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.     

The role of macroinvertebrates and fish in regulating the provision by macrophytes of refugia for zooplankton in a warm temperate shallow lake

1. The zooplankton often undergoes diel horizontal migration (DHM) from the open water to the littoral of shallow lakes, thus avoiding predators in the former. This behaviour has functional impacts within the lake, as it enhances zooplankton survival, increases their control of phytoplankton and tends to stabilise the clear water state. However, most of the evidence supporting this migration pattern comes from cold north temperate lakes, and more evidence from tropical and subtropical areas, as well as from southern temperate areas, is needed. 2. We conducted a field study of the diel horizontal and vertical migration of zooplankton, and the horizontal distribution of potential predatory macroinvertebrates and fish, over two consecutive days in the summer in a temperate lake in the southern hemisphere. We took zooplankton samples at two depths, at three sampling stations (inside beds of aquatic macrophytes, at their edge and in open water) along three transects running from the centre of a bed of Ceratophyllum demersum to open water. At each sampling station, we also took samples of macroinvertebrates and fish and measured physical and chemical environmental variables. 3. Zooplankton (pelagic cladocerans, calanoid copepods and rotifers) avoided the shore, probably because of the greater risk from predators there. Larger and more vulnerable cladocerans, such as Diaphanosoma brachyurum and Moina micrura, were two to four times more abundant in open water than at the edge of or inside beds of macrophytes, respectively, by both day and night. Less vulnerable zooplankton [i.e. of medium body size (Ceriodaphnia dubia) or with the ability to swim fast (calanoid copepods)] were distributed evenly between open water and the edge of the plant beds. Small zooplankton, Bosmina huaronensis and pelagic rotifers, showed an even distribution among the three sampling stations. Accordingly, no DHM of zooplankton occurred, although larger organisms migrated vertically inside C. demersum stands. 4. Macrophytes contained high densities of predatory macroinvertebrates and fish. The predator assemblage, composed of large‐bodied macroinvertebrates (including odonates and shrimps) and small littoral fish, was permanently associated with submerged macrophytes. None of these groups moved outside the plant beds or changed their population structure (fish) over the diel cycle. 5. Submerged macrophyte beds do not represent a refuge for zooplankton in lakes where predators are numerous among the plants, implying a weaker top‐down control of phytoplankton biomass by zooplankton and, consequently, a more turbid lake. The effectiveness of macrophytes as a refuge for zooplankton depends on the associated assemblage of predatory macroinvertebrates and fish among the plants.     

Does roach behaviour differ between shallow lakes of different environmental state?

The diel activity levels and spatial distribution of roach Rutilus rutilus differed markedly between two shallow lakes of different environmental state. The movements of roach (12–25 cm L _T), with surgically implanted mini‐radio transmitters, were monitored regularly during several 48 h tracking sessions in a clearwater and in a turbid lake. In both lakes, the roach in general were most active during dawn and dusk and least active during the night. Activity level in midsummer was lowest around noon in the clear lake and high around noon in the turbid lake. In summer, roach in the clear lake stayed passively in a restricted area of water lilies during the day and moved into the central part of the lake during the night. In the turbid lake, roach were dispersed all over the lake during the day and moved close to the shoreline at night. Predator : prey fish ratios did not differ in the two lakes, however the observed behaviour of roach in the clearwater lake may be explained by a larger predation pressure from fish and birds, both being favoured in the clear water.     

