Natural and constructed littoral zones as nutrient traps in eutrophicated shallow lakes

It is generally known that the water quality of shallow lakes can be influenced significantly by marginal wetlands. In order to study the efficacy of constructed littoral wetlands in the IJsselmeer area (The Netherlands) for water quality improvement, a field survey was carried out in 2003. Vegetation, soil, pore water and surface water characteristics were measured in spring and summer in two types of littoral zones: natural and constructed for 8–16 years. The study showed that constructed wetlands perform well and are suitable to enlarge the vegetated littoral zone in the IJsselmeer area. In both natural and constructed sites vegetation biomass varied between 2,200 g m⁻² for helophyte vegetation and 1,300 g m⁻² for low herbaceous vegetation. Nutrient concentrations in the pore water of constructed sites tended to be higher than in natural sites. and concentrations in pore water were much lower when vegetation was present, probably as a result of plant uptake. The N and P accumulation rate in the soil of constructed wetlands was 20 g N m⁻² y⁻¹ and 3 g P m⁻² y⁻¹ in vegetated plots; without vegetation the rate was much lower (8 g N m⁻² y⁻¹ and 1.8 g P m⁻² y⁻¹). We conclude that concerning their effect on water quality, constructed sites may replace natural sites, at least after 8–16 years. Principal component analysis showed a relationship between vegetation biomass and flooding, and nutrient concentrations in soil and pore water. Biomass was negatively correlated with extractable nutrients and positively with soil total N and P content. Flooding duration was negatively related to pore water salinity and positively to pore water nutrients. Due to their high biomass, helophyte stands retained significantly more nutrients than low pioneer vegetation and are therefore more suitable for improving water quality. Handling editor: S. Declerck     

Phycological investigations of pelagic vs. littoral zones of shallow lake ecosystems

The main goal of this study was to examine the influence of two aquatic plants representing various ecological types (Chara tomentosa and Typha angustifolia) and comparatively the pelagic zone, on the phytoplankton communities in three shallow and macrophyte-dominated lakes.     

Substantial differences in littoral fish community structure and dynamics in subtropical and temperate shallow lakes

1. Fish play a key role in the functioning of temperate shallow lakes by affecting nutrient exchange among habitats as well as lake trophic structure and dynamics. These processes are, in turn, strongly influenced by the abundance of submerged macrophytes, because piscivorous fish are often abundant at high macrophyte density. Whether this applies to warmer climates as well is virtually unknown. 2. To compare fish community structure and dynamics in plant beds between subtropical and temperate shallow lakes we conducted experiments with artificial submerged and free‐floating plant beds in a set of 10 shallow lakes in Uruguay (30°–35°S) and Denmark (55°–57°N), paired along a gradient of limnological characteristics. 3. The differences between regions were more pronounced than differences attributable to trophic state. The subtropical littoral fish communities were characterised by higher species richness, higher densities, higher biomass, higher trophic diversity (with predominance of omnivores and lack of true piscivores) and smaller body size than in the comparable temperate lakes. On average, fish densities were 93 ind. m⁻² (±10 SE) in the subtropical and 10 ind. m⁻² (±2 SE) in the temperate lakes. We found a twofold higher total fish biomass per unit of total phosphorus in the subtropical than in the temperate lakes, and as fish size is smaller in the former, the implication is that more energy reaches the littoral zone fish community of the warmer lakes. 4. Plant architecture affected the spatial distribution of fish within each climate zone. Thus, in the temperate zone fish exhibited higher densities among the artificial free‐floating plants while subtropical fish were denser in the artificial submerged plant beds. These patterns appeared in most lakes, regardless of water turbidity or trophic state. 5. The subtropical littoral fish communities resembled the fish communities typically occurring in temperate eutrophic and hypertrophic lakes. Our results add to the growing evidence that climate warming may lead to more complex and omnivory‐dominated food webs and higher density and dominance of smaller‐sized fish. This type of community structure may lead to a weakening of the trophic cascading effects commonly observed in temperate shallow lakes and a higher risk of eutrophication.     

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.     

