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.     

Can warm climate‐related structure of littoral predator assemblies weaken the clear water state in shallow lakes?

Shallow lakes, the most abundant lake type in the world, are very sensitive to climatic changes. The structure and functioning of shallow lakes are greatly impacted by submerged plants, and these may be affected by climate warming in various, contrasting, ways. Following a space‐for‐time substitution approach, we aimed to analyse the role of aquatic (submerged and free‐floating) plants in shallow lakes under warm climates. We introduced artificial submerged and free‐floating plant beds in five comparable lakes located in the temperate zone (Denmark, 55–57 °N) and in the subtropical zone (Uruguay, 30–35 °S), with the aim to study the structure and dynamics of the main associated communities. Regardless of differences in environmental variables, such as area, water transparency and nutrient status, we found consistent patterns in littoral community dynamics and structure (i.e. densities and composition of fish, zooplankton, macroinvertebrates, and periphyton) within, but substantial differences between, the two regions. Subtropical fish communities within the macrophyte beds exhibited higher diversity, higher density, smaller size, lower relative abundance of potentially piscivores, and a preference for submerged plants, compared with otherwise similar temperate lakes. By contrast, macroinvertebrates and cladocerans had higher taxon richness and densities, and periphyton higher biomass, in the temperate lakes. Several indicators suggest that the fish predation pressure was much stronger among the plants in the subtropical lakes. The antipredator behaviour of cladocerans also differed significantly between climate zones. Submerged and free‐floating plants exerted different effects on the spatial distribution of the main communities, the effects differing between the climate zones. In the temperate lakes, submerged plants promoted trophic interactions with potentially positive cascading effects on water transparency, in contrast to the free‐floating plants, and in strong contrast to the findings in the subtropical lakes. The higher impact of fish may result in higher sensitivity of warm lakes to external changes (e.g. increase in nutrient loading or water level changes). The current process of warming, particularly in temperate lakes, may entail an increased sensitivity to eutrophication, and a threat to the high diversity, clear water state.     

Sediments,not plants,offer the preferred refuge for Daphnia against fish predation in Mediterranean shallow lakes: an experimental demonstration

1. Different behavioural responses of planktonic animals to their main predators, fish, have been reported from shallow lakes. In north temperate lakes, large‐bodied zooplankton may seek refuge from predation among macrophytes, whereas in subtropical lakes, avoidance of macrophytes has been observed. The prevalent behaviour probably depends on the characteristics of the fish community, which in Mediterranean lakes is typically dispersed in both the open water zone and in the littoral, as in temperate lakes, and is dominated by small size classes, as in subtropical lakes. 2. We performed ‘habitat choice’ experiments to test the response of Daphnia magna to predation cues at both the horizontal and vertical level by mimicking a ‘shallow littoral’ zone with plants and a ‘deeper pelagic’ zone with sediments. 3. Initial separate response experiments showed that natural plants, artificial plants and predation cues all repelled D. magna in the absence of other stimuli, while sediments alone did not trigger any significant response by D. magna. 4. The habitat choice experiments showed that, in the presence of predation cues and absence of plants, Daphnia moved towards areas with sediment. In the presence of both plants and sediments, Daphnia moved away from the plants towards the sediments under both shallow and deep water treatment conditions. 5. Based on these results, we suggest that Daphnia in Mediterranean shallow lakes avoid submerged macrophytes and instead prefer to hide near the sediment when exposed to predation risk, as also observed in subtropical shallow lakes. This pattern is not likely to change with water level alterations, a common feature of lakes in the region, even if the effectiveness of the refuge may be reduced.     

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.     

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.     

Impact of fish predation on cladoceran body weight distribution and zooplankton grazing in lakes during winter

