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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.
Species can adjust their traits in response to selection which may strongly influence species coexistence. Nevertheless, current theory mainly assumes distinct and time-invariant trait values. We examined the combined effects of the range and the speed of trait adaptation on species coexistence using an innovative multispecies predator–prey model. It allows for temporal trait changes of all predator and prey species and thus simultaneous coadaptation within and among trophic levels. We show that very small or slow trait adaptation did not facilitate coexistence because the stabilizing niche differences were not sufficient to offset the fitness differences. In contrast, sufficiently large and fast trait adaptation jointly promoted stable or neutrally stable species coexistence. Continuous trait adjustments in response to selection enabled a temporally variable convergence and divergence of species traits; that is, species became temporally more similar (neutral theory) or dissimilar (niche theory) depending on the selection pressure, resulting over time in a balance between niche differences stabilizing coexistence and fitness differences promoting competitive exclusion. Furthermore, coadaptation allowed prey and predator species to cluster into different functional groups. This equalized the fitness of similar species while maintaining sufficient niche differences among functionally different species delaying or preventing competitive exclusion. In contrast to pre-
vious studies, the emergent feedback between biomass and trait dynamics enabled supersaturated coexistence for a broad range of potential trait adaptation and parameters. We conclude that accounting for trait adaptation may explain stable and supersaturated species coexistence for a broad range of environmental conditions in natural systems when the absence of such adaptive changes would preclude it. Small trait changes, coincident with those that may occur within many natural populations, greatly enlarged the number of coexisting species.
Species can adjust their traits in response to selection which may strongly influence species coexistence. Nevertheless, current theory mainly assumes distinct and time-invariant trait values. We examined the combined effects of the range and the speed of trait adaptation on species coexistence using an innovative multispecies predator–prey model. It allows for temporal trait changes of all predator and prey species and thus simultaneous coadaptation within and among trophic levels. We show that very small or slow trait adaptation did not facilitate coexistence because the stabilizing niche differences were not sufficient to offset the fitness differences. In contrast, sufficiently large and fast trait adaptation jointly promoted stable or neutrally stable species coexistence. Continuous trait adjustments in response to selection enabled a temporally variable convergence and divergence of species traits; that is, species became temporally more similar (neutral theory) or dissimilar (niche theory) depending on the selection pressure, resulting over time in a balance between niche differences stabilizing coexistence and fitness differences promoting competitive exclusion. Furthermore, coadaptation allowed prey and predator species to cluster into different functional groups. This equalized the fitness of similar species while maintaining sufficient niche differences among functionally different species delaying or preventing competitive exclusion. In contrast to previous studies, the emergent feedback between biomass and trait dynamics enabled supersaturated coexistence for a broad range of potential trait adaptation and parameters. We conclude that accounting for trait adaptation may explain stable and supersaturated species coexistence for a broad range of environmental conditions in natural systems when the absence of such adaptive changes would preclude it. Small trait changes, coincident with those that may occur within many natural populations, greatly enlarged the number of coexisting species.
Investigating the mechanisms which underlie the biomass fluctuations of populations and communities is important to better understand the processes which buffer community biomass in a variable environment. Based on long- term data of plankton biomass in Lake Constance (Bodensee), this study aims at explaining the different degree of synchrony among populations observed within two freshwater plankton groups, phytoplankton and ciliates. Established measures of temporal variability such as the variance ratio and cross-correlation coefficients were combined with first- order autoregressive models that allow estimating species interactions from time-series data. We found that predation was an important driver of the observed seasonal variability patterns in phytoplankton and ciliates, and that competitive interactions only played a subordinate role. In Lake Constance copepods and cladocerans, two major invertebrate predator groups, focus their grazing pressure at different times of the season. Model results suggested that compensatory dynamics detected in phytoplankton originate from the differential vulnerability of species to either one of these two predator groups. For ciliates model results advocated that synchrony among species occurs because ciliates tend to be vulnerable to both predator groups. Our findings underline the necessity of extending studies of community variability to multiple trophic levels because accounting for predator-prey interactions may often be more important than accounting for competitive interactions at one trophic level
Functional groups with diverse responses to environmental factors sum to produce communities with less temporal variability in their biomass than those lacking this diversity. The detection of these compensatory dynamics can be complicated by a spatio-temporal alternation in the environmental factors limiting growth (both abiotic and biotic), which restricts the occurrence of compensatory dynamics to certain periods or locations. Hence, resolving the spatio- temporal scale may uncover important spatial and/or temporal components in community variability. Using long-term data from Lake Constance (Bodensee), we find that a reduction in grazing pressure and relaxed competition for nutrients during winter and spring generates coherent dynamics among edible and less edible phytoplankton. During summer and fall, when both grazing pressure and nutrient limitation are present, edible and less edible phytoplankton exhibit compensatory dynamics. This study supports recent work suggesting that both abiotic and biotic interactions promote compensatory dynamics and to our knowledge, this is the first example of a system where compensatory and coherent dynamics seasonally alternate.
