TY - JOUR A1 - van Velzen, Ellen T1 - Predator coexistence through emergent fitness equalization JF - Ecology : a publication of the Ecological Society of America N2 - The competitive exclusion principle is one of the oldest ideas in ecology and states that without additional self-limitation two predators cannot coexist on a single prey. The search for mechanisms allowing coexistence despite this has identified niche differentiation between predators as crucial: without this, coexistence requires the predators to have exactly the same R* values, which is considered impossible. However, this reasoning misses a critical point: predators' R* values are not static properties, but affected by defensive traits of their prey, which in turn can adapt in response to changes in predator densities. Here I show that this feedback between defense and predator dynamics enables stable predator coexistence without ecological niche differentiation. Instead, the mechanism driving coexistence is that prey adaptation causes defense to converge to the value where both predators have equal R* values ("fitness equalization"). This result is highly general, independent of specific model details, and applies to both rapid defense evolution and inducible defenses. It demonstrates the importance of considering long-standing ecological questions from an eco-evolutionary viewpoint, and showcases how the effects of adaptation can cascade through communities, driving diversity on higher trophic levels. These insights offer an important new perspective on coexistence theory. KW - coexistence KW - competition KW - competitive exclusion KW - defense KW - eco-evolutionary feedbacks KW - emergent facilitation KW - predator KW - prey KW - dynamics Y1 - 2020 U6 - https://doi.org/10.1002/ecy.2995 SN - 0012-9658 SN - 1939-9170 VL - 101 IS - 5 PB - Wiley CY - Hoboken ER - TY - GEN A1 - Raatz, Michael A1 - van Velzen, Ellen A1 - Gaedke, Ursula T1 - Co‐adaptation impacts the robustness of predator–prey dynamics against perturbations T2 - Postprints der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe N2 - 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. T3 - Zweitveröffentlichungen der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe - 809 KW - disturbance KW - evolutionary rescue KW - population dynamics KW - stability KW - trait adaptation Y1 - 2020 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-442489 SN - 1866-8372 IS - 809 ER - TY - JOUR A1 - van Velzen, Ellen A1 - Gaedke, Ursula T1 - Reversed predator-prey cycles are driven by the amplitude of prey oscillations JF - Ecology and evolution N2 - 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. KW - coevolution KW - ecoevolutionary dynamics KW - predator-prey dynamics KW - top-down control Y1 - 2018 U6 - https://doi.org/10.1002/ece3.4184 SN - 2045-7758 VL - 8 IS - 12 SP - 6317 EP - 6329 PB - Wiley CY - Hoboken ER - TY - JOUR A1 - Bengfort, Michael A1 - van Velzen, Ellen A1 - Gaedke, Ursula T1 - Slight phenotypic variation in predators and prey causes complex predator-prey oscillations JF - Ecological Complexity N2 - 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. KW - Predator-prey cycles KW - Phase relationships KW - Intermittent cycles KW - Adaptive traits KW - Eco-evolutionary dynamics KW - Complex dynamics Y1 - 2017 U6 - https://doi.org/10.1016/j.ecocom.2017.06.003 SN - 1476-945X SN - 1476-9840 VL - 31 SP - 115 EP - 124 PB - Elsevier CY - Amsterdam ER - TY - JOUR A1 - Yamamichi, Masato A1 - Klauschies, Toni A1 - Miner, Brooks E. A1 - van Velzen, Ellen T1 - Modelling inducible defences in predator-prey interactions BT - assumptions and dynamical consequences of three distinct approaches JF - Ecology letters N2 - Inducible defences against predation are widespread in the natural world, allowing prey to economise on the costs of defence when predation risk varies over time or is spatially structured. Through interspecific interactions, inducible defences have major impacts on ecological dynamics, particularly predator-prey stability and phase lag. Researchers have developed multiple distinct approaches, each reflecting assumptions appropriate for particular ecological communities. Yet, the impact of inducible defences on ecological dynamics can be highly sensitive to the modelling approach used, making the choice of model a critical decision that affects interpretation of the dynamical consequences of inducible defences. Here, we review three existing approaches to modelling inducible defences: Switching Function, Fitness Gradient and Optimal Trait. We assess when and how the dynamical outcomes of these approaches differ from each other, from classic predator-prey dynamics and from commonly observed eco-evolutionary dynamics with evolving, but non-inducible, prey defences. We point out that the Switching Function models tend to stabilise population dynamics, and the