1. It is well accepted that fish, if abundant, can have a major impact on the zooplankton community structure during summer, which, particularly in eutrophic lakes, may cascade to phytoplankton and ultimately influence water clarity. Fish predation affects mean size of cladocerans and the zooplankton grazing pressure on phytoplankton. Little is, however, known about the role of fish during winter. 2. We analysed data from 34 lakes studied for 8–9 years divided into three seasons: summer, autumn/spring and winter, and four lake classes: all lakes, shallow lakes without submerged plants, shallow lakes with submerged plants and deep lakes. We recorded how body weight of Daphnia and then cladocerans varied among the three seasons. For all lake types there was a significant positive correlation in the mean body weight of Daphnia and all cladocerans between the different seasons, and only in lakes with macrophytes did the slope differ significantly from one (winter versus summer for Daphnia). 3. These results suggest that the fish predation pressure during autumn/spring and winter is as high as during summer, and maybe even higher during winter in macrophyte‐rich lakes. It could be argued that the winter zooplankton community structure resembles that of the summer community because of low specimen turnover during winter mediated by low fecundity, which, in turn, reflects food shortage, low temperatures and low winter hatching from resting eggs. However, we found frequent major changes in mean body weight of Daphnia and cladocerans in three fish‐biomanipulated lakes during the winter season. 4. The seasonal pattern of zooplankton : phytoplankton biomass ratio showed no correlation between summer and winter for shallow lakes with abundant vegetation or for deep lakes. For the shallow lakes, the ratio was substantially higher during summer than in winter and autumn/spring, suggesting a higher zooplankton grazing potential during summer, while the ratio was often higher in winter in deep lakes. Direct and indirect effects of macrophytes, and internal P loading and mixing, all varying over the season, might weaken the fish signal on this ratio. 5. Overall, our data indicate that release of fish predation may have strong cascading effects on zooplankton grazing on phytoplankton and water clarity in temperate, coastal situated eutrophic lakes, not only during summer but also during winter.     

An experimental study of habitat choice by Daphnia: plants signal danger more than refuge in subtropical lakes

1. In shallow temperate lakes, submerged plants often provide refuge for pelagic zooplankton against fish predation, a mechanism with potential strong cascading effects on water transparency and on the entire ecosystem. In (sub)tropical lakes, however, the interaction between aquatic plants and predation may be more complex, particularly because fish density is high within the plant beds in such systems. 2. Using laboratory ‘habitat choice’ experiments, we determined the effects of three (sub)tropical free‐floating plants, Eichhornia crassipes, Pistia stratiotes and Salvinia auriculata and the cosmopolitan submerged Ceratophyllum demersum, on horizontal movement by the water flea Daphnia obtusa. We tested for avoidance of plants in the absence and presence of alarm signals from crushed conspecifics and chemical cues from the fish Cnesterodon decemmaculatus, the fish have been subjected to different feeding regimes. 3. In the absence of other stimuli, D. obtusa strongly avoided the plants and the crushed conspecifics, as expected. However, the response to fish was insignificant regardless of their previous feeding regime. The avoidance of free‐floating plants was more pronounced than that of the submerged plant. Contrary to predictions based on research in temperate lakes, Daphnia did not take refuge among the plants but rather swam away from them when exposed simultaneously to plants and alarm signals. 4. We hypothesise that the avoidance of plants by D. obtusa may ultimately be attributable to an expectedly higher predation risk within the plants than in the pelagic, because of a high density of associated zooplanktivorous fish in the former. In the (sub)tropics, therefore, aquatic plants and particularly the free‐floating ones, may not promote cascading effects via Daphnia grazing on phytoplankton as seen in temperate eutrophic lakes.     

Vertical, lateral and longitudinal movement of zooplankton in a large river

1. The spatial distribution and movement patterns of zooplankton in large rivers are little known compared with those in lake environments. We conducted a series of studies in the Ohio River (U.S.A.) during the low flow period to assess diel vertical (DVM), longitudinal and lateral movement of crustacean zooplankton. 2. The dominant large zooplankter, the copepod Eurytemora affinis, showed a consistent vertical migration pattern of daytime ascent and night‐time descent during all sampling periods – the reverse of the most common migratory pattern of zooplankton in lakes. The cladoceran Bosmina migrated in a similar way in two of the three sampling periods. Surveys taken longitudinally in the river showed similar trends for both taxa. 3. During the lateral surveys, E. affinis was significantly more abundant in the shallow littoral zone during the night than in the daytime. The combination of vertical and lateral movement patterns along with the diel distribution of zooplanktivorous fish suggest that these movements are a predator‐avoidance mechanism. 4. Sampling programmes in large rivers should consider that larger zooplankton such as E. affinis may not be randomly distributed in the river channel and behaviours such as diel vertical migration may be just as evident in river habitats as in lakes.     

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.     

Applicability of planktonic biomanipulation for managing eutrophication in the subtropics

Although large-bodied cladocerans such asDaphnia can reduce algal biomass significantly in temperate lakes if freed from fish predation, the applicability of such biomanipulation techniques for eutrophication management in the subtropics and tropics has been examined only recently. Subtropical cladoceran assemblages differ from those of temperate lakes by their low species richness, early summer gameogenesis, and greatly reduced body size. Eutrophic Florida lakes are dominated by pump-filter feeding fish rather than by size selective planktivores as a temperate lakes. Cladocerans in Florida lakes can increase in abundance significantly if freed from fish but fail to have an impact on algal biomass or composition. The greatest potential for using biomanipulation to manage phytoplankton-dominated lakes in the subtropics and tropics lies with phytophagous fish. Future research should concentrate on defining the role of individual fish taxa on phytoplankton composition and community structure, nutrient cycling, and planktonic productivity before embarking on whole lake manipulation projects.     

