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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.

Efficient zooplankton grazing is a prerequisite for establishing a cascading food web control over phytoplankton in a lake. We studied if the top-down impact of fish could reach phytoplankton in a lake where the grazing pressure of small-sized zooplankton on filamentous phytoplankton is considered weak. We analysed >30 years of data on plankton, fish catches, hydrochemistry, hydrology, and meteorology from Vortsjarv, a large and shallow eutrophic lake in Estonia with intensive commercial fisheries. The lake’s unregulated water level has been considered the strongest factor affecting the ecosystem through modifying sediment resuspension, internal loading of nutrients, and underwater light conditions and spawning conditions for fish. We found a negative relationship between phytoplankton biomass and pikeperch biomass indicating a potential top-down cascading effect in the food web. Top-down control of phytoplankton by zooplankton was reflected in a negative relationship between phyto- and zooplankton biomasses. A decrease of the individual weight of crustacean zooplankton with increasing biomass of small fish suggested top-down control of zooplankton by planktivorous fish. In contrast, we could not demonstrate a direct linkage between piscivorous fish and small fish. The top-down food web impact of piscivores, however, was manifested at zooplankton level in a positive correlation of pikeperch biomass with the biomass of dominating cladoceran species Bosmina coregoni and the individual weight of copepods. At high biomasses of small fish, ciliate domination over metazooplankton increased and thus enhanced the strength of the microbial food web. According to our results, fishery management measures that increase small plankti- and benthivorous fish biomass have to be avoided as they have a cascading negative effect on the ecosystem health.

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.

We analysed changes in the abundance, biomass and cell size of the microbial food web community (bacteria, heterotrophicnanoflagellates, ciliates) at contrasting nutrient concentrations and temperatures during a simulated heat wave. We used 24mesocosms mimicking shallow lakes in which two nutrient levels (unenriched and enriched by adding nitrogen and phosphorus)and three different temperature scenarios (ambient, IPCC A2 scenario and A2+%50) are simulated (4 replicates of each).Experiments using the mesocosms have been running un-interrupted since 2003. A 1-month heat wave was imitated by anextra 5◦C increase in the previously heated mesocosms (from 1st July to 1st August 2014). Changes in water temperatureinduced within a few days a strong effect on the microbial food web functioning, demonstrating a quick response of microbialcommunities to the changes in environment, due to their short generation times. Warming and nutrients showed synergisticeffects. Microbial assemblages of heterotrophic nanoflagellates and ciliates responded positively to the heating, the increasebeing largest in the enriched mesocosms. The results indicate that warming and nutrients in combination can set off complexinteractions in the microbial food web functioning. This work was supported by MARS project (ManagingAquatic ecosystems and water Resources under multipleStress, EU 7th Framework Programme, contract no.: 603378)and by institutional grant IUT 21-2 of the Estonian Min-istry of Education and Research. The Danish group wasalso supported by PROGNOS (Predicting in-lake RespOnsesto chanGe using Near real time mOdelS – Water JointProgramme Initiative) and AQUACOSM (Network of Lead-ing European AQUAtic MesoCOSM Facilities ConnectingMountains to Oceans from the Arctic to the Mediterranean),the latter also supporting the Turkish group. Yu Cao issupported by NSF-China (31500296) and CSC (The ChinaScholarship Council). This work was supported by MARS project (ManagingAquatic ecosystems and water Resources under multipleStress, EU 7th Framework Programme, contract no.: 603378)and by institutional grant IUT 21-2 of the Estonian Min-istry of Education and Research. The Danish group wasalso supported by PROGNOS (Predicting in-lake RespOnsesto chanGe using Near real time mOdelS – Water JointProgramme Initiative) and AQUACOSM (Network of Lead-ing European AQUAtic MesoCOSM Facilities ConnectingMountains to Oceans from the Arctic to the Mediterranean),the latter also supporting the Turkish group. Yu Cao issupported by NSF-China (31500296) and CSC (The ChinaScholarship Council).

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.

