@article{EhrlichBecksGaedke2017, author = {Ehrlich, Elias and Becks, Lutz and Gaedke, Ursula}, title = {Trait-fitness relationships determine how trade-off shapes affect species coexistence}, series = {Ecology : a publication of the Ecological Society of America}, volume = {98}, journal = {Ecology : a publication of the Ecological Society of America}, publisher = {Wiley}, address = {Hoboken}, issn = {0012-9658}, doi = {10.1002/ecy.2047}, pages = {3188 -- 3198}, year = {2017}, abstract = {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.}, 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{vanVelzen2020, author = {van Velzen, Ellen}, title = {Predator coexistence through emergent fitness equalization}, series = {Ecology : a publication of the Ecological Society of America}, volume = {101}, journal = {Ecology : a publication of the Ecological Society of America}, number = {5}, publisher = {Wiley}, address = {Hoboken}, issn = {0012-9658}, doi = {10.1002/ecy.2995}, pages = {10}, year = {2020}, abstract = {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.}, 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}, address = {Hoboken}, 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{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{TanentzapLeeSchulz2013, author = {Tanentzap, Andrew J. and Lee, William G. and Schulz, Katharina A. C.}, title = {Niches drive peaked and positive relationships between diversity and disturbance in natural ecosystems}, series = {Ecosphere : the magazine of the International Ecology University}, volume = {4}, journal = {Ecosphere : the magazine of the International Ecology University}, number = {11}, publisher = {Wiley}, address = {Washington}, issn = {2150-8925}, doi = {10.1890/ES13-00102.1}, pages = {28}, year = {2013}, abstract = {A unified understanding of the relationship between disturbance and biodiversity is needed to predict biotic responses to global change. Recent advances have identified the need to deconstruct traditional models of disturbance into intensity and frequency to reconcile empirical studies that appear to generate contradictory associations between species diversity and disturbance. We integrate results from theoretical simulation modelling, field-based surveys of 5176 vegetation plots from 48 transects across 6 sites, and experimental pot-based manipulations of flooding to identify how disturbance drives species diversity within ephemeral wetlands in South Island, New Zealand. We find empirical, hump-shaped and positive relationships between species diversity and both disturbance intensity and frequency, mirroring patterns from a simulation model in which species differed in their demographic responses to disturbance. More generally, our simulations show that the relationships between diversity and disturbance shift from positive to hump-shaped to negative as species that are favored at low disturbance because of their resistance strategies, defined by low mortality and recruitment, decline within communities relative to resilient species. Resilient species with higher mortality and recruitment rates are instead favored as disturbance intensity and frequency intensify. Our theoretical findings suggest that sites must also have a third group of unique species with intermediate resilience and resistance. Analyses of community composition along our disturbance gradients support this prediction, emphasizing that shifts in community-level resistance and resilience drive empirical associations between diversity and disturbance. Overall, terrestrial plants may be unable to resist intense and frequent flooding, even with specialized traits. Only fast-growing species with high regeneration from seed may respond once flooding subsides and dominate community composition in these situations, especially on nutrient-rich soils. However, different strategies can co-occur at intermediate disturbance, ultimately increasing species richness. As disturbances become more pervasive globally, our results suggest that differences in the niches of species, rather than demographic stochasticity, drive biodiversity patterns. These niche-based processes may especially prevail, without accompanying losses in species richness, where sites are initially dominated by resistant taxa or life history strategies that balance resistance and resilience.}, language = {en} } @article{SchirmerHoffmannEccardetal.2020, author = {Schirmer, Annika and Hoffmann, Julia and Eccard, Jana and Dammhahn, Melanie}, title = {My niche}, series = {Proceedings of the Royal Society of London : B, Biological sciences}, volume = {287}, journal = {Proceedings of the Royal Society of London : B, Biological sciences}, number = {1918}, publisher = {Royal Society}, address = {London}, issn = {0962-8452}, doi = {10.1098/rspb.2019.2211}, pages = {9}, year = {2020}, abstract = {Intraspecific trait variation is an important determinant of fundamental ecological interactions. Many of these interactions are mediated by behaviour. Therefore, interindividual differences in behaviour should contribute to individual niche specialization. Comparable with variation in morphological traits, behavioural differentiation between individuals should limit similarity among competitors and thus act as a mechanism maintaining within-species variation in ecological niches and facilitating species coexistence. Here, we aimed to test whether interindividual differences in boldness covary with spatial interactions within and between two ecologically similar, co-occurring rodent species (Myodes glareolus, Apodemus agrarius). In five subpopulations in northeast Germany, we quantified individual differences in boldness via repeated standardized tests and spatial interaction patterns via capture-mark- recapture (n = 126) and automated VHF telemetry (n = 36). We found that boldness varied with space use in both species. Individuals of the same population occupied different spatial niches, which resulted in non-random patterns of within- and between-species spatial interactions. Behavioural types mainly differed in the relative importance of intra- versus interspecific competition. Within-species variation along this competition gradient could contribute to maintaining individual niche specialization. Moreover, behavioural differentiation between individuals limits similarity among competitors, which might facilitate the coexistence of functionally equivalent species and, thus, affect community dynamics and local biodiversity.