Effects of mixing on the pelagic food web in shallow lakes

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Effects of habitat complexity on community structure and predator avoidance behaviour of littoral zooplankton in temperate versus subtropical shallow lakes

1. Structural complexity may stabilise predator–prey interactions and affect the outcome of trophic cascades by providing prey refuges. In deep lakes, vulnerable zooplankton move vertically to avoid fish predation. In contrast, submerged plants often provide a diel refuge against fish predation for large‐bodied zooplankton in shallow temperate lakes, with consequences for the whole ecosystem. 2. To test the extent to which macrophytes serve as refuges for zooplankton in temperate and subtropical lakes, we introduced artificial plant beds into the littoral area of five pairs of shallow lakes in Uruguay (30°–35°S) and Denmark (55°–57°N). We used plants of different architecture (submerged and free‐floating) along a gradient of turbidity over which the lakes were paired. 3. We found remarkable differences in the structure (taxon‐richness at the genus level, composition and density) of the zooplankton communities in the littoral area between climate zones. Richer communities of larger‐bodied taxa (frequently including Daphnia spp.) occurred in the temperate lakes, whereas small‐bodied taxa characterised the subtropical lakes. More genera and a higher density of benthic/plant‐associated cladocerans also occurred in the temperate lakes. The density of all crustaceans, except calanoid copepods, was significantly higher in the temperate lakes (c. 5.5‐fold higher). 4. Fish and shrimps (genus Palaemonetes) seemed to exert a stronger predation pressure on zooplankton in the plant beds in the subtropical lakes, while the pelagic invertebrate Chaoborus sp. was slightly more abundant than in the temperate lakes. In contrast, plant‐associated predatory macroinvertebrates were eight times more abundant in the temperate than in the subtropical lakes. 5. The artificial submerged plants hosted significantly more cladocerans than the free‐floating plants, which were particularly avoided in the subtropical lakes. Patterns indicating diel horizontal migration were frequently observed for both overall zooplankton density and individual taxa in the temperate, but not the subtropical, lakes. In contrast, patterns of diel vertical migration prevailed for both the overall zooplankton and for most individual taxa in the subtropics, irrespective of water turbidity. 6. Higher fish predation probably shapes the general structure and dynamics of cladoceran communities in the subtropical lakes. Our results support the hypothesis that horizontal migration is less prevalent in the subtropics than in temperate lakes, and that no predator‐avoidance behaviour effectively counteracts predation pressure in the subtropics. Positive effects of aquatic plants on water transparency, via their acting as a refuge for zooplankton, may be generally weak or rare in warm lakes.     

Benthic and pelagic food resources for zooplankton in shallow high-latitude lakes and ponds

1. Shallow lakes and ponds are a major component of the northern landscape and often contain a high zooplankton biomass despite clear waters that are poor in phytoplankton. 2. In this study we quantified zooplankton food sources and feeding rates in the shallow waters of two contrasting high‐latitude biomes: subarctic forest tundra (Kuujjuarapik, Quebec) and high arctic polar desert (Resolute, Nunavut). Five substrate types were tested (beads, bacteria, picophytoplankton, filamentous plankton and microbial mats). Special attention was given to the role of benthos, a component that is usually poorly integrated into models of aquatic foodwebs. 3. Consistent with observations elsewhere in the circumpolar region, high concentrations of adult macrozooplankton occurred in all sites (up to 17 100 crustaceans m^?3) while phytoplankton concentrations and primary productivity were low. The communities were composed of multiple species, including Daphnia middendorfiana, Hesperodiaptomus arcticus, Leptodiaptomus minutus, Artemiopsis stefanssoni and Branchinecta paludosa. 4. Detritus made 89–98% of the planktonic resource pool and bacteria contributed the highest biomass (up to 29 mg C m^?3) of the planktonic food particles available to zooplankton. Benthic resources were dominated by microbial mats that grew in nutrient‐rich conditions at the base of the ponds and which dominated overall ecosystem biomass and productivity. 5. All species were flexible in their feeding but there were large, order of magnitude differences in clearance rates among taxa. These differences likely resulted from different grazing strategies among cladocerans, copepods and fairy shrimps, and possibly also from adaptation to specific food types and size ranges that occur locally in these waters. 6. The subarctic cladocerans Ceriodaphnia quadrangula and D. middendorfiana, and the arctic fairy shrimp B. paludosa were observed to graze directly on the microbial mats and the feeding experiments confirmed their assimilation of benthic substrates. The other zooplankton species showed a more pelagic feeding mode but were capable of using microbial mat filaments, thus may be indirectly linked to benthic processes via resuspension. 7. Our study indicates that the classical aquatic food web in which phytoplankton provide the sole production base for grazers does not apply to northern shallow lakes and ponds. Instead, microbial mats increase the physical complexity of these high latitude ecosystems and likely play a role in sustaining their high zooplankton biomass.     