Inorganic carbon limitation and mixotrophic growth in Chlamydomonas from an acidic mining lake
(2005)
Plankton communities in acidic mining lakes (pH 2.5-3.3) are species-poor because they face extreme environmental conditions, e.g. 150 mg l(-1) Fe2++Fe3+. We investigated the growth characteristics of the dominant pigmented species, the flagellate Chlamydomonas acidophila, in semi-continuous culture experiments under in situ conditions. The following hypotheses were tested: (1) Low inorganic carbon (IC) concentrations in the epilimnion (e.g. 0.3 mg l(-1)) arising from the low pH limit phototrophic growth (H-1); (2) the additional use of dissolved organic carbon (mixotrophy) leads to higher growth rates under IC-limitation (H-2), and (3) phagotrophy is not relevant (H-3). H- 1 was supported as the culture experiments, in situ PAR and IC concentrations indicated that IC potentially limited phototrophic growth in the mixed surface layers. H-2 was also supported: mixotrophic growth always exceeded pure phototrophic growth even when photosynthesis was saturated. Dark growth in filtered lake water illuminated prior to inoculation provided evidence that Chlamydomonas was able to use the natural DOC. The alga did not grow on bacteria, thus confirming H-3. Chlamydomonas exhibited a remarkable resistance to starvation in the dark. The compensation light intensity (ca. 20 mu mol photons m(-2) s(-1)) and the maximum phototrophic growth (1.50 d(-1)) fell within the range of algae from non-acidic waters. Overall, Chlamydomonas, a typical r-strategist in circum-neutral systems, showed characteristics of a K-strategist in the stable, acidic lake environment in achieving moderate growth rates and minimizing metabolic losses. (c) 2005 Elsevier GmbH. All rights reserved
During 1987-1998, the ciliates and their prey and predator communities in large, deep, mesotrophic Lake Constance were intensively studied as it underwent re-oligotrophication. Ciliate biomass exhibited the bimodal seasonal distribution typical for meso-eutrophic lakes, with high biomass in spring and summer and low biomass in winter and during the clear-water phase. Cluster analysis produced nine groups of temporally co-occurring ciliate morphotypes with potentially similar ecological characteristics. The clusters exhibited a larger seasonality than found in the size distribution, showing that similarly-sized ciliates had seasonally compensatory dynamics. Ciliate biomass declined by approx. 30 % during the 12 years of study, i.e. considerably less than daphnids (and total phosphorus). This yielded a significant increase in the ratio between summer ciliate and daphnid biomass as re-oligotrophication progressed, in contrast to previous studies. Few indications for a mechanistic link between phosphorus concentrations (which declined threefold during the study period) and ciliate biomass or community composition via group-specific food concentrations were found. The relative contribution of three of the nine clusters changed as re-oligotrophication progressed. Ciliate size distribution was related to reoligotrophication and daphnid biomass in summer. The smallest and largest ciliates gained in importance when daphnids decreased whereas large ciliates declined. Overall, summer daphnid biomas had a greater predictive power for attributes of the ciliate community than the other factors studied (phosphorus, prey biomass, copepod biomass). The extent of bottom-up and top-down control of ciliates appeared to be time and group specific. Overall, the ciliate community exhibited remarkably recurrent seasonal patterns despite major alternations in abiotic and biotic conditions.
Biomass size spectra collate structural and functional attributes of plankton communities enabling standardised temporal and cross-system comparisons and may be rapidly obtained by automated particle counters. To examine how differences in plankton communities from highly eutrophic and more oligotrophic lakes are reflected in size spectra, a three-year time series of biomass size spectra was established for polymictic, eutrophic Lake Müggelsee, based on approximately weekly sampling and microscopic enumeration. The continuous but often bumpy size spectra reflected appropriately the seasonal and trophy-related variations in the plankton composition and growth conditions and the potential impact of daphnids on smaller plankton. We tested the hypothesis that more diverse plankton communities have smoother size spectra than impoverished ones. The spectra of L. Müggelsee and other more less eutrophic lakes covaried roughly with the functional diversity in total plankton composition but were unrelated to taxonomical diversity within the phyto- or mesozooplankton. The slopes of the normalised size spectra of Lake Müggelsee were generally more negative than -1, exhibited a recurrent seasonal pattern, and were strongly correlated with crustacean biomass. In contrast to less eutrophic systems, slopes could not be used to quantify energy fluxes within the foodweb due to highly variable algal P/B ratios and frequently bumpy size distributions. The latter indicated stronger deviations from the ideal concept of a steady energy flow along the size gradient than found in e. g. large, mesotrophic Lake Constance.
Clear-water phase (CWP) is an important event in seasonal plankton succession. We examined the influence of all herbivorous zooplankton on its initiation under different weather and climatic conditions using up to 19 years of observations from the large, deep Lake Constance (Europe) and estimates of relative clearance rates. A CWP occurred regularly, even if daphnid biomass was still very low. CWP was attributed to strong grazing either by a daphnid- dominated zooplankton community or by a diverse assemblage consisting of micro- and meso-zooplankton. Both types of zooplankton communities occurred with approximately the same frequency. The timing of the CWP was unrelated to the North Atlantic Oscillation (NAO) but correlated with the wind-dependent intensity of deep vertical mixing 3 months earlier, during early spring. Less mixing enabled early growth of phytoplankton, ciliates and rotifers despite low temperatures, which prevented daphnid development at this time. This resulted in enhanced grazing of ciliates and rotifers, which increased the importance of phytoplankton less edible for most ciliates, rotifers and daphnids. Ciliates clearly dominated the grazing pressure on phytoplankton throughout spring, maintaining high biomasses together with the phytoplankton for up to 2 months. A CWP was observed when herbivores grazing on larger phytoplankton developed in addition to ciliates