TY - JOUR A1 - Ehrlich, Elias A1 - Becks, Lutz A1 - Gaedke, Ursula T1 - Trait-fitness relationships determine how trade-off shapes affect species coexistence JF - Ecology : a publication of the Ecological Society of America N2 - Trade-offs between functional traits are ubiquitous in nature and can promote species coexistence depending on their shape. Classic theory predicts that convex trade-offs facilitate coexistence of specialized species with extreme trait values (extreme species) while concave trade-offs promote species with intermediate trait values (intermediate species). We show here that this prediction becomes insufficient when the traits translate non-linearly into fitness which frequently occurs in nature, e.g., an increasing length of spines reduces grazing losses only up to a certain threshold resulting in a saturating or sigmoid trait-fitness function. We present a novel, general approach to evaluate the effect of different trade-off shapes on species coexistence. We compare the trade-off curve to the invasion boundary of an intermediate species invading the two extreme species. At this boundary, the invasion fitness is zero. Thus, it separates trait combinations where invasion is or is not possible. The invasion boundary is calculated based on measurable trait-fitness relationships. If at least one of these relationships is not linear, the invasion boundary becomes non-linear, implying that convex and concave trade-offs not necessarily lead to different coexistence patterns. Therefore, we suggest a new ecological classification of trade-offs into extreme-favoring and intermediate-favoring which differs from a purely mathematical description of their shape. We apply our approach to a well-established model of an empirical predator-prey system with competing prey types facing a trade-off between edibility and half-saturation constant for nutrient uptake. We show that the survival of the intermediate prey depends on the convexity of the trade-off. Overall, our approach provides a general tool to make a priori predictions on the outcome of competition among species facing a common trade-off in dependence of the shape of the trade-off and the shape of the trait-fitness relationships. KW - coexistence KW - competition KW - fitness KW - functional traits KW - invasion boundary KW - neutrality KW - predator-prey model KW - shape KW - trade-offs Y1 - 2017 U6 - https://doi.org/10.1002/ecy.2047 SN - 0012-9658 SN - 1939-9170 VL - 98 SP - 3188 EP - 3198 PB - Wiley CY - Hoboken ER - TY - JOUR A1 - Ehrlich, Elias A1 - Gaedke, Ursula T1 - Not attackable or not crackable BT - How pre- and post-attack defenses with different competition costs affect prey coexistence and population dynamics JF - Ecology and evolution N2 - 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. KW - coexistence KW - competition-defense trade-off KW - defense against predation KW - functional response KW - indirect facilitation KW - predator-prey cycles Y1 - 2018 U6 - https://doi.org/10.1002/ece3.4145 SN - 2045-7758 VL - 8 IS - 13 SP - 6625 EP - 6637 PB - Wiley CY - Hoboken ER - TY - JOUR A1 - Ehrlich, Elias A1 - Gaedke, Ursula T1 - Not attackable or not crackable BT - How pre-and post-attack defenses with different competition costs affect prey coexistence and population dynamics JF - Ecology and Evolution N2 - 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. KW - coexistence KW - competition–defense trade‐off KW - defense against predation KW - functional response KW - indirect facilitation KW - predator–prey cycles Y1 - 2018 U6 - https://doi.org/10.1002/ece3.4145 SN - 2045-7758 VL - 8 IS - 13 SP - 6625 EP - 6637 PB - Wiley ER - TY - GEN A1 - Ehrlich, Elias A1 - Gaedke, Ursula T1 - Not attackable or not crackable BT - How pre-and post-attack defenses with different competition costs affect prey coexistence and population dynamics T2 - Ecology and Evolution N2 - 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. T3 - Zweitveröffentlichungen der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe - 466 KW - coexistence KW - competition–defense trade‐off KW - defense against predation KW - functional response KW - indirect facilitation KW - predator–prey cycles Y1 - 2018 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-417391 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 - TY - JOUR A1 - Ehrlich, Elias A1 - Gaedke, Ursula T1 - Coupled changes in traits and biomasses cascading through a tritrophic plankton food web JF - Limnology and oceanography N2 - 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. KW - community ecology KW - cyclops vicinus KW - dynamics KW - functional traits KW - lake KW - life-cycle KW - natural rotifer KW - phytoplankton KW - trophic cascades KW - zooplankton Y1 - 2020 U6 - https://doi.org/10.1002/lno.11466 SN - 0024-3590 SN - 1939-5590 VL - 65 IS - 10 SP - 2502 EP - 2514 PB - Wiley CY - Hoboken ER - TY - JOUR A1 - Ehrlich, Elias A1 - Kath, Nadja Jeanette A1 - Gaedke, Ursula T1 - The shape of a defense-growth trade-off governs seasonal trait dynamics in natural phytoplankton JF - The ISME journal N2 - 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. KW - coexistence KW - community ecology KW - diversity KW - evolution KW - fitness KW - functional traits KW - lake KW - maintenance KW - mechanisms KW - plankton Y1 - 2020 U6 - https://doi.org/10.1038/s41396-020-0619-1 SN - 1751-7362 SN - 1751-7370 VL - 14 IS - 6 SP - 1451 EP - 1462 PB - Nature Publishing Group CY - London ER - TY - GEN A1 - Ehrlich, Elias A1 - Kath, Nadja Jeanette A1 - Gaedke, Ursula T1 - The shape of a defense-growth trade-off governs seasonal trait dynamics in natural phytoplankton T2 - Zweitveröffentlichungen der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe N2 - 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. T3 - Zweitveröffentlichungen der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe - 1390 KW - functional traits KW - community ecology KW - evolution KW - lake KW - mechanisms KW - diversity KW - plankton KW - fitness KW - maintenance KW - coexistence Y1 - 2020 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-513956 SN - 1866-8372 IS - 6 ER - TY - JOUR A1 - Ehrlich, Elias A1 - Thygesen, Uffe Høgsbro A1 - Kiørboe, Thomas T1 - Evolution of toxins as a public good in phytoplankton JF - Proceedings of the Royal Society of London : B, Biological sciences N2 - Toxic phytoplankton blooms have increased in many waterbodies worldwide with well-known negative impacts on human health, fisheries and ecosystems. However, why and how phytoplankton evolved toxin production is still a puzzling question, given that the producer that pays the costs often shares the benefit with other competing algae and thus provides toxins as a 'public good' (e.g. damaging a common competitor or predator). Furthermore, blooming phytoplankton species often show a high intraspecific variation in toxicity and we lack an understanding of what drives the dynamics of coexisting toxic and non-toxic genotypes. Here, by using an individual-based two-dimensional model, we show that small-scale patchiness of phytoplankton strains caused by demography can explain toxin evolution in phytoplankton with low motility and the maintenance of genetic diversity within their blooms. This patchiness vanishes for phytoplankton with high diffusive motility, suggesting different evolutionary pathways for different phytoplankton groups. In conclusion, our study reveals that small-scale spatial heterogeneity, generated by cell division and counteracted by diffusive cell motility and turbulence, can crucially affect toxin evolution and eco-evolutionary dynamics in toxic phytoplankton species. This contributes to a better understanding of conditions favouring toxin production and the evolution of public goods in asexually reproducing organisms in general. KW - toxic algal blooms KW - evolution of cooperation KW - coexistence KW - patchiness in KW - phytoplankton KW - eco-evolutionary feedback KW - spatial pattern formation Y1 - 2022 U6 - https://doi.org/10.1098/rspb.2022.0393 SN - 0962-8452 SN - 1471-2954 VL - 289 IS - 1977 PB - Royal Society CY - London ER -