021}, abstract = {Biodiversity decline causes a loss of functional diversity, which threatens ecosystems through a dangerous feedback loop: This loss may hamper ecosystems' ability to buffer environmental changes, leading to further biodiversity losses. In this context, the increasing frequency of human-induced excessive loading of nutrients causes major problems in aquatic systems. Previous studies investigating how functional diversity influences the response of food webs to disturbances have mainly considered systems with at most two functionally diverse trophic levels. We investigated the effects of functional diversity on the robustness, that is, resistance, resilience, and elasticity, using a tritrophic—and thus more realistic—plankton food web model. We compared a non-adaptive food chain with no diversity within the individual trophic levels to a more diverse food web with three adaptive trophic levels. The species fitness differences were balanced through trade-offs between defense/growth rate for prey and selectivity/half-saturation constant for predators. We showed that the resistance, resilience, and elasticity of tritrophic food webs decreased with larger perturbation sizes and depended on the state of the system when the perturbation occurred. Importantly, we found that a more diverse food web was generally more resistant and resilient but its elasticity was context-dependent. Particularly, functional diversity reduced the probability of a regime shift toward a non-desirable alternative state. The basal-intermediate interaction consistently determined the robustness against a nutrient pulse despite the complex influence of the shape and type of the dynamical attractors. This relationship was strongly influenced by the diversity present and the third trophic level. Overall, using a food web model of realistic complexity, this study confirms the destructive potential of the positive feedback loop between biodiversity loss and robustness, by uncovering mechanisms leading to a decrease in resistance, resilience, and potentially elasticity as functional diversity declines.}, language = {en} } @misc{CeulemansGaedkeKlauschiesetal.2019, author = {Ceulemans, Ruben and Gaedke, Ursula and Klauschies, Toni and Guill, Christian}, title = {The effects of functional diversity on biomass production, variability, and resilience of ecosystem functions in a tritrophic system}, series = {Postprints der Universit{\"a}t Potsdam Mathematisch-Naturwissenschaftliche Reihe}, journal = {Postprints der Universit{\"a}t Potsdam Mathematisch-Naturwissenschaftliche Reihe}, number = {744}, issn = {1866-8372}, doi = {10.25932/publishup-43543}, url = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus4-435439}, pages = {16}, year = {2019}, abstract = {Diverse communities can adjust their trait composition to altered environmental conditions, which may strongly influence their dynamics. Previous studies of trait-based models mainly considered only one or two trophic levels, whereas most natural system are at least tritrophic. Therefore, we investigated how the addition of trait variation to each trophic level influences population and community dynamics in a tritrophic model. Examining the phase relationships between species of adjacent trophic levels informs about the strength of top-down or bottom-up control in non-steadystate situations. Phase relationships within a trophic level highlight compensatory dynamical patterns between functionally different species, which are responsible for dampening the community temporal variability. Furthermore, even without trait variation, our tritrophic model always exhibits regions with two alternative states with either weak or strong nutrient exploitation, and correspondingly low or high biomass production at the top level. However, adding trait variation increased the basin of attraction of the high-production state, and decreased the likelihood of a critical transition from the high- to the lowproduction state with no apparent early warning signals. Hence, our study shows that trait variation enhances resource use efficiency, production, stability, and resilience of entire food webs.}, language = {en} } @article{SperfeldSchmidtkeGaedkeetal.2010, author = {Sperfeld, Erik and Schmidtke, Andrea and Gaedke, Ursula and Weithoff, Guntram}, title = {Productivity, herbivory, and species traits rather than diversity influence invasibility of experimental phytoplankton communities}, issn = {0029-8549}, doi = {10.1007/s00442-010-1594-4}, year = {2010}, abstract = {Biological invasions are a major threat to natural biodiversity; hence, understanding the mechanisms underlying invasibility (i.e., the susceptibility of a community to invasions by new species) is crucial. Invasibility of a resident community may be affected by a complex but hitherto hardly understood interplay of (1) productivity of the habitat, (2) diversity, (3) herbivory, and (4) the characteristics of both invasive and resident species. Using experimental phytoplankton microcosms, we investigated the effect of nutrient supply and species diversity on the invasibility of resident communities for two functionally different invaders in the presence or absence of an herbivore. With increasing nutrient supply, increased herbivore abundance indicated enhanced phytoplankton biomass production, and the invasion success of both invaders showed a unimodal pattern. At low nutrient supply (i.e., low influence of herbivory), the invasibility depended mainly on the competitive abilities of the invaders, whereas at high nutrient supply, the susceptibility to herbivory dominated. This resulted in different optimum nutrient levels for invasion success of the two species due to their individual functional traits. To test the effect of diversity on invasibility, a species richness gradient was generated by random selection from a resident species pool at an intermediate nutrient level. Invasibility was not affected by species richness; instead, it was driven by the functional traits of the resident and/or invasive species mediated by herbivore density. Overall, herbivory was the driving factor for invasibility of phytoplankton communities, which implies that other factors affecting the intensity of herbivory (e.g., productivity or edibility of primary producers) indirectly influence invasions.