@article{KlauschiesCoutinhoGaedke2018, author = {Klauschies, Toni and Coutinho, Renato Mendes and Gaedke, Ursula}, title = {A beta distribution-based moment closure enhances the reliability of trait-based aggregate models for natural populations and communities}, series = {Ecological modelling : international journal on ecological modelling and engineering and systems ecolog}, volume = {381}, journal = {Ecological modelling : international journal on ecological modelling and engineering and systems ecolog}, publisher = {Elsevier}, address = {Amsterdam}, issn = {0304-3800}, doi = {10.1016/j.ecolmodel.2018.02.001}, pages = {46 -- 77}, year = {2018}, abstract = {Ecological communities are complex adaptive systems that exhibit remarkable feedbacks between their biomass and trait dynamics. Trait-based aggregate models cope with this complexity by focusing on the temporal development of the community's aggregate properties such as its total biomass, mean trait and trait variance. They are based on particular assumptions about the shape of the underlying trait distribution, which is commonly assumed to be normal. However, ecologically important traits are usually restricted to a finite range, and empirical trait distributions are often skewed or multimodal. As a result, normal distribution-based aggregate models may fail to adequately represent the biomass and trait dynamics of natural communities. We resolve this mismatch by developing a new moment closure approach assuming the trait values to be beta-distributed. We show that the beta distribution captures important shape properties of both observed and simulated trait distributions, which cannot be captured by a Gaussian. We further demonstrate that a beta distribution-based moment closure can strongly enhance the reliability of trait-based aggregate models. We compare the biomass, mean trait and variance dynamics of a full trait distribution (FD) model to the ones of beta (BA) and normal (NA) distribution-based aggregate models, under different selection regimes. This way, we demonstrate under which general conditions (stabilizing, fluctuating or disruptive selection) different aggregate models are reliable tools. All three models predicted very similar biomass and trait dynamics under stabilizing selection yielding unimodal trait distributions with small standing trait variation. We also obtained an almost perfect match between the results of the FD and BA models under fluctuating selection, promoting skewed trait distributions and ongoing oscillations in the biomass and trait dynamics. In contrast, the NA model showed unrealistic trait dynamics and exhibited different alternative stable states, and thus a high sensitivity to initial conditions under fluctuating selection. Under disruptive selection, both aggregate models failed to reproduce the results of the FD model with the mean trait values remaining within their ecologically feasible ranges in the BA model but not in the NA model. Overall, a beta distribution-based moment closure strongly improved the realism of trait-based aggregate models.}, language = {en} } @article{MooijTrolleJeppesenetal.2010, author = {Mooij, Wolf M. and Trolle, Dennis and Jeppesen, Erik and Arhonditsis, George B. and Belolipetsky, Pavel V. and Chitamwebwa, Deonatus B. R. and Degermendzhy, Andrey G. and DeAngelis, Donald L. and Domis, Lisette Nicole de Senerpont and Downing, Andrea S. and Elliott, J. Alex and Fragoso Jr, Carlos Ruberto and Gaedke, Ursula and Genova, Svetlana N. and Gulati, Ramesh D. and H{\aa}kanson, Lars and Hamilton, David P. and Hipsey, Matthew R. and 't Hoen, Jochem and H{\"u}lsmann, Stephan and Los, F. Hans and Makler-Pick, Vardit and Petzoldt, Thomas and Prokopkin, Igor G. and Rinke, Karsten and Schep, Sebastiaan A. and Tominaga, Koji and Van Dam, Anne A. and Van Nes, Egbert H. and Wells, Scott A. and Janse, Jan H.}, title = {Challenges and opportunities for integrating lake ecosystem modelling approaches}, series = {Aquatic ecology}, volume = {44}, journal = {Aquatic ecology}, publisher = {Springer Science + Business Media B.V.}, address = {Dordrecht}, issn = {1573-5125}, doi = {10.1007/s10452-010-9339-3}, pages = {633 -- 667}, year = {2010}, abstract = {A large number and wide variety of lake ecosystem models have been developed and published during the past four decades. We identify two challenges for making further progress in this field. One such challenge is to avoid developing more models largely following the concept of others ('reinventing the wheel'). The other challenge is to avoid focusing on only one type of model, while ignoring new and diverse approaches that have become available ('having tunnel vision'). In this paper, we aim at improving the awareness of existing models and knowledge of concurrent approaches in lake ecosystem modelling, without covering all possible model tools and avenues. First, we present a broad variety of modelling approaches. To illustrate these approaches, we give brief descriptions of rather arbitrarily selected