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Ecosystems are generally linked via fluxes of nutrients and energy across their boundaries. For example, freshwater ecosystems in temperate regions may receive significant inputs of terrestrially derived carbon via autumnal leaf litter. This terrestrial particulate organic carbon (POC) is hypothesized to subsidize animal production in lakes, but direct evidence is still lacking. We divided two small eutrophic lakes each into two sections and added isotopically distinct maize litter to the treatment sections to simulate increased terrestrial POC inputs via leaf litter in autumn. We quantified the reliance of aquatic consumers on terrestrial resources (allochthony) in the year subsequent to POC additions by applying mixing models of stable isotopes. We also estimated lake-wide carbon (C) balances to calculate the C flow to the production of the major aquatic consumer groups: benthic macroinvertebrates, crustacean zooplankton, and fish. The sum of secondary production of crustaceans and benthic macroinvertebrates supported by terrestrial POC was higher in the treatment sections of both lakes. In contrast, total secondary and tertiary production (supported by both autochthonous and allochthonous C) was higher in the reference than in the treatment sections of both lakes. Average aquatic consumer allochthony per lake section was 27-40%, although terrestrial POC contributed less than about 10% to total organic C supply to the lakes. The production of aquatic consumers incorporated less than 5% of the total organic C supply in both lakes, indicating a low ecological efficiency. We suggest that the consumption of terrestrial POC by aquatic consumers facilitates a strong coupling with the terrestrial environment. However, the high autochthonous production and the large pool of autochthonous detritus in these nutrient-rich lakes make terrestrial POC quantitatively unimportant for the C flows within food webs.
Climate change will alter the forces of predation and competition in temperate ectotherm food webs. This may increase local extinction rates, change the fate of invasions and impede species reintroductions into communities. Invasion success could be modulated by traits (e.g., defenses) and adaptations to climate. We studied how different temperatures affect the time until extinction of species, using bitrophic and tritrophic planktonic food webs to evaluate the relative importance of predatory overexploitation and competitive exclusion, at 15 and 25 A degrees C. In addition, we tested how inclusion of a subtropical as opposed to a temperate strain in this model food web affects times until extinction. Further, we studied the invasion success of the temperate rotifer Brachionus calyciflorus into the planktonic food web at 15 and 25 A degrees C on five consecutive introduction dates, during which the relative forces of predation and competition differed. A higher temperature dramatically shortened times until extinction of all herbivore species due to carnivorous overexploitation in tritrophic systems. Surprisingly, warming did not increase rates of competitive exclusion among the tested herbivore species in bitrophic communities. Including a subtropical herbivore strain reduced top-down control by the carnivore at high temperature. Invasion attempts of temperate B. calyciflorus into the food web always succeeded at 15 A degrees C, but consistently failed at 25 A degrees C due to voracious overexploitation by the carnivore. Pre-induction of defenses (spines) in B. calyciflorus before the invasion attempt did not change its invasion success at the high temperature. We conclude that high temperatures may promote local extinctions in temperate ectotherms and reduce their chances of successful recovery.
Indoor mesocosm experiments were conducted to test for potential climate change effects on the spring succession of Baltic Sea plankton. Two different temperature (Delta 0 A degrees C and Delta 6 A degrees C) and three light scenarios (62, 57 and 49 % of the natural surface light intensity on sunny days), mimicking increasing cloudiness as predicted for warmer winters in the Baltic Sea region, were simulated. By combining experimental and modeling approaches, we were able to test for a potential dietary mismatch between phytoplankton and zooplankton. Two general predator-prey models, one representing the community as a tri-trophic food chain and one as a 5-guild food web were applied to test for the consequences of different temperature sensitivities of heterotrophic components of the plankton. During the experiments, we observed reduced time-lags between the peaks of phytoplankton and protozoan biomass in response to warming. Microzooplankton peak biomass was reached by 2.5 day A degrees C-1 earlier and occurred almost synchronously with biomass peaks of phytoplankton in the warm mesocosms (Delta 6 A degrees C). The peak magnitudes of microzooplankton biomass remained unaffected by temperature, and growth rates of microzooplankton were higher at Delta 6 A degrees C (mu(a dagger 0 A degrees C) = 0.12 day(-1) and mu(a dagger 6 A degrees C) = 0.25 day(-1)). Furthermore, warming induced a shift in microzooplankton phenology leading to a faster species turnover and a shorter window of microzooplankton occurrence. Moderate differences in the light levels had no significant effect on the time-lags between autotrophic and heterotrophic biomass and on the timing, biomass maxima and growth rate of microzooplankton biomass. Both models predicted reduced time-lags between the biomass peaks of phytoplankton and its predators (both microzooplankton and copepods) with warming. The reduction of time-lags increased with increasing Q(10) values of copepods and protozoans in the tritrophic food chain. Indirect trophic effects modified this pattern in the 5-guild food web. Our study shows that instead of a mismatch, warming might lead to a stronger match between protist grazers and their prey altering in turn the transfer of matter and energy toward higher trophic levels.
Standing stocks are typically easier to measure than process rates such as production. Hence, stocks are often used as indicators of ecosystem functions although the latter are generally more strongly related to rates than to stocks. The regulation of stocks and rates and thus their variability over time may differ, as stocks constitute the net result of production and losses. Based on long-term high frequency measurements in a large, deep lake we explore the variability patterns in primary and bacterial production and relate them to those of the corresponding standing stocks, i.e. chlorophyll concentration, phytoplankton and bacterial biomass. We employ different methods (coefficient of variation, spline fitting and spectral analysis) which complement each other for assessing the variability present in the plankton data, at different temporal scales. In phytoplankton, we found that the overall variability of primary production is dominated by fluctuations at low frequencies, such as the annual, whereas in stocks and chlorophyll in particular, higher frequencies contribute substantially to the overall variance. This suggests that using standing stocks instead of rate measures leads to an under- or overestimation of food shortage for consumers during distinct periods of the year. The range of annual variation in bacterial production is 8 times greater than biomass, showing that the variability of bacterial activity (e.g. oxygen consumption, remineralisation) would be underestimated if biomass is used. The P/B ratios were variable and although clear trends are present in both bacteria and phytoplankton, no systematic relationship between stock and rate measures were found for the two groups. Hence, standing stock and process rate measures exhibit different variability patterns and care is needed when interpreting the mechanisms and implications of the variability encountered.
