Decoupling of pelagic and littoral food webs in oligotrophic Canadian Shield lakes

The importance of top-down effects of piscivorous fish on phytoplankton in natural oligotrophic lakes is still debated. In this study, we analyzed patterns in phytoplankton and zooplankton abundance in 37 oligotrophic Canadian Shield lakes in relation to variations in both piscivorous fish predation and resources (total phosphorus; TP). Zooplankton community structure (but not total biomass) was partially affected by the variation in fish predation while the phytoplankton community structure and total biomass showed no response. Carbon isotope analyses revealed that the lack of top-down effects is due to the uncoupling of the littoral and the pelagic food webs. We found that the fish community depends mostly on benthic resources, suggesting that only low planktivory occurred in our study lakes. Due to the absence of specialized zooplanktivorous fish, zooplankton is poorly exploited in these lakes and thus able to control phytoplankton by grazing. A comparison of our data with published studies on the TP–chlorophyll a relationships in both natural and manipulated systems shows that the phytoplankton biomass per unit of TP is relatively low in Canadian Shield lakes.     

OPINION Manipulating lake community structure: where do we go from here?

SUMMARY. 1 More than 10 years experience with whole lake pelagic manipulation has suggested some general trends applicable to all freshwater pelagic communities and some specific trends related to lake depth.
2 Among the general trends is the observation that the trophic cascade is strongly damped. This means that changes in phytoplankton biomass can be assured only when the fish community is strongly manipulated.
3 Among the depth related trends is the observation that in shallow lakes, changes in fish community structure are more likely to have cascading impacts on phytoplankton than are changes in deep lakes.
4 In shallow lakes, fish removal frequently results in decreased turbidity which is associated with the development of dense macrophyte populations and significant reductions of algal standing stocks. The mechanisms involve: increased grazing by zooplankton, the removal of fish induced bioturbation and nutrient recycling, and direct and indirect macrophyte effects (shading, zooplankton refuges and competition for nutrients).
5 In shallow lakes, where planktivore biomass can be regulated and macrophyte development is acceptable, fish biomanipulalions are likely to result in reduced algal populations and improved water quality.
6 In deep lakes, where macrophytes are not as important, long-term effects of fish manipulations are strongly dependent upon the probability of non-grazable algal bloom development. This is determined by many factors (chemical, physical and grazer related) which modify the impact that grazers have on phytoplankton biomass.
7 In deep lakes, successful fish biomanipulations may only be effective when chemical and physical factors are altered to produce algal species compositions that permit strong top-down control of prey by predators.     

Spatial insurance against a heatwave differs between trophic levels in experimental aquatic communities

Climate change-related heatwaves are major threats to biodiversity and ecosystem functioning. However, our current understanding of the mechanisms governing community resistance to and recovery from extreme temperature events is still rudimentary. The spatial insurance hypothesis postulates that diverse regional species pools can buffer ecosystem functioning against local disturbances through the immigration of better-adapted taxa. Yet, experimental evidence for such predictions from multi-trophic communities and pulse-type disturbances, like heatwaves, is largely missing. We performed an experimental mesocosm study to test whether species dispersal from natural lakes prior to a simulated heatwave could increase the resistance and recovery of plankton communities. As the buffering effect of dispersal may differ among trophic groups, we independently manipulated the dispersal of organisms from lower (phytoplankton) and higher (zooplankton) trophic levels. The experimental heatwave suppressed total community biomass by having a strong negative effect on zooplankton biomass, probably due to a heat-induced increase in metabolic costs, resulting in weaker top-down control on phytoplankton. While zooplankton dispersal did not alleviate the negative heatwave effects on zooplankton biomass, phytoplankton dispersal enhanced biomass recovery at the level of primary producers, providing partial evidence for spatial insurance. The differential responses to dispersal may be linked to the much larger regional species pool of phytoplankton than of zooplankton. Our results suggest high recovery capacity of community biomass independent of dispersal. However, community composition and trophic structure remained altered due to the heatwave, implying longer-lasting changes in ecosystem functioning.     