sets of specific models. We deal with static models (steady state and regression models), complex dynamic models (CAEDYM, CE-QUAL-W2, Delft 3D-ECO, LakeMab, LakeWeb, MyLake, PCLake, PROTECH, SALMO), structurally dynamic models and minimal dynamic models. We also discuss a group of approaches that could all be classified as individual based: super-individual models (Piscator, Charisma), physiologically structured models, stage-structured models and traitbased models. We briefly mention genetic algorithms, neural networks, Kalman filters and fuzzy logic. Thereafter, we zoom in, as an in-depth example, on the multi-decadal development and application of the lake ecosystem model PCLake and related models (PCLake Metamodel, Lake Shira Model, IPH-TRIM3D-PCLake). In the discussion, we argue that while the historical development of each approach and model is understandable given its 'leading principle', there are many opportunities for combining approaches. We take the point of view that a single 'right' approach does not exist and should not be strived for. Instead, multiple modelling approaches, applied concurrently to a given problem, can help develop an integrative view on the functioning of lake ecosystems. We end with a set of specific recommendations that may be of help in the further development of lake ecosystem models.}, 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} } @article{RaatzvanVelzenGaedke2019, author = {Raatz, Michael and van Velzen, Ellen and Gaedke, Ursula}, title = {Co-adaptation impacts the robustness of predator-prey dynamics against perturbations}, series = {Ecology and Evolution}, volume = {9}, journal = {Ecology and Evolution}, number = {7}, publisher = {John Wiley \& Sons}, address = {Hoboken, NJ}, issn = {2045-7758}, doi = {10.1002/ece3.5006}, pages = {3823 -- 3836}, year = {2019}, abstract = {Global change threatens the maintenance of ecosystem functions that are shaped by the persistence and dynamics of populations. It has been shown that the persistence of species increases if they possess larger trait adaptability. Here, we investigate whether trait adaptability also affects the robustness of population dynamics of interacting species and thereby shapes the reliability of ecosystem functions that are driven by these dynamics. We model co-adaptation in a predator-prey system as changes to predator offense and prey defense due to evolution or phenotypic plasticity. We investigate how trait adaptation affects the robustness of population dynamics against press perturbations to environmental parameters and against pulse perturbations targeting species abundances and their trait values. Robustness of population dynamics is characterized by resilience, elasticity, and resistance. In addition to employing established measures for resilience and elasticity against pulse perturbations (extinction probability and return time), we propose the warping distance as a new measure for resistance against press perturbations, which compares the shapes and amplitudes of pre- and post-perturbation population dynamics. As expected, we find that the robustness of population dynamics depends on the speed of adaptation, but in nontrivial ways. Elasticity increases with speed of adaptation as the system returns more rapidly to the pre-perturbation state. Resilience, in turn, is enhanced by intermediate speeds of adaptation, as here trait adaptation dampens biomass oscillations. The resistance of population dynamics strongly depends on the target of the press perturbation, preventing a simple relationship with the adaptation speed. In general, we find that low robustness often coincides with high amplitudes of population dynamics. Hence, amplitudes may indicate the robustness against perturbations also in other natural systems with similar dynamics. Our findings show that besides counteracting extinctions, trait adaptation indeed strongly affects the robustness of population dynamics against press and pulse perturbations.}, language = {en} } @article{SpijkermanBehrendFachetal.2018, author = {Spijkerman, Elly and Behrend, Hella and Fach, Bettina and Gaedke, Ursula}, title = {Decreased phosphorus incorporation explains the negative effect of high iron concentrations in the green microalga Chlamydomonas acidophila}, series = {The science of the total environment : an international journal for scientific research into the environment and its relationship with man}, volume = {626}, journal = {The science of the total environment : an international journal for scientific research into the environment and its relationship with man}, publisher = {Elsevier}, address = {Amsterdam}, issn = {0048-9697}, doi = {10.1016/j.scitotenv.2018.01.188}, pages = {1342 -- 1349}, year = {2018}, abstract = {The green microalga Chlamydomonas acidophila is an important primary producer in very acidic lakes (pH 2.0-3.5), characterized by high concentrations of ferric iron (up to 1 g total Fe L-1) and low rates of primary production. It was previously suggested that these