1. The polyunsaturated fatty acid eicosapentaenoic acid (EPA) plays an important role in aquatic food webs, in particular at the primary producerconsumer interface where keystone species such as daphnids may be constrained by its dietary availability. Such constraints and their seasonal and interannual changes may be detected by continuous measurements of EPA concentrations. However, such EPA measurements became common only during the last two decades, whereas long-term data sets on plankton biomass are available for many well-studied lakes. Here, we test whether it is possible to estimate EPA concentrations from abiotic variables (light and temperature) and the biomass of prey organisms (e.g. ciliates, diatoms and cryptophytes) that potentially provide EPA for consumers. 2. We used multiple linear regression to relate size- and taxonomically resolved plankton biomass data and measurements of temperature and light intensity to directly measured EPA concentrations in Lake Constance during a whole year. First, we tested the predictability of EPA concentrations from the biomass of EPA-rich organisms (diatoms, cryptophytes and ciliates). Secondly, we included the variables mean temperature and mean light intensity over the sampling depth (020 m) and depth (08 and 820 m) as factors in our model to check for large-scale seasonal- and depth-dependent effects on EPA concentrations. In a third step, we included the deviations of light and temperature from mean values in our model to allow for their potential influence on the biochemical composition of plankton organisms. We used the Akaike Information Criterion to determine the best models. 3. All approaches supported our proposition that the biomasses of specific plankton groups are variables from which seston EPA concentrations can be derived. The importance of ciliates as an EPA source in the seston was emphasised by their high weight in our models, although ciliates are neglected in most studies that link fatty acids to seston taxonomic composition. The large-scale seasonal variability of light intensity and its interaction with diatom biomass were significant predictors of EPA concentrations. The deviation of temperature from mean values, accounting for a depth-dependent effect on EPA concentrations, and its interaction with ciliate biomass were also variables with high predictive power. 4. The best models from the first and second approaches were validated with measurements of EPA concentrations from another year (1997). The estimation with the best model including only biomass explained 80%, and the best model from the second approach including mean temperature and depth explained 87% of the variability in EPA concentrations in 1997. 5. We show that it is possible to predict EPA concentrations reliably from plankton biomass, while the inclusion of abiotic factors led to results that were only partly consistent with expectations from laboratory studies. Our approach of including biotic predictors should be transferable to other systems and allow checking for biochemical constraints on primary consumers.
BISSINGER, V.; TITTEL, J.: Process rates and growth limiting factors of planktonic algae (Chlamydomonas sp.) from extremely acidic (pH 2,5 - 3) mining lakes in Germany ; BORK, H.-R. et al.: Erodierte Autos und Brunnen in Oregon, USA ; BRONSTERT, A. et al.: Bewirtschaftunsmöglichkeiten im Einzugsgebiet der Havel ; JELTSCH, F. et al.: Beweidung als Degradationsfaktor in ariden und semiariden Weidesystemen ; JELTSCH, F. et al.: Entstehung und Bedeutung räumlicher Vegetationsstrukturen in Trockensavannen: Baum-Graskoexistenz und Artenvielfalt ; JESSEL, B. et al.: Bodenbewertung für Planungs- und Zulassungsverfahren in Brandenburg ; JESSEL, B.; ZSCHALICH, A.: Erarbeitung von Ausgleichs- und Ersatzmaßnahmen für die Wert- und Funktionselemente des Landschaftsbildes ; RÖßLING, H. et al.: Umsetzung von Ausgleichs- und Ersatzmaßnahmen beim Ausbau der Bundesautobahn A 9 ; SPINDLER, J.; GAEDKE, U.: Estimating production in plankton food webs from biomass size spectra and allometric relationships ; TIELBÖRGER, K. et al.: Sukzessionsprozesse in einem Sanddünengebiet nach Ausschluß von Beweidung ; TIELBÖRGER, K. et al.: Populationsdynamische Funktionen von Ausbreitung und Dormanz ; TIELBÖRGER, K. et al.: Raum-zeitliche Populationsdynamik von einjährigen Wüstenpflanzen ; TITTEL, J. et al.: Ressourcennutzung und -weitergabe im planktischen Nahrungsnetz eines extrem sauren (pH 2,7) Tagebausees ; WALLSCHLÄGER, D.; WIEGLEB, G.: Offenland-Management auf ehemaligen und in Nutzung befindlichen Truppenübungsplätzen im pleistozänen Flachland Nordostdeutschlands: Naturschutzfachliche Grundlagen und praktische Anwendungen ; WEITHOFF, G.; GAEDKE, U.: Planktische Räuber-Beute-Systeme: Experimentelle Untersuchung von ökologischen Synchronisationen
Trophic transfer efficiency (TTE) is usually calculated as the ratio of production rates between two consecutive trophic levels. Although seemingly simple, TTE estimates from lakes are rare. In our review, we explore the processes and structures that must be understood for a proper lake TTE estimate.
We briefly discuss measurements of production rates and trophic positions and mention how ecological efficiencies, nutrients (N, P) and other compounds (fatty acids) affect energy transfer between trophic levels and hence TTE.
Furthermore, we elucidate how TTE estimates are linked with size-based approaches according to the Metabolic Theory of Ecology, and how food-web models can be applied to study TTE in lakes.
Subsequently, we explore temporal and spatial heterogeneity of production and TTE in lakes, with a particular focus on the links between benthic and pelagic habitats and between the lake and the terrestrial environment.
We provide an overview of TTE estimates from lakes found in the published literature. Finally, we present two alternative approaches to estimating TTE. First, TTE can be seen as a mechanistic quantity informing about the energy and matter flow between producer and consumer groups.
This approach is informative with respect to food-web structure, but requires enormous amounts of data. The greatest uncertainty comes from the proper consideration of basal production to estimate TTE of omnivorous organisms.