Effects of different fish species and biomass on plankton interactions in a shallow lake

Thirty-six enclosures, surface area 4 m², were placed in Little Mere, a shallow fertile lake in Cheshire, U.K. The effects of different fish species (common carp, common bream, tench and roach) of zooplanktivorous size, and their biomass (0, 200 and 700 kg ha^–1) on water chemistry, zooplankton and phytoplankton communities were investigated. Fish biomass had a strong effect on mean zooplankton size and abundance. When fish biomass rose, larger zooplankters were replaced by more numerous smaller zooplankters. Consequently phytoplankton abundance rose in the presence of higher densities of zooplanktivorous fish, as zooplankton grazing was reduced. Fish species were also significant in determining zooplankton community size structure. In enclosures with bream there were significantly greater densities of small zooplankters than in enclosures stocked with either carp, tench and, in part, roach. When carp or roach were present, the phytoplankton had a greater abundance of Cyanophyta than when bream or tench were present. Whilst top-down effects of fish predation controlled the size partitioning of the zooplankton community, this, in turn apparently controlled the bottom-up regeneration of nutrients for the phytoplankton community. At the zooplankton–phytoplankton interface, both top-down and bottom-up processes were entwined in a reciprocal feedback mechanism with the extent and direction of that relationship altered by changes in fish species. This has consequences for the way that top-down and bottom-up processes are generalised.     

Cascading effects of generalist fish introduction in oligotrophic lakes

Introduction of strictly planktivorous fish to lakes can alter plankton communities via cascading interactions in food webs. Less is known about the large-scale and long-term effects resulting from the introduction of fish with more generalist feeding habits, and the extent to which these effects depend on lake trophic status. Paleolimnological records of three oligotrophic lakes in Maine, USA were used to analyze the response of plankton communities to the introduction of white perch (Morone americana), a fish that often numerically dominates fish assemblages and switches from strict planktivory to a more generalist diet during ontogeny. After white perch introduction, cladoceran ephippia size increased up to 50 %, suggesting that the most important role of this generalist fish, with respect to water quality, is as a piscivorous trophic link. Algal standing crop declined by a quarter to over a half of pre-introduction levels, suggesting that top-down effects of white perch reduced phytoplankton biomass. In comparing these oligotrophic lakes to prior work in a eutrophic system, white perch introduction had similar effects on zooplankton body size; however, cascading effects to phytoplankton were only observed in low productivity lakes.     

Cascading Trophic Interactions in an Oligotrophic Species-poor Alpine Lake

Non-native brook trout (Salvelinus fontinalis) were eradicated from alpine Bighorn Lake, Alberta, Canada, to test whether strong cascading trophic interactions (CTI) can occur in oligotrophic, high seston C:P, species-poor lakes. Fishless alpine Pipit Lake was used as a reference ecosystem. Bighorn Lake zooplankton biomass increased from 0.14:1 relative to Pipit Lake before fish removal began in 1997 to 0.6:1 afterwards due to an increase in the abundance of adult cyclopoid copepods beginning in 1997 and the reappearance of Daphnia middendorffiana in 1998. Following the reappearance of Daphnia, Bighorn Lake total phytoplankton biomass fell from 64:1 relative to Pipit Lake to 0.9:1. Over the same periods Bighorn Lake:Pipit Lake chlorophyll-a ratios declined from 2.4:1 to 1.6:1, although the decrease was not statistically significant. Mid-summer Secchi disc depth in Bighorn Lake increased from 3.1 m before manipulation to 9.2 m, the maximum depth of the lake, in 2001 and 2002. Increased transparency was most likely due to increased filtration of suspended inorganic particles from the water column by higher abundances of large zooplankton. Post-manipulation increases in dissolved inorganic nitrogen (DIN), DIN:total dissolved phosphorus (TDP) ratio and declines in TDP in Bighorn Lake were not attributable to ecosystem manipulation, similar changes were observed in reference Pipit Lake. We conclude that strong pelagic CTI, expressed as change in total phytoplankton biomass and largely mediated by Daphnia, can occur in oligotrophic, high seston C:P, species-poor ecosystems. However, strong CTI responses in phytoplankton biomass may lag trophic manipulation by several years.     