high iron concentrations result in high iron accumulation and inhibit photosynthesis in C. acidophila. To test this, the alga was grown in sterilized lake water and in medium with varying total iron concentrations under limiting and sufficient inorganic phosphorus (Pi) supply, because Pi is an important growth limiting nutrient in acidic waters. Photosynthesis and growth of C. acidophila as measured over 5 days were largely unaffected by high total iron concentrations and only decreased if free ionic Fe3+ concentrations exceeded 100 mg Fe3+ L-1. Although C. acidophila was relatively rich in iron (up to 5 mmol Fe: mol C), we found no evidence of iron toxicity. In contrast, a concentration of 260 mg total Fe L-1 (i.e. 15 mg free ionic Fe3+ L-1), which is common in many acidic lakes, reduced Pi-incorporation by 50\% and will result in Pi-limited photosynthesis. The resulting Pi-limitation present at high iron and Pi concentrations was illustrated by elevated maximum Pi-uptake rates. No direct toxic effects of high iron were found, but unfavourable chemical Pi-speciation reduced growth of the acidophile alga.}, 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{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{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{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} } @article{vanVelzenThieserBerendonketal.2018, author = {van Velzen, Ellen and Thieser, Tamara and Berendonk, Thomas U. and Weitere, Markus and Gaedke, Ursula}, title = {Inducible defense destabilizes predator-prey dynamics}, series = {Oikos}, volume = {127}, journal = {Oikos}, number = {11}, publisher = {Wiley}, address = {Hoboken}, issn = {0030-1299}, doi = {10.1111/oik.04868}, pages = {1551 -- 1562}, year = {2018}, abstract = {Phenotypic plasticity in prey can have a dramatic impact on predator-prey dynamics, e.g. by inducible defense against temporally varying levels of predation. Previous work has overwhelmingly shown that this effect is stabilizing: inducible defenses dampen the amplitudes of population oscillations or eliminate them altogether. However, such studies have neglected scenarios where being protected against one predator increases vulnerability to another (incompatible defense). Here we develop a model for such a scenario, using two distinct prey phenotypes and two predator species. Each prey phenotype is defended against one of the predators, and vulnerable to the other. In strong contrast with previous studies on the dynamic effects of plasticity involving a single predator, we find that increasing the level of plasticity consistently destabilizes the system, as measured by the amplitude of oscillations and the coefficients of variation of both total prey and total predator biomasses. We explain this unexpected and seemingly counterintuitive result by showing that plasticity causes synchronization between the two prey phenotypes (and, through this, between the predators), thus increasing the temporal variability in biomass dynamics. These results challenge the common view that plasticity should always have a stabilizing effect on biomass dynamics: adding a single predator-prey interaction to an established model structure gives rise to a system where different mechanisms may be at play, leading to dramatically different outcomes.}, language = {en} } @article{FloederYongKlauschiesetal.2021, author = {Fl{\"o}der, Sabine and Yong, Joanne and Klauschies, Toni and Gaedke, Ursula and Poprick, Tobias and Brinkhoff, Thorsten and Moorthi, Stefanie}, title = {Intraspecific trait variation alters the outcome of competition in freshwater ciliates}, series = {Ecology and evolution}, volume = {11}, journal = {Ecology and evolution}, number = {15}, publisher = {Wiley}, address = {Hoboken}, issn = {2045-7758}, doi = {10.1002/ece3.7828}, pages = {10225 -- 10243}, year = {2021}, abstract = {Trait variation among heterospecific and conspecific organisms may substantially affect community and food web dynamics. While the relevance of competition and feeding traits have been widely studied for different consumer species, studies on intraspecific differences are more scarce, partly owing to difficulties in distinguishing different clones of the same species. Here, we investigate how intraspecific trait variation affects the competition between the freshwater ciliates Euplotes octocarinatus and Coleps hirtus in a nitrogen-limited chemostat system. The ciliates competed for the microalgae Cryptomonas sp. (Cry) and Navicula pelliculosa (Nav), and the bacteria present in the cultures over a period of 33 days. We used monoclonal Euplotes and three different Coleps clones (Col 1, Col 2, and Col 3) in the experiment that could be distinguished by a newly developed rDNA-based molecular assay based on the internal transcribed spacer (ITS) regions. While Euplotes feeds on Cry and on bacteria, the Coleps clones cannot survive on bacteria alone but feed on both Cry and Nav with clone-specific rates. Experimental treatments comprised two-species mixtures of Euplotes and one or all of the three different Coleps clones, respectively. We found intraspecific variation in the traits "selectivity" and "maximum ingestion rate" for the different algae to significantly affect the competitive outcome between the two ciliate species. As Nav quickly escaped top-down control and likely reached a state of low food quality, ciliate competition was strongly determined by the preference of different Coleps clones for Cry as opposed to feeding on Nav. In addition, the ability of Euplotes to use bacteria as an alternative food source strengthened its persistence once Cry was depleted. Hence, trait variation at both trophic levels codetermined the population dynamics and the outcome of species competition.