An alternative approach is estimating food-chain and food-web efficiencies, by comparing the heterotrophic production of single consumer levels or the total sum of all heterotrophic production including that of heterotrophic bacteria to the total sum of primary production.
We close the review by pointing to a few research questions that would benefit from more frequent and standardized estimates of TTE in lakes.
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.
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.
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.
The shape of a defense-growth trade-off governs seasonal trait dynamics in natural phytoplankton
(2020)
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.
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
Large (472 km2) and deep (zmean=101 m) Lake Constance is undergoing re-oligotrophication. Total phosphorus during winter mixing (TPmix) decreased from >80 during 1975-1981 to 22 ;g/l in 1996. Average summer values of secchi and euphotic depth increased significantly from 4.5 to 6.5 m and from 10.5 to 13 m, respectively. The algal species composition changed and, during summer, total algal biomass decreased by 50 % and primary production by 25 %. Standing stocks of well-edible algae, rotifers, and herbivorous and carnivorous crustaceans did not exhibit a trend with TPmix, whereas their species compositions or egg-ratios were partially altered. The age-at-capture of planktivorous whitefish increased slightly. I tested the hypotheses that (1) changes should first be observed at the level of individuals or within species (altering e. g. C:P or egg-ratios) prior to changes within communities (affecting e. g. the taxonomic composition) and at the community level (affecting e. g. total biomass or production). This would imply that it is more appropriate to conceptualize step-wise responses along a hierarchical gradient of increasing aggregation as suggested by hierarchy theory, rather than simultaneous changes at all hierarchical levels. (2) Responses become dampened along the food chain and with increasing body size, i. e. bottom-up control is most important for autotrophs. All communities studied (phytoplankton, crustaceans, fish) reacted at the individual level (e. g. by changes of (re)production rates), and/or within the community (e. g. altered taxonomic composition) whereas changes of bulk parameters of the entire community were restricted to phytoplankton. Hence, the first hypothesis is partially supported by the observed reactions and demands further testing. The second hypothesis is clearly supported by our data when comparing autotrophs and consumers, but not when comparing crustaceans and fish. The testing of these hypotheses is complicated by the large differences in size and, consequently, in reaction times of pelagic organisms on the one hand and the rather fixed time scale of limnological research on the other hand. The different time scales imply a selective perception of the various potential responses of the differently sized organisms as the time scales of the responses depend on body size and the level of aggregation. For example, we are more likely to establish physiological or behaviourial changes of fish, and taxonomical or biomass changes of phytoplankton. Acknowledging the scale dependence and level of aggregation is also crucial for cross-system comparisons.
The intrinsic predictability of ecological time series and its potential to guide forecasting
(2019)
Phytoplankton dynamics in a shallow eutrophic lake were investigated over a 3-year period with respect to environmental forces which drive species composition and diversity. Diversity was calculated on the basis of species as well as on the basis of their functional properties (the C-R-S-concept). Stratification and water column mixing had a strong impact on phytoplankton composition. Application of a similarity-diversity model revealed that a high diversity was a transient non-stable state, whereas drastic changes or long-lasting stable environmental conditions are characterized by low diversity. This effect was more pronounced when the diversity was calculated on the basis of the phytoplankton species functional properties. Thus, this functional approach supports the intermediate disturbance hypothesis from field data.
Long-term measurements (1979-1994) of meteorological parameters and of algal and crustacean biomass were used in conjunction with a comprehensive hydrodynamic model to evaluate the impact of weather conditions on plankton dynamics in a large, deep, temperate lake (Upper Lake Constance), and to identify potential causal mechanisms. The natural variability of weather conditions, including the exceptionally mild winters during the late eighties, allowed us the investigation of the covariation of meteorological parameters such as irradiance, air temperature, and wind with vernal algal and crustacean population growth. Crustacean zooplankton responded strongly to differences in surface water temperature, but not to mixing depth or algal biomass. Clear relationships between changes of algal biomass and meteorological factors were only found during the rare occasions when acted together to favour or hamper algal development. Otherwise, the impact of meterological conditions on the physical conditions which were most likely conducive to phytoplankton development, could not be followed by this simple approach. This problem was overcome with a one-dimensional hydrodynamic turbulent exchange model driven by the meteorological boundary conditions at the water surface. It was used to simulate the development of the vernal density stratification and to investigate the relationships between meteorological conditions and exchange rates from the euphotic to the aphotic zone. The beginning of the spring algal bloom was shown to depend on the stabilization of the upper part of the water column. As soon as mixing below 20 m was inhibited, confining the algae to the euphotic zone for prolonged periods of time, substantial increases in algal standing stock occurred consistently. In contrast, during periods when high vertical mixing rates were computed with the model no substantial increases of algal biomass were found. This tight coupling between the estimates of vertical mixing intensity and observed algal development, combined with knowledge about the impact of individual meteorological factors on mixing, enabled predictions about the response of algae to different weather conditions during spring.
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.
Spring algal development in deep temperate lakes is thought to be strongly influenced by surface irradiance, vertical mixing and temperature, all of which are expected to be altered by climate change. Based on long-term data from Lake Constance, we investigated the individual and combined effects of these variables on algal dynamics using descriptive statistics, multiple regression models and a processoriented dynamic simulation model. The latter considered edible and less-edible algae and was forced by observed or anticipated irradiance, temperature and vertical mixing intensity. Unexpectedly, irradiance often dominated algal net growth rather than vertical mixing for the following reason: algal dynamics depended on algal net losses from the euphotic layer to larger depth due to vertical mixing. These losses strongly depended on the vertical algal gradient which, in turn, was determined by the mixing intensity during the previous days, thereby introducing a memory effect. This observation implied that during intense mixing that had already reduced the vertical algal gradient, net losses due to mixing were small. Consequently, even in deep Lake Constance, the reduction in primary production due to low light was often more influential than the net losses due to mixing. In the regression model, the dynamics of small, fast-growing algae was best explained by vertical mixing intensity and global irradiance, whereas those of larger algae were best explained by their biomass 1 week earlier. The simulation model additionally revealed that even in late winter grazing may represent an important loss factor during calm periods when losses due to mixing are small. The importance of losses by mixing and grazing changed rapidly as it depended on the variable mixing intensity. Higher temperature, lower global irradiance and enhanced mixing generated lower algal biomass and primary production in the dynamic simulation model. This suggests that potential consequences of climate change may partly counteract each other.
