TY - JOUR A1 - Ceulemans, Ruben A1 - Guill, Christian A1 - Gaedke, Ursula T1 - Top predators govern multitrophic diversity effects in tritrophic food webs JF - Ecology : a publication of the Ecological Society of America N2 - It is well known that functional diversity strongly affects ecosystem functioning. However, even in rather simple model communities consisting of only two or, at best, three trophic levels, the relationship between multitrophic functional diversity and ecosystem functioning appears difficult to generalize, because of its high contextuality. In this study, we considered several differently structured tritrophic food webs, in which the amount of functional diversity was varied independently on each trophic level. To achieve generalizable results, largely independent of parametrization, we examined the outcomes of 128,000 parameter combinations sampled from ecologically plausible intervals, with each tested for 200 randomly sampled initial conditions. Analysis of our data was done by training a random forest model. This method enables the identification of complex patterns in the data through partial dependence graphs, and the comparison of the relative influence of model parameters, including the degree of diversity, on food-web properties. We found that bottom-up and top-down effects cascade simultaneously throughout the food web, intimately linking the effects of functional diversity of any trophic level to the amount of diversity of other trophic levels, which may explain the difficulty in unifying results from previous studies. Strikingly, only with high diversity throughout the whole food web, different interactions synergize to ensure efficient exploitation of the available nutrients and efficient biomass transfer to higher trophic levels, ultimately leading to a high biomass and production on the top level. The temporal variation of biomass showed a more complex pattern with increasing multitrophic diversity: while the system initially became less variable, eventually the temporal variation rose again because of the increasingly complex dynamical patterns. Importantly, top predator diversity and food-web parameters affecting the top trophic level were of highest importance to determine the biomass and temporal variability of any trophic level. Overall, our study reveals that the mechanisms by which diversity influences ecosystem functioning are affected by every part of the food web, hampering the extrapolation of insights from simple monotrophic or bitrophic systems to complex natural food webs. KW - food-web efficiency KW - functional diversity KW - machine learning KW - nutrient KW - exploitation KW - production KW - random forest KW - temporal variability KW - top KW - predator KW - trait diversity Y1 - 2021 U6 - https://doi.org/10.1002/ecy.3379 SN - 0012-9658 SN - 1939-9170 VL - 102 IS - 7 PB - Wiley CY - Hoboken ER - TY - JOUR A1 - Ceulemans, Ruben A1 - Gaedke, Ursula A1 - Klauschies, Toni A1 - Guill, Christian T1 - The effects of functional diversity on biomass production, variability, and resilience of ecosystem functions in a tritrophic system JF - Scientific Reports N2 - Diverse communities can adjust their trait composition to altered environmental conditions, which may strongly influence their dynamics. Previous studies of trait-based models mainly considered only one or two trophic levels, whereas most natural system are at least tritrophic. Therefore, we investigated how the addition of trait variation to each trophic level influences population and community dynamics in a tritrophic model. Examining the phase relationships between species of adjacent trophic levels informs about the strength of top-down or bottom-up control in non-steadystate situations. Phase relationships within a trophic level highlight compensatory dynamical patterns between functionally different species, which are responsible for dampening the community temporal variability. Furthermore, even without trait variation, our tritrophic model always exhibits regions with two alternative states with either weak or strong nutrient exploitation, and correspondingly low or high biomass production at the top level. However, adding trait variation increased the basin of attraction of the high-production state, and decreased the likelihood of a critical transition from the high- to the lowproduction state with no apparent early warning signals. Hence, our study shows that trait variation enhances resource use efficiency, production, stability, and resilience of entire food webs. KW - early-warning signals KW - top-down control KW - community ecology KW - regime