}, language = {en} } @article{MittlerBlasiusGaedkeetal.2018, author = {Mittler, Udo and Blasius, Bernd and Gaedke, Ursula and Ryabov, Alexey B.}, title = {Length-volume relationship of lake phytoplankton}, series = {Limnology and Oceanography: Methods}, volume = {17}, journal = {Limnology and Oceanography: Methods}, number = {1}, publisher = {Wiley}, address = {Hoboken}, issn = {1541-5856}, doi = {10.1002/lom3.10296}, pages = {58 -- 68}, year = {2018}, abstract = {The shapes of phytoplankton units (unicellular organisms and colonies) are extremely diverse, and no unique relationship exists between their volume, V, and longest linear dimension, L. However, an approximate scaling between these parameters can be found because the shape variations within each size class are constrained by cell physiology, grazing pressure, and optimality of resource acquisition. To determine this scaling and to test for its seasonal and interannual variation under changing environmental conditions, we performed weighted regression analysis of time-dependent length-volume relations of the phytoplankton community in large deep Lake Constance from 1979 to 1999. We show that despite a large variability in species composition, the V(L) relationship can be approximated as a power law, V similar to L-alpha, with a scaling exponent alpha = 3 for small cells (L < 25 mu m) and alpha = 1.7 if the fitting is performed over the entire length range, including individual cells and colonies. The best description is provided by a transitional power function describing a regime change from a scaling exponent of 3 for small cells to an exponent of 0.4 in the range of large phytoplankton. Testing different weighted fitting approaches we show that remarkably the best prediction of the total community biovolume from measurements of L and cell density is obtained when the regression is weighted with the squares of species abundances. Our approach should also be applicable to other systems and allows converting phytoplankton length distributions (e.g., obtained with automatic monitoring such as flow cytometry) into distributions of biovolume and biovolume-related phytoplankton traits.}, language = {en} } @article{BlasiusRudolfWeithoffetal.2019, author = {Blasius, Bernd and Rudolf, Lars and Weithoff, Guntram and Gaedke, Ursula and Fussmann, Gregor F.}, title = {Long-term cyclic persistence in an experimental predator-prey system}, series = {Nature : the international weekly journal of science}, volume = {577}, journal = {Nature : the international weekly journal of science}, number = {7789}, publisher = {Nature Publ. Group}, address = {London}, issn = {0028-0836}, doi = {10.1038/s41586-019-1857-0}, pages = {226 -- 230}, year = {2019}, abstract = {Predator-prey cycles rank among the most fundamental concepts in ecology, are predicted by the simplest ecological models and enable, theoretically, the indefinite persistence of predator and prey(1-4). However, it remains an open question for how long cyclic dynamics can be self-sustained in real communities. Field observations have been restricted to a few cycle periods(5-8) and experimental studies indicate that oscillations may be short-lived without external stabilizing factors(9-19). Here we performed microcosm experiments with a planktonic predator-prey system and repeatedly observed oscillatory time series of unprecedented length that persisted for up to around 50 cycles or approximately 300 predator generations. The dominant type of dynamics was characterized by regular, coherent oscillations with a nearly constant predator-prey phase difference. Despite constant experimental conditions, we also observed shorter episodes of irregular, non-coherent oscillations without any significant phase relationship. However, the predator-prey system showed a strong tendency to return to the dominant dynamical regime with a defined phase relationship. A mathematical model suggests that stochasticity is probably responsible for the reversible shift from coherent to non-coherent oscillations, a notion that was supported by experiments with external forcing by pulsed nutrient supply. Our findings empirically demonstrate the potential for infinite persistence of predator and prey populations in a cyclic dynamic regime that shows resilience in the presence of stochastic events.