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Biomass-weighted mean traits of a community's constituent species are a useful tool to assess environmental filtering in community function in response to environmental change. We show that annually averaged phytoplankton community function, expressed by the community mean traits phosphate and light affinity, responded strongly and reversibly to long-term changes in nutrient supply over a 42-year period of eutrophication and re-oligotrophication of Lake Constance. Within the lake's species pool, phosphate and light affinities were weakly negatively correlated, suggesting a weak physiological trade-off. Yet, a strong trade-off between these traits emerged when species were weighted by their biomass, suggesting species sorting along the trade-off line across years of shifting nutrient status. Emergent trade-offs, that is, trade-offs that become apparent first when trait combinations are weighted by the contributions of the trait-bearing organisms to community biomass, may be a useful, novel concept in trait-based ecology of potentially similar importance as commonly considered physiological trade-offs.

Within aquatic ecosystems, heterotrophic, mixotrophic and autotrophic plankton are entangled in a complex network of competitive, predatory and mutualistic interactions. "Browning," the increase of colored dissolved organic matter (CDOM) from terrestrial catchments, can affect this network of interactions by simultaneously decreasing light availability and increasing organic carbon and nutrients supplies. Here, we introduce a conceptual, process-based numerical model to investigate the effects of browning on a microbial food web consisting of heterotrophic bacterioplankton, bacterivorous phago-mixoplankton, autotrophic phytoplankton and the resources light, inorganic phosphorus and DOM. Additionally, we explore how the investment in autotrophic vs. phagotrophic resource acquisition influences mixoplankton performance. Several model predictions are in broad agreement with empirical observations under increasing CDOM supply, including increased bacterial biomass and inorganic phosphorous, decreased light penetration, the potential for a unimodal phytoplankton biomass response and a local minimum in mixoplankton biomass. Our results also suggest that mixoplankton with a high investment in phototrophy perform best in many conditions but that phosphorous acquisition via prey is crucial under high light-low nutrient conditions. Overall, our model analyses suggest that responses to altered CDOM supply are largely determined by systematic changes in the relative importance of nutrient vs. energy limitation of each plankton group.

A vertical separation in light and nutrient availability is observed in many terrestrial and aquatic ecosystems. In lakes and oceans, the opposing vertical gradients of light and nutrients typically observed are believed to promote phagomixotrophy, a generalist strategy that combines resource acquisition through photoautotrophic and phagoheterotrophic pathways. While phagomixotrophy is widespread, it is not well understood how this strategy performs against pure specialist strategies in a resource competition context. We simulate the dynamics of three competitors (pure photoautotroph, phagomixotroph, pure phagoheterotroph) and bacterial prey over the vertical dimension of a water column to investigate what conditions of resource availability favor mixotrophy and how the presence of the phagomixotroph alters community dynamics. Since mixotrophs can be more or less photoautotrophic, we incorporated this variability into our model. Under weak vertical mixing, mixotrophs persist under most light and nutrient conditions and negatively affect specialists. Mixotrophs can even be dominant competitors when they display an optimal degree of phototrophy, which is positively related to water transparency and negatively related to nutrient supply. The model indicates that the spatial organization of nanophytoplankton communities in water columns could arise through vertical niche partitioning of multiple resource acquisition strategies and that phagomixotrophy can promote overall community production.

Une séparation verticale de la disponibilité de la lumière et des nutriments peut être observée dans de nombreux écosystèmes terrestres et aquatiques. Dans les lacs et les océans, on observe des gradients verticaux opposés de lumière et de nutriments qui sont considérés comme des facteurs favorisant la phago-mixotrophie, une stratégie de nutrition généraliste qui combine la photo-autotrophie et la phago-hétérotrophie pour l'acquisition des ressources. Bien que la phago-mixotrophie soit répandue, les performances de cette stratégie dans le cadre de la compétition pour les ressources contre des stratégies spécialisées sont mal comprises. Nous avons simulé la dynamique de trois compétiteurs (photo-autotrophe pur, phago-mixotrophe, phago-hétérotrophe pur) et de bactéries sur la dimension verticale d'une colonne d’eau pour étudier quelles conditions de disponibilité des ressources favorisent la mixotrophie et comment la présence d'un phago-mixotrophe modifie la dynamique de la communauté. Les mixotrophes pouvant être plus ou moins photo-autotrophes, nous avons intégré cette variabilité fonctionnelle dans notre modèle. Quand la colonne d'eau est peu mélangée, les mixotrophes subsistent dans la plupart des conditions de lumière et de nutriments et affectent négativement les spécialistes. Les mixotrophes peuvent même dominer la communauté lorsque leur degré de phototrophie, qui est positivement lié à la transparence de l'eau et négativement à l'apport de nutriments, est optimal. Le modèle montre que l'organisation spatiale des communautés nanophytoplanctoniques dans les colonnes d'eau pourrait résulter de la partition verticale des niches de compétiteurs présentant des stratégies d'acquisition de ressources contrastées, et que la phago-mixotrophie peut favoriser la production globale de la communauté.