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.     

The potential of artificial refugia for maintaining a community of large-bodied cladocera against fish predation in a shallow eutrophic lake

The Norfolk Broads are a series of shallow lakes which are highly eutrophic and typified by dense populations of phytoplankton and an absence of submerged aquatic plants. The zooplankton community is subject to intense predation pressure by young fish and is dominated by small-bodied organisms which have a low potential for reducing phytoplankton populations through grazing. Various designs and densities of artificial refugia for zooplankton against fish predation were established in Hoveton Great Broad in order to enhance populations of large-bodied Cladocera. Initially some of the refuges contained higher densities and larger individuals ofDaphnia andCeriodaphnia than the surrounding open water. However, towards the end of the first season and throughout the subsequent two years, population densities and size-structure were similar both within and outside the refuges, although there was still evidence of enhanced body-size ofDaphnia within the refuges compared with the open water. The provision of habitat structures designed as refugia from fish predation did not enhance large-bodied cladoceran populations enough to promote this restoration strategy as feasible for eutrophic and shallow lakes.     

Mesocosm experiments on nutrient and fish effects on shallow lake food webs in a Mediterranean climate

1. Nutrient and fish manipulations in mesocosms were carried out on food‐web interactions in a Mediterranean shallow lake in south‐east Spain. Nutrients controlled biomass of phytoplankton and periphyton, while zooplankton, regulated by planktivorous fish, influenced the relative percentages of the dominant phytoplankton species. 2. Phytoplankton species diversity decreased with increasing nutrient concentration and planktivorous fish density. Cyanobacteria grew well in both turbid and clear‐water states. 3. Planktivorous fish increased concentrations of soluble reactive phosphorus (SRP). Larger zooplankters (mostly Ceriodaphnia and copepods) were significantly reduced when fish were present, whereas rotifers increased, after fish removal of cyclopoid predators and other filter feeders (cladocerans, nauplii). The greatest biomass and diversity of zooplankton was found at intermediate nutrient levels, in mesocosms without fish and in the presence of macrophytes. 4. Water level decrease improved underwater light conditions and favoured macrophyte persistence. Submerged macrophytes (Chara spp.) outcompeted algae up to an experimental nutrient loading equivalent to added concentrations of 0.06 mg L^?1 PO₄‐P and 0.6 mg L^?1 NO₃‐N, above which an exponential increase in periphyton biomass and algal turbidity caused characean biomass to decline. 5. Declining water levels during summer favoured plant‐associated rotifer species and chroococcal cyanobacteria. High densities of chroococcal cyanobacteria were related to intermediate nutrient enrichment and the presence of small zooplankton taxa, while filamentous cyanobacteria were relatively more abundant in fishless mesocosms, in which Crustacea were more abundant, and favoured by dim underwater light. 6. Benthic macroinvertebrates increased significantly at intermediate nutrient levels but there was no relationship with planktivorous fish density. 7. The thresholds of nutrient loading and in‐lake P required to avoid a turbid state and maintain submerged macrophytes were lower than those reported from temperate shallow lakes. Mediterranean shallow lakes may remain turbid with little control of zooplankton on algal biomass, as observed in tropical and subtropical lakes. Nutrient loading control and macrophyte conservation appear to be especially important in these systems to maintain high water quality.     

High predation is of key importance for dominance of small-bodied zooplankton in warm shallow lakes: evidence from lakes, fish exclosures and surface sediments