}, language = {en} } @article{EhrlichGaedke2018, author = {Ehrlich, Elias and Gaedke, Ursula}, title = {Not attackable or not crackable}, series = {Ecology and Evolution}, volume = {8}, journal = {Ecology and Evolution}, number = {13}, publisher = {Wiley}, issn = {2045-7758}, doi = {10.1002/ece3.4145}, pages = {6625 -- 6637}, year = {2018}, abstract = {It is well-known that prey species often face trade-offs between defense against predation and competitiveness, enabling predator-mediated coexistence. However, we lack an understanding of how the large variety of different defense traits with different competition costs affects coexistence and population dynamics. Our study focusses on two general defense mechanisms, that is, pre-attack (e.g., camouflage)and post-attack defenses (e.g., weaponry) that act at different phases of the predator—prey interaction. We consider a food web model with one predator, two prey types and one resource. One prey type is undefended, while the other one is pre-or post-attack defended paying costs either by a higher half-saturation constant for resource uptake or a lower maximum growth rate. We show that post-attack defenses promote prey coexistence and stabilize the population dynamics more strongly than pre-attack defenses by interfering with the predator's functional response: Because the predator spends time handling "noncrackable" prey, the undefended prey is indirectly facilitated. A high half-saturation constant as defense costs promotes coexistence more and stabilizes the dynamics less than a low maximum growth rate. The former imposes high costs at low resource concentrations but allows for temporally high growth rates at predator-induced resource peaks preventing the extinction of the defended prey. We evaluate the effects of the different defense mechanisms and costs on coexistence under different enrichment levels in order to vary the importance of bottom-up and top-down control of the prey community.}, language = {en} } @article{VelzenGaedkeKlauschies2022, author = {Velzen, Ellen van and Gaedke, Ursula and Klauschies, Toni}, title = {Quantifying the capacity for contemporary trait changes to drive intermittent predator-prey cycles}, series = {Ecological monographs : a publication of the Ecological Society of America}, volume = {92}, journal = {Ecological monographs : a publication of the Ecological Society of America}, number = {2}, publisher = {Wiley}, address = {New York}, issn = {1557-7015}, doi = {10.1002/ecm.1505}, pages = {29}, year = {2022}, abstract = {A large and growing body of theory has demonstrated how the presence of trait variation in prey or predator populations may affect the amplitude and phase of predator-prey cycles. Less attention has been given to so-called intermittent cycles, in which predator-prey oscillations recurrently disappear and re-appear, despite such dynamics being observed in empirical systems and modeling studies. A comprehensive understanding of the conditions under which trait changes may drive intermittent predator-prey dynamics, as well as their potential ecological implications, is therefore missing. Here we provide a first systematic analysis of the eco-evolutionary conditions that may give rise to intermittent predator-prey cycles, investigating 16 models that incorporate different types of trait variation within prey, predators, or both. Our results show that intermittent dynamics often arise through predator-prey coevolution, but only very rarely when only one trophic level can adapt. Additionally, the frequency of intermittent cycles depends on the source of trait variation (genetic variation or phenotypic plasticity) and the genetic architecture (Mendelian or quantitative traits), with intermittency occurring most commonly through Mendelian evolution, and very rarely through phenotypic plasticity. Further analysis identified three necessary conditions for when trait variation can drive intermittent cycles. First, the intrinsic stability of the predator-prey system must depend on the traits of prey, predators, or both. Second, there must be a mechanism causing the recurrent alternation between stable and unstable states, leading to a "trait" cycle superimposed on the population dynamics. Finally, these trait dynamics must be significantly slower than the predator-prey cycles. We show how these conditions explain all the abovementioned patterns. We further show an important unexpected consequence of these necessary conditions: they are most easily met when intraspecific trait variation is at high risk of being lost. As trait diversity is positively associated with ecosystem functioning, this can have potentially severe negative consequences. This novel result highlights the importance of identifying and understanding intermittent cycles in theoretical studies and natural systems. The new approach for detecting and quantifying intermittency we develop here will be instrumental in enabling future study.