Lake size and fish diversity determine resource use and trophic position of a top predator in high‐latitude lakes

Prey preference of top predators and energy flow across habitat boundaries are of fundamental importance for structure and function of aquatic and terrestrial ecosystems, as they may have strong effects on production, species diversity, and food‐web stability. In lakes, littoral and pelagic food‐web compartments are typically coupled and controlled by generalist fish top predators. However, the extent and determinants of such coupling remains a topical area of ecological research and is largely unknown in oligotrophic high‐latitude lakes. We analyzed food‐web structure and resource use by a generalist top predator, the Arctic charr Salvelinus alpinus (L.), in 17 oligotrophic subarctic lakes covering a marked gradient in size (0.5–1084 km²) and fish species richness (2–13 species). We expected top predators to shift from littoral to pelagic energy sources with increasing lake size, as the availability of pelagic prey resources and the competition for littoral prey are both likely to be higher in large lakes with multispecies fish communities. We also expected top predators to occupy a higher trophic position in lakes with greater fish species richness due to potential substitution of intermediate consumers (prey fish) and increased piscivory by top predators. Based on stable carbon and nitrogen isotope analyses, the mean reliance of Arctic charr on littoral energy sources showed a significant negative relationship with lake surface area, whereas the mean trophic position of Arctic charr, reflecting the lake food‐chain length, increased with fish species richness. These results were supported by stomach contents data demonstrating a shift of Arctic charr from an invertebrate‐dominated diet to piscivory on pelagic fish. Our study highlights that, because they determine the main energy source (littoral vs. pelagic) and the trophic position of generalist top predators, ecosystem size and fish diversity are particularly important factors influencing function and structure of food webs in high‐latitude lakes.     

Trophic dynamics in a whole lake experiment: size-structured interactions and recruitment variation

Horizontal and vertical heterogeneity as a result of size‐structured processes are important factors influencing indirect effects in food webs. In a whole‐lake experiment covering 5 years, we added the intermediate consumer roach (Rutilus rutilus) to two out of four lakes previously inhabited by the omnivorous top predator perch (Perca fluviatilis). We focused our study on the direct consumption effect of roach presence on zooplankton (and indirectly phytoplankton) versus the indirect effect of roach on zooplankton (and phytoplankton) mediated via effects on perch reproductive performance. The patterns in zooplankton and phytoplankton abundances were examined in relation to population density of roach and perch including young‐of‐the‐year (YOY) perch in the light of non‐equilibrium dynamics. The presence of roach resulted in changed seasonal dynamics of zooplankton with generally lower biomasses in May–June and higher biomasses in July–August in roach lakes compared to control lakes. Roach presence affected perch recruitment negatively and densities of YOY perch were on average higher in control lakes than in treatment lakes. In years when perch recruitment did not differ between lakes as a result of experimental addition of perch eggs, total zooplankton biomass was lower in treatment lakes than in control lakes. Phytoplankton biomass showed a tendency to increase in roach lakes compared to control lakes. Within treatment variation in response variables was related to differences in lake morphometry in treatment lakes. Analyses of the trophic dynamics of each lake separately showed strong cascading effects of both roach and YOY perch abundance on zooplankton and phytoplankton dynamics. Consideration of the long transients in the dynamics of top predators (fish) in aquatic systems that are related to their long life span involving ontogenetic niche shifts is essential for making relevant interpretations of experimental perturbations. This conclusion is further reinforced by the circumstance that the intrinsic dynamics of fish populations may in many cases involve high amplitude dynamics with long time lags.     

Effects of piscivore-mediated habitat use on growth, diet and zooplankton consumption of roach: an individual-based modelling approach

SUMMARY 1. We used an individual based modelling approach for roach to (i) simulate observed diel habitat shifts between the pelagic and littoral zone of a mesotrophic lake; (ii) analyse the relevance of these habitat shifts for the diet, activity costs and growth of roach; and (iii) quantify the effects of a hypothetical piscivore-mediated (presence of pikeperch) confinement of roach to the littoral zone on roach diet, activity costs and growth.
2. The model suggests that in the presence of pikeperch, roach shifts from zooplankton as the primary diet to increased consumption of less nutritious food items such as macrophytes, filamentous algae and detritus.
3. The growth of roach between May and October was predicted to be significantly higher in the absence of pikeperch, although the net activity costs were about 60% higher compared with the scenario where pikeperch were present.
4. These modelling results provide quantitative information for interpreting diel horizontal migrations of roach as a result from a trade-off between food availability and predation risk in different habitats of a lake.
5. Altering the habitat selection mode of planktivorous roach by piscivore stocking has the potential to reduce zooplankton consumption by fish substantially, and could therefore be used as a biomanipulation technique complementing the reduction of zooplanktivorous fish.     

