@article{KlauschiesCoutinhoGaedke2018, author = {Klauschies, Toni and Coutinho, Renato Mendes and Gaedke, Ursula}, title = {A beta distribution-based moment closure enhances the reliability of trait-based aggregate models for natural populations and communities}, series = {Ecological modelling : international journal on ecological modelling and engineering and systems ecolog}, volume = {381}, journal = {Ecological modelling : international journal on ecological modelling and engineering and systems ecolog}, publisher = {Elsevier}, address = {Amsterdam}, issn = {0304-3800}, doi = {10.1016/j.ecolmodel.2018.02.001}, pages = {46 -- 77}, year = {2018}, abstract = {Ecological communities are complex adaptive systems that exhibit remarkable feedbacks between their biomass and trait dynamics. Trait-based aggregate models cope with this complexity by focusing on the temporal development of the community's aggregate properties such as its total biomass, mean trait and trait variance. They are based on particular assumptions about the shape of the underlying trait distribution, which is commonly assumed to be normal. However, ecologically important traits are usually restricted to a finite range, and empirical trait distributions are often skewed or multimodal. As a result, normal distribution-based aggregate models may fail to adequately represent the biomass and trait dynamics of natural communities. We resolve this mismatch by developing a new moment closure approach assuming the trait values to be beta-distributed. We show that the beta distribution captures important shape properties of both observed and simulated trait distributions, which cannot be captured by a Gaussian. We further demonstrate that a beta distribution-based moment closure can strongly enhance the reliability of trait-based aggregate models. We compare the biomass, mean trait and variance dynamics of a full trait distribution (FD) model to the ones of beta (BA) and normal (NA) distribution-based aggregate models, under different selection regimes. This way, we demonstrate under which general conditions (stabilizing, fluctuating or disruptive selection) different aggregate models are reliable tools. All three models predicted very similar biomass and trait dynamics under stabilizing selection yielding unimodal trait distributions with small standing trait variation. We also obtained an almost perfect match between the results of the FD and BA models under fluctuating selection, promoting skewed trait distributions and ongoing oscillations in the biomass and trait dynamics. In contrast, the NA model showed unrealistic trait dynamics and exhibited different alternative stable states, and thus a high sensitivity to initial conditions under fluctuating selection. Under disruptive selection, both aggregate models failed to reproduce the results of the FD model with the mean trait values remaining within their ecologically feasible ranges in the BA model but not in the NA model. Overall, a beta distribution-based moment closure strongly improved the realism of trait-based aggregate models.}, language = {en} } @article{GaedkeKlauschies2017, author = {Gaedke, Ursula and Klauschies, Toni}, title = {Analyzing the shape of observed trait distributions enables a data-based moment closure of aggregate models}, series = {Limnology and Oceanography: Methods}, volume = {15}, journal = {Limnology and Oceanography: Methods}, publisher = {Wiley}, address = {Hoboken}, issn = {1541-5856}, doi = {10.1002/lom3.10218}, pages = {979 -- 994}, year = {2017}, abstract = {The shape of trait distributions may inform about the selective forces that structure ecological communities. Here, we present a new moment-based approach to classify the shape of observed biomass-weighted trait distributions into normal, peaked, skewed, or bimodal that facilitates spatio-temporal and cross-system comparisons. Our observed phytoplankton trait distributions exhibited substantial variance and were mostly skewed or bimodal rather than normal. Additionally, mean, variance, skewness und kurtosis were strongly correlated. This is in conflict with trait-based aggregate models that often assume normally distributed trait values and small variances. Given these discrepancies between our data and general model assumptions we used the observed trait distributions to test how well different aggregate models with first- or second-order approximations and different types of moment closure predict the biomass, mean trait, and trait variance dynamics using weakly or moderately nonlinear fitness functions. For weakly non-linear fitness functions aggregate models with a second-order approximation and a data-based moment closure that relied on the observed correlations between skewness and mean, and kurtosis and variance predicted biomass and often also mean trait changes fairly well and better than models with first-order approximations or a normal-based moment closure. In contrast, none of the models reflected the changes of the trait variances reliably. Aggregate model performance was often also poor for moderately nonlinear fitness functions. This questions a general applicability of the normal-based approach, in particular for predicting variance dynamics determining the speed of trait changes and maintenance of biodiversity. We evaluate in detail how and why better approximations can be obtained.