Biological indicators track differential responses of pelagic and littoral areas to nutrient load reductions in German lakes

External nutrient loading was reduced over the past decades as a measure for improving the water quality of eutrophic lakes in western Europe, and has since been accelerated by the adoption of the European Water Framework Directive (WFD) in 2000 (EC, 2000). A variety of eutrophication-related metrics have indicated that the response of biological communities to this decreased nutrient loading has been diverse. Phytoplankton, a major component of the pelagic community, often responded rapidly, whereas a significant delay was observed for submerged macrophytes colonizing littoral areas. In this study we tested whether assessment methods developed for phytoplankton and macrophytes in lakes during Germany's implementation of the WFD reflect this differential response. An assessment of 263 German lakes confirmed that a lower ecological state was recorded when based on the biological quality element (BQE) for macrophytes than the BQE for phytoplankton during the investigated period (2003–2012). On average, lakes had a moderate ecological status for both phytoplankton and macrophyte BQEs, but differences of up to three classes were observed in single cases. Long-term data were available for five lowland lakes subject to strong reductions in phosphorus loading. Their phytoplankton-based assessments indicated a constant improvement of the ecological status in parallel to decreasing water phosphorus concentrations. In contrast, macrophyte-based assessments indicated a 10–20 year delay in their ecological recovery following nutrient load reduction. This delay was confirmed by detailed data on the temporal development of macrophyte species diversity and maximum colonization depths of two lakes after nutrient load reduction. We conclude that the available WFD assessment methods for phytoplankton and macrophyte BQEs are suitable to track the differential response of pelagic and littoral areas to nutrient load reductions in German lakes.     

Trophic-based diversification in benthivorous charrs (Salvelinus) dwelling littoral zones of Northern lakes

Charrs of the genus Salvelinus form distinct trophic morphs living in sympatry in numerous postglacial lakes. Here we demonstrate that charrs can diversify into amphipod foraging specialists and sedentary macrobenthos consumers in the shallow-water ecosystems. This pattern was revealed in three out of six lakes under exploration supported by differences in stomach content, trophic-transmitted parasite, and stable isotope ratio analyzes. The body shape and growth rate comparison indicates that this kind of trophic-based diversification emerges at a juvenile stage and is maintained throughout the whole life. The restriction in gene flow found between the morphs allows to propose the possibility for the hereditable-based specialization to evolve. We found that those diversification phenomena are possible only in the lakes situated in vicinity of the ocean coastline, while no evidence of this split was found for inland mountain lakes. We suggest that the trophic-based diversification in the littoral ecosystems is accounted for the heterogeneity in the ecological conditions and food resources’ distribution linked to coastal wind action. This phenomenon was previously reported for the charr in Lake Fjellfrosvatn, Scandinavia, so it seems to be some universal yet poorly described kind of disruptive selection pressure for northern latitude fishes.

     

Bacterioplankton production in freshwater Antarctic lakes

1. Bacterioplankton production was measured in the water columns of two ultra‐oligotrophic, freshwater Antarctic lakes (Crooked Lake and Lake Druzhby) during an annual cycle. In both lakes bacterial production, measured by the incorporation of [³H] thymidine, continued in winter and showed a cycle over the year. The range of production was between 0 and 479 ng C L^?1 h^?1 in Crooked Lake and 0–354 ng L^?1 h^?1 in Lake Druzhby. 2. Abundance and mean cell volume both varied, producing marked changes in biomass during the year, with highest biomass occurring in the winter and early spring. Biomass showed similar seasonal trends in both lakes. 3. For most of the year inorganic forms of nitrogen and phosphorus were detectable in the water columns of the lakes and were unlikely to have limited bacterial production. Dissolved organic carbon (DOC) was below 3000 μg L^?1. Dissolved amino acids and carbohydrates contributed 5–25% of the DOC pool in Crooked Lake and 5–64% in Lake Druzhby. Dissolved carbohydrates were consistently low, suggesting that this may have been the preferred carbon substrate for bacterioplankton. 4. Aggregate associated bacteria had higher mean cell volume, abundances and production than freely suspended bacteria in Lake Druzhby, while in Crooked Lake aggregate associated bacteria consistently had higher mean cell volumes than free bacteria, but abundance and production were on occasion higher in free bacteria compared with aggregate associated communities. 5. The data indicated that production is limited by continuous low temperatures and the limited availability of suitable DOC substrate. However, the bacterioplankton functions year round, responding to factors other than temperature.     

