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 - Velzen, Ellen van A1 - Gaedke, Ursula A1 - Klauschies, Toni T1 - Quantifying the capacity for contemporary trait changes to drive intermittent predator-prey cycles JF - Ecological monographs : a publication of the Ecological Society of America N2 - 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. KW - eco-evolutionary feedbacks KW - ecosystem functioning KW - intraspecific trait KW - variation KW - population cycles KW - predator-prey dynamics KW - trait dynamics Y1 - 2022 U6 - https://doi.org/10.1002/ecm.1505 SN - 1557-7015 SN - 0012-9615 VL - 92 IS - 2 PB - Wiley CY - New York 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 - 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 - 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 - THES A1 - Ehrlich, Elias T1 - On the role of trade-offs in predator-prey interactions T1 - Trade-offs und ihre Bedeutung in Räuber-Beute Interaktionen N2 - Predation drives coexistence, evolution and population dynamics of species in food webs, and has strong impacts on related ecosystem functions (e.g. primary production). The effect of predation on these processes largely depends on the trade-offs between functional traits in the predator and prey community. Trade-offs between defence against predation and competitive ability, for example, allow for prey speciation and predator-mediated coexistence of prey species with different strategies (defended or competitive), which may stabilize the overall food web dynamics. While the importance of such trade-offs for coexistence is widely known, we lack an understanding and the empirical evidence of how the variety of differently shaped trade-offs at multiple trophic levels affect biodiversity, trait adaptation and biomass dynamics in food webs. Such mechanistic understanding is crucial for predictions and management decisions that aim to maintain biodiversity and the capability of communities to adapt to environmental change ensuring their persistence. In this dissertation, after a general introduction to predator-prey interactions and tradeoffs, I first focus on trade-offs in the prey between qualitatively different types of defence (e.g. camouflage or escape behaviour) and their costs. I show that these different types lead to different patterns of predator-mediated coexistence and population dynamics, by using a simple predator-prey model. In a second step, I elaborate quantitative aspects of trade-offs and demonstrates that the shape of the trade-off curve in combination with trait-fitness relationships strongly affects competition among different prey types: Either specialized species with extreme trait combinations (undefended or completely defended) coexist, or a species with an intermediate defence level dominates. The developed theory on trade-off shapes and coexistence is kept general, allowing for applications apart from defence-competitiveness trade-offs. Thirdly, I tested the theory on trade-off shapes on a long-term field data set of phytoplankton from Lake Constance. The measured concave trade-off between defence and growth governs seasonal trait changes of phytoplankton in response to an altering grazing pressure by zooplankton, and affects the maintenance of trait variation in the community. In a fourth step, I analyse the interplay of different tradeoffs at multiple trophic levels with plankton data of Lake Constance and a corresponding tritrophic food web model. The results show that the trait and biomass dynamics of the different three trophic levels are interrelated in a trophic biomass-trait cascade, leading to unintuitive patterns of trait changes that are reversed in comparison to predictions from bitrophic systems. Finally, in the general discussion, I extract main ideas on trade-offs in multitrophic systems, develop a graphical theory on trade-off-based coexistence, discuss the interplay of intra- and interspecific trade-offs, and end with a management-oriented view on the results of the dissertation, describing how food webs may respond to future global changes, given their trade-offs. N2 - Trophische Interaktionen sind von entscheidender Bedeutung für die Biodiversität in Ökosystemen und die daran gekoppelten Ökosystemfunktionen (z.B. Primärproduktion, Nährstoffkreislauf). Außerdem beeinflussen sie die Evolution und Populationsdynamiken von Arten. Die Wirkungsweise von trophischen