}, language = {en} } @article{CoutinhoKlauschiesGaedke2016, author = {Coutinho, Renato Mendes and Klauschies, Toni and Gaedke, Ursula}, title = {Bimodal trait distributions with large variances question the reliability of trait-based aggregate models}, series = {Theoretical ecology}, volume = {9}, journal = {Theoretical ecology}, publisher = {Springer}, address = {Heidelberg}, issn = {1874-1738}, doi = {10.1007/s12080-016-0297-9}, pages = {389 -- 408}, year = {2016}, abstract = {Functionally diverse communities can adjust their species composition to altered environmental conditions, which may influence food web dynamics. Trait-based aggregate models cope with this complexity by ignoring details about species identities and focusing on their functional characteristics (traits). They describe the temporal changes of the aggregate properties of entire communities, including their total biomasses, mean trait values, and trait variances. The applicability of aggregate models depends on the validity of their underlying assumptions that trait distributions are normal and exhibit small variances. We investigated to what extent this can be expected to work by comparing an innovative model that accounts for the full trait distributions of predator and prey communities to a corresponding aggregate model. We used a food web structure with well-established trade-offs among traits promoting mutual adjustments between prey edibility and predator selectivity in response to selection. We altered the shape of the trade-offs to compare the outcome of the two models under different selection regimes, leading to trait distributions increasingly deviating from normality. Their biomass and trait dynamics agreed very well for stabilizing selection and reasonably well for directional selection, under which different trait values are favored at different times. However, for disruptive selection, the results of the aggregate model strongly deviated from the full trait distribution model that showed bimodal trait distributions with large variances. Hence, the outcome of aggregate models is reliable under ideal conditions but has to be questioned when confronted with more complex selection regimes and trait distributions, which are commonly observed in nature.}, language = {en} } @article{KlauschiesBauerAberleMalzahnetal.2012, author = {Klauschies, Toni and Bauer, Barbara and Aberle-Malzahn, Nicole and Sommer, Ulrich and Gaedke, Ursula}, title = {Climate change effects on phytoplankton depend on cell size and food web structure}, series = {Marine biology : international journal on life in oceans and coastal waters}, volume = {159}, journal = {Marine biology : international journal on life in oceans and coastal waters}, number = {11}, publisher = {Springer}, address = {New York}, issn = {0025-3162}, doi = {10.1007/s00227-012-1904-y}, pages = {2455 -- 2478}, year = {2012}, abstract = {We investigated the effects of warming on a natural phytoplankton community from the Baltic Sea, based on six mesocosm experiments conducted 2005-2009. We focused on differences in the dynamics of three phytoplankton size groups which are grazed to a variable extent by different zooplankton groups. While small-sized algae were mostly grazer-controlled, light and nutrient availability largely determined the growth of medium- and large-sized algae. Thus, the latter groups dominated at increased light levels. Warming increased mesozooplankton grazing on medium-sized algae, reducing their biomass. The biomass of small-sized algae was not affected by temperature, probably due to an interplay between indirect effects spreading through the food web. Thus, under the higher temperature and lower light levels anticipated for the next decades in the southern Baltic Sea, a higher share of smaller phytoplankton is expected. We conclude that considering the size structure of the phytoplankton community strongly improves the reliability of projections of climate change effects.