Climate warming alters the seasonal timing of biological events. This raises concerns that species-specific responses to warming may de-synchronize co-evolved consumer-resource phenologies, resulting in trophic mismatch and altered ecosystem dynamics. We explored the effects of warming on the synchrony of two events: the onset of the phytoplankton spring bloom and the spring/summer maximum of the grazer Daphnia. Simulation of 16 lake types over 31 years at 1907 North African and European locations under 5 climate scenarios revealed that the current median phenological delay between the two events varies greatly (20–190 days) across lake types and geographic locations. Warming moves both events forward in time and can lengthen or shorten the delay between them by up to ±60 days. Our simulations suggest large geographic and lake-specific variations in phenological synchrony, provide quantitative predictions of its dependence on physical lake properties and geographic location and highlight research needs concerning its ecological consequences.

In many ecosystems, consumers respond to warming differently than their resources, sometimes leading to temporal mismatches between seasonal maxima in consumer demand and resource availability. A potentially equally pervasive, but less acknowledged threat to the temporal coherence of consumer-resource interactions is mismatch in food quality. Many plant and algal communities respond to warming with shifts toward more carbon-rich species and growth forms, thereby diluting essential elements in their biomass and intensifying the stoichiometric mismatch with herbivore nutrient requirements. Here we report on a mesocosm experiment on the spring succession of an assembled plankton community in which we manipulated temperature (ambient vs. +3.6°C) and presence versus absence of two types of grazers (ciliates and Daphnia), and where warming caused a dramatic regime shift that coincided with extreme stoichiometric mismatch. At ambient temperatures, a typical spring succession developed, where a moderate bloom of nutritionally adequate phytoplankton was grazed down to a clear-water phase by a developing Daphnia population. While warming accelerated initial Daphnia population growth, it speeded up algal growth rates even more, triggering a massive phytoplankton bloom of poor food quality. Consistent with the predictions of a stoichiometric producer–grazer model, accelerated phytoplankton growth promoted the emergence of an alternative system attractor, where the extremely low phosphorus content of the abundant algal food eventually drove Daphnia to extinction. Where present, ciliates slowed down the phytoplankton bloom and the deterioration of its nutritional value, but this only delayed the regime shift. Eventually, phytoplankton also grew out of grazer control in the presence of ciliates, and the Daphnia population crashed. To our knowledge, the experiment is the first empirical demonstration of the “paradox of energy enrichment” (grazer starvation in an abundance of energy-rich but nutritionally imbalanced food) in a multispecies phytoplankton community. More generally, our results support the notion that warming can exacerbate the stoichiometric mismatch at the plant–herbivore interface and limit energy transfer to higher trophic levels.

A key phenological event in the annual cycle of many pelagic ecosystems is the onset of the spring algal bloom (OAB). Descriptions of the factors controlling the OAB in temperate to polar lakes have been limited to isolated studies of single systems and conceptual models. Here we present a validated modelling approach that, for the first time, enables a quantitative prediction of the OAB and a systematic assessment of the processes controlling its timing on a continental scale. We used a weather-driven, one-dimensional lake model to simulate the seasonal dynamics of the underwater light climate in 16 lake types characterized by the factorial combination of four lake depths with four levels of water transparency. We did so at 1962 locations across Western Europe and over 31 years (1979–2009). Assuming that phytoplankton production is light-limited in winter, we identified four patterns of OAB control across lake types and climate zones. OAB timing is controlled by (i) the timing of ice-off in ice-covered clear or shallow lakes, (ii) the onset of thermal stratification in sufficiently deep and turbid lakes and (iii) the seasonal increase in incident radiation in all other lakes, except for (iv) ice-free, shallow and clear lakes in the south, where phytoplankton is not light-limited. The model predicts that OAB timing should respond to two pervasive environmental changes, global warming and browning, in opposite ways. OAB timing should be highly sensitive to warming in lakes where it is controlled by either ice-off or the onset of stratification, but resilient to warming in lakes where it is controlled by incident radiation. Conversely, OAB timing should be most sensitive to browning where it is controlled by incident radiation, but resilient to browning where it is controlled by ice-off or the onset of stratification. Available lake data are consistent with our findings.