The mean body size of limnetic cladocerans decreases from cold temperate to tropical regions, in both the northern and the southern hemisphere. This size shift has been attributed to both direct (e.g. physiological) or indirect (especially increased predation) impacts. To provide further information on the role of predation, we compiled results from several studies of subtropical Uruguayan lakes using three different approaches: (i) field observations from two lakes with contrasting fish abundance, Lakes Rivera and Rodó, (ii) fish exclusion experiments conducted in in-lake mesocosms in three lakes, and (iii) analyses of the Daphnia egg bank in the surface sediment of eighteen lakes. When fish predation pressure was low due to fish kills in Lake Rivera, large-bodied Daphnia appeared. In contrast, small-sized cladocerans were abundant in Lake Rodó, which exhibited a typical high abundance of fish. Likewise, relatively large cladocerans (e.g. Daphnia and Simocephalus) appeared in fishless mesocosms after only 2 weeks, most likely hatched from resting egg banks stored in the surface sediment, but their abundance declined again after fish stocking. Moreover, field studies showed that 9 out of 18 Uruguayan shallow lakes had resting eggs of Daphnia in their surface sediment despite that this genus was only recorded in three of the lakes in summer water samples, indicating that Daphnia might be able to build up populations at low risk of predation. Our results show that medium and large-sized zooplankton can occur in subtropical lakes when fish predation is removed. The evidence provided here collectively confirms the hypothesis that predation, rather than high-temperature induced physiological constraints, is the key factor determining the dominance of small-sized zooplankton in warm lakes.     

The structuring role of free-floating versus submerged plants in a subtropical shallow lake

In shallow temperate lakes many ecological processes depend on submerged macrophytes. In subtropical and tropical lakes, free-floating macrophytes may be equally or more important. We tested the hypothesis that different macrophyte growth forms would be linked with different bottom-up and top-down mechanisms in out-competing phytoplankton. We compared experimentally the effects of submerged and free-floating plants on water chemistry, phytoplankton biomass, zooplankton and fish community structure in a shallow hypertrophic lake (Lake Rodó, 34°55S 56°10W, Uruguay). Except for the retention of suspended solids, we found no other significant bottom-up process connected with either Eichhornia crassipes or Potamogeton pectinatus. Free-floating plants had a lower abundance of medium-sized zooplankton than any other microhabitat and submerged plants were apparently preferred by microcrustaceans. Fish showed a differential habitat use according to species, size-class and feeding habits. Dominant omnivore-planktivores, particularly the smallest size classes, preferred submerged plants. In contrast, omnivore-piscivores were significantly associated with free-floating plants. The density of omnivorous-planktivorous fish, by size class, significantly explained the distribution of medium-sized zooplankton, the high number of size 0 fish being the main factor. The abiotic environment and the structure of the zooplankton community explained little of the fish distribution pattern. Our results suggest that bottom-up effects of free-floating plants are weak when cover is low or intermediate. Top-down effects are complex, as effects on zooplankton and fish communities seem contradictory. The low piscivores:planktivores ratio in all microhabitats suggests, however, that cascading effects on phytoplankton through free-floating plant impacts on piscivorous fish are unlikely to be strong.     

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)     

Trophic structure, species richness and biodiversity in Danish lakes: changes along a phosphorus gradient

1. Using data from 71, mainly shallow (an average mean depth of 3 m), Danish lakes with contrasting total phosphorus concentrations (summer mean 0.02–1.0 mg P L?l), we describe how species richness, biodiversity and trophic structure change along a total phosphorus (TP) gradient divided into five TP classes (class 1–5: <0.05, 0.05–0.1, 0.1–0.2, 0.2–0.4,> 0.4 mg P L?1).
2. With increasing TP, a significant decline was observed in the species richness of zooplankton and submerged macrophytes, while for fish, phytoplankton and floating‐leaved macrophytes, species richness was unimodally related to TP, all peaking at 0.1–0.4 mg P L?1. The Shannon–Wiener and the Hurlbert probability of inter‐specific encounter (PIE) diversity indices showed significant unimodal relationships to TP for zooplankton, phytoplankton and fish. Mean depth also contributed positively to the relationship for rotifers, phytoplankton and fish.
3. At low nutrient concentrations, piscivorous fish (particularly perch, Perca fluviatilis) were abundant and the biomass ratio of piscivores to plankti‐benthivorous cyprinids was high and the density of cyprinids low. Concurrently, the zooplankton was dominated by large‐bodied forms and the biomass ratio of zooplankton to phytoplankton and the calculated grazing pressure on phytoplankton were high. Phytoplankton biomass was low and submerged macrophyte abundance high.
4. With increasing TP, a major shift occurred in trophic structure. Catches of cyprinids in multiple mesh size gill nets increased 10‐fold from class 1 to class 5 and the weight ratio of piscivores to planktivores decreased from 0.6 in class 1 to 0.10–0.15 in classes 3–5. In addition, the mean body weight of dominant cyprinids (roach, Rutilus rutilus, and bream, Abramis brama) decreased two–threefold. Simultaneously, small cladocerans gradually became more important, and among copepods, a shift occurred from calanoid to cyclopoids. Mean body weight of cladocerans decreased from 5.1 μg in class 1 to 1.5 μg in class 5, and the biomass ratio of zooplankton to phytoplankton from 0.46 in class 1 to 0.08–0.15 in classes 3–5. Conversely, phytoplankton biomass and chlorophyll a increased 15‐fold from class 1 to 5 and submerged macrophytes disappeared from most lakes.
5. The suggestion that fish have a significant structuring role in eutrophic lakes is supported by data from three lakes in which major changes in the abundance of planktivorous fish occurred following fish kill or fish manipulation. In these lakes, studied for 8 years, a reduction in planktivores resulted in a major increase in cladoceran mean size and in the biomass ratio of zooplankton to phytoplankton, while chlorophyll a declined substantially. In comparison, no significant changes were observed in 33 ‘control’ lakes studied during the same period.     