The diet and growth of roach (Rutilus rutilus (L.)) in Lake Vesijärvi and possible changes in the course of biomanipulation

Previous studies have suggested that the roach Rutilus rutilus (L.) stock of Lake Vesijärvi is one of the main factors delaying the recovery of the lake after sewage diversion. This study is concerned with the documentation of the diet of roach in the lake. In total, 531 roach were examined. Both in the pelagial and in the littoral the roach had mixed diets in May and in September—October. The importance of zooplankton decreased and the importance of benthos and plants increased with increasing size of roach. In July, in the pelagic zone all sizes of roach fed exclusively on zooplankton (Bosmina spp.), while in the littoral zooplankton had the highest volume proportions only in the smallest (<130 mm) roach. The frequent use of plant food and slow growth rate of large roach indicate a low availability of animal prey. As the fish densities decrease due to the mass removal taking place in the lake, the percentage of plant food in the diets of roach will probably decrease and the growth of roach will increase. Additionally, the tendency of the roach to migrate into the pelagic zone in early summer may be reduced, which would decrease their predation on the zooplankton.     

Depth gradients in food‐web processes linking habitats in large lakes: Lake Superior as an exemplar ecosystem

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Rapid and unexpected effects of piscivore introduction on trophic position and diet of perch (Perca flavescens) in lakes recovering from acidification and metal contamination

1. Yellow perch (Perca flavescens) are often the only surviving fish species in acidified lakes. We studied four lakes along a gradient of recovery from acidification and that had different food web complexities. All had abundant yellow perch, two had low piscivore abundance, one had a well‐established piscivore population and one was manipulated by introducing piscivorous smallmouth bass (Micropterus dolomieu). We hypothesised that there would be strong effects on perch abundance, behaviour and diet induced by the presence of piscivores. 2. In the manipulated lake, the bass reduced yellow perch abundance by 75% over a 2‐year period. Concomitantly, perch use of the pelagic habitat fell from 48 to 40%. 3. In contrast to findings from less disturbed systems, yellow perch in the littoral zone of the manipulated lake did not strongly shift from zooplankton to benthic food sources after the arrival of piscivores. Diet analysis using stable carbon isotopes revealed a strong continued reliance on zooplankton in all lakes, independent of the degree of piscivory. The failure to switch to benthos in the refuge area of the littoral zone is most likely related to the depauperate benthos communities in these formerly acidified lakes. 4. Yellow perch in lakes recovering from acidification face a considerable ecological challenge as the necessary switch to benthic diet is hindered by a low abundance of benthos. The arrival of piscivores in these recovering lakes imposes further restrictions on perch access to food items. We infer that future recovery of perch populations (and higher trophic levels) will have to be preceded by the re‐establishment of diverse benthic macroinvertebrate communities in these lakes.     

Determinants of the distribution of juvenile fish in the littoral area of a shallow lake

1. Research has often focused on the pelagic areas of lakes; the littoral zone has received less attention. The few studies concerning fish distribution in littoral habitats have concentrated on stands of submersed macrophytes, whereas other littoral habitat types have seldom been investigated. 2. This study aimed to predict the occurrence of juvenile fish in several littoral habitats of a shallow lake as a function of food availability, complexity of habitat structure, water depth and substrate. Habitats comprising reed, woody structures, and two open water areas differing in depth were sampled for fish and invertebrate biomasses on two shores, over 6 months and during both daylight and at night. 3. The juvenile fish community consisted almost exclusively of 0+ and 1+ roach and perch. There was a strong diel component in habitat use, with a predominant occurrence of fish in complex habitats (mainly woody structures) during the day, and a partial migration towards the open habitats at night, more strongly expressed in roach than in perch. 4. The diet of all fish groups was relatively constant over the seasonal cycle, and was independent of habitat. There was a higher degree of planktivory in roach than in perch, but both species fed on benthic macroinvertebrates to a substantial extent. 5. According to a logistic regression model, the biomass of potential food organisms in the different habitats had little predictive effect on the spatial distribution of the fish, whereas the structural complexity of the habitats combined with the diel cycle explained about 28% of the occurrence patterns in 0+ and 1+ perch and 1+ roach.     