}, language = {en} } @article{LiYangSunetal.2021, author = {Li, Xiaoxiao and Yang, Wei and Sun, Tao and Gaedke, Ursula}, title = {Quantitative food web structure and ecosystem functions in a warm-temperate seagrass bed}, series = {Marine biology : international journal on life in oceans and coastal waters}, volume = {168}, journal = {Marine biology : international journal on life in oceans and coastal waters}, number = {5}, publisher = {Springer}, address = {Berlin ; Heidelberg}, issn = {0025-3162}, doi = {10.1007/s00227-021-03878-z}, pages = {17}, year = {2021}, abstract = {Seagrass beds are important habitats in coastal areas but increasingly decline in area and quality, thus conservation measures are urgently needed. Quantitative food webs, describing the biomass distribution and energy fluxes among trophic groups, reveal structural and functional aspects of ecosystems. Their knowledge can improve ecological conservation. For the recently discovered large warm-temperate seagrass (Zostera japonica) habitat in China's Yellow River Delta wetland, we used delta C-13 and delta N-15 measurements and a Bayesian isotope mixing model to construct its food web diagram with quantitative estimations of consumer diet compositions, comprising detritus and 14 living trophic groups from primary producers to fish. We then estimated the quantitative food web fluxes based on biomass measurements and calculated corresponding ecosystem functions. Pelagic producers were significantly C-13-depleted compared to benthic sources. Consumers (except zooplankton) were increasingly C-13-depleted with increasing trophic positions even though the consumed benthic production surpassed the pelagic one. Bivalves dominated consumer biomasses and fluxes and were the first to connect the pelagic and benthic pathways, whereas zooplankton and gastropods were specialized on the two pathways, respectively. We found flat biomass and production pyramids indicating low trophic transfer efficiencies. Generally, the energetic structure of the quantitative food web was consistent with the stable isotope analysis, and the estimated net primary production and most estimated production to biomass ratios of the trophic groups fell within literature ranges. This study provides a systematical understanding of the quantitative trophic ecology of a seagrass bed and facilitates synergistic knowledge on management, conservation, and restoration.}, language = {en} } @article{MuhlRoelkeZoharyetal.2018, author = {Muhl, Rika M. W. and Roelke, Daniel L. and Zohary, Tamar and Moustaka-Gouni, Maria and Sommer, Ulrich and Borics, Gabor and Gaedke, Ursula and Withrow, Frances G. and Bhattacharyya, Joydeb}, title = {Resisting annihilation}, series = {Ecology letters}, volume = {21}, journal = {Ecology letters}, number = {9}, publisher = {Wiley}, address = {Hoboken}, issn = {1461-023X}, doi = {10.1111/ele.13109}, pages = {1390 -- 1400}, year = {2018}, abstract = {Allelopathic species can alter biodiversity. Using simulated assemblages that are characterised by neutrality, lumpy coexistence and intransitivity, we explore relationships between within-assemblage competitive dissimilarities and resistance to allelopathic species. An emergent behaviour from our models is that assemblages are more resistant to allelopathy when members strongly compete exploitatively (high competitive power). We found that neutral assemblages were the most vulnerable to allelopathic species, followed by lumpy and then by intransitive assemblages. We find support for our modeling in real-world time-series data from eight lakes of varied morphometry and trophic state. Our analysis of this data shows that a lake's history of allelopathic phytoplankton species biovolume density and dominance is related to the number of species clusters occurring in the plankton assemblages of those lakes, an emergent trend similar to that of our modeling. We suggest that an assemblage's competitive power determines its allelopathy resistance.}, language = {en} } @article{vanVelzenGaedke2018, author = {van Velzen, Ellen and Gaedke, Ursula}, title = {Reversed predator}, series = {Ecology and Evolution}, journal = {Ecology and Evolution}, publisher = {Wiley}, address = {Hoboken}, issn = {2045-7758}, doi = {10.1002/ece3.4184}, pages = {1 -- 13}, year = {2018}, abstract = {Ecoevolutionary feedbacks in predator-prey systems have been shown to qualitatively alter predator-prey dynamics. As a striking example, defense-offense coevolution can reverse predator-prey cycles, so predator peaks precede prey peaks rather than vice versa. However, this has only rarely been shown in either model studies or empirical systems. Here, we investigate whether this rarity is a fundamental feature of reversed cycles by exploring under which conditions they should be found. For this, we first identify potential conditions and parameter ranges most likely to result in reversed cycles by developing a new measure, the effective prey biomass, which combines prey biomass with prey and predator traits, and