Bottom-Up Effects on Biomass Versus Top-Down Effects on Identity: A Multiple-Lake Fish Community Manipulation Experiment

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.     

Biomanipulation development in Norway

Since 1974 several studies have been carried out in Norway to investigate the interactions between planktivorous fish, zooplankton, phytoplankton and water chemistry. Since 1978 a long-term national research program has been conducted by the Norwegian Council for Scientific and Industrial Research (NTNF). In this program several whole lake manipulations of the fish stocks have been performed to test hypotheses about trophic interactions. It was predicted that manipulations of planktivorous fish populations, might also improve water quality in lakes undergoing eutrophication. Two examples are given to illustrate the achieved results. I: Whole lake fertilization experiment (1974–1978) carried out by Langeland and Reinertsen. The results revealed the importance of top-down effects in the lake ecosystem. When cladocerans dominated, the zooplankton community was able to maintain a more or less constant phytoplankton biomass and a rather low phytoplankton production even when nutrient levels were increased. During years with rotifer dominance, algal biomass and productivity increased, despite the low amounts of added nutrients. II: Experiment performed by Reinertsen, Jensen, Koksvik, Langeland and Olsen in the eutrophic Lake Haugatjern, total elimination of the fish populations by rotenone in late 1980, resulted in a 4-fold decrease in the algal biomass. The species composition changed from the dominance of large-sizedAnabaena flos-aquae andStaurastrum luetkemuelleri to smaller, fastgrowing species and gelatinous green algae. The results are discussed in relation to management of inland waters by combined techniques of biomanipulation and reduced external nutrient supply which increase food-chain efficiency.     

Zooplankton biomass dynamics in oligotrophic versus eutrophic conditions: a test of the PEG model

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Lake Chapo: a baseline study of a deep, oligotrophic North Patagonian lake prior to its use for hydroelectricity generation: II. Biological properties

The present paper focuses on the biological properties of Lake Chapo prior to the installation of a hydroelectric power station. Most of the biological parameters indicate that the lake is oligotrophic, i.e., it has very low values of chlorophyll a and primary production. The phytoplankton community of 73 species shows a predominance of only a few species in terms of density and biomass. The zooplankton community is poor in species, with one of the lowest densities measured in the North Patagonian lakes. The fish community is dominated by two introduced salmonid species. Comparison with other North Patagonian lakes reveals that Lake Chapo is the most oligotrophic.     

Water level and fish-mediated cascading effects on the microbial community in eutrophic warm shallow lakes: a mesocosm experiment

Information on the effects of water level changes on microbial planktonic communities in lakes is limited but vital for understanding ecosystem dynamics in Mediterranean lakes subjected to major intra- and inter-annual variations in water level. We performed an in situ mesocosm experiment in an eutrophic Turkish lake at two different depths crossed with presence/absence of fish in order to explore the effects of water level variations and the role of top-down regulation at contrasting depths. Strong effects of fish were found on zooplankton, weakening through the food chain to ciliates, HNF and bacterioplankton, whereas the effect of water level variations was overall modest. Presence of fish resulted in lower biomass of zooplankton and higher biomasses of phytoplankton, ciliates and total plankton. The cascading effects of fish were strongest in the shallow mesocosms as evidenced by a lower zooplankton contribution to total plankton biomass and lower zooplankton:ciliate and HNF:bacteria biomass ratios. Our results suggest that a lowering of the water level in warm shallow lakes will enhance the contribution of bacteria, HNF and ciliates to the plankton biomass, likely due to increased density of submerged macrophytes (less phytoplankton); this effect will, however, be less pronounced in the presence of fish.     