shifts KW - food webs KW - compensatory dynamics KW - consumer diversity KW - metabolic theory KW - rapid evolution KW - stable states Y1 - 2019 U6 - https://doi.org/10.1038/s41598-019-43974-1 SN - 2045-2322 VL - 9 PB - Macmillan Publishers Limited CY - London ER - TY - JOUR A1 - Ryser, Remo A1 - Häussler, Johanna A1 - Stark, Markus A1 - Brose, Ulrich A1 - Rall, Björn C. A1 - Guill, Christian T1 - The biggest losers: habitat isolation deconsructs complex food webs from top to bottom JF - Proceedings of the Royal Society of London : B, Biological sciences N2 - Habitat fragmentation threatens global biodiversity. To date, there is only limited understanding of how the different aspects of habitat fragmentation (habitat loss, number of fragments and isolation) affect species diversity within complex ecological networks such as food webs. Here, we present a dynamic and spatially explicit food web model which integrates complex food web dynamics at the local scale and species-specific dispersal dynamics at the landscape scale, allowing us to study the interplay of local and spatial processes in metacommunities. We here explore how the number of habitat patches, i.e. the number of fragments, and an increase of habitat isolation affect the species diversity patterns of complex food webs (alpha-,beta-,gamma-, diversities). We specifically test whether there is a trophic dependency in the effect of these two factors on species diversity. In our model, habitat isolation is the main driver causing species loss and diversity decline. Our results emphasize that large-bodied consumer species at high trophic positions go extinct faster than smaller species at lower trophic levels, despite being superior dispersers that connect fragmented landscapes better. We attribute the loss of top species to a combined effect of higher biomass loss during dispersal with increasing habitat isolation in general, and the associated energy limitation in highly fragmented landscapes, preventing higher trophic levels to persist. To maintain trophic-complex and species-rich communities calls for effective conservation planning which considers the interdependence of trophic and spatial dynamics as well as the spatial context of a landscape and its energy availability. KW - food webs KW - allometry KW - bioenergetic model KW - metacommunity dynamics KW - dispersal mortality KW - landscape structure Y1 - 2019 U6 - https://doi.org/10.1098/rspb.2019.1177 SN - 0962-8452 SN - 1471-2954 VL - 286 IS - 1908 PB - Royal Society CY - London ER - TY - JOUR A1 - Guill, Christian A1 - Hülsemann, Janne A1 - Klauschies, Toni T1 - Self-organised pattern formation increases local diversity in metacommunities JF - Ecology letters N2 - 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. KW - biomass-trait feedback KW - fitness gradient KW - food chain KW - functional KW - diversity KW - metacommunity KW - self-organisation KW - source-sink dynamics KW - spatio-temporal pattern KW - trait-based aggregate model KW - Turing instability Y1 - 2021 U6 - https://doi.org/10.1111/ele.13880 SN - 1461-023X SN - 1461-0248 VL - 24 IS - 12 SP - 2624 EP - 2634 PB - Wiley-Blackwell CY - Oxford ER - TY - JOUR A1 - Stark, Markus A1 - Bach, Moritz A1 - Guill, Christian T1 - Patch isolation and periodic environmental disturbances have idiosyncratic effects on local and regional population variabilities in meta-food chains JF - Theoretical ecology N2 - While habitat loss is a known key driver of biodiversity decline, the impact of other landscape properties, such as patch isolation, is far less clear. When patch isolation is low, species may benefit from a broader range of foraging opportunities, but are at the same time adversely affected by higher predation pressure from mobile predators. Although previous approaches have successfully linked such effects to biodiversity, their impact on local and metapopulation dynamics has largely been ignored. Since population dynamics may also be affected by environmental disturbances that temporally change the degree of patch isolation, such as periodic changes in habitat availability, accurate assessment of its link with isolation is highly challenging. To analyze the effect of patch isolation on the population dynamics on different spatial scales, we simulate a three-species meta-food chain on complex networks of habitat patches and assess the average variability of local populations and metapopulations, as well as the level of synchronization among patches. To evaluate the impact of periodic environmental disturbances, we contrast simulations of static landscapes with simulations of dynamic