Niche segregation of coexisting Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) constrains food web coupling in subarctic lakes

1. Generalist fish species are recognised as important couplers of benthic and pelagic food‐web compartments in lakes. However, interspecific niche segregation and individual specialisation may limit the potential for generalistic feeding behaviour. 2. We studied summer habitat use, stomach contents and stable isotopic compositions of the generalist feeder Arctic charr coexisting with its common resource competitor brown trout in five subarctic lakes in northern Norway to reveal population‐level and individual‐level niche plasticity. 3. Charr and trout showed partial niche segregation in all five lakes. Charr used all habitat types and a wide variety of invertebrate prey including zooplankton, whereas trout fed mainly on insects in the littoral zone. Hence, charr showed a higher potential to promote habitat and food‐web coupling compared to littoral‐dwelling trout. 4. The level of niche segregation between charr and trout and between pelagic‐caught and littoral‐caught charr depended on the prevailing patterns of interspecific and intraspecific resource competition. The two fish species had partially overlapping trophic niches in one lake where charr numerically dominated the fish community, whereas the most segregated niches occurred in lakes where trout were more abundant. 5. In general, pelagic‐caught charr had substantially narrower dietary and isotopic niches and relied less on littoral carbon sources compared to littoral‐caught conspecifics that included generalist as well as specialised benthivorous and planktivorous individuals. Despite the partially specialised planktivorous niche and thus reduced potential of pelagic‐dwelling charr to promote benthic–pelagic coupling, the isotopic compositions of both charr subpopulations suggested a significant reliance on both littoral and pelagic carbon sources in all five study lakes. 6. Our study demonstrates that both interspecific niche segregation between and individual trophic specialisation within generalist fish species can constrain food‐web coupling and alter energy mobilisation to top consumers in subarctic lakes. Nevertheless, pelagic and littoral habitats and food‐web compartments may still be highly integrated due to the potentially plastic foraging behaviour of top consumers.     

Contrasting Zooplankton Assemblages in Two Oxbow Lakes with Low Transparencies and Narrow Emergent Macrophyte Belts (Krapina River,Croatia)

The aim of this study was to examine the combined effect of water transparency and narrow macrophyte belts on zooplankton assemblages in two oxbow lakes (Krapina River, Croatia). Samples were collected in open water and among helophytes in the littoral zone from April until September 2008. Rotifers were the most abundant group of zooplankton in both lakes, and dominated in the Krapina oxbow lake 1 (KO1). Lake KO1 had significantly lower transparency, lower percentage macrophyte cover and higher chlorophyll a concentration than Krapina oxbow lake 2 (KO2). In lake KO1, variation in the horizontal distribution of cladocerans and rotifers in terms of their abundance seemed to be determined by competition between Bosmina longirostris and Keratella cochlearis, initiated by oscillation in transparency and detritus availability. In lake KO2, with higher transparency and higher percentage macrophyte cover, the abundance of small‐ and large‐bodied cladocerans increased in the littoral zone simultaneously with higher transparency, suggesting fish predation. Results of this study indicated that small differences in transparencies between the two lakes caused significant differences in horizontal distribution of the zooplankton assemblage. Even narrow helophyte belts offered a refuge to zooplankton, although lower transparencies reduced the effectiveness of macrophytes as a refuge from predators. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

The role of phosphorus release by roach [Rutilus rutilus (L.)] in the water quality changes of a biomanipulated lake