Fitness Gradient models should be carefully used, as the difference with evolutionary dynamics is important. We discuss advantages of each approach for applications to ecological systems with particular features, with the goal of providing guidelines for future researchers to build on. KW - Adaptive dynamics KW - fitness gradient KW - inducible defence KW - optimal trait KW - phenotypic plasticity KW - predator-prey dynamics KW - reaction norm KW - switching function Y1 - 2019 U6 - https://doi.org/10.1111/ele.13183 SN - 1461-023X SN - 1461-0248 VL - 22 IS - 2 SP - 390 EP - 404 PB - Wiley CY - Hoboken ER - TY - JOUR A1 - van Velzen, Ellen A1 - Thieser, Tamara A1 - Berendonk, Thomas U. A1 - Weitere, Markus A1 - Gaedke, Ursula T1 - Inducible defense destabilizes predator–prey dynamics BT - the importance of multiple predators JF - Oikos N2 - 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. KW - phenotypic plasticity KW - inducible defense KW - stability KW - synchronization KW - predator-prey dynamics Y1 - 2018 U6 - https://doi.org/10.1111/oik.04868 SN - 0030-1299 SN - 1600-0706 VL - 127 IS - 11 SP - 1551 EP - 1562 PB - Wiley CY - Hoboken ER - TY - JOUR A1 - Kath, Nadja Jeanette A1 - Gaedke, Ursula A1 - van Velzen, Ellen T1 - The double-edged sword of inducible defences: costs and benefits of maladaptive switching from the individual to the community level JF - Scientific Reports N2 - Phenotypic plasticity can increase individual fitness when environmental conditions change over time. Inducible defences are a striking example, allowing species to react to fluctuating predation pressure by only expressing their costly defended phenotype under high predation risk. Previous theoretical investigations have focused on how this affects predator–prey dynamics, but the impact on competitive outcomes and broader community dynamics has received less attention. Here we use a small food web model, consisting of two competing plastic autotrophic species exploited by a shared consumer, to study how the speed of inducible defences across three trade-off constellations affects autotroph coexistence, biomasses across trophic levels, and temporal variability. Contrary to the intuitive idea that faster adaptation increases autotroph fitness, we found that higher switching rates reduced individual fitness as it consistently provoked more maladaptive switching towards undefended phenotypes under high predation pressure. This had an unexpected positive impact on the consumer, increasing consumer biomass and lowering total autotroph biomass. Additionally, maladaptive switching strongly reduced autotroph coexistence through an emerging source-sink dynamic between defended and undefended phenotypes. The striking impact of maladaptive switching on species and food web dynamics indicates that this mechanism may be of more critical importance than previously recognized. Y1 - 2022 U6 - https://doi.org/10.1038/s41598-022-13895-7 SN - 2045-2322 VL - 12 SP - 1 EP - 14 PB - Springer Nature CY - London ER - TY - GEN A1 - van Velzen, Ellen A1 - Gaedke, Ursula T1 - Reversed predator BT - prey cycles are driven by the amplitude of prey oscillations N2 - 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. T3 - Zweitveröffentlichungen der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe - 433 KW - coevolution KW - ecoevolutionary dynamics KW - predator-prey dynamics KW - top-down control Y1 - 2018 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-411652 ER - TY - JOUR A1 - van Velzen, Ellen A1 - Gaedke, Ursula T1 - Reversed predator BT - prey cycles are driven by the amplitude of prey oscillations JF - Ecology and Evolution N2 - 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. KW - coevolution KW - ecoevolutionary dynamics KW - predator-prey dynamics KW - top-down control Y1 - 2018 U6 - https://doi.org/10.1002/ece3.4184 SN - 2045-7758 SP - 1 EP - 13 PB - Wiley CY - Hoboken ER - TY - JOUR A1 - Raatz, Michael A1 - van Velzen, Ellen A1 - Gaedke, Ursula T1 - Co‐adaptation impacts the robustness of predator–prey dynamics against perturbations JF - Ecology and Evolution N2 - 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. KW - disturbance KW - evolutionary rescue KW - population dynamics KW - stability KW - trait adaptation Y1 - 2019 U6 - https://doi.org/10.1002/ece3.5006 SN - 2045-7758 VL - 9 IS - 7 SP - 3823 EP - 3836 PB - John Wiley & Sons CY - Hoboken, NJ ER - TY - JOUR A1 - van Velzen, Ellen A1 - Gaedke, Ursula T1 - Disentangling eco-evolutionary dynamics of predator-prey coevolution: the case of antiphase cycles JF - Scientific reports N2 - The impact of rapid predator-prey coevolution on predator-prey dynamics remains poorly understood, as previous modelling studies have given rise to contradictory conclusions and predictions. Interpreting and reconciling these contradictions has been challenging due to the inherent complexity of