Role of macrophytes in the variability of zooplankton community structure in the littoral zone of shallow lakes

Distribution, diurnal variability, aggregation of zooplankton in the littoral zone of lakes and effect of various macrophyte species on the structure of its community are considered. It is shown that the horizontal migrations of zooplankton, both direct and reverse ones, are caused mainly by the pressure of fish. The effect of predacious zooplankton is less important and is directed mainly at small-sized species. The intensity of horizontal migrations of zooplankton decreases with depth, while the effect of shore avoidance is observed for the large-sized zooplankton species and is caused not only by the pressure of fish but also by other factors, most likely abiotic. The problem of interaction between macrophytes and zooplankton cannot be reduced to the role of macrophytes as a refuge. Allelopathic properties of macrophytes, competitive relations between separate zooplankton species in macrophyte thickets, as well as the effect of predacious invertebrates associated with macrophytes on zooplankton remain unclear. The role of macrophytes as a factor causing changes in hydrodynamic processes in the littoral regions of lakes is also unknown.     

Seasonal changes in the diets and relative abundances of perch and roach in the littoral and pelagic zones of a large lake

In the Enonselkä and Laitialanselkä basins of Lake Vesijärvi, perch Perca fluviatilis and roach Rutilus rutilus were abundant in the littoral and in the pelagic zones throughout the summer. In the littoral zone, roach was always more numerous than perch, while perch dominated in the open water. Intraspecific diet overlap values were higher than interspecific values. In the pelagic zone, perch <155 mm fed mainly on the cladoceran Leptodora kindtii , while small bosminids were most important food items for roach. Large perch were piscivorous, feeding mainly on smelt Osmerus eperlanus . In the littoral zone small perch foraged on zooplankton and chironomid larvae and large perch on chironomids and fish (small perch). Small roach fed mainly on bosminids and detritus, while for roach <185 mm macrophytes ( Elodea Canadensis, Lemna trisulca ) were also of importance. Detritus was more common in the food of roach in Laitialanselkä than in Enonselkä. The slower growth rate of roach in Laitialanselkä compared with Enonselkä was probably connected with this. However, considering the latitude of the lake, the growth rate of both roach and perch was relatively fast in both basins. The results indicated that in a large lake both perch and roach are able to utilize effectively the different habitats and diverse food resources. By segregation in food resource utilization they are able to co-exist in large quantities, at the same time maintaining a relatively fast growth rate.     

Distribution and growth of benthic macroinvertebrates among different patch types of the littoral zones of two arctic lakes

1. To evaluate the effect of habitat patch heterogeneity on abundance and growth of macroinvertebrates in arctic lakes, macroinvertebrate abundance, individual biomass, and potential food resources were studied in three patch types in two arctic lakes on the Alaskan North Slope near the Toolik Lake Field Station. An experiment was conducted to determine which sediment patch type supported higher growth rates for Chironomus sp., a commonly occurring macroinvertebrate. 2. Potential organic matter (OM) resources were significantly higher in both rock and macrophyte patches than in open‐mud patches. Total macroinvertebrate densities in both lakes were highest in rock patches, intermediate in macrophytes and lowest in open‐mud. The open‐mud patches also had lower species richness compared with other patch types. Additionally, individual biomass for one clam species and two chironomid species was significantly greater in rock patches than in open‐mud. 3. In a laboratory experiment, Chironomus showed two to three times greater mass increase in sediments from macrophyte and rock patches than from open‐mud patches. Rock and macrophyte experimental sediments had at least 1.5 × the percentage OM as open‐mud sediments. 4. Chlorophyll a appeared to be the best predictor for invertebrate abundances across all patch types measured, whereas OM content appeared to be the variable most closely associated with Chironomus growth. 5. Our results combined with previous studies show that the relationships between macroinvertebrate community structure, individual growth, and habitat heterogeneity are complex, reflecting the interaction of multiple resources, and biotic interactions, such as the presence or absence of a selective vertebrate predator (lake trout, Salvelinus namaycush).     

Spatial scale and benthic community organisation in the littoral zones of large oligotrophic lakes: potential for cross-scale interactions

1. We studied the spatial organisation of littoral, benthic invertebrate communities in two large oligotrophic lakes with very extensive heterogeneous littoral zones, and report that littoral communities show significant variation in their structure at multiple spatial scales. 2. At coarser scales, littoral zone topography, riparian inputs and exposure may be important drivers of community spatial structure. At finer scales, the composition of substrates, particularly macrophytes and inorganic sediments, may be drivers of community spatial structure. 3. The effects of substrates on community spatial patterns were non‐additive across scales; community‐habitat correspondence became very noisy when we attempted to extrapolate results throughout the littoral. We present some evidence to suggest that this non‐additivity is because of the interaction among structuring processes that operate at different scales.     