}, language = {en} } @article{MillesDammhahnGrimm2020, author = {Milles, Alexander and Dammhahn, Melanie and Grimm, Volker}, title = {Intraspecific trait variation in personality-related movement behavior promotes coexistence}, series = {Oikos}, volume = {129}, journal = {Oikos}, number = {10}, publisher = {Wiley}, address = {Hoboken}, issn = {0030-1299}, doi = {10.1111/oik.07431}, pages = {1441 -- 1454}, year = {2020}, abstract = {Movement behavior is an essential element of fundamental ecological processes such as competition and predation. Although intraspecific trait variation (ITV) in movement behaviors is pervasive, its consequences for ecological community dynamics are still not fully understood. Using a newly developed individual-based model, we analyzed how given and constant ITVs in foraging movement affect differences in foraging efficiencies between species competing for common resources under various resource distributions. Further, we analyzed how the effect of ITV on emerging differences in competitive abilities ultimately affects species coexistence. The model is generic but mimics observed patterns of among-individual covariation between personality, movement and space use in ground-dwelling rodents. Interacting species differed in their mean behavioral types along a slow-fast continuum, integrating consistent individual variation in average behavioral expression and responsiveness (i.e. behavioral reaction norms). We found that ITV reduced interspecific differences in competitive abilities by 5-35\% and thereby promoted coexistence via an equalizing mechanism. The emergent relationships between behavioral types and foraging efficiency are characteristic for specific environmental contexts of resource distribution and population density. As these relationships are asymmetric, species that were either 'too fast' or 'too slow' benefited differently from ITV. Thus, ITV in movement behavior has consequences for species coexistence but to predict its effect in a given system requires intimate knowledge on how variation in movement traits relates to fitness components along an environmental gradient.}, language = {en} } @article{SoliveresMaestreUlrichetal.2015, author = {Soliveres, Santiago and Maestre, Fernando T. and Ulrich, Werner and Manning, Peter and Boch, Steffen and Bowker, Matthew A. and Prati, Daniel and Delgado-Baquerizo, Manuel and Quero, Jose L. and Sch{\"o}ning, Ingo and Gallardo, Antonio and Weisser, Wolfgang W. and M{\"u}ller, J{\"o}rg and Socher, Stephanie A. and Garcia-Gomez, Miguel and Ochoa, Victoria and Schulze, Ernst-Detlef and Fischer, Markus and Allan, Eric}, title = {Intransitive competition is widespread in plant communities and maintains their species richness}, series = {Ecology letters}, volume = {18}, journal = {Ecology letters}, number = {8}, publisher = {Wiley-Blackwell}, address = {Hoboken}, issn = {1461-023X}, doi = {10.1111/ele.12456}, pages = {790 -- 798}, year = {2015}, abstract = {Intransitive competition networks, those in which there is no single best competitor, may ensure species coexistence. However, their frequency and importance in maintaining diversity in real-world ecosystems remain unclear. We used two large data sets from drylands and agricultural grasslands to assess: (1) the generality of intransitive competition, (2) intransitivity-richness relationships and (3) effects of two major drivers of biodiversity loss (aridity and land-use intensification) on intransitivity and species richness. Intransitive competition occurred in >65\% of sites and was associated with higher species richness. Intransitivity increased with aridity, partly buffering its negative effects on diversity, but was decreased by intensive land use, enhancing its negative effects on diversity. These contrasting responses likely arise because intransitivity is promoted by temporal heterogeneity, which is enhanced by aridity but may decline with land-use intensity. We show that intransitivity is widespread in nature and increases diversity, but it can be lost with environmental homogenisation.}, language = {en} } @article{JeltschGrimmReegetal.2019, author = {Jeltsch, Florian and Grimm, Volker and Reeg, Jette and Schl{\"a}gel, Ulrike E.}, title = {Give chance a chance}, series = {Ecosphere}, volume = {10}, journal = {Ecosphere}, number = {5}, publisher = {ESA}, address = {Ithaca, NY}, issn = {2150-8925}, doi = {10.1002/ecs2.2700}, pages = {19}, year = {2019}, abstract = {A large part of biodiversity theory is driven by the basic question of what allows species to coexist in spite of a confined number of niches. A substantial theoretical background to this question is provided by modern coexistence theory (MCT), which rests on mathematical approaches of invasion analysis to categorize underlying mechanisms into factors that reduce either niche overlap (stabilizing mechanisms) or the average fitness differences of species (equalizing mechanisms). While MCT has inspired biodiversity theory in the search for these underlying mechanisms, we feel that the strong focus on coexistence causes a bias toward the most abundant species and neglects the plethora of species that are less abundant and often show high local turnover. Given the more stochastic nature of their occurrence, we advocate a complementary cross-level approach that links individuals, small populations, and communities and explicitly takes into account (1) a more complete inclusion of environmental and demographic stochasticity affecting small populations, (2) intraspecific trait variation and behavioral plasticity, and (3) local heterogeneities, interactions, and feedbacks. Focusing on mechanisms that drive the temporary coviability of species rather than infinite coexistence, we suggest a new approach that could be dubbed coviability analysis (CVA). From a modeling perspective, CVA builds on the merged approaches of individual-based modeling and population viability analysis but extends them to the community level. From an empirical viewpoint, CVA calls for a stronger integration of spatiotemporal data on variability and noise, changing drivers, and interactions at the level of individuals. The resulting large volumes of data from multiple sources could be strongly supported by novel techniques tailored to the discovery of complex patterns in high-dimensional data. By complementing MCT through a stronger focus on the coviability of less common species, this approach can help make modern biodiversity theory more comprehensive, predictive, and relevant for applications.}, language = {en} }