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Relationships between biomass and ecological factors including trophic interactions were examined to understand the dynamics of six fish species in Lake Võrtsjärv, a large shallow eutrophic lake located in Estonia (north-eastern Europe). The database contained initially 31 predictive variables that were monitored in situ for nearly forty years. The strongest predictive variables were selected by three parallel approaches: single correlation (Pearson), a multivariate method (Co-inertia analyses), and a machine learning algorithm (Random Forests), followed by a Generalized Least Squares model to determine meaningful relationships with fish biomass. Models with both additive and interactive effects were constructed. The results revealed that the indicators of degraded ecological conditions (high cyanobacteria biomass and their proportion in total phytoplankton, high summer temperature, high nutrient concentration) were negatively correlated to fish biomass. Benthic macroinvertebrates and other biotic predictors (biomass of specific fish prey and predators) were also important contributors to fish biomass dynamics. Together, abiotic and biotic factors explained 40–60% of the variance of fish biomass, depending of the species. Our findings suggest that both abiotic and biotic factors control fish biomass changes in this eutrophic lake. This research was funded by Estonian Research Council grants (PSG 32), institutional research funding IUT 21-2 of the Estonian Ministry of Education and Research and by European Union Regional Development Fund program DoRa+ and ASTRA. The authors are thankful to Dr. Külli Kangur (Estonian University of Life Sciences) for participating in the sampling effort of benthic macroinvertebrates. This research was funded by Estonian Research Council grants (PSG 32), institutional research funding IUT 21-2 of the Estonian Ministry of Education and Research and by European Union Regional Development Fund program DoRa+ and ASTRA. The authors are thankful to Dr. Külli Kangur (Estonian University of Life Sciences) for participating in the sampling effort of benthic macroinvertebrates.

We numerically explored the effects of long-term water level changes on biotic biomass and spatial distribution of fish in a large shallow lake. We calibrated Ecospace model (Ecopath with Ecosim modelling suite) with data from various functional groups (ranging from phytoplankton to piscivorous fish), and considered 14 different habitats. Two scenarios representing, respectively, a long-term water-level increase and decrease by 1 m were constructed and run for a period of thirty eight years (1979–2016). The results showed a very uneven spatial distribution of fish biomass in the lake, with the highest con- centration in the southern basin. The 1 m decrease scenario caused a diminution in the biomass of all groups but piscivorous fish. The 1 m increase scenario saw a weak decrease in most species biomass. Consequently, in both scenarios, long-term water level changes would be generally detrimental to the lake biota. In the context of more frequent climate-induced hydrological fluctuations, we encourage the use of these simulations as effective tools for future prediction and assessment of ecosystem-based fisheries management and ecological status maintenance of shallow lakes. This research was supported by Estonian Research Council grant PSG32 to FC and UB, by Environmental Investment Center project-T180209PKKH, by IUT 21-2 of the Estonian Ministry of Education and by European Union Regional Development Fund program DoRa+ and ASTRA. This research was supported by Estonian Research Council grant PSG32 to FC and UB, by Environmental Investment Center project-T180209PKKH, by IUT 21-2 of the Estonian Ministry of Education and by European Union Regional Development Fund program DoRa+ and ASTRA.

We aimed to investigate the influence of environmental factors and predict zooplankton biomass and abundance in shallow eutrophic lakes. We employed time series of zoo- plankton and environmental parameters that were measured monthly during 38 years in a large, shallow eutrophic lake in Estonia to build estimates of zooplankton community metrics (cladocerans, copepods, rotifers, ciliates). The analysis of historical time series revealed that air temperature was by far the most important variable for explaining zooplankton biomass and abundance, followed, in decreasing order of importance, by pH, phytoplankton biomass and nitrate concentration. Models constructed with the best predicting variables explained up to 71% of zooplankton biomass variance. Most of the predictive variables had opposing or antagonistic interactions, often mitigating the effect of temperature. In the second part of the study, three future climate scenarios were developed following different Intergovernmental Panel on Climate Change (IPCC) tem- perature projections and entered into an empirical model. Simulation results showed that only a scenario in which air temperature stabilizes would curb total metazooplankton biomass and abundance. In other scenarios, metazooplankton biomass and abundance would likely exceed historical ranges whereas ciliates would not expand. Within the metazooplankton community, copepods would increase in biomass and abundance, whereas cladocerans would lose in biomass but not in abundance. These changes in the zooplankton community will have important consequences for lake trophic structure and ecosystem functioning. This research was supported by the Estonian Research Council Grants PSG32, PRG709 and institutional research funding IUT 21-2 of the Estonian Ministry of Education and Research. This research was supported by the Estonian Research Council Grants PSG32, PRG709 and institutional research funding IUT 21-2 of the Estonian Ministry of Education and Research.