Phenotypic plasticity can increase individual fitness when environmental conditions change over time. Inducible defences are a striking example, allowing species to react to fluctuating predation pressure by only expressing their costly defended phenotype under high predation risk. Previous theoretical investigations have focused on how this affects predator–prey dynamics, but the impact on competitive outcomes and broader community dynamics has received less attention. Here we use a small food web model, consisting of two competing plastic autotrophic species exploited by a shared consumer, to study how the speed of inducible defences across three trade-off constellations affects autotroph coexistence, biomasses across trophic levels, and temporal variability. Contrary to the intuitive idea that faster adaptation increases autotroph fitness, we found that higher switching rates reduced individual fitness as it consistently provoked more maladaptive switching towards undefended phenotypes under high predation pressure. This had an unexpected positive impact on the consumer, increasing consumer biomass and lowering total autotroph biomass. Additionally, maladaptive switching strongly reduced autotroph coexistence through an emerging source-sink dynamic between defended and undefended phenotypes. The striking impact of maladaptive switching on species and food web dynamics indicates that this mechanism may be of more critical importance than previously recognized.
Phenotypic plasticity can increase individual fitness when environmental conditions change over time. Inducible defences are a striking example, allowing species to react to fluctuating predation pressure by only expressing their costly defended phenotype under high predation risk. Previous theoretical investigations have focused on how this affects predator–prey dynamics, but the impact on competitive outcomes and broader community dynamics has received less attention. Here we use a small food web model, consisting of two competing plastic autotrophic species exploited by a shared consumer, to study how the speed of inducible defences across three trade-off constellations affects autotroph coexistence, biomasses across trophic levels, and temporal variability. Contrary to the intuitive idea that faster adaptation increases autotroph fitness, we found that higher switching rates reduced individual fitness as it consistently provoked more maladaptive switching towards undefended phenotypes under high predation pressure. This had an unexpected positive impact on the consumer, increasing consumer biomass and lowering total autotroph biomass. Additionally, maladaptive switching strongly reduced autotroph coexistence through an emerging source-sink dynamic between defended and undefended phenotypes. The striking impact of maladaptive switching on species and food web dynamics indicates that this mechanism may be of more critical importance than previously recognized.
1. The in situ abundance, biomass and mean cell volume of Actinophrys sol (Sarcodina: Heliozoa), the top predator in an extremely acidic German mining lake (Lake 111; pH 2.65), were determined over three consecutive years (spring to autumn, 2001-03). 2. Actinophrys sol exhibited pronounced temporal and vertical patterns in abundance, biomass and mean cell volume. Increasing from very low spring densities, maxima in abundance and biomass were observed in late June/early July and September. The highest mean abundance recorded during the study was 7 x 10(3) Heliozoa L-1. Heliozoan abundance and biomass were higher in the epilimnion than in the hypolimnion. Actinophrys sol cells from this acidic lake were smaller than individuals of the same species found in other aquatic systems. 3. We determined the growth rate of A. sol using all potential prey items available in, and isolated and cultured from, Lake 111. Prey items included: single-celled and filamentous bacteria of unknown taxonomic affinity, the mixotrophic flagellates Chlamydomonas acidophila and Ochromonas sp., the ciliate Oxytricha sp. and the rotifers Elosa worallii and Cephalodella hoodi. Actinophrys sol fed over a wide-size spectrum from bacteria to metazoans. Positive growth was not supported by all naturally available prey. Actinophrys sol neither increased in cell number (k) nor biomass (k(b)) when starved, with low concentrations of single-celled bacteria or with the alga Ochromonas sp. Positive growth was achieved with single- celled bacteria (k = 0.22 +/- 0.02 d(-1); k(b) = -0.06 +/- 0.02 d(-1)) and filamentous bacteria (k = 0.52 +/- < 0.01 d(- 1); k(b) = 0.66 d(-1)) at concentrations greater than observed in situ, and the alga C. acidophila (up to k = 0.43 +/- 0.03 d(-1); k(b) = 0.44 +/- 0.04 d(-1)), the ciliate Oxytricha sp. (k = 0.34 +/- 0.01 d(-1)) and in mixed cultures containing rotifers and C. acidophila (k = 0.23 +/- 0.02-0.32 +/- 0.02 d(-1); maximum k(b) = 0.42 +/- 0.05 d(-1)). The individual- and biomass-based growth of A. sol was highest when filamentous bacteria were provided. 4. Existing quantitative carbon flux models for the Lake 111 food web can be updated in light of our results. Actinophrys sol are omnivorous predators supported by a mixed diet of filamentous bacteria and C. acidophila in the epilimnion. Heliozoa are important components in the planktonic food webs of 'extreme' environments
To test the consequences of decreased diversity and exclusion of keystone species, we compared the planktonic food webs in two acidic (pH <= 3), species-poor mining lakes with those in two species-rich, neutral lakes. The ratio of heterotrophic to autotrophic biomass (HIA) was similar in acidic and neutral lakes with comparable productivity. However, food webs in both acidic lakes were largely restricted to two trophic levels in contrast to the four levels found in neutral lakes. This restriction in food chain length was attributed to the absence of efficient secondary consumers, rather than to productivity or lake size which resulted in unusually low predator-prey weight ratios, with small top predators hardly exceeding their pry in size. In contrast to the neutral lakes, plankton biomass size spectra of acidic lakes were discontinuous due to a lack of major functional groups. The unique size-dependence of feeding modes in pelagic food webs, with bacteria in the smallest size classes followed by autotrophs, herbivores and carnivores, was maintained for bacteria but the other feeding modes strongly overlapped in size. Thus, their characteristic succession along the size gradient was roughly preserved under extreme conditions but the flow of energy along the size gradient was truncated in the acidic lakes. For most but not all attributes studied, differences were larger between acidic and neutral lakes than between neutral lakes of different trophic state
