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The extent to which ecosystems are regulated by top-down relative to bottom-up control has been a dominant paradigm in ecology for many decades. For lakes, it has been shown that predation by fish is an important determinant of variation in zooplankton and phytoplankton community characteristics. Effects of fish are expected to not only be a function of total fish biomass, but also of functional composition of the fish community. Previous research on the importance of trophic cascades in lakes has largely focused on the role of zooplanktivorous and piscivorous fish. We conducted a large-scale multiple-lake fish community manipulation experiment to test for the effect of differences in fish functional community composition on the trophic structure of lakes. We examine the effect of top-down and bottom-up factors on phytoplankton and zooplankton biomass as well as on their community composition. We put our data in a broader perspective by comparing our results to data of a survey that also included ponds with low fish densities as well as ponds with very high densities of fish. Our results indicate that the overall food web structure under relative high fish densities is primarily structured by bottom-up factors, whereas community characteristics seem to be primarily regulated by top-down factors. Our results suggest a subtle interplay between bottom-up and top-down factors, in which bottom-up factors dominate in determining quantities while top-down effects are important in determining identities of the communities.

© 2016 John Wiley & Sons Ltd The tropical Andes has a high density of glacial lakes that are situated in the high-altitude páramo (3500–4500 m). Ecological information about such lakes is scant despite the fact that these lakes are an important source of water for drinking, irrigation and electricity generation and feed several major tributaries of the Amazon. In this study, we provide data on a survey of 31 lakes in Cajas National Park (Ecuador). Two of the lakes were monitored monthly during one year. In situ nutrient addition experiments were carried out in three of the lakes. Seasonal monitoring in two lakes revealed a thermal stratification of the water column between October and June, with a small temperature difference between epi- and hypolimnion (2–3 °C). Oxygen depletion of the hypolimnion towards the end of the stratification period indicated that no complete mixing of the water column occurred during stratification. There was no evidence of depletion of nutrients in the epilimnion or accumulation in the hypolimnion during stratification. There were also no clear seasonal changes in chlorophyll-a (Chl-a) concentration nor in phytoplankton community composition in the two lakes. Inputs of dissolved organic carbon (DOC) from the vegetated catchment resulted in high DOC concentrations (median 2.9 mg L−1) compared to temperate mountain lakes. Water transparency was relatively low, with a median extinction coefficient for photosynthetic active radiation of 0.50 m−1 and for UV-B radiation of 10.13 m−1. Although the thermocline was deep and water transparency was low, estimates of the critical depth for photosynthesis were deeper than the mean water depth in all lakes, suggesting that phytoplankton was not light limited. The phytoplankton community was dominated by chlorophytes (e.g. Oocystis), diatoms (small Cyclotella spp.) or small colonial cyanobacteria (Aphanocapsa, Merismopedia). The zooplankton community was either dominated by large cladocerans and cyclopoid copepods, or by the calanoid copepod Boeckella occidentalis. Total concentrations of phosphorus (P) and nitrogen (N) were comparable to those in temperate mountain lakes (4–35 μg P L−1 and 162–758 μg N L−1) while Chl-a concentrations were in the lower range (

Keywords: allelochemicals; chemical ecology; competition; nutrient limitation; shallow lakes Summary 1.It is well known that submerged macrophytes can suppress phytoplankton blooms in lakes and thus promote water quality and biodiversity. One of the possible mechanisms through which submerged macrophytes control phytoplankton is by producing allelochemicals that suppress phytoplankton growth rates. The in situ importance of allelopathy, however, is often questioned because it is assumed that phytoplankton communities can rapidly evolve resistance to allelochemicals. 2.Here, we present the results of two mesocosm experiments in which we evaluated whether the submerged macrophyte Elodea nuttallii is capable of controlling phytoplankton biomass over periods of 4 to 8 weeks. Such a timescale is long relative to the generation time of phytoplankton and is therefore expected to allow the development of resistance through compositional shifts at both population and community levels. 3.Although the mesocosms were inoculated with a diverse phytoplankton inoculum including species that had previously been exposed to Elodea, phytoplankton biomass remained consistently low during the course of the experiments in the treatments with Elodea. As zooplankton grazing and competition for nutrients and light by macrophytes were excluded in our experiments, this suggests that phytoplankton was controlled by allelopathy. 4.Dialysis bag assays, performed at the end of each mesocosm experiment, showed that phytoplankton communities from mesocosms with Elodea were equally sensitive to exudates from Elodea than phytoplankton communities from mesocosms without Elodea. 5.These results suggest that phytoplankton communities do not evolve resistance to allelochemicals from Elodea. This may allow Elodea to control phytoplankton in natural ecosystems over prolonged time periods through allelopathy.