Zooplankton and environmental factors of a recovering eutrophic lake (Lysimachia Lake, Western Greece)

The present study investigates the zooplankton community dynamics and the abiotic environment in the eutrophic Lake Lysimachia (western Greece). The lake is considered to be recovering from eutrophication after the termination of an urban sewage inflow in 2000, and its waters are replenished constantly from the nearby oligotrophic Lake Trichonis. The results show that, although a decrease in nutrient concentrations was observed compared to the past, the lake still has eutrophic characteristics. This was reflected in the zooplankton community which is typical of those found in eutrophic lakes where rotifers prevail. Similarities among this lake and other nearby lakes were found considering the zooplankton community composition and seasonal variation. However, Lake Lysimachia is inhabited also by a number of different and even unique species (e.g., Moina micrura), suggesting that this ecosystem may be an important biodiversity refuge. Most of the zooplankton species were correlated with water temperature and, to a lesser extent, eutrophication key-water quality variables. Although there are few available data on the zooplankton of the lake, the abundance and composition of the community presenting characteristics indicative of intermediate trophic conditions and suggesting that the lake is probably under a kind of “biological” recovery.     

Spatial patterns and relationships between phytoplankton, zooplankton and water quality in the Saimaa lake system, Finland

The interactions between phytoplankton and zooplankton were studied in two large lakes in the Saimaa lake system, Finland. Both are subjected to substantial waste water loading, and exhibit a clear trophic gradient between the loaded and unloaded areas. The phytoplankton and zooplankton were compared in terms of composition, abundance and biomass at 34–39 stations located in different parts of the lakes. At least four mechanisms were thought to affect the composition of plankton communities: (1) the amount of nutrients (trophic gradient), (2) grazing of algae by herbivores, (3) the effect of the algal species composition on feeding by zooplankters (large, colonial algae in the more loaded parts of the lakes) and (4) the regeneration and reorganization of nutrients.     

Phytoplankton communities in the littoral zone of lakes: Observations on structure and dynamics in oligotrophic and eutrophic systems

Summary The structure and seasonal dynamics of phytoplankton communities in the littoral zone were compared between oligotrophic and eutrophic lakes in the southeastern United States. Differences in diversity and species composition between lakes could be ascribed to long-term variation in nutrients corresponding to trophic status. However, significant within-lake variation could not be accounted for by microstratification of nutrients or other abiotic variables. Local biotic factors, perhaps dominated by the spawning activities of centrarchid fishes, resuspend periphyton and generate tychoplankton which becomes a persistent and integral part of the phytoplankton community in eutrophic systems. The patchy distribution of these biotic factors and resultant tychoplankton may lead to the observed variation. Grazing by herbivorous zooplankton was considered to be the major factor affecting the relative abundance of phytoplankton in the littoral zone, completely overriding the effects of nutrient concentration and biotic interactions between phytoplankton species during spring and summer.The relative importance of tychoplankton and grazing as regulatory factors operates independently of the trophic status or geographical location of a lake, making comparisons of different studies difficult and perhaps meaningless if traditional analyses based only on nutrients and interactions between species of phytoplankton are used. Limnetic as well as littoral components must be considered in future studies of phytoplankton communities in the littoral zone.     

Temporal dynamics and relationship between climate,limnological variables and zooplankton composition in climate-sensitive Lake Bosumtwi,Ghana