1. While 13 C-depleted carbon derived from biogenic methane can substantially contribute to the benthic secondary production in deep stratified lakes, its role in shallow lakes is less clear. We investigated the dynamics of δ13 C and δ15 N in the larvae of Chironomus plumosus throughout an annual cycle in two ecologically distinct basins (open-water plankton-dominated and sheltered macrophyte- covered) of a large (270 km2 ), shallow, polymictic and eutrophic lake (Võrtsjärv, Estonia, North Europe). The larval stable isotopic compositions were linked to the presence of methane-oxidising bacteria (MOB) in larval guts and sediments. 2. Molecular detection of MOB revealed their presence in various sediment types, but stable isotope (SI) analysis revealed clear differences in the feeding of chironomid larvae between the plankton- and macrophyte-dominated habitats. 3. In the plankton-dominated habitat, the mean δ13 C values of larvae remained relatively constant (−38.3‰ to −35.5‰) and corresponded closely to the sediment δ13 C values. Mean δ13 C values of chironomid larvae were generally lower in macrophyte-dominated habitats (−43.4‰ to −33.0‰), and both seasonal and individual variation in larval δ13 C values were more pronounced. MOB presence in larval guts proved a dietary contribution from biogenic methane in macrophyte-dominated habitats. Both the SI and molecular results indicated that MOB could help support larvae even during the cold temperature-limited and ice- covered periods. 4. Our study indicates that methane-derived carbon makes a low but steady contri- bution to the larval chironomids throughout an annual cycle in large shallow Võrtsjärv. However, this contribution can be substantially higher in the lake habitats with abundant macrophytes. The study provides further evidence that a carbon flow pathway from biogenic methane can contribute to the benthic food web under variable habitat conditions in a shallow polymictic lake. Funding information: Estonian Research Council, Grant/Award Number: IUT 21-02, PUT 134, PUT 1389 Funding information: Estonian Research Council, Grant/Award Number: IUT 21-02, PUT 134, PUT 1389

The coexistence of potentially toxic bloom-forming cyanobacteria (CY) and generally smaller-sized grazer communities has raised the question of zooplankton (ZP) ability to control harmful cyanobacterial blooms and highlighted the need for species-specific research on ZP-CY trophic interactions in naturally occurring communities. A combination of HPLC, molecular and stable isotope analyses was used to assess in situ the importance of CY as a food source for dominant crustacean ZP species and to quantify the grazing on potentially toxic strains of Microcystis during bloom formation in large eutrophic Lake Peipsi (Estonia). Aphanizomenon, Dolichospermum, Gloeotrichia and Microcystis dominated bloom-forming CY, while Microcystis was the major genus producing cyanotoxins all over the lake. Grazing studies showed that CY, and especially colonial CY, formed a significant, and also preferred component of algae ingested by the cladocerans Bosmina spp. and Daphnia spp. while this was not the case for the more selective calanoid copepod Eudiaptomus gracilis. Molecular analyses confirmed the presence of CY, including Microcystis, in ZP guts. Further analyses using qPCR targeting cyanobacterial genusspecific mcyE synthase genes indicated that potentially toxic strains of Microcystis can be ingested directly or indirectly by all the dominant crustacean grazers. However, stable isotope analyses indicated that little, if any, assimilation from ingested bloom-forming CY occurred. The study suggests that CY, and particularly Microcystis with both potentially toxic and non-toxic strains, can be widely ingested by cladoceran grazers during a bloom event with implications for control of CY abundance and for transfer of CY toxins through the food web. Funding for this research was provided by institutional research funding IUT 21-2 of the Estonian Ministry of Education and Research and by base-financed project P180023PKKH of Estonian University of Life Sciences. The authors are grateful to Tuula Sinisalo and Mikko Kiljunen for assistance with the stable isotope analyses at Jyväskylä University. COST Action ES 1105 “CYANOCOST – Cyanobacterial blooms and toxins in water resources: Occurrence, impacts and management” is acknowledged for networking and sharing of knowledge. [CG] Funding for this research was provided by institutional research funding IUT 21-2 of the Estonian Ministry of Education and Research and by base-financed project P180023PKKH of Estonian University of Life Sciences. The authors are grateful to Tuula Sinisalo and Mikko Kiljunen for assistance with the stable isotope analyses at Jyväskylä University. COST Action ES 1105 “CYANOCOST – Cyanobacterial blooms and toxins in water resources: Occurrence, impacts and management” is acknowledged for networking and sharing of knowledge. [CG]