represents the prey biomass as perceived by the predator. We show that predator dynamics always follow the dynamics of the effective prey biomass with a classic ¼-phase lag. From this key insight, it follows that in reversed cycles (i.e., ¾-lag), the dynamics of the actual and the effective prey biomass must be in antiphase with each other, that is, the effective prey biomass must be highest when actual prey biomass is lowest, and vice versa. Based on this, we predict that reversed cycles should be found mainly when oscillations in actual prey biomass are small and thus have limited impact on the dynamics of the effective prey biomass, which are mainly driven by trait changes. We then confirm this prediction using numerical simulations of a coevolutionary predator-prey system, varying the amplitude of the oscillations in prey biomass: Reversed cycles are consistently associated with regions of parameter space leading to small-amplitude prey oscillations, offering a specific and highly testable prediction for conditions under which reversed cycles should occur in natural systems.}, language = {en} } @article{vanVelzenGaedke2018, author = {van Velzen, Ellen and Gaedke, Ursula}, title = {Reversed predator-prey cycles are driven by the amplitude of prey oscillations}, series = {Ecology and evolution}, volume = {8}, journal = {Ecology and evolution}, number = {12}, publisher = {Wiley}, address = {Hoboken}, issn = {2045-7758}, doi = {10.1002/ece3.4184}, pages = {6317 -- 6329}, year = {2018}, abstract = {Ecoevolutionary feedbacks in predator-prey systems have been shown to qualitatively alter predator-prey dynamics. As a striking example, defense-offense coevolution can reverse predator-prey cycles, so predator peaks precede prey peaks rather than vice versa. However, this has only rarely been shown in either model studies or empirical systems. Here, we investigate whether this rarity is a fundamental feature of reversed cycles by exploring under which conditions they should be found. For this, we first identify potential conditions and parameter ranges most likely to result in reversed cycles by developing a new measure, the effective prey biomass, which combines prey biomass with prey and predator traits, and represents the prey biomass as perceived by the predator. We show that predator dynamics always follow the dynamics of the effective prey biomass with a classic 1/4-phase lag. From this key insight, it follows that in reversed cycles (i.e., -lag), the dynamics of the actual and the effective prey biomass must be in antiphase with each other, that is, the effective prey biomass must be highest when actual prey biomass is lowest, and vice versa. Based on this, we predict that reversed cycles should be found mainly when oscillations in actual prey biomass are small and thus have limited impact on the dynamics of the effective prey biomass, which are mainly driven by trait changes. We then confirm this prediction using numerical simulations of a coevolutionary predator-prey system, varying the amplitude of the oscillations in prey biomass: Reversed cycles are consistently associated with regions of parameter space leading to small-amplitude prey oscillations, offering a specific and highly testable prediction for conditions under which reversed cycles should occur in natural systems.}, language = {en} } @article{BengfortvanVelzenGaedke2017, author = {Bengfort, Michael and van Velzen, Ellen and Gaedke, Ursula}, title = {Slight phenotypic variation in predators and prey causes complex predator-prey oscillations}, series = {Ecological Complexity}, volume = {31}, journal = {Ecological Complexity}, publisher = {Elsevier}, address = {Amsterdam}, issn = {1476-945X}, doi = {10.1016/j.ecocom.2017.06.003}, pages = {115 -- 124}, year = {2017}, abstract = {Predator-prey oscillations are expected to show a 1/4-phase lag between predator and prey. However, observed dynamics of natural or experimental predator-prey systems are often more complex. A striking but hardly studied example are sudden interruptions of classic 1/4-lag cycles with periods of antiphase oscillations, or periods without any regular predator-prey oscillations. These interruptions occur for a limited time before the system reverts to regular 1/4-lag oscillations, thus yielding intermittent cycles. Reasons for this behaviour are often difficult to reveal in experimental systems. Here we test the hypothesis that such complex dynamical behaviour may result from minor trait variation and trait adaptation in both the prey and predator, causing recurrent small changes in attack rates that may be hard to capture by empirical measurements. Using a model structure where the degree of trait variation in the predator can be explicitly controlled, we show that a very limited amount of adaptation resulting in 10-15\% temporal variation in attack rates is already sufficient to generate these intermittent dynamics. Such minor variation may be present in experimental predator-prey systems, and may explain disruptions in regular 1/4-lag oscillations.