Vertical differences in food web structure were examined in an extremely acidic, iron-rich mining lake in Germany (Lake 111; pH 2.6, total Fe 150mg L-1) during the period of stratification. We tested whether or not the seasonal variation of the plankton composition is less pronounced than the differences observed over depth. The lake was strongly stratified in summer, and concentrations of dissolved organic carbon and inorganic carbon were consistently low in the epilimnion but high in the hypolimnion. Oxygen concentrations declined in the hypolimnion but were always above 2mg L-1. Light attenuation did not change over depth and time and was governed by dissolved ferric iron. The plankton consisted mainly of single-celled and filamentous bacteria, the two mixotrophic flagellates Chlamydomonas sp. and Ochromonas sp., the two rotifer species Elosa worallii and Cephalodella hoodi, and Heliozoa as top predators. We observed very few ciliates and rhizopods, and no heterotrophic flagellates, crustaceans or fish. Ochromonas sp., bacterial filaments, Elosa and Heliozoa dominated in the epilimnion whereas Chlamydomonas sp., single-celled bacteria and Cephalodella dominated in the hypolimnion. Single-celled bacteria were controlled by Ochromonas sp. whereas the lack of large consumers favoured a high proportion of bacterial filaments. The primarily phototrophic Chlamydomas sp. was limited by light and CO2 and may have been reduced due to grazing by Ochromonas sp. in the epilimnion. The distribution of the primarily phagotrophic Ochromonas sp. and of the animals seemed to be controlled by prey availability. Differences in the plankton composition were much higher between the epilimnion and hypolimnion than within a particular stratum over time. The food web in Lake 111 was extremely species-poor enabling no functional redundancy. This was attributed to the direct exclusion of species by the harsh environmental conditions and presumably enforced by competitive exclusion. The latter was promoted by the low diversity at the first trophic level which, in turn, was attributed to relatively stable growth conditions and the independence of resource availability (inorganic carbon and light) from algal density. Ecological theory suggests that low functional redundancy promotes low stability in ecosystem processes which was not supported by our data.
Chlamydomonas acidophila faces high heavy-metal concentrations in acidic mining lakes, where it is a dominant phytoplankton species. To investigate the importance of metals to C. acidophila in these lakes, we examined the response of growth, photosynthesis, cell structure, heat-shock protein (Hsp) accumulation, and metal adsorption after incubation in metal-rich lake water and artificial growth medium enriched with metals (Fe, Zn). Incubation in both metal-rich lake water and medium caused large decreases in photosystem II function (though no differences among lakes), but no decrease in growth rate (except for medium + Fe). Concentrations of small Hsps were higher in algae incubated in metal-rich lake- water than in metal-enriched medium, whereas Hsp60 and Hsp70A were either less or equally expressed. Cellular Zn and Fe contents were lower, and metals adsorbed to the cell surface were higher, in lake-water-incubated algae than in medium- grown cells. The results indicate that high Zn or Fe levels are likely not the main or only contributor to the low primary production in mining lakes, and multiple adaptations of C. acidophila (e.g., high Hsp levels, decreased metal accumulation) increase its tolerance to metals and permit survival under such adverse environmental conditions. Supposedly, the main stress factor present in the lake water is an interaction between low P and high Fe concentrations.
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
Predator-prey oscillations are expected to show a 1/4-phase lag between predator and prey. However, observed dynamics of natural or experimental predator-prey systems are often more complex. A striking but hardly studied example are sudden interruptions of classic 1/4-lag cycles with periods of antiphase oscillations, or periods without any regular predator-prey oscillations. These interruptions occur for a limited time before the system reverts to regular 1/4-lag oscillations, thus yielding intermittent cycles. Reasons for this behaviour are often difficult to reveal in experimental systems. Here we test the hypothesis that such complex dynamical behaviour may result from minor trait variation and trait adaptation in both the prey and predator, causing recurrent small changes in attack rates that may be hard to capture by empirical measurements. Using a model structure where the degree of trait variation in the predator can be explicitly controlled, we show that a very limited amount of adaptation resulting in 10-15% temporal variation in attack rates is already sufficient to generate these intermittent dynamics. Such minor variation may be present in experimental predator-prey systems, and may explain disruptions in regular 1/4-lag oscillations.
Gaining understanding of food-web processes often requires a simplified representation of natural diversity. One such simplification can be based on functional traits, as functionally similar species may provide a similar contribution to ecosystem level-processes. However, understanding how similarity in functional traits actually translates into similar contributions to ecosystem-level properties remains a challenge due to the complex ways in which traits can influence species' dynamics. Moreover, in many communities, seasonality alters the abiotic and biotic forcing regime, causing ongoing changes to patterns of species' dominance; groups of species do not stay intact but are rather continuously subjected to changes throughout the year. Using long-term high frequency measurements of phytoplankton in Lake Constance, we investigated the effect of seasonal changes on the relationship between functional similarity and temporal dynamics similarity of 36 morphotypes, and the relative contribution of different functional traits during the different parts of the year. Our results revealed seasonal differences in the overall degree of synchronization of morphotypes' temporal dynamics and how combinations of functional traits influence the relationship between functional trait similarity and temporal dynamics similarity, showing that different forcing regimes change how species cope with their environment based on their functional traits. Moreover, we show that the individual functional traits matter at different periods of the year indicating that species which are dynamically similar at certain parts of the year may not be at others. The differential strength of the overall and individual impact of functional traits on species' temporal dynamics makes the cohesion of a pair of functionally similar species dependent on the different forcing. Hence, simplifying food webs based solely on functional traits may not provide consistent estimates of functional groups over all seasons.