With increasing primary productivity, ciliates may become the most important members of the microbial loop and form a central linkage in the transformation of microbial production to upper trophic levels. How metazooplankters, especially copepods, regulate ciliate community structure in shallow eutrophic waters is not completely clear. We carried out mesocosm experiments with different cyclopoid copepod enrichments in a shallow eutrophic lake to examine the responses of ciliate community structure and abundance to changes in cyclopoid copepod biomass and to detect any cascading effects on bacterioplankton and edible phytoplankton. Our results indicate that an increase in copepod zooplankton biomass favours the development of small-sized bacterivorous ciliates. This effect is unleashed by the decline of predaceous ciliate abundance, which would otherwise graze effectively on the small-sized ciliates. The inverse relationship between crustacean zooplankton and large predaceous ciliates is an important feature adjusting not only the structure of the ciliate community but also the energy transfer between meta- and protozooplankton. Still we could not detect any cascading effects on bacterio- or phytoplankton that would be caused by the structural changes in the ciliate community.

The study of lake sediments and archived biological remains is a promising approach to better under- stand the impacts of climate change on aquatic ecosystems. Small lakes have been shown to be strongly sensitive to past climate change, but similar information is lacking for large lakes. By identifying re- sponses to climate change of carbon flows through benthic food web in two different sized lakes, we aimed to understand how lake morphometry can mediate the effects of climate change. We recon- structed the dynamics of phytoplankton community composition and carbon resources sustaining chironomid biomass during the Holocene from the combined analysis of sedimentary pigment quanti- fication and carbon stable isotopic composition of subfossil chironomid head capsules (d13 C HC) in a large lake in the Baltic area (Estonia). Our results showed that chironomid biomass in the large lake was mainly sustained by phytoplankton, with no significant relationship between d13 C HC values and tem- perature fluctuations. We suggest that lake morphometry (including distance of the sampling zone to the shoreline, and lake volume for primary producers) mediates the effects of climate change, making large lakes less sensitive to climate change. Complementary studies are needed to better understand differ- ences in organic matter dynamics in different sized lakes and to characterize the response of the aquatic carbon cycle to past climate change. Financial support was provided by institutional research grants IUT1-8 and IUT21-2 from the Estonian Science Agency and by Estonian University of Life Sciences, project No. 8M160101PKLJ. We are grateful to Christian Hossann (INRA Nancy, Champenoux) for assistance in carbon stable isotope analysis. The PTEF facility is supported by the French National Research Agency through the Laboratory of Excellence ARBRE (ANR-11-LABX-0002-01). We acknowledge the contributions of Atko Heinsalu, Siim Veski and Jüri Vassiljev for coring and sediment dating. We also thank G. L. Simpson for valuable comments on statistical analyses. Editing services were provided by Sea Pen Scientific Writing. Financial support was provided by institutional research grants IUT1-8 and IUT21-2 from the Estonian Science Agency and by Estonian University of Life Sciences, project No. 8M160101PKLJ. We are grateful to Christian Hossann (INRA Nancy, Champenoux) for assistance in carbon stable isotope analysis. The PTEF facility is supported by the French National Research Agency through the Laboratory of Excellence ARBRE (ANR-11-LABX-0002-01). We acknowledge the contributions of Atko Heinsalu, Siim Veski and Jüri Vassiljev for coring and sediment dating. We also thank G. L. Simpson for valuable comments on statistical analyses. Editing services were provided by Sea Pen Scientific Writing.