}, language = {en} } @article{PennekampIlesGarlandetal.2019, author = {Pennekamp, Frank and Iles, Alison C. and Garland, Joshua and Brennan, Georgina and Brose, Ulrich and Gaedke, Ursula and Jacob, Ute and Kratina, Pavel and Matthews, Blake and Munch, Stephan and Novak, Mark and Palamara, Gian Marco and Rall, Bjorn C. and Rosenbaum, Benjamin and Tabi, Andrea and Ward, Colette and Williams, Richard and Ye, Hao and Petchey, Owen L.}, title = {The intrinsic predictability of ecological time series and its potential to guide forecasting}, series = {Ecological monographs : a publication of the Ecological Society of America.}, volume = {89}, journal = {Ecological monographs : a publication of the Ecological Society of America.}, number = {2}, publisher = {Wiley}, address = {Hoboken}, issn = {0012-9615}, doi = {10.1002/ecm.1359}, pages = {17}, year = {2019}, language = {en} } @article{EhrlichKathGaedke2020, author = {Ehrlich, Elias and Kath, Nadja Jeanette and Gaedke, Ursula}, title = {The shape of a defense-growth trade-off governs seasonal trait dynamics in natural phytoplankton}, series = {The ISME journal}, volume = {14}, journal = {The ISME journal}, number = {6}, publisher = {Nature Publishing Group}, address = {London}, issn = {1751-7362}, doi = {10.1038/s41396-020-0619-1}, pages = {1451 -- 1462}, year = {2020}, abstract = {Theory predicts that trade-offs, quantifying costs of functional trait adjustments, crucially affect community trait adaptation to altered environmental conditions, but empirical verification is scarce. We evaluated trait dynamics (antipredator defense, maximum growth rate, and phosphate affinity) of a lake phytoplankton community in a seasonally changing environment, using literature trait data and 21 years of species-resolved high-frequency biomass measurements. The trait data indicated a concave defense-growth trade-off, promoting fast-growing species with intermediate defense. With seasonally increasing grazing pressure, the community shifted toward higher defense levels at the cost of lower growth rates along the trade-off curve, while phosphate affinity explained some deviations from it. We discuss how low fitness differences of species, inferred from model simulations, in concert with stabilizing mechanisms, e.g., arising from further trait dimensions, may lead to the observed phytoplankton diversity. In conclusion, quantifying trade-offs is key for predictions of community trait adaptation and biodiversity under environmental change.}, 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{KlauschiesVasseurGaedke2016, author = {Klauschies, Toni and Vasseur, David A. and Gaedke, Ursula}, title = {Trait adaptation promotes species coexistence in diverse predator and prey communities}, series = {Ecology and evolution}, journal = {Ecology and evolution}, publisher = {John Wiley \& Sons, Inc.}, issn = {2045-7758}, doi = {10.1002/ece3.2172}, pages = {19}, year = {2016}, abstract = {Species can adjust their traits in response to selection which may strongly influence species coexistence. Nevertheless, current theory mainly assumes distinct and time-invariant trait values. We examined the combined effects of the range and the speed of trait adaptation on species coexistence using an innovative multispecies predator-prey model. It allows for temporal trait changes of all predator and prey species and thus simultaneous coadaptation within and among trophic levels. We show that very small or slow trait adaptation did not facilitate coexistence because the stabilizing niche differences were not sufficient to offset the fitness differences. In contrast, sufficiently large and fast trait adaptation jointly promoted stable or neutrally stable species coexistence. Continuous trait adjustments in response to selection enabled a temporally variable convergence and divergence of species traits; that is, species became temporally more similar (neutral theory) or dissimilar (niche theory) depending on the selection pressure, resulting over time in a balance between niche differences stabilizing coexistence and fitness differences promoting competitive exclusion. Furthermore, coadaptation allowed prey and predator species to cluster into different functional groups. This equalized the fitness of similar species while maintaining sufficient niche differences among functionally different species delaying or preventing competitive exclusion. In contrast to pre- vious studies, the emergent feedback between biomass and trait dynamics enabled supersaturated coexistence for a broad range of potential trait adaptation and parameters. We conclude that accounting for trait adaptation may explain stable and supersaturated species coexistence for a broad range of environmental conditions in natural systems when the absence of such adaptive changes would preclude it. Small trait changes, coincident with those that may occur within many natural populations, greatly enlarged the number of coexisting species.}, language = {en} }