landscapes in which 30 percent of the patches periodically become unavailable as habitat. We find that increasing mean patch isolation often leads to more asynchronous population dynamics, depending on the parameterization of the food chain. However, local population variability also increases due to indirect effects of increased dispersal mortality at high mean patch isolation, consequently destabilizing metapopulation dynamics and increasing extinction risk. In dynamic landscapes, periodic changes of patch availability on a timescale much slower than ecological interactions often fully synchronize the dynamics. Further, these changes not only increase the variability of local populations and metapopulations, but also mostly overrule the effects of mean patch isolation. This may explain the often small and inconclusive impact of mean patch isolation in natural ecosystems. KW - Metacommunity dynamics KW - Dispersal KW - Patch isolation KW - Stability KW - Synchronization KW - Disturbance Y1 - 2021 U6 - https://doi.org/10.1007/s12080-021-00510-0 SN - 1874-1738 SN - 1874-1746 VL - 14 IS - 3 SP - 489 EP - 500 PB - Springer CY - Dordrecht ER - TY - JOUR A1 - Miele, Vincent A1 - Guill, Christian A1 - Ramos-Jiliberto, Rodrigo A1 - Kéfi, Sonia T1 - Non-trophic interactions strengthen the diversity-functioning relationship in an ecological bioenergetic network model JF - PLoS Computational Biology : a new community journal N2 - Ecological communities are undeniably diverse, both in terms of the species that compose them as well as the type of interactions that link species to each other. Despite this long recognition of the coexistence of multiple interaction types in nature, little is known about the consequences of this diversity for community functioning. In the ongoing context of global change and increasing species extinction rates, it seems crucial to improve our understanding of the drivers of the relationship between species diversity and ecosystem functioning. Here, using a multispecies dynamical model of ecological communities including various interaction types (e.g. competition for space, predator interference, recruitment facilitation in addition to feeding), we studied the role of the presence and the intensity of these interactions for species diversity, community functioning (biomass and production) and the relationship between diversity and functioning. Taken jointly, the diverse interactions have significant effects on species diversity, whose amplitude and sign depend on the type of interactions involved and their relative abundance. They however consistently increase the slope of the relationship between diversity and functioning, suggesting that species losses might have stronger effects on community functioning than expected when ignoring the diversity of interaction types and focusing on feeding interactions only. Y1 - 2020 U6 - https://doi.org/10.1371/journal.pcbi.1007269 SN - 1553-7358 VL - 15 IS - 8 PB - PLoS CY - San Fransisco ER - TY - JOUR A1 - Gross, Thilo A1 - Allhoff, Korinna Theresa A1 - Blasius, Bernd A1 - Brose, Ulrich A1 - Drossel, Barbara A1 - Fahimipour, Ashkaan K. A1 - Guill, Christian A1 - Yeakel, Justin D. A1 - Zeng, Fanqi T1 - Modern models of trophic meta-communities JF - Philosophical transactions of the Royal Society of London : B, Biological sciences N2 - Dispersal and foodweb dynamics have long been studied in separate models. However, over the past decades, it has become abundantly clear that there are intricate interactions between local dynamics and spatial patterns. Trophic meta-communities, i.e. meta-foodwebs, are very complex systems that exhibit complex and often counterintuitive dynamics. Over the past decade, a broad range of modelling approaches have been used to study these systems. In this paper, we review these approaches and the insights that they have revealed. We focus particularly on recent papers that study trophic interactions in spatially extensive settings and highlight the common themes that emerged in different models. There is overwhelming evidence that dispersal (and particularly intermediate levels of dispersal) benefits the maintenance of biodiversity in several different ways. Moreover, some insights have been gained into the effect of different habitat topologies, but these results also show that the exact relationships are much more complex than previously thought, highlighting the need for further research in this area. This article is part of the theme issue 'Integrative research perspectives on marine conservation'. KW - dispersal KW - meta-community KW - foodweb Y1 - 2020 U6 - https://doi.org/10.1098/rstb.2019.0455 SN - 0962-8436 SN - 1471-2970 VL - 375 IS - 1814 PB - Royal Society CY - London ER - TY - JOUR A1 - Binzer, Amrei A1 - Guill, Christian A1 - Rall, Björn C. A1 - Brose, Ulrich T1 - Interactive effects of warming, eutrophication and size structure: impacts on biodiversity and food-web structure JF - Global change biology N2 - Warming and eutrophication are two of the most important global change stressors for natural ecosystems, but their interaction is poorly understood. We used a dynamic model of complex, size-structured food webs to assess interactive effects on diversity and network structure. We found antagonistic impacts: Warming increases diversity in eutrophic systems and decreases it in oligotrophic systems. These effects interact with the community size structure: Communities of similarly sized species such as parasitoid-host systems are stabilized by warming and destabilized by eutrophication, whereas the diversity of size-structured predator-prey networks decreases strongly with warming, but decreases only weakly with eutrophication. Nonrandom extinction risks for generalists and specialists lead to higher connectance in networks without size structure and lower connectance in size-structured communities. Overall, our results unravel interactive impacts of warming and eutrophication and suggest that size structure may serve as an important proxy for predicting the community sensitivity to these global change stressors. KW - complex food webs KW - extinctions KW - generalists KW - global change KW - size structure KW - specialists Y1 - 2016 U6 - https://doi.org/10.1111/gcb.13086 SN - 1354-1013 SN - 1365-2486 VL - 22 SP - 220 EP - 227 PB - Wiley-Blackwell CY - Hoboken ER - TY - JOUR A1 - Allhoff, Korinna Theresa A1 - Ritterskamp, Daniel A1 - Rall, Björn C. A1 - Drossel, Barbara A1 - Guill, Christian T1 - Evolutionary food web model based on body masses gives realistic networks with permanent species turnover JF - Scientific reports N2 - The networks of predator-prey interactions in ecological systems are remarkably complex, but nevertheless surprisingly stable in terms of long term persistence of the system as a whole. In order to understand the mechanism driving the complexity and stability of such food webs, we developed an eco-evolutionary model in which new species emerge as modifications of existing ones and dynamic ecological interactions determine which species are viable. The food-web structure thereby emerges from the dynamical interplay between speciation and trophic interactions. The proposed model is less abstract than earlier evolutionary food web models in the sense that all three evolving traits have a clear biological meaning, namely the average body mass of the individuals, the preferred prey body mass, and the width of their potential prey body mass spectrum. We observed networks with a wide range of sizes and structures and high similarity to natural food webs. The model networks exhibit a continuous species turnover, but massive extinction waves that affect more than 50% of the network are not observed. Y1 - 2015 U6 - https://doi.org/10.1038/srep10955 SN - 2045-2322 VL - 5 PB - Nature Publ. Group CY - London ER - TY - JOUR A1 - Schneider, Florian D. A1 - Brose, Ulrich A1 - Rall, Björn C. A1 - Guill, Christian T1 - Animal diversity and ecosystem functioning in dynamic food webs JF - Nature Communications N2 - Species diversity is changing globally and locally, but the complexity of ecological communities hampers a general understanding of the consequences of animal species loss on ecosystem functioning. High animal diversity increases complementarity of herbivores but also increases feeding rates within the consumer guild. Depending on the balance of these counteracting mechanisms, species-rich animal communities may put plants under top-down control or may release them from grazing pressure. Using a dynamic food-web model with body-mass constraints, we simulate ecosystem functions of 20,000 communities of varying animal diversity. We show that diverse animal communities accumulate more biomass and are more exploitative on plants, despite their higher rates of intra-guild predation. However, they do not reduce plant biomass because the communities are composed of larger, and thus energetically more efficient, plant and animal species. This plasticity of community body-size structure reconciles the debate on the consequences of animal species loss for primary productivity. Y1 - 2016 U6 - https://doi.org/10.1038/ncomms12718 SN - 2041-1723 VL - 7 SP - 3129 EP - 3138 PB - Nature Publ. Group CY - London ER -