SUMMARY 1. Following fish removal, the water quality in biomanipulated lakes often improves concomitant with decreased phosphorus (P) levels. Because the decrease in P concentrations derives most probably either directly or indirectly from fish, which are the main target of biomanipulation, this study examined the P release of 0+, 1+ and 2+ roach [Rutilus rutilus (L.)] and changes in the P release during summer in a shallow eutrophic lake in Finland. 2. The P release was separated into P derived from benthic and littoral food items and into recycled P derived from feeding on zooplankton, to estimate the contribution of net P additions to the water column by the fish to the increase in P concentrations of the lake water (75–110 mg P m^?3) in summer 1991–96. 3. Individual P release of roach by both egestion and excretion was estimated with a bioenergetics model. The size of the roach population was estimated with a depletion method and the proportions of different age groups from catch samples, using a programme separating mixtures of normal distributions. The sensitivity of the release estimates to variation in the growth data was estimated with the jackknife technique. 4. The biomass‐specific P release by 0+ roach (0.36–0.54 mg P g^?1 day^?1) was higher than that by older roach (0.07–0.16 mg P g^?1 day^?1) throughout the summer. The P release by the whole roach population deriving from benthic and littoral food items (0.7–2.7 mg m^?3 during July to August, representing a net addition to the water column) was 5–19 times lower in 1991–96 than the recycled P release deriving from zooplankton (8.9–25.7 mg m^?3), and too low to explain the increase in the P concentration of the lake water during the summer. Because the biomass‐specific P release and roach diet composition vary with fish age, it is important to consider the age structure of fish populations to obtain correct estimates of P release and net additions to the water column. 5. The removal of roach by fishing diminished the roach stock greatly, but the fish‐mediated P release to the water column changed little. This effect was because of the high compensation capacity of the roach population, leading to high recruitment of young fish with higher biomass‐specific P release rates. 6. External loading is very low during summer months and therefore it cannot explain the increase in the P concentration of water during that time. Internal loading from the sediment might be as high as 10.2 mg P m^?2 day^?1, i.e. 50 times higher than the maximum net P addition by the total roach population.     

Timing of the diet shift from zooplankton to macroinvertebrates and size at maturity determine whether normally piscivorous fish can persist in otherwise fishless lakes

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Community structure and diel migration of zooplankton in shallow brackish lakes: role of salinity and predators

Diel horizontal migration (DHM), where zooplankton moves towards macrophytes during daytime to avoid planktivorous fish, has been reported as a common migration pattern of zooplankton in shallow temperate freshwater lakes. However, in shallow eutrophic brackish lakes, macrophytes seem not to have the same refuge effect, as these lakes may remain turbid even at relatively high macrophyte abundances. To investigate the extent to which macrophytes serve as a refuge for zooplankton at different salinities, we introduced artificial plants mimicking submerged macrophytes in the littoral zone of four shallow lakes, with salinities ranging from almost freshwater (0.3) to oligohaline waters (3.8). Furthermore, we examined the effects of different salinities on the community structure. Diel samples of zooplankton were taken from artificial plants, from areas where macrophytes had been removed (intermediate areas) and, in two of the lakes, also in open water. Fish and macroinvertebrates were sampled amongst the artificial plants and in intermediate areas to investigate their influence on zooplankton migration. Our results indicated that diel vertical migration (DVM) was the most frequent migration pattern of zooplankton groups, suggesting that submerged macrophytes were a poor refuge against predation at all salinities under study. Presumably, this pattern was the result of the relatively high densities of small planktivorous fish and macroinvertebrate predators within the submerged plants. In addition, we found major differences in the composition of zooplankton, fish and macroinvertebrate communities at the different salinities and species richness and diversity of zooplankton decreased with increasing salinity. At low salinities both planktonic/free-swimming and benthic/plant-associated cladocerans occurred, whilst only benthic ones occurred at the highest salinity. The low zooplankton biomass and overall smaller-bodied zooplankton specimens may result in a lower grazing capacity on phytoplankton, and enhance the turbid state in nutrient rich shallow brackish lakes.     