Distribution and coexistence patterns of phytoplankton in subtropical shallow lakes and the role of niche-based and spatial processes

Phytoplankton is an important microbial component in many ecosystems, and thus, knowing the predictive variables of its distribution is very valuable. In this study, we evaluated the roles of niche-based and spatial processes on phytoplankton communities of nine lakes in southern Brazil (spatial distance covered of 220 km). Variation partitioning analysis was employed to determine the relative contributions of environmental and spatial variables on the distribution of prokaryotic (PRP) and eukaryotic (EUP) components of phytoplankton at multispatial extents. C-score analysis was carried out, at intra- and interlake scales, to examine whether phytoplankton species coexist less or more than expected by chance (taxa segregation and aggregation, respectively). Our results showed that, at broader spatial scales, both PRP and EUP distributions are mainly influenced by pure space, while at finer spatial scales, the two groups are mainly influenced by pure environment. PRP distribution at broader spatial scales was also significantly influenced by spatially structured environment, and was in general more affected by spatial variables than by EUP. Finally, a taxa segregation pattern was observed at the interlake scale. In general, our results suggest that niche-based processes are more important in structuring phytoplankton on the local scale while spatial processes on the regional scale.     

Relationships between microcystins and environmental parameters in 30 subtropical shallow lakes along the Yangtze River, China

1. A survey of 30 subtropical shallow lakes in the middle and lower reaches of the Yangtze River area in China was conducted during July–September in 2003–2004 to study how environmental and biological variables were associated with the concentration of the cyanobacterial toxin microcystin (MC). 2. Mean MC concentration in seasonally river‐connected lakes (SL) was nearly 33 times that in permanently river‐connected lakes (RL), and more than six times that in city lakes (NC) and non‐urban lakes (NE) which were not connected to the Yangtze River. The highest MC (8.574 μg L^?1) was detected in Dianshan Lake. 3. MC‐RR and MC‐LR were the primary toxin variants in our data. MC‐RR, MC‐YR and MC‐LR were significantly correlated with Chl a, biomass of cyanobacteria, Microcystis and Anabaena, indicating that microcystins were mainly produced by Microcystis and Anabaena sp. in these lakes. 4. Nonlinear interval maxima regression indicated that the relationships of Secchi depth, total nitrogen (TN) : total phosphorus (TP) and NH with MC were characterised by negative exponential curves. The relationships between MC and TN, TP, NO + NO were fitted well with a unimodal curve. 5. Multivariate analyses by principal component and classifying analysis indicated that MC was mainly affected by Microcystis among the biological factors, and was closely related with temperature among physicochemical factors.     

Autochthonous production in shallow littoral zones of five floodplain rivers: effects of flow,turbidity and nutrients

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Seasonal fluctuations in macrophyte cover and water transparency of four brown-water lakes: implications for crustacean zooplankton in littoral and pelagic habitats

The dynamics of crustacean zooplankton in the littoral and pelagic zones of four forest lakes having variable water qualities (colour range 130–340 mg Pt l⁻¹, Secchi depth 70–160 cm) were studied. The biomass of zooplankton was higher in the littoral zone than in the pelagic zone only in the lake having the highest transparency. In the three other lakes, biomass was significantly higher in the pelagic zone than in the littoral zone. In the two lakes with highest transparency, the littoral biomass of cladocerans significantly followed the development of macrophyte vegetation, and cladoceran biomass reached the maximum value at the time of highest macrophyte coverage. In lakes with lowest transparency, littoral zooplankton biomass developed independently of macrophyte density and decreased when macrophyte beds were densest. The seasonal development of the littoral copepod biomass did not follow the development of macrophytes in any of the lakes. The mean size of cladocerans in the pelagic zone decreased with increasing Secchi depth of the lake, whereas in the littoral zone no such phenomenon was detected. Seasonally, when water transparency increased temporarily in two of the lakes, the mean size of cladocerans in the pelagic zone decreased steeply. For copepods, no relationship between water transparency and body size was observed. The results suggested that in humic lakes the importance of the littoral zone as a refuge decreases with decreasing transparency of the water and that low water transparency protects cladocerans from fish predation. All the observed between-lake differences could not be explained by fish predation, but were probably attributed to the presence of chaoborid larvae with variable densities. Feeding efficiency of chaoborids is not affected by visibility and thus they can obscure the relationship between water quality, fish density, and the structure of crustacean zooplankton assemblages. Handling editor: S. I. Dodson