The ecology of chroococcoid picocyanobacteria was studied from 1987 to 1997 in large, deep, mesotrophic Lake Constance in relation to various abiotic and biotic factors that may influence their population dynamics. Picocyanobacteria dominated the autotrophic picoplankton (APP) numerically in this lake at all depths and times. Their abundances did not respond unequivocally to the decline of winterly phosphorus concentrations by a factor of 2.5 during the decade of investigation. They showed a recurrent seasonal pattern with peaks in spring and late summer, interspersed by a pronounced minimum during and after the clear-water phase around June. The magnitude, timing, and number of peaks and troughs which varied interannually, could in part be related to weather conditions or the impact of other plankton groups. Larger phytoplankton and picocyanobacteria exhibited a distinct and predictable response to the vertical mixing intensity during early spring. Except for 1993, picocyanobacteria and larger phytoplankton decreased simultaneously during the mass development of daphnids in late May or June which gave rise to the clear-water phase. As the daphnid development depends more strongly on surface water temperature than on vertical mixing intensity an early onset of stratification may imply a longer spring development which contributed to a higher seasonal average of picocyanobacterial abundances in 1989-1991. The decline in picocyanobacteria around the clear-water phase was often more pronounced and lasted longer than did the decline in larger algae. The rate of decrease may be related to daphnid abundance, however, no such relationship existed in respect to its duration. Summer peaks of picocyanobacteria were recorded despite the presence of relatively high densities of daphnids. We conclude that with the exception of the clear- water phase, grazing control by nano- and microzooplankton may be more important for controlling picocyanobacterial numbers than is grazing by daphnids. Picocyanobacteria declined in autumn prior to or concomitant with larger algae without any obvious relationship to phytoplankton biovolume or the extent of vertical mixing within the uppermost 20 m. The as yet unexplained variation in the population dynamics of picocyanobacteria points to the significance of species- specific protist grazing and to shifts in picocyanobacterial species composition which should be tackled in future studies.
Ecoevolutionary feedbacks in predator-prey systems have been shown to qualitatively alter predator-prey dynamics. As a striking example, defense-offense coevolution can reverse predator-prey cycles, so predator peaks precede prey peaks rather than vice versa. However, this has only rarely been shown in either model studies or empirical systems. Here, we investigate whether this rarity is a fundamental feature of reversed cycles by exploring under which conditions they should be found. For this, we first identify potential conditions and parameter ranges most likely to result in reversed cycles by developing a new measure, the effective prey biomass, which combines prey biomass with prey and predator traits, and represents the prey biomass as perceived by the predator. We show that predator dynamics always follow the dynamics of the effective prey biomass with a classic 1/4-phase lag. From this key insight, it follows that in reversed cycles (i.e., -lag), the dynamics of the actual and the effective prey biomass must be in antiphase with each other, that is, the effective prey biomass must be highest when actual prey biomass is lowest, and vice versa. Based on this, we predict that reversed cycles should be found mainly when oscillations in actual prey biomass are small and thus have limited impact on the dynamics of the effective prey biomass, which are mainly driven by trait changes. We then confirm this prediction using numerical simulations of a coevolutionary predator-prey system, varying the amplitude of the oscillations in prey biomass: Reversed cycles are consistently associated with regions of parameter space leading to small-amplitude prey oscillations, offering a specific and highly testable prediction for conditions under which reversed cycles should occur in natural systems.
Reversed predator
(2018)
Ecoevolutionary feedbacks in predator–prey systems have been shown to qualitatively alter predator–prey dynamics. As a striking example, defense–offense coevolution can reverse predator–prey cycles, so predator peaks precede prey peaks rather than vice versa. However, this has only rarely been shown in either model studies or empirical systems. Here, we investigate whether this rarity is a fundamental feature of reversed cycles by exploring under which conditions they should be found. For this, we first identify potential conditions and parameter ranges most likely to result in reversed cycles by developing a new measure, the effective prey biomass, which combines prey biomass with prey and predator traits, and represents the prey biomass as perceived by the predator. We show that predator dynamics always follow the dynamics of the effective prey biomass with a classic ¼‐phase lag. From this key insight, it follows that in reversed cycles (i.e., ¾‐lag), the dynamics of the actual and the effective prey biomass must be in antiphase with each other, that is, the effective prey biomass must be highest when actual prey biomass is lowest, and vice versa. Based on this, we predict that reversed cycles should be found mainly when oscillations in actual prey biomass are small and thus have limited impact on the dynamics of the effective prey biomass, which are mainly driven by trait changes. We then confirm this prediction using numerical simulations of a coevolutionary predator–prey system, varying the amplitude of the oscillations in prey biomass: Reversed cycles are consistently associated with regions of parameter space leading to small‐amplitude prey oscillations, offering a specific and highly testable prediction for conditions under which reversed cycles should occur in natural systems.
We investigated the response of the microbial components of the pelagic food web to re-oligotrophication of large, deep Lake Constance where total phosphorus concentrations during mixing decreased from a maximum of 2.81 mu mol L- 1 in 1979 via 1.87 mu mol L-1 in 1987 to 0.26 mu mol L-1 in 2007. Measurements of heterotrophic bacteria, autotrophic picoplankton (APP) and heterotrophic nanoflagellates (HNF) in 2006 and 2007 were compared to values from 1987 to 1997. We hypothesized that the biomass and seasonal variability of all groups will decrease under more oligotrophic conditions due to reduced resource availability, particularly for APP and HNF but less for the competitively stronger bacteria. Average bacterial biomass between spring and autumn was unrelated to phosphorus, whereas the ratio of bacterial biomass to chlorophyll a concentration increased with decreasing trophy due to declining chlorophyll concentrations. In contrast, a unimodal relationship was found between APP and phosphorus with low biomass at low and high phosphorus concentrations and maximum biomass in between. Average HNF biomass decreased strongly by a factor of 10-30 with decreasing trophy, and chlorophyll-specific HNF biomass was unimodally related to phosphorus. The relative seasonal biomass variability did not change for any group during re-oligotrophication. To conclude, HNF responded much more strongly and bacteria less so than chlorophyll concentrations to oligotrophication, whereas APP exhibited a more complex pattern.