The feeding impact of planktivorous fish on microbial organisms is still poorly understood. We followed the seasonal dynamics of the food web in two natural fishponds for two years: one was stocked with planktivorous whitefish while the other had no planktivorous fish. The aim of the study was the simultaneous assessment of the feeding behaviours of planktivorous fish and of bacterivorous meta-/protozooplankters. We hypothesized that in the presence of planktivorous fish there would be fewer metazooplankton, more protozoans and decreased numbers of bacteria. Our results showed that the amount of metazooplankton eaten by the fish was indeed negatively correlated with metazooplankton biomass. The feeding impact of planktivorous fish in shaping the microbial loop was remarkable. The main grazers of bacteria in the fishpond were ciliates, whereas in the pond without fish these were heterotrophic nanoflagellates. In the fishless pond the role of the top predator shifted to the predaceous metazooplankter Leptodora kindtii which controlled the abundance of herbivorous metazooplankters. We found a negative relationship between the number of bacteria and flagellates in the fishless pond, while the number of bacterivorous ciliates was suppressed by predaceous ciliates. Therefore the bacteria-grazing activity was higher in the absence of planktivorous fish.

Summary Biogenic methane as an alternative carbon and energy source for freshwater organisms has been receiving increasing attention, but the phenomenon is still poorly understood for shallow lakes. We measured the carbon and nitrogen stable isotope signatures (δ13C, δ15N) for key groups of pelagic and benthic organisms, including crustacean zooplankton, chironomid larvae, young-of-the-year and adult fish, to assess whether biogenic methane contributes to pelagic and benthic food webs in a large, shallow lake, Lake Vortsjarv, Estonia. In the southern part of the lake, covered by macrophytes, crustacean zooplankton showed strong seasonal variation of δ13C, with the lowest values occurring in the period of oxygen depletion. Chironomid (Chironomus plumosus) larvae showed high isotopic variability within the lake, with strongly 13C-depleted and 15N-depleted signatures (down to δ13C −64.0‰ and δ15N −2.6‰) in the macrophyte-covered area. Our results indicate that carbon derived from biogenic methane can contribute seasonally to the benthic food web and, to some extent, also to the pelagic food web, where the lake is covered by macrophytes. Moreover, δ13C values for roach (Rutilus rutilus), perch (Perca fluviatilis) and pike (Esox lucius) from the macrophyte-dominated area were on average 3.5‰ lower than those for the same fish from the plankton-dominated lake area, suggesting some carbon derived from methane is transferred up through the food web. Although no direct evidence is available, our results, together with previous studies of the lake, suggest that protozoans could be a potentially important link from methane-oxidising bacteria to animals higher in the web.

Summary 1. As grazers on picoplankton and nanoplankton, planktonic ciliates form an important link in pelagic food webs. Ciliate communities may be controlled by predation by metazooplankton. In eutrophic systems, however, where the number of large crustaceans is often low, the mechanisms that regulate ciliate dynamics have rarely been described. 2. We conducted an enclosure experiment with natural and screened (145 μm) summer plankton communities to investigate the effect of the small-sized crustacean zooplankton on ciliate community structure and the microbial loop in a shallow eutrophic lake. 3. The removal of the larger fraction of crustaceans initiated a decrease in total ciliate abundance. At the community level, we observed a substantial increase in large-sized predacious ciliates (>100 μm) and a simultaneous decrease in the abundance of smaller ciliates (

Seasonal population dynamics and community composition of planktonic ciliates were studied for over a decade in a naturally eutrophic shallow lake, Lake Vortsjarv, which has a large open surface area (270 km 2 on average) and highly turbid water (Secchi depth 300 µm) gymnostomes. There was a positive correlation between ciliates and phyto- and bacterioplankton, implying that the ciliates were clearly bottom-up or food-controlled. This was further confi rmed by the positive correlation between ciliates and metazooplankton. As both bacteria and phytoplankton are coupled with resuspension of lake sediments, the large fl uctuations in water level infl uenced the ciliate community biomass annually.