The community composition and the factors affecting seasonal and interannual dynamics of zooplankton in Lake Bosumtwi were studied biweekly at a central index station during 2005 and 2006. The lake zooplankton community was species poor. Mesocyclops bosumtwii was numerically superior seasonally and interannually and was endemic to the lake. Minor constituents included Moina micrura, six rotifer species (except for Hexarthra intermedia) and Chaoborus ceratopogones larvae. Low variance of cyanobacteria-dominated phytoplankton biomass underlined stable zooplankton community structure. Emergence of rare species of rotifers occurred seasonally. The climatic signature on the lake’s stratification and mixing regime was strongly influenced by atmospheric temperature, but weakly by wind strength, because of sheltering of the lake by high crater walls. Increasing mixing depth entrained high TP concentrations from below the thermocline seasonally, but reflected poorly in the phytoplankton biomass behaviour. Total zooplankton abundance did not differ seasonally, but varied markedly from year to year in its timing and magnitude. Herbivores were squeezed between food limitation and high predation pressure from Chaoborus all year round. The low fish planktivory (high fishing pressure) on Chaoborus may create a trophic bottleneck restricting energy transfer efficiency from zooplankton to fish.     

Distribution of planktonic ciliates in highly coloured subtropical lakes: comparison with clearwater ciliate communities and the contribution of myxotrophic taxa to total autotrophic biomass

SUMMARY 1. The planktonic ciliate communities of eleven organically coloured north and central Florida lakes representing a variety of trophic conditions were examined during 1979–80. The total abundance and biomass of ciliates were not significantly different from comparable clearwater lakes and only minor taxonomic replacements were noted at the order level.
2. Timing of population peaks of oligotrophic lakes was dissimilar to clearwater lakes of the same trophic state, but seasonality in meso-trophic and eutrophic lakes resembled patterns described for comparable clearwater lakes.
3. Various ciliate components were strongly correlated with chlorophyll a concentrations, but only moderately correlated to dominant phytoplankton groups. No significant correlations were found between ciliate components and bacterial abundance.
4. Myxotrophic taxa numerically dominated oligotrophic systems, particularly during midsummer, and accounted for a large percentage of the total ciliate biomass. Estimates of the ciliate contribution to total autotrophic biomass indicate that these zoochlorellae-bearing protozoa may account for much of the autotrophic biomass during midsummer periods in coloured lakes, and thus may lead to an overestimation of phytoplankton standing crops available to zooplankton grazers if chlorophyll a is used as a surrogate measure of algal biomass.     

Effects of water-column mixing on bacteria,phytoplankton, and rotifers under different levels of herbivory in a shallow eutrophic lake

Water-column mixing is known to have a decisive impact on plankton communities. The underlying mechanisms depend on the size and depth of the water body, nutrient status and the plankton community structure, and they are well understood for shallow polymictic and deep stratified lakes. Two consecutive mixing events of similar intensity under different levels of herbivory were performed in enclosures in a shallow, but periodically stratified, eutrophic lake, in order to investigate the effects of water-column mixing on bacteria abundance, phytoplankton abundance and diversity, and rotifer abundance and fecundity. When herbivory by filter-feeding zooplankton was low, water-column mixing that provoked a substantial nutrient input into the euphotic zone led to a strong net increase of bacteria and phytoplankton biomass. Phytoplankton diversity was lower in the mixed enclosures than in the undisturbed ones because of the greater contribution of a few fast-growing species. After the second mixing event, at a high biomass of filter-feeding crustaceans, the increase of phytoplankton biomass was lower than after the first mixing, and diversity remained unchanged because enhanced growth of small fast-growing phytoplankton was prevented by zooplankton grazing. Bacterial abundance did not increase after the second mixing, when cladoceran biomass was high. Changes in rotifer fecundity indicated a transmission of the phytoplankton response to the next trophic level. Our results suggest that water-column mixing in shallow eutrophic lakes with periodic stratification has a strong effect on the plankton community via enhanced nutrient availability rather than resuspension or reduced light availability. This fuels the basis of the classic and microbial food chain via enhanced phytoplankton and bacterial growth, but the effects on biomass may be damped by high levels of herbivory. Received: 3 May 1999 / Accepted: 13 April 2000     

Top-down control of natural phyto- and bacterioplankton prey communities by Daphnia magna and by the natural zooplankton community of the hypertrophic Lake Blankaart