}, language = {en} } @article{CeulemansGuillGaedke2021, author = {Ceulemans, Ruben and Guill, Christian and Gaedke, Ursula}, title = {Top predators govern multitrophic diversity effects in tritrophic food webs}, series = {Ecology : a publication of the Ecological Society of America}, volume = {102}, journal = {Ecology : a publication of the Ecological Society of America}, number = {7}, publisher = {Wiley}, address = {Hoboken}, issn = {0012-9658}, doi = {10.1002/ecy.3379}, pages = {16}, year = {2021}, abstract = {It is well known that functional diversity strongly affects ecosystem functioning. However, even in rather simple model communities consisting of only two or, at best, three trophic levels, the relationship between multitrophic functional diversity and ecosystem functioning appears difficult to generalize, because of its high contextuality. In this study, we considered several differently structured tritrophic food webs, in which the amount of functional diversity was varied independently on each trophic level. To achieve generalizable results, largely independent of parametrization, we examined the outcomes of 128,000 parameter combinations sampled from ecologically plausible intervals, with each tested for 200 randomly sampled initial conditions. Analysis of our data was done by training a random forest model. This method enables the identification of complex patterns in the data through partial dependence graphs, and the comparison of the relative influence of model parameters, including the degree of diversity, on food-web properties. We found that bottom-up and top-down effects cascade simultaneously throughout the food web, intimately linking the effects of functional diversity of any trophic level to the amount of diversity of other trophic levels, which may explain the difficulty in unifying results from previous studies. Strikingly, only with high diversity throughout the whole food web, different interactions synergize to ensure efficient exploitation of the available nutrients and efficient biomass transfer to higher trophic levels, ultimately leading to a high biomass and production on the top level. The temporal variation of biomass showed a more complex pattern with increasing multitrophic diversity: while the system initially became less variable, eventually the temporal variation rose again because of the increasingly complex dynamical patterns. Importantly, top predator diversity and food-web parameters affecting the top trophic level were of highest importance to determine the biomass and temporal variability of any trophic level. Overall, our study reveals that the mechanisms by which diversity influences ecosystem functioning are affected by every part of the food web, hampering the extrapolation of insights from simple monotrophic or bitrophic systems to complex natural food webs.}, language = {en} } @article{EhrlichGaedke2020, author = {Ehrlich, Elias and Gaedke, Ursula}, title = {Coupled changes in traits and biomasses cascading through a tritrophic plankton food web}, series = {Limnology and oceanography}, volume = {65}, journal = {Limnology and oceanography}, number = {10}, publisher = {Wiley}, address = {Hoboken}, issn = {0024-3590}, doi = {10.1002/lno.11466}, pages = {2502 -- 2514}, year = {2020}, abstract = {Trait-based approaches have broadened our understanding of how the composition of ecological communities responds to environmental drivers. This research has mainly focussed on abiotic factors and competition determining the community trait distribution, while effects of trophic interactions on trait dynamics, if considered at all, have been studied for two trophic levels at maximum. However, natural food webs are typically at least tritrophic. This enables indirect interactions of traits and biomasses among multiple trophic levels leading to underexplored effects on food web dynamics. Here, we demonstrate the occurrence of mutual trait adjustment among three trophic levels in a natural plankton food web (Lake Constance) and in a corresponding mathematical model. We found highly recurrent seasonal biomass and trait dynamics, where herbivorous zooplankton increased its size, and thus its ability to counter phytoplankton defense, before phytoplankton defense actually increased. This is contrary to predictions from bitrophic systems where counter-defense of the consumer is a reaction to prey defense. In contrast, counter-defense of carnivores by size adjustment followed the defense of herbivores as expected. By combining observations and model simulations, we show how the reversed trait dynamics at the two lower trophic levels result from a "trophic biomass-trait cascade" driven by the carnivores. Trait adjustment between two trophic levels can therefore be altered by biomass or trait changes of adjacent trophic levels. Hence, analyses of only pairwise trait adjustment can be misleading in natural food webs, while multitrophic trait-based approaches capture indirect biomass-trait interactions among multiple trophic levels.}, language = {en} }