Interaktionen auf diese Prozesse hängt dabei von den Trade-offs ab, denen Räuber und Beute z.B. auf Grund physiologischer Beschränkungen unterliegen. Als Trade-off wird die Kosten-Nutzen-Beziehung zwischen zwei oder mehr funktionellen Eigenschaften eines Organismus bezeichnet, so zum Beispiel das Einhergehen einer höheren Verteidigung gegen Fraß mit einer geringeren Konkurrenzfähigkeit um Ressourcen. Solche Trade-offs zwischen Verteidigung und Konkurrenzfähigkeit ermöglichen die Koexistenz von Beutearten mit verschiedenen Strategien (verteidigt oder konkurrenzfähig), was sich stabilisierend auf die gesamten Dynamiken im Nahrungsnetz auswirken kann. Obwohl die Annahme weit verbreitet ist, dass Trade-offs die Koexistenz von Arten fördern, mangelt es am Verständnis und an empirischen Nachweisen, wie sich die Vielzahl unterschiedlich geformter Trade-offs von Arten verschiedener trophischer Ebenen auf die Biodiversität, die Anpassung von funktionellen Eigenschaften und die Biomassedynamik in Nahrungsnetzen auswirkt. Solch ein Verständnis ist jedoch entscheidend für die Vorhersagen und Managemententscheidungen bezüglich des Erhalts von Biodiversität, die das Anpassungspotential von Artengemeinschaften an zukünftige Veränderung in der Umwelt und damit das Überdauern von Artengemeinschaften langfristig sicherstellt. Die hier vorliegende Dissertation startet mit einer kurzen Einführung in die Rolle von Räuber-Beute-Beziehungen und Trade-offs in Ökosystemen. In einem ersten Schritt, lege ich den Fokus zunächst auf Trade-offs in Beutegemeinschaften zwischen qualitativ verschiedenen Verteidigungsmechanismen (z.B. Tarnung oder Fluchtverhalten) und -kosten, und zeige anhand von einfachen Räuber-Beute Modellen, wie sich diese Mechanismen hinsichtlich ihrer Wirkungsweise auf die Koexistenz und die Populationsdynamiken von Beutearten unterscheiden. Als Zweites konzentriert sich die Dissertation dann auf quantitative Aspekte der Trade-offs. So wird aufgezeigt, wie die Form der Trade-off-Kurve bei verschiedenen Beziehungen zwischen funktionellen Eigenschaften und der Fitness den Ausgang von Konkurrenzprozessen innerhalb von Beutegemeinschaften beeinflusst. Dabei kann es in Abhängigkeit von der Form der Trade-off-Kurve entweder zu Koexistenz von spezialisierten Arten kommen (unverteidigt oder komplett verteidigt) oder aber zur Dominanz einer Art mit mittlerer Verteidigung. Der dritte Schwerpunkt dieser Arbeit liegt dann auf dem Test der Theorie zur Trade-off-Kurve und Koexistenz anhand von Langzeitfelddaten des Phytoplanktons im Bodensee. Es zeigt sich hierbei, dass der gefundene konkave Trade-off zwischen Verteidung und Wachstumsrate in Kombination mit einem sich verändernden Fraßdruck durch das Zooplankton die Anpassung von funktionellen Eigenschaften und den Erhalt von Variation dieser Eigenschaften innerhalb der Phytoplanktongemeinschaft steuert. In einem vierten Schritt, analysiere ich das Zusammenspiel von Trade-offs auf mehreren trophischen Ebenen, basierend auf Phyto- und Zooplanktondaten aus dem Bodensee und einem dafür entwickelten tritrophischen Nahrungsnetzmodell. Die Ergebnisse zeigen, dass die Dynamiken der funktionellen Eigenschaften und Biomassen durch eine Kaskade über die drei trophischen Ebenen hinweg gekoppelt sind, die zu unintuitiven Mustern in den Anpassungen der funktionellen Eigenschaften zwischen den Ebenen führt. In der generellen Diskussion bringe ich \textit{abschließend} die Ideen zur Wirkung von Trade-offs in multitrophischen System in einen breiteren Kontext. Zudem entwickle ich eine generelle graphische Theorie zur Trade-off basierten Koexistenz in Abhängigkeit von der Fitnesslandschaft, diskutiere das mögliche Zusammenspiel von intra- und interspezifischen Trade-offs, und gebe schlussendlich einen Management-orientierten Einblick in die Relevanz der Ergebnisse dieser Dissertation für das Verhalten von Nahrungsnetzen im Zuge des Globalen Wandels unter der Wirkung von Trade-offs. KW - trade-offs between functional traits KW - predator-prey dynamics KW - food web KW - coexistence KW - trait variation KW - theoretical ecology KW - phytoplankton and zooplankton KW - Trade-offs zwischen funktionellen Eigenschaften KW - Räuber-Beute Dynamiken KW - Nahrungsnetz KW - Koexistenz KW - Variation in funktionellen Eigenschaften KW - theoretische Ökologie KW - Phytoplankton und Zooplankton Y1 - 2019 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-430631 ER -