Restoring lakes by using artificial plant beds: habitat selection of zooplankton in a clear and a turbid shallow lake

1. Return of large‐bodied zooplankton populations is of key importance for creating a shift from a turbid to a clear‐water state in shallow lakes after a nutrient loading reduction. In temperate lakes, recovery is promoted by submerged macrophytes which function as a daytime refuge for large zooplankton. However, recovery of macrophytes is often delayed and use of artificial plant beds (APB) has been suggested as a tool to enhance zooplankton refuges, thereby reinforcing the shift to a clear‐water state and, eventually, colonisation of natural plants. 2. To further evaluate the potential of APB in lake restoration, we followed the day–night habitat choices of zooplankton throughout summer in a clear and a turbid lake. Observations were made in the pelagic and littoral zones and in APB in the littoral representing three different plant densities (coverage 0%, 40% and 80%). 3. In the clear lake, the zooplankton (primarily Daphnia) were mainly found in the pelagic area in spring, but from mid‐May they were particularly abundant in the APB and almost exclusively so in mid‐June and July, where they appeared in extremely high densities during day (up to 2600 ind. L⁻¹). During night Daphnia densities were overall more equally distributed between the five habitats. Ceriodaphnia was proportionally more abundant in the APB during most of the season. Cyclopoids were more abundant in the high APB during day but were equally distributed between the five habitats during night. 4. In the turbid lake, however, no clear aggregation was observed in the APB for either of the pelagic genera (Daphnia and Bosmina). This may reflect a higher refuge effect in the open water due to the higher turbidity, reduced ability to orient to plant beds and a significantly higher fish density (mainly of roach, Rutilus rutilus, and perch, Perca fluviatilis) in the plant beds than in the clear lake. Chydorus was found in much higher proportions among the plants, while cyclopoids, particularly the pelagic Cyclops vicinus, dominated in the pelagic during day and in the pelagic and high density plants during night. 5. Our results suggest that water clarity is decisive for the habitat choice of large‐bodied zooplankton and that introduction of APB as a restoration measure to enhance zooplankton survival is only a useful tool when water clarity increases following loading reduction. Our results indicate that dense APB will be the most efficient.     

Extensive diel fish migrations in a deep ultraoligotrophic lake of Patagonia Argentina

Andean Patagonian lakes are ultraoligotrophic, highly transparent water bodies with very low densities of organisms at the different levels of their trophic food webs. Little is known about spatial distribution and habitat use by fish in these lakes. Hydroacoustic and active sampling techniques were used to study the distribution, composition, and displacement of organisms in an Andean lake throughout the diel cycle on two different moon phase days. Sound scattering layers (SSLs) were found both in the near-shore and pelagic habitats at different times of the day. These SSLs, formed by galaxiid larvae and adults, underwent displacement during the time periods of dawn and dusk, giving rise to the redistribution of organisms in the different habitats. The organisms show high sensitivity to light intensity, displaying different behaviors depending on moon phases. Extensive diel vertical migrations show that the deep pelagic habitats provide diurnal refuge for native galaxiids in Andean Patagonian lakes. Due to the high densities of galaxiid larvae and adults, it is likely that diel migrations generate an active flow of energy and matter between habitats, which could have a profound influence on whole lake dynamics.     

Top-down and bottom-up impacts of juvenile fish in a littoral reed stand

1. Research has often focused on pelagic food chains and processes of lakes; less is known about the contribution of benthic energy flows to whole‐lake ecosystem energetics. This stems from the fact that the shoreline and littoral habitats, which provide a key linkage between sediment and water column, have only recently become a significant focus for study. 2. This study aimed to quantify the feeding and phosphorus allocation of a juvenile fish community in a littoral zone of a shallow lake in response to the biomass succession of the invertebrate prey community. Habitats comprising reed and adjacent open water were sampled over two consecutive years during day and night. 3. Although there were substantial year‐to‐year differences in the biomass of invertebrates, the fish community composition, diet consumption rates and phosphorus allocations were very similar in both study years. Biomasses and predation impacts by juvenile fish on prey groups were substantially higher within the reeds than in the adjacent open water habitat. This may be explained by the refuge‐seeking behaviour of the fish. 4. In general, invertebrates were negligibly influenced by fish feeding, with the exception for a strong top‐down control of large cladocerans. In response to the resulting low Daphnia biomass, fish were forced to switch to a higher degree of benthivory. Consequently, juvenile fish in littoral reed stands may shift benthic‐derived energy and phosphorus via the excretion of soluble reactive phosphorus into the open water.     