model dynamics, defying mathematical analysis and mechanistic understanding. We develop a new approach here, based on the Geber method for deconstructing eco-evolutionary dynamics, for gaining such understanding. We apply this approach to a co-evolutionary predator-prey model to disentangle the processes leading to either antiphase or 1/4-lag cycles. Our analysis reveals how the predator-prey phase relationship is driven by the temporal synchronization between prey biomass and defense dynamics. We further show when and how prey biomass and trait dynamics become synchronized, resulting in antiphase cycles, allowing us to explain and reconcile previous modelling and empirical predictions. The successful application of our proposed approach provides an important step towards a comprehensive theory on eco-evolutionary feedbacks in predator-prey systems. Y1 - 2017 U6 - https://doi.org/10.1038/s41598-017-17019-4 SN - 2045-2322 VL - 7 PB - Nature Publ. Group CY - London ER - TY - JOUR A1 - Kath, Nadja Jeanette A1 - Gaedke, Ursula A1 - van Velzen, Ellen T1 - The double-edged sword of inducible defences: costs and benefits of maladaptive switching from the individual to the community level JF - Zweitveröffentlichungen der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe N2 - Phenotypic plasticity can increase individual fitness when environmental conditions change over time. Inducible defences are a striking example, allowing species to react to fluctuating predation pressure by only expressing their costly defended phenotype under high predation risk. Previous theoretical investigations have focused on how this affects predator–prey dynamics, but the impact on competitive outcomes and broader community dynamics has received less attention. Here we use a small food web model, consisting of two competing plastic autotrophic species exploited by a shared consumer, to study how the speed of inducible defences across three trade-off constellations affects autotroph coexistence, biomasses across trophic levels, and temporal variability. Contrary to the intuitive idea that faster adaptation increases autotroph fitness, we found that higher switching rates reduced individual fitness as it consistently provoked more maladaptive switching towards undefended phenotypes under high predation pressure. This had an unexpected positive impact on the consumer, increasing consumer biomass and lowering total autotroph biomass. Additionally, maladaptive switching strongly reduced autotroph coexistence through an emerging source-sink dynamic between defended and undefended phenotypes. The striking impact of maladaptive switching on species and food web dynamics indicates that this mechanism may be of more critical importance than previously recognized. T3 - Zweitveröffentlichungen der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe - 1288 Y1 - 2022 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-572006 SN - 1866-8372 IS - 1288 ER - TY - JOUR A1 - Seiler, Claudia A1 - van Velzen, Ellen A1 - Neu, Thomas R. A1 - Gaedke, Ursula A1 - Berendonk, Thomas U. A1 - Weitere, Markus T1 - Grazing resistance of bacterial biofilms: a matter of predators’ feeding trait JF - FEMS microbiology ecology N2 - Biofilm formation in bacteria is considered to be one strategy to avoid protozoan grazing. However, this assumption is largely based on experiments with suspension-feeding protozoans. Here we test the hypothesis that grazing resistance depends on both the grazers’ feeding trait and the bacterial phenotype, rather than being a general characteristic of bacterial biofilms. We combined batch experiments with mathematical modelling, considering the bacterium Pseudomonas putida and either a suspension-feeding (i.e. the ciliate Paramecium tetraurelia) or a surface-feeding grazer (i.e. the amoeba Acanthamoeba castellanii). We find that both plankton and biofilm phenotypes were consumed, when exposed to their specialised grazer, whereas the other phenotype remained grazing-resistant. This was consistently shown in two experiments (starting with either only planktonic bacteria or with additional pre-grown biofilms) and matches model predictions. In the experiments, the plankton feeder strongly stimulated the biofilm biomass. This stimulation of the resistant prey phenotype was not predicted by the model and it was not observed for the biofilm feeders, suggesting the existence of additional mechanisms that stimulate biofilm formation besides selective feeding. Overall, our results confirm our hypothesis that grazing resistance is a matter of the grazers’ trait (i.e. feeding type) rather than a biofilm-specific property. KW - protozoa KW - biofilm KW - plankton KW - predator-prey model KW - grazing defence KW - feeding trait Y1 - 2017 U6 - https://doi.org/10.1093/femsec/fix112 SN - 0168-6496 SN - 1574-6941 VL - 93 PB - Oxford Univ. Press CY - Oxford ER -