Lentic ecosystems act as sentinels of climate change, and evidence exists that their sensitivity to warming varies along a latitudinal gradient. We assessed the effects of nutrient and water level variability on zooplankton community composition, taxonomic diversity and size structure in different climate zones by running a standardised controlled 6-months (May to November) experiment in six countries along a European north-south latitudinal temperature gradient. The mesocosms were established with two different depths and nutrient levels. We took monthly zooplankton samples during the study period and pooled a subsample from each sampling to obtain one composite sample per mesocosm. We found a significant effect of temperature on the community composition and size structure of the zooplankton, whereas no effects of water depth or nutrient availability could be traced. The normalised size spectrum became flatter with increasing temperature reflecting higher zooplankton size diversity due to higher abundance of calanoid copepods, but did not differ among depths or nutrient levels. Large-bodied cladocerans such as Daphnia decreased with temperature. Taxonomic diversity was positively related to size diversity, but neither of the two diversity measures demonstrated a clear pattern along the temperature gradient nor with nutrient and water levels. However, genus richness decreased at the warm side of the temperature gradient. Our experiment generally supports recent empirically based findings that a continuing temperature increase may result in lower genus richness and lower abundance of large-sized zooplankton grazers, the latter likely resulting in reduced control of phytoplankton.

Small-bodied cladocerans and cyclopoid copepods are becoming increasingly dominant over large crustacean zooplankton in eutrophic waters where they often coexist with cyanobacterial blooms. However, relatively little is known about their algal diet preferences. We studied grazing selectivity of small crustaceans (the cyclopoid copepods Mesocyclops leuckarti, Thermocyclops oithonoides, Cyclops kolensis, and the cladocerans Daphnia cucullata, Chydorus sphaericus, Bosmina spp.) by liquid chromatographic analyses of phytoplankton marker pigments in the shallow, highly eutrophic Lake Võrtsjärv (Estonia) during a seasonal cycle. Copepods (mainly C. kolensis) preferably consumed cryptophytes (identified by the marker pigment alloxanthin in gut contents) during colder periods, while they preferred small non-filamentous diatoms and green algae (identified mainly by diatoxanthin and lutein, respectively) from May to September. All studied cladoceran species showed highest selectivity towards colonial cyanobacteria (identified by canthaxanthin). For small C. sphaericus, commonly occuring in the pelagic zone of eutrophic lakes, colonial cyanobacteria can be their major food source, supporting their coexistence with cyanobacterial blooms. Pigments characteristic of filamentous cyanobacteria and diatoms (zeaxanthin and fucoxanthin, respectively), algae dominating in Võrtsjärv, were also found in the grazers' diet but were generally avoided by the crustaceans commonly dominating the zooplankton assemblage. Together these results suggest that the co-occurring small-bodied cyclopoid and cladoceran species have markedly different algal diets and that the cladocera represent the main trophic link transferring cyanobacterial carbon to the food web in a highly eutrophic lake.

We modeled energy transfer and trophic position of fish, plankton, and macroinvertebrates and the relative importance of top-down versus bottom-up processes in Lake Võrtsjärv, a large shallow eutrophic lake in Estonia (northeastern Europe). We employed input values based on 37 years of biomass and fishing activity monitoring for calibrating the Ecopath with Ecosim (EwE) model. Energy flows from primary producers and detritus, represented by total system throughput, were nearly equal (51 and 49%, respec- tively). Simulation revealed that top-down and bot- tom-up forces were at play, metazooplankton was not efficiently grazing phytoplankton production, and a trophic cascade proceeded through macroinverte- brates rather than through zooplankton. Detritivory was responsible for the relatively low trophic position of Võrtsjärv fish compared to other lakes. Bottom-up processes were the main drivers for the dual, primary production- and detritus-based pathways in energy flow. Our findings suggest that the predicted biomass increase of cyanobacteria in shallow lakes in the future will strengthen the reliance of consumers on the detrital pathway at the expense of the primary production pathway. This research was supported by the Start-Up Personal Research Grant PUT 777 to FC and UB, by IUT 21-2 of the Estonian Ministry of Education and Research, and by MARS project (Managing Aquatic ecosystems and water Resources under multiple Stress) funded under the 7th EU Framework Programme, Theme 6 (Environment including Climate Change), Contract No.: 603378 (http://www.mars-project.eu). This research was supported by the Start-Up Personal Research Grant PUT 777 to FC and UB, by IUT 21-2 of the Estonian Ministry of Education and Research, and by MARS project (Managing Aquatic ecosystems and water Resources under multiple Stress) funded under the 7th EU Framework Programme, Theme 6 (Environment including Climate Change), Contract No.: 603378 (http://www.mars-project.eu).