Describing how resource consumption rates depend on resource density, conventionally termed "functional responses," is crucial to understanding the population dynamics of trophically interacting organisms. Yet, accurately determining the functional response for any given pair of predator and prey remains a challenge. Moreover, functional responses are potentially complex surfaces in multidimensional space, where resource density is only one of several factors determining consumption rates. We explored how three sources of error can be addressed in the design and statistical analysis of functional response experiments: ill-chosen spacing of prey densities, heteroscedastic variance in consumption rates, and non-independence of parameters of the function describing prey consumption in relation to prey density and additional environmental factors. We generated extensive, virtual data sets that simulated feeding experiments in which both prey density and environmental temperature were varied, and for which the true, deterministic functional response surface was known and realistic variance had been added. We compared eight different methods of functional response fitting, one of which stood out as best performing. We subsequently tested several conclusions from the simulation study against experimental data of zooplankton feeding on algae across a broad range of temperatures. We summarize our main findings in three best practice guidelines for the experimental estimation of functional response surfaces, of which the second is the most important: (1) space prey densities logarithmically, starting from very low densities; (2) log-transform prey consumption data prior to fitting; and (3) fit a multivariate functional response surface to all data (including all prey densities and other factors, in our case temperature) in a single step. We also observed that functional response surfaces were fitted more accurately and precisely than their component parameters. The latter occurred because parameter estimates were non-independent, which is an inevitable feature of fitting complex nonlinear functions to data: A given response surface can often be described with near-equal accuracy by multiple parameter combinations. We therefore conclude that fitted functional response models perform better at optimizing the fit of the overall response surface than at determining how component parameters, such as the attack rate or handling time, depend on environmental factors such as temperature.

The productivity and trophic structure of aquatic ecosystems is the result of an interplay between bottom-up and top-down forces that operate both within and across the benthic and pelagic compartments of lake food webs. Contemporary and projected climate changes urge the question how this interplay will be affected by increasing inputs of terrestrial derived, dissolved organic matter (‘browning’) and warming. We addressed this issue by exploring how browning and warming affect the behavior of a relatively simple, conceptual model of a shallow lake food web that is compartmentalized into, dynamically coupled, benthic and pelagic components (abiotic resources, primary producers, grazers, and carnivores). We compared model expectations with the results of a factorial manipulation of browning and warming in a replicated, large-scale field experiment. Both the model and the experiment suggest that browning affects the food web from the bottom-up by reducing light supply to the benthic habitat and increasing nutrient supply to the pelagic habitat, with concomitant decreases of benthic and increases of pelagic primary and secondary production. The model also predicts that warming effects should primarily operate via relaxed top-down control by top consumers in the more productive of the two habitats. The latter was only partially supported by the experimental data, possibly because the model still lacks one or two important trophic links, such as the one from pelagic producers to benthic deposit feeders. We propose that our coupled benthic-pelagic food web model provides a useful conceptual starting point for future theoretical and empirical studies of the impacts of environmental changes on shallow lakes.

Spatial environmental heterogeneity coupled with dispersal can promote ecological persistence of diverse metacommunities. Does this premise hold when metacommunities evolve? Using a two-resource competition model, we studied the evolution of resource-uptake specialisation as a function of resource type (substitutable to essential) and shape of the trade-off between resource uptake affinities (generalist- to specialist-favouring). In spatially homogeneous environments, evolutionarily stable coexistence of consumers is only possible for sufficiently substitutable resources and specialist-favouring trade-offs. Remarkably, these same conditions yield comparatively low diversity in heterogeneous environments, because they promote sympatric evolution of two opposite resource specialists that, together, monopolise the two resources everywhere. Consumer diversity is instead maximised for intermediate trade-offs and clearly substitutable or clearly essential resources, where evolved metacommunities are characterised by contrasting selection regimes. Taken together, our results present new insights into resource-competition-mediated evolutionarily stable diversity in homogeneous and heterogeneous environments, which should be applicable to a wide range of systems.

Disentangling the process information contained in a diatom sediment signature is crucial for reliable future predictions based on paleolimnological records. In this study, we combine limnological and paleolimnological monitoring to address the fundamental question: Which environmental information is contained in a diatom sediment signal? We compared annual diatom sequential sediment trap records with the diatom record of the annually varved lake sediment of Nylandssjon (northern Sweden) from three meteorologically different years (2012-2014). The seasonal patterns in diatom sedimentation were strikingly different in varve years 2012 and 2014 compared to varve year 2013. In 2012 and 2014, up to 70% of the annual flux occurred in a single spring month and was dominated by Cyclotella glomerata. In contrast, in 2013, peak fluxes were much lower and more annually integrated. Next, we compared the full-year diatom trap results with year round in-lake physical, chemical, and biological monitored parameters, as well as meteorological variables. Annual averages of environmental conditions did not explain the interannual variability in diatom sedimentation. Instead, the seasonality of diatom sedimentation was determined by the timing of the spring diatom bloom relative to lake over-turn in winters with warm vs. cold air temperature. With our combined limnological and paleolimnological monitoring approach, we find that an annual diatom signal can either contain primarily seasonal climate information from a short time period or be annually integrated. We synthesize our results in a novel conceptual model, which describes the response of sediment diatom signals to two distinct sequences of late-winter conditions.