Resisting annihilation
(2018)
Allelopathic species can alter biodiversity. Using simulated assemblages that are characterised by neutrality, lumpy coexistence and intransitivity, we explore relationships between within-assemblage competitive dissimilarities and resistance to allelopathic species. An emergent behaviour from our models is that assemblages are more resistant to allelopathy when members strongly compete exploitatively (high competitive power). We found that neutral assemblages were the most vulnerable to allelopathic species, followed by lumpy and then by intransitive assemblages. We find support for our modeling in real-world time-series data from eight lakes of varied morphometry and trophic state. Our analysis of this data shows that a lake's history of allelopathic phytoplankton species biovolume density and dominance is related to the number of species clusters occurring in the plankton assemblages of those lakes, an emergent trend similar to that of our modeling. We suggest that an assemblage's competitive power determines its allelopathy resistance.
Regulation of planktonic ciliate dynamics and functional composition during spring in Lake Constance
(2007)
Protozoans are among the most important grazers of phytoplankton and remineralizers of nutrients in marine and freshwater ecosystems, but less is known about the regulation of their population dynamics. We analyzed a 12 yr data set of ciliate biomass and species composition in large, deep Lake Constance to understand the factors influencing ciliate spring development. The start of ciliate net growth in spring was closely linked to that of edible algae, chlorophyll a and the vertical mixing intensity, but independent of water temperature. During ciliate spring growth, the relative contribution of ciliated interception feeders was positively related to that of cryptomonads, whereas the relative contribution of filter feeders correlated positively with that of non-cryptomonads. The duration of ciliate dominance in spring was largely controlled by the highly variable onset of the phytoplankton bloom, as the termination of the ciliate bloom was less variable. During years with an extended spring bloom of algae and ciliates, internally forced species shifts were observed in both communities. Interception feeders alternated with filter feeders in their relative importance as did cryptomonads and non-cryptomonads. Extended spring blooms were observed when vertical mixing intensity was low at low temperatures during early spring, which will become less likely under the anticipated climate change scenarios. The termination of the ciliate spring bloom occurred prior to a reduction in food concentration and mostly also prior to the mass development of daphnids alone, but coincided with increased grazing by various predators together, such as rotifers, copepods and daphnids in late May/early June.
Seagrass beds are important habitats in coastal areas but increasingly decline in area and quality, thus conservation measures are urgently needed. Quantitative food webs, describing the biomass distribution and energy fluxes among trophic groups, reveal structural and functional aspects of ecosystems. Their knowledge can improve ecological conservation. For the recently discovered large warm-temperate seagrass (Zostera japonica) habitat in China's Yellow River Delta wetland, we used delta C-13 and delta N-15 measurements and a Bayesian isotope mixing model to construct its food web diagram with quantitative estimations of consumer diet compositions, comprising detritus and 14 living trophic groups from primary producers to fish. We then estimated the quantitative food web fluxes based on biomass measurements and calculated corresponding ecosystem functions. Pelagic producers were significantly C-13-depleted compared to benthic sources. Consumers (except zooplankton) were increasingly C-13-depleted with increasing trophic positions even though the consumed benthic production surpassed the pelagic one. Bivalves dominated consumer biomasses and fluxes and were the first to connect the pelagic and benthic pathways, whereas zooplankton and gastropods were specialized on the two pathways, respectively. We found flat biomass and production pyramids indicating low trophic transfer efficiencies. Generally, the energetic structure of the quantitative food web was consistent with the stable isotope analysis, and the estimated net primary production and most estimated production to biomass ratios of the trophic groups fell within literature ranges. This study provides a systematical understanding of the quantitative trophic ecology of a seagrass bed and facilitates synergistic knowledge on management, conservation, and restoration.
Quantifying the capacity for contemporary trait changes to drive intermittent predator-prey cycles
(2022)
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.
Biological invasions are a major threat to natural biodiversity; hence, understanding the mechanisms underlying invasibility (i.e., the susceptibility of a community to invasions by new species) is crucial. Invasibility of a resident community may be affected by a complex but hitherto hardly understood interplay of (1) productivity of the habitat, (2) diversity, (3) herbivory, and (4) the characteristics of both invasive and resident species. Using experimental phytoplankton microcosms, we investigated the effect of nutrient supply and species diversity on the invasibility of resident communities for two functionally different invaders in the presence or absence of an herbivore. With increasing nutrient supply, increased herbivore abundance indicated enhanced phytoplankton biomass production, and the invasion success of both invaders showed a unimodal pattern. At low nutrient supply (i.e., low influence of herbivory), the invasibility depended mainly on the competitive abilities of the invaders, whereas at high nutrient supply, the susceptibility to herbivory dominated. This resulted in different optimum nutrient levels for invasion success of the two species due to their individual functional traits. To test the effect of diversity on invasibility, a species richness gradient was generated by random selection from a resident species pool at an intermediate nutrient level. Invasibility was not affected by species richness; instead, it was driven by the functional traits of the resident and/or invasive species mediated by herbivore density. Overall, herbivory was the driving factor for invasibility of phytoplankton communities, which implies that other factors affecting the intensity of herbivory (e.g., productivity or edibility of primary producers) indirectly influence invasions.
Neglecting the naturally existing functional diversity of communities and the resulting potential to respond to altered conditions may strongly reduce the realism and predictive power of ecological models. We therefore propose and study a predator-prey model that describes mutual feedback via species shifts in both predator and prey, using a dynamic trait approach. Species compositions of the two trophic levels were described by mean functional traits-prey edibility and predator food-selectivity- and functional diversities by the variances. Altered edibility triggered shifts in food-selectivity so that consumers continuously respond to the present prey composition, and vice versa. This trait-mediated feedback mechanism resulted in a complex dynamic behavior with ongoing oscillations in the mean trait values, reflecting continuous reorganization of the trophic levels. The feedback was only possible if sufficient functional diversity was present in both trophic levels. Functional diversity was internally maintained on the prey level as no niche existed in our system, which was ideal under any composition of the predator level due to the trade-offs between edibility, growth and carrying capacity. The predators were only subject to one trade-off between food-selectivity and grazing ability and in the absence of immigration, one predator type became abundant, i.e., functional diversity declined to zero. In the lack of functional diversity the system showed the same dynamics as conventional models of predator-prey interactions ignoring the potential for shifts in species composition. This way, our study identified the crucial role of trade-offs and their shape in physiological and ecological traits for preserving diversity.