}, language = {en} } @article{BauerVosKlauschiesetal.2014, author = {Bauer, Barbara and Vos, Matthijs and Klauschies, Toni and Gaedke, Ursula}, title = {Diversity, functional similarity, and top-down control drive synchronization and the reliability of ecosystem function}, series = {The American naturalist : a bi-monthly journal devoted to the advancement and correlation of the biological sciences}, volume = {183}, journal = {The American naturalist : a bi-monthly journal devoted to the advancement and correlation of the biological sciences}, number = {3}, publisher = {Univ. of Chicago Press}, address = {Chicago}, issn = {0003-0147}, doi = {10.1086/674906}, pages = {394 -- 409}, year = {2014}, abstract = {The concept that diversity promotes reliability of ecosystem function depends on the pattern that community-level biomass shows lower temporal variability than species-level biomasses. However, this pattern is not universal, as it relies on compensatory or independent species dynamics. When in contrast within--trophic level synchronization occurs, variability of community biomass will approach population-level variability. Current knowledge fails to integrate how species richness, functional distance between species, and the relative importance of predation and competition combine to drive synchronization at different trophic levels. Here we clarify these mechanisms. Intense competition promotes compensatory dynamics in prey, but predators may at the same time increasingly synchronize, under increasing species richness and functional similarity. In contrast, predators and prey both show perfect synchronization under strong top-down control, which is promoted by a combination of low functional distance and high net growth potential of predators. Under such conditions, community-level biomass variability peaks, with major negative consequences for reliability of ecosystem function.}, language = {en} } @article{FloederYongKlauschiesetal.2021, author = {Fl{\"o}der, Sabine and Yong, Joanne and Klauschies, Toni and Gaedke, Ursula and Poprick, Tobias and Brinkhoff, Thorsten and Moorthi, Stefanie}, title = {Intraspecific trait variation alters the outcome of competition in freshwater ciliates}, series = {Ecology and evolution}, volume = {11}, journal = {Ecology and evolution}, number = {15}, publisher = {Wiley}, address = {Hoboken}, issn = {2045-7758}, doi = {10.1002/ece3.7828}, pages = {10225 -- 10243}, year = {2021}, abstract = {Trait variation among heterospecific and conspecific organisms may substantially affect community and food web dynamics. While the relevance of competition and feeding traits have been widely studied for different consumer species, studies on intraspecific differences are more scarce, partly owing to difficulties in distinguishing different clones of the same species. Here, we investigate how intraspecific trait variation affects the competition between the freshwater ciliates Euplotes octocarinatus and Coleps hirtus in a nitrogen-limited chemostat system. The ciliates competed for the microalgae Cryptomonas sp. (Cry) and Navicula pelliculosa (Nav), and the bacteria present in the cultures over a period of 33 days. We used monoclonal Euplotes and three different Coleps clones (Col 1, Col 2, and Col 3) in the experiment that could be distinguished by a newly developed rDNA-based molecular assay based on the internal transcribed spacer (ITS) regions. While Euplotes feeds on Cry and on bacteria, the Coleps clones cannot survive on bacteria alone but feed on both Cry and Nav with clone-specific rates. Experimental treatments comprised two-species mixtures of Euplotes and one or all of the three different Coleps clones, respectively. We found intraspecific variation in the traits "selectivity" and "maximum ingestion rate" for the different algae to significantly affect the competitive outcome between the two ciliate species. As Nav quickly escaped top-down control and likely reached a state of low food quality, ciliate competition was strongly determined by the preference of different Coleps clones for Cry as opposed to feeding on Nav. In addition, the ability of Euplotes to use bacteria as an alternative food source strengthened its persistence once Cry was depleted. Hence, trait variation at both trophic levels codetermined the population dynamics and the outcome of species competition.}, language = {en} } @misc{YamamichiKlauschiesMineretal.2019, author = {Yamamichi, Masato and Klauschies, Toni and Miner, Brooks E. and van Velzen, Ellen}, title = {Modelling inducible defences in predator-prey interactions}, series = {Ecology letters}, volume = {22}, journal = {Ecology letters}, number = {2}, publisher = {Wiley}, address = {Hoboken}, issn = {1461-023X}, doi = {10.1111/ele.13183}, pages = {390 -- 404}, year = {2019}, abstract = {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.