Biomanipulation, through the reduction of fish abundance resulting in an increase of large filter feeders and a stronger top-down control on algae, is commonly used as a lake restoration tool in eutrophic lakes. However, cyanobacteria, often found in eutrophic ponds, can influence the grazing capacity of filter feeding zooplankton. We performed grazing experiments in hypertrophic Lake Blankaart during two consecutive summers (1998, with and 1999, without cyanobacteria) to elucidate the influence of cyanobacteria on the grazing pressure of zooplankton communities. We compared the grazing pressure of the natural macrozooplankton community (mainly small to medium-sized cladocerans and copepods) with that of large Daphnia magna on the natural bacterioplankton and phytoplankton prey communities. Our results showed that in the absence of cyanobacteria, Daphnia magna grazing pressure on bacteria was higher compared to the grazing pressure of the natural zooplankton community. However, Daphnia grazing rates on phytoplankton were not significantly different compared to the grazing rates of the natural zooplankton community. When cyanobacteria were abundant, grazing pressure of Daphnia magnaseemed to be inhibited, and the grazing pressure on bacteria and phytoplankton was similar to that of the natural macrozooplankton community. Our results suggest that biomanipulation may not always result in a more effective top-down control of the algal biomass.     

Analysis of the community structure of planktonic ciliated protozoa relative to trophic state in Florida lakes

The planktonic ciliate populations of 30 Florida lakes constituting a broad trophic gradient were examined to determine the response of protozoan community structure to increasing eutrophication. Both ciliate abundance and biomass were strongly related to lake trophic state. Comparison of the Florida data base with a comparable north temperate lake group indicated that subtropical lakes generally possess higher ciliate abundance and biomass at a given trophic state than temperate lakes. However the equations derived for each data base were not significantly different. Community diversity and species richness increased with increasing lake productivity. Highly acidic lakes displayed significantly reduced diversity and numbers of species when contrasted with nonacidic oligotrophic lakes. Small-bodied (< 30 um) ciliates dominated all lakes but were proportionally less important in oligotrophic lakes. Presence-absence data produced three assemblages: an ubiquitous association of primarily small ciliate taxa, a group of large ciliates mainly restricted to eutrophic-hypereutrophic lakes, and a very large ciliate,Stentor niger, which dominated the protozoan communities of acidic oligotrophic lakes.     

Weedbeds and big bugs: the importance of scale in detecting the influence of nutrients and predation on macroinvertebrates in plant‐dominated shallow lakes

1. The scale of investigations influences the interpretation of results. Here, we investigate the influence of fish and nutrients on biotic communities in shallow lakes, using studies at two different scales: (i) within‐lake experimental manipulation and (ii) comparative, among‐lake relationships. 2. At both scales, fish predation had an overriding influence on macroinvertebrates; fish reduced macroinvertebrate biomass and altered community composition. Prey selection appeared to be size based. Fish influenced zooplankton abundance and light penetration through the water column also, but there was no indication that fish caused increased resuspension of sediment. 3. There were effects of nutrients at both scales, but these effects differed with the scale of the investigation. Nutrients increased phytoplankton and periphyton at the within‐lake scale, and were associated with increased periphyton at the among‐lake scale. No significant effect of nutrients on macroinvertebrates was observed at the within‐lake scale. However, at the among‐lake scale, nutrients positively influenced the biomass and density of macroinvertebrates, and ameliorated the effect of fish on macroinvertebrates. 4. Increased prey availability at higher nutrient concentrations would be expected to cause changes in the fish community. However, at the among‐lake scale, differences were not apparent in fish biomass among lakes with different nutrient conditions, suggesting that stochastic events influence the fish community in these small and relatively isolated shallow lakes. 5. The intensity of predation by fish significantly influences macroinvertebrate community structure of shallow lakes, but nutrients also play a role. The scale of investigation influences the ability to detect the influence of nutrients on the different components of shallow lake communities, particularly for longer lived organisms such as macroinvertebrates, where the response takes longer to manifest.