Body size has been widely recognised as a key factor determining community structure in ecosystems. We analysed size diversity patterns of phytoplankton, zooplankton and fish assemblages in 13 data sets from freshwater and marine sites with the aim to assess whether there is a general trend in the effect of predation and resource competition on body size distribution across a wide range of aquatic ecosystems. We used size diversity as a measure of the shape of size distribution. Size diversity was computed based on the Shannon-Wiener diversity expression, adapted to a continuous variable, i.e. as body size. Our results show that greater predation pressure was associated with reduced size diversity of prey at all trophic levels. In contrast, competition effects depended on the trophic level considered. At upper trophic levels (zooplankton and fish), size distributions were more diverse when potential resource availability was low, suggesting that competitive interactions for resources promote diversification of aquatic communities by size. This pattern was not found for phytoplankton size distributions where size diversity mostly increased with low zooplankton grazing and increasing nutrient availability. Relationships we found were weak, indicating that predation and competition are not the only determinants of size distribution. Our results suggest that predation pressure leads to accumulation of organisms in the less predated sizes, while resource competition tends to favour a wider size distribution.
Effects of plant community diversity on ecosystem processes have recently received major attention. In contrast, effects of species richness and functional richness on individual plant performance, and their magnitude relative to effects of community composition, have been largely neglected. Therefore, we examined height, aboveground biomass, and inflorescence production of individual plants of all species present in 82 large plots of the Jena Experiment, a large grassland biodiversity experiment in Germany. These plots differed in species richness (1-60), functional richness (1-4), and community composition. On average, in more species-rich communities, plant individuals grew taller, but weighed less, were less likely to flower, and had fewer inflorescences. In plots containing legumes, non-legumes were higher and weighed more than in plots without legumes. In plots containing grasses, non-grasses were less likely to flower than in plots without grasses. This indicates that legumes positively and grasses negatively affected the performance of other species. Species richness and functional richness effects differed systematically between functional groups. The magnitude of the increase in plant height with increasing species richness was greatest in grasses and was progressively smaller in legumes, small herbs, and tall herbs. Individual aboveground biomass responses to increasing species richness also differed among functional groups and were positive for legumes, less pronouncedly positive for grasses, negative for small herbs, and more pronouncedly negative for tall herbs. Moreover, these effects of species richness differed strongly between species within these functional groups. We conclude that individual plant performance largely depends on the diversity of the surrounding community, and that the direction and magnitude of the effects of species richness and functional richness differs largely between species. Our study suggests that diversity of the surrounding community needs to be taken into account when interpreting drivers of the performance of individual plants.
Planktische Räuber-Beute-Systeme : experimentelle Untersuchungen von ökologischen Synchronisationen
(2000)
In Lake Constance, phytoplankton productivity, together with parameters relevant for the production process, was assessed year-round at about 500 dates between 1980 and 1995/1996. During this period, the concentration of total phosphorus during winter circulation decreased from more than 80 to 22 ;g/l as a consequence of sewage diversion and waste water treatment within the catchment area. By contrast, annual photosynthetic rates remained virtually unchanged for about 10 more years following phosphorus decline (mean value 288 " 21 g C m-2 a-1), and thereafter decreased only by about 25 % until 1996. The aim of this study is to analyze factors responsible for this pronounced resilience.
Bacterivory by mixotrophic flagellates may contribute to their nutrient supply, providing a competitive advantage in oligotrophic waters. We hypothesized an increase in Dinobryon biomass during the re-oligotrophication process in the large and deep Lake Constance. To estimate whether bacterivory contributed substantially to the flagellates' phosphorus supply, we determined ingestion rates. Dinobryon biomass increased with decreasing total phosphorus concentrations in the lake over a period of 17 years (P = 0.0005). The promotion of Dinobryon biomass during re-oligotrophication may be explained by the increasing light availability due to the decreasing biomass of other phytoplankton yielding a release from competition. The date of the Dinobryon abundance maximum shifted to earlier time points in the year, probably because a smaller phosphorus pool was depleted more quickly. Ingestion rates of Dinobryon ranged between 0.5 and 13 bacteria cell(-1) h(-1) (0.2-5.4 fg C pg C-1 h(-1)), and clearance rates varied between 0.2 and 3.2 nL cell(-1) h(-1) (4-78 pL pg C-1 h(-1)), leading to bacterial losses of up to 30% day(-1) of bacterial standing stock. The ingestion of bacteria covered 77% of the phosphorus need of the flagellate during the period of maximum growth in 1996 (net growth rate 0.34 day(-1)), and it fully covered the need at all other times.
Chemostat experiments are employed to study predator-prey and other trophic interactions, frequently using phytoplankton-zooplankton systems. These experiments often use population dynamics as fingerprints of ecological and evolutionary processes, assuming that the contributions of all major actors to these dynamics are known. However, bacteria are often neglected although they are frequently present. We argue that even without external carbon input bacteria may affect the experimental outcomes depending on experimental conditions and the physiological traits of bacteria, phytoplankton, and zooplankton. Using a static carbon flux model and a dynamic simulation model, we predict the minimum and maximum impact of bacteria on phytoplankton-zooplankton population dynamics. Under bacteria-suppressing conditions, we find that the effect of bacteria is indeed negligible and their omission justified. Under bacteria-favoring conditions, however, bacteria may strongly affect average biomasses of phytoplankton and zooplankton. The population dynamics may become highly complex, which may result in wrong interpretations when inferring processes (e.g., trait changes) from population dynamic patterns without considering bacteria. We provide suggestions to reduce the bacterial impact experimentally. Besides optimizing experimental conditions (e.g., the dilution rate) the appropriate choice of the zooplankton predator is decisive. Counterintuitively, bacteria have a larger impact if the predator is not bacterivorous as high bacterial biomasses and complex population dynamics arise via competition for nutrients with the phytoplankton. Only at least partial bacterivory minimizes the impact of bacteria. Our results help to improve the design of chemostat experiments and their interpretation, and advance the study of ecological and evolutionary processes in aquatic food webs.
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.
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.