1. In temperate regions, submerged macrophytes can hamper phytoplankton blooms. Such an effect could arise directly, for instance via allelopathy, or indirectly, via competition for nutrients or the positive interaction between submerged macrophytes and zooplankton grazing. However, there is some evidence that the positive interaction between submerged macrophytes and zooplankton grazing is less marked in warmer regions, where the interaction is less well studied, and that negative effects of higher water plants on phytoplankton biomass are weaker. 2. We carried out two consecutive mesocosm experiments in Uruguay (subtropical South America) to study the effects of two common submerged macrophytes from this region (Egeria densa and Potamogeton illinoensis) on phytoplankton biomass, in the absence of zooplankton grazing. We compared phytoplankton development between different macrophyte treatments (no macrophytes, artificial macrophytes, real Egeria and real Potamogeton). We used artificial macrophytes to differentiate between physical effects (i.e. shading, sedimentation and competition with periphyton) and biological effects (i.e. nutrient competition and allelopathy). 3. In Experiment 1, we found no evidence for physical effects of macrophytes on phytoplankton biomass, but both macrophyte species seemed to exert strong biological effects on phytoplankton biomass. Only Egeria affected phytoplankton community structure, particularly tempering the dominance of Scenedesmus. Nutrient addition assays revealed that only Egeria suppressed phytoplankton through nutrient competition. 4. We performed a second mesocosm experiment with the same design, but applying saturating nutrient conditions as a way of excluding the effects of competition for nutrients. This experiment showed that both macrophytes were still able to suppress phytoplankton through biological mechanisms, providing evidence for allelopathic effects. Our results indicate that both common macrophytes are able to keep phytoplankton biomass low, even in the absence of zooplankton grazing.

Low phytoplankton biomass usually occurs in the presence of submerged macrophytes, possibly because submerged macrophytes enhance top-down control of phytoplankton by offering a refuge for efficient grazers like Daphnia against fish predation. However, other field studies also suggest that submerged macrophytes suppress phytoplankton in the absence of Daphnia. In order to investigate these mechanisms further, we conducted an outdoor mesocosm experiment to study the effect of submerged macrophytes (Elodea nuttallii) on phytoplankton and zooplankton biomass. The experiment combined four nutrient addition levels (0, 10, 100, and 1000 μg P l−1; N/P ratio: 16) with three macrophyte levels (no macrophytes, artificial macrophytes, and real macrophytes). We inoculated the tanks with species-rich inocula of phytoplankton and zooplankton but excluded fish or macro-invertebrates. Probably due to the lack of predators in the mesocosms, potential grazing rates of pelagic zooplankton (estimated from zooplankton biomass) did not differ between the macrophyte treatment combinations. Compared to the treatment combinations without macrophytes, lower phytoplankton biomass occurred in the treatment combinations with real macrophytes at all the nutrient addition levels and in those with artificial macrophytes at all the nutrient levels except the highest. Significantly, higher abundances of plant-associated filter feeders (Simocephalus vetulus and Ceriodaphnia spp.) occurred in the treatment combinations with real and artificial macrophytes. The estimated potential grazing rate of these plant-associated filter feeders indicated that these filter feeders could be responsible for the lower phytoplankton biomass in the presence of real and artificial macrophytes. Our results suggest that the plant-associated filter feeders may be significant grazers in vegetated shallow lakes.

In order to evaluate latitudinal differences in the relationship of phytoplankton biomass and diversity with environmental conditions in shallow lakes, we sampled 98 shallow lakes from three European regions: Denmark (DK), Belgium/The Netherlands (BNL) and southern Spain (SP). Phytoplankton biomass increased with total phosphorus (TP) concentrations and decreased with submerged macrophyte cover across the three regions. Generic richness was significantly negatively related to submerged macrophyte cover and related environmental variables. Zooplankton:phytoplankton biomass ratios were positively related to submerged macrophyte cover and negatively to phytoplankton generic richness in DK and BNL, suggesting that the low generic richness in lakes with submerged macrophytes was due to a higher zooplankton grazing pressure in these regions. In SP, phytoplankton generic richness was not influenced by zooplankton grazing pressure but related to conductivity. We observed no relationship between phytoplankton generic richness and TP concentration in any of the three regions. The three regions differed significantly with respect to mean local and regional generic richness, with BNL being more diverse than the other two regions. Our observations suggest that phytoplankton diversity in European shallow lakes is influenced by submerged macrophyte cover indirectly by modulating zooplankton grazing. This influence of submerged macrophytes and zooplankton grazing on phytoplankton diversity decreases from north to south.