To migrate, or not to migrate: partial diel horizontal migration of fish in a temperate freshwater reservoir

The diel horizontal migration (DHM) of fish between the inshore and offshore zones of the ?ímov Reservoir (Czech Republic, deep, stratified, meso-eutrophic) was investigated by a combination of horizontal and vertical hydroacoustic surveys at 3-h intervals over 48 h and day/night purse seining in August 2007. An overwhelming majority of fish were aggregated within the epilimnetic layer. Considering only the horizontal surveys, cyclic diel fish movements between inshore and offshore habitats were apparent, while the total fish biomass remained constant between day and night. A higher fish biomass was detected in the offshore zone during daytime by both hydroacoustics and purse seining. In contrast, a higher fish biomass was recorded at night in the inshore zone. Bream Abramis brama, roach Rutilus rutilus, and perch Perca fluviatilis dominated the daytime offshore fish assemblage whereas bleak Alburnus alburnus prevailed at night. Bream and roach decreased in abundance at night while perch completely disappeared from the offshore habitat. The diel differences in size distributions and direct catches suggested the population-wide horizontal offshore migration of bleak and inshore migration of all perch during dusk. On the other hand, partial inshore migration of bream and roach adults was observed during dusk (52 and 80%, respectively). The different proportions of offshore residents among species and size classes suggested that differences in size, and, therefore, predation vulnerability, contributed to the observed migration patterns.     

Habitat selection and diel distribution of the crustacean zooplankton from a shallow Mediterranean lake during the turbid and clear water phases

1. The fish fauna of many shallow Mediterranean Lakes is dominated by small‐bodied exotic omnivores, with potential implications for fish–zooplankton interactions still largely unknown. Here we studied diel variation in the vertical and horizontal distribution of the crustacean plankton in Lake Vela, a shallow polymictic and eutrophic lake. Diel sampling was carried out on three consecutive days along a horizontal transect, including an open‐water station and a macrophyte (Nymphaea alba) bed. Since transparency is a key determinant of the predation risk posed by fish, the zooplankton sampling campaigns were conducted in both the turbid (autumn) and clear water (spring) phases. 2. In the turbid phase, most taxa were homogeneously distributed along the vertical and horizontal axes in the three consecutive days. The only exception was for copepod nauplii, which showed vertical heterogeneity, possibly as a response to invertebrate predators. 3. In the clear water phase, most zooplankton taxa displayed habitat selection. Vertically, the general response consisted of a daily vertical migration (DVM), despite the limited depth (1.6 m). Horizontally, zooplankters showed an overall preference for the pelagic zone, independent of the time of the day. Such evidence is contrary to the postulated role of macrophytes as an anti‐predator refuge for the zooplankton. 4. These vertical (DVM) and horizontal (macrophyte‐avoidance) patterns were particularly conspicuous for large Daphnia, suggesting that predation risk from size‐selective predators (fish) was the main factor behind the spatial heterogeneity of zooplankton in the spring. Thus, the difference in the zooplankton spatial distribution pattern and habitat selection among seasons (turbid and clear water phases) seems to be mediated the predation risk from fish, which is directly related to water transparency. 5. The zooplankton in Lake Vela have anti‐predator behaviour that minimises predation from fish. We hypothesise that, due to the distinct fish community of shallow Mediterranean lakes, aquatic macrophytes may not provide adequate refuge to zooplankters, as seen in northern temperate lakes.