A quantitative test of the size efficiency hypothesis by means of a physiologically structured model

According to the size‐efficiency hypothesis (SEH) larger bodied cladocerans are better competitors for food than small bodied species. In environments with fish, however, the higher losses of the large bodied species due to size‐selective predation may shift the balance in favor of the small bodied species. Here we present a theoretical framework for the analysis of the competitive abilities of zooplankton species that takes both competition and predation into account in one coherent analysis. By applying the conceptually well‐understood framework of physiologically structured population models we were able to predict the relative difference in predation rates necessary to cause a shift in dominance of the large‐bodied species (Daphnia pulicaria) to the small‐bodied species (D. galeata). These predictions depend only on seven easily interpretable parameters per species: size at birth, size at maturity and maximum size, age at maturity, maximal clutch size, egg development time and finally the half‐saturation constant for food. The critical equilibrium mortality of D. pulicaria was 0.16 d^?1 at food concentrations close to the critical food concentration of D. galeata, i.e. D. pulicaria will win the competition as long as its mortality rate is below 0.16 d^?1. At higher food concentrations the differential mortality curve (plotting equilibrium mortalities of both species against each other) approached a linear function with a slope of one and an intercept equal to the difference in maximal population birth rates. The prediction of critical predation rates was independent of the ingestion rate of the cladocerans and the algal carrying capacity and food regeneration rate of the environment although the mechanism works through competition for a shared algal food resource. We interpret these findings in terms of the relative predation risk large and small‐bodied cladocerans will face in various freshwater ecosystems.     

Diel zooplankton community feeding activity and filtration rates of Pseudoboeckella volucris and Daphniopsis studeri on sub-antarctic Marion Island

Experiments measuring zooplankton filter-feeding rates using a ¹⁴C-labelled green alga were carred out in situ for the first time in three shallow sub-antarctic lakes. The number and biomass of zooplankton were high in samples from two lakes enriched by penguin and seal faeces, and low in samples from a peat-lined lava-lakelet not biotically enriched. A diel survey of the lava-lakelet over 48 h showed that no vertical migration or diel variation in filter-feeding rate occured in the strongly pigmented copepod Pseudoboeckella volucris Kiefer population, the most abundant entomostracan present. The unpigmented cladoceran Daphniopsis studeri Rühe had a high in situ filter-feeding rate, exhibited very marked vertical migration, and was primarily a nocturnal grazer.     

Zooplankton response to extreme drought in a large subtropical lake

Plankton data from 1997 to 2005 were used to examine impacts of a managed draw-down, subsequent drought and resulting historic low water levels (during 2000 and 2001) on the zooplankton of Lake Okeechobee, Florida. Prior to the drought the lake supported less than 150 ha of submerged vegetation. Following the drought, over 15,000 ha of submerged vegetation developed around the lake shore and conditions favored greater survival of age 0 fish. The zooplankton changed significantly from the pre- to post-drought period, including: (a) a near-complete loss of all dominant species of cladocerans and rotifers; and (b) an abrupt transition to a community with over 80% of total biomass comprised of Arctodiaptomus dorsalis, a calanoid copepod previously described as being resistant to fish predation. These changes persisted over a 5 year post-drought sampling period. In contrast, there were no systematic changes in biomass of bacteria, phytoplankton, inedible cyanobacteria, algal cell size, suspended solids, or any other physical or chemical attributes known to affect zooplankton in shallow lakes. Evidence points towards increased predation by fish, and perhaps invertebrates, as factors responsible for loss of cladocerans and rotifers following the drought, and indicates a need for future research to link changes in water level to shifts in predation pressure in this and other shallow lakes. Handling editor: S.I. Dodson