The abundance and biomass of ciliates, rotifers, cladocerans and copepods were studied in Lake Peipsi and Lake Vortsjarv, both of which are shallow, turbid and large. Our hypothesis was that in a large shallow eutrophic lake, the ciliates could be the most important zooplankton group. The mean metazooplankton biomass was higher in Peipsi than in Vortsjarv (mean values and SD, 1.8 ± 0.7 and 1.3 ± 0.6 mg WM l−1). In Peipsi, the metazooplankton biomass was dominated by filtrators that feed on large-sized phytoplankton and are characteristic of oligo-mesotrophic waters. In Vortsjarv, the metazooplankton was dominated by species characteristic of eutrophic waters. The planktonic ciliates in both lakes were dominated by oligotrichs. The biomass of ciliates was much greater in Vortsjarv (mean 2.3 ± 1.4 mg WM l−1) than in Peipsi (0.1 ± 0.08 mg WM l−1). Ciliates formed about 60% of the total zooplankton biomass in Vortsjarv but only 6% in Peipsi. Thus, the food chains in the two lakes differ: a grazing food chain in Peipsi and a detrital food-chain in Vortsjarv. Consequently, top-down control of phytoplankton can be assumed to be much more important in Peipsi than in Vortsjarv. When the detrital food chain prevails, the planktonic ciliates become the most important zooplankton group in shallow, eutrophic and large lake. Neglecting protozooplankton can result in serious underestimates of total zooplankton biomass since two-thirds of the zooplankton biomass in Vortsjarv comprises ciliates.

The grazing impact of different sized zooplankton on ‘edible’ and total phytoplankton biomass and primary production was measured in L. Vortsjarv during a seasonal study in 1998 and 2000. The organisms of 48–100 μm size class, composed of ciliates and rotifers, contributed significantly to the total grazing of zooplankton community throughout the study period (average 68%). The average daily filtering and grazing rate of the whole zooplankton community (micro- and macro-zooplankton) remained low, corresponding to a filtration of 44% of the water volume, 4% of the total phytoplankton biomass and 29% of primary production. However, a strong grazing pressure on small-sized phytoplankton (

The vertical distribution of planktonic ciliates in eight strongly stratified temperate lakes was studied in summer 1998. Ciliate abundance and biomass were highest (mean 39.9 cells ml−1 and 181.9 μg C l−1) in the epi-, and lowest (mean 8.2 cells ml−1 and 97.6 μg C l−1) in the hypolimnion. The community of ciliates was dominated by five orders: Oligotrichida, Haptorida, Prostomatida, Scuticociliatida and Peritrichida. The community composition varied greatly with depth. In the epilimnion, the ciliate numbers were dominated by oligotrichs but small algivorous prostomatids, peritrichs and haptorids were also numerous. In the metalimnion, these groups were replaced by scuticociliates and mixotrophic prostomatids. In the hypolimnion species known as benthic migrants appeared. We found a positive significant correlation (p < 0.05) between ciliate numbers and Chl a and bacterial densities. Only in the hypolimnion, the correlation between ciliates numbers and Chl a was not significant.