}, language = {en} } @article{VelzenGaedkeKlauschies2022, author = {Velzen, Ellen van and Gaedke, Ursula and Klauschies, Toni}, title = {Quantifying the capacity for contemporary trait changes to drive intermittent predator-prey cycles}, series = {Ecological monographs : a publication of the Ecological Society of America}, volume = {92}, journal = {Ecological monographs : a publication of the Ecological Society of America}, number = {2}, publisher = {Wiley}, address = {New York}, issn = {1557-7015}, doi = {10.1002/ecm.1505}, pages = {29}, year = {2022}, abstract = {A large and growing body of theory has demonstrated how the presence of trait variation in prey or predator populations may affect the amplitude and phase of predator-prey cycles. Less attention has been given to so-called intermittent cycles, in which predator-prey oscillations recurrently disappear and re-appear, despite such dynamics being observed in empirical systems and modeling studies. A comprehensive understanding of the conditions under which trait changes may drive intermittent predator-prey dynamics, as well as their potential ecological implications, is therefore missing. Here we provide a first systematic analysis of the eco-evolutionary conditions that may give rise to intermittent predator-prey cycles, investigating 16 models that incorporate different types of trait variation within prey, predators, or both. Our results show that intermittent dynamics often arise through predator-prey coevolution, but only very rarely when only one trophic level can adapt. Additionally, the frequency of intermittent cycles depends on the source of trait variation (genetic variation or phenotypic plasticity) and the genetic architecture (Mendelian or quantitative traits), with intermittency occurring most commonly through Mendelian evolution, and very rarely through phenotypic plasticity. Further analysis identified three necessary conditions for when trait variation can drive intermittent cycles. First, the intrinsic stability of the predator-prey system must depend on the traits of prey, predators, or both. Second, there must be a mechanism causing the recurrent alternation between stable and unstable states, leading to a "trait" cycle superimposed on the population dynamics. Finally, these trait dynamics must be significantly slower than the predator-prey cycles. We show how these conditions explain all the abovementioned patterns. We further show an important unexpected consequence of these necessary conditions: they are most easily met when intraspecific trait variation is at high risk of being lost. As trait diversity is positively associated with ecosystem functioning, this can have potentially severe negative consequences. This novel result highlights the importance of identifying and understanding intermittent cycles in theoretical studies and natural systems. The new approach for detecting and quantifying intermittency we develop here will be instrumental in enabling future study.}, language = {en} } @phdthesis{Klauschies2016, author = {Klauschies, Toni}, title = {Revealing causes and consequences of functional diversity using trait-based models}, school = {Universit{\"a}t Potsdam}, pages = {231}, year = {2016}, language = {en} } @article{GuillHuelsemannKlauschies2021, author = {Guill, Christian and H{\"u}lsemann, Janne and Klauschies, Toni}, title = {Self-organised pattern formation increases local diversity in metacommunities}, series = {Ecology letters}, volume = {24}, journal = {Ecology letters}, number = {12}, publisher = {Wiley-Blackwell}, address = {Oxford}, issn = {1461-023X}, doi = {10.1111/ele.13880}, pages = {2624 -- 2634}, year = {2021}, abstract = {Self-organised formation of spatial patterns is known from a variety of different ecosystems, yet little is known about how these patterns affect the diversity of communities. Here, we use a food chain model in which autotroph diversity is described by a continuous distribution of a trait that affects both growth and defence against heterotrophs. On isolated patches, diversity is always lost over time due to stabilising selection, and the local communities settle on one of two alternative stable community states that are characterised by a dominance of either defended or undefended species. In a metacommunity context, dispersal can destabilise these states and complex spatio-temporal patterns in the species' abundances emerge. The resulting biomass-trait feedback increases local diversity by an order of magnitude compared to scenarios without self-organised pattern formation, thereby maintaining the ability of communities to adapt to potential future changes in biotic or abiotic environmental conditions.}, language = {en} }