An analysis of plankton seasonal succession in large shallow eutrophic lake Vortsjarv (270 km 2 , mean depth 2.8 m, max. depth 6 m) is presented. Weekly samples for 1995 have been analysed using the PEG model approach. In winter, light was the main factor controlling phytoplankton growth. In early spring phytoplankton was mainly resource-controlled, competition for phosphorus being the main driving force. Ciliates (Paradileptus sp., Strobilidium sp. and Vorticella sp.) were the first herbivores which started to increase in April causing a twofold decline of phytoplankton biomass. The annual maximum of primary production (PP) in early May was probably caused by soluble reactive phosphorus (SRP), regenerated by herbivores, and stirred up from bottom sediments as a result of strong wind stress. This primary production peak provided substrate for further increase of bacterial biomass. The more abundant food supply supported the development of the second spring peak of herbivores (ciliates, rotifers, juvenile copepods) which was followed by the second modest clear water phase in late May. Silicon was depleted by the end of May causing a strong decrease in primary production of the diatom-dominated community. whereas the biomass of the cyanophytes increased under the improved nutrient conditions. Some weeks later, inorganic N was depleted and the period of N limitation with the appearance of N-fixing cyanophytes began. The ciliate collapse at the beginning of June coincided with the start of the cladoceran development and with the increase of other metazooplankton groups. This explains the further decrcase of the biomass of phytoplankton and bacteria in spite of their high production. Beginning from late June, silicon appeared again and SRP started to occur periodically. while inorganic N remained close to zero until November. During this period, phytoplankton development relied to a great extent on the N-fixation and N-regeneration potential. The collapse of the ciliate community in September removed the top-down control from bacteria and their biomass increased, while the development of cladocerans still suppressed phytoplankton biomass in spite of a quite high PP. In October phytoplankton biomass and chlorophyll a (Chla) increased, SRP was completely depleted by the middle of October reflecting a slow regeneration due to the declined activity of zooplankton in cold water. In November nitrates appeared again. and silicon reached the same level as in spring. The biomass of N-fixing Aphanizomenon skujae decreased while Limnothrix redekei and L. planctonica were quite abundant together with diatoms.

The wastewaters of an iron mining company in NW Russia have changed the water quality in some forest lakes to be hard with a high pH, in contrast to the soft water and low pH in natural lakes. Two impacted lakes and a reference lake were sampled once in early August in three successive years for water quality, plankton communities, and fish. The concentrations of potassium, lithium, and sulphate were high in the impacted lakes. Total phosphorus was low (~ 10 µg L -1 ) in all lakes. The primary producers' biomass and chlorophyll a content in the impacted lakes were high because of high densities of autotrophic picocyanobacteria (Synechococcus spp.). All planktonic communities showed a changed taxa composition and lower species richness. Zooplankton was predominantly Rotatoria in all the lakes. In the impacted lakes, Cladocera was represented by Bosmina, and Copepoda by small cyclopoids. The most obvious substance to be harmful to some planktonic species was potassium. The high pH and high mineral content of the water obviously lower both the toxicity and the bioavailability of heavy metals in the impacted lakes.

We studied the relationships between the planktonic food base and feeding patterns of juvenile mid-summer/ early autumn Eurasian perch Perca fluviatilis L., a common predatory freshwater fish in large parts of Europe and Asia. The feeding of 0+ perch was studied during summer and autumn in littoral habitats of seven lakes with different environmental conditions –four Latvian (Auciema, Riebinu, Vārzgūnes, Laukezers) and three Estonian (Kaiavere, Prossa and Akste) lakes. Simultaneously, the abundance, biomass and structure of zooplankton communities were examined. We focused on the littoral areas because many studies in lakes suggest that littoral habitats are particularly important for 0+ fish growth and survival. We were interested in the question: can the diet and growth of 0+ perch be explained by zooplankton community structure? We also presumed that if the amount of zooplankton is low, more benthic invertebrates will be consumed by 0+ perch. Opposite to expectations, we found that zooplankton always counted for over 90% of diet biomass in perch. There were also clear correlations between the zooplankton biomass in a given lake, the zooplankton biomass in 0+ perch stomachs, and the fish growth rate. The study also suggested that nutrient enrichment can positively impact the 0+ perch feeding conditions in lakes. The project is financed by the European Regional Development Fund, the State budget of the Republic of Latvia and the foundation “Institute for Environmental Solutions”. Funding for this project was also provided by the European Union‘s Horizon 2020 research and innovation programme under Grant Agreement No. 951963 (TREICLAKE “Towards Research Excellence and Innovation Capacity in Studying Lake Ecosystems Functional Structures and Climate Change Impact”). The project is financed by the European Regional Development Fund, the State budget of the Republic of Latvia and the foundation “Institute for Environmental Solutions”. Funding for this project was also provided by the European Union‘s Horizon 2020 research and innovation programme under Grant Agreement No. 951963 (TREICLAKE “Towards Research Excellence and Innovation Capacity in Studying Lake Ecosystems Functional Structures and Climate Change Impact”).

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).