Piscivory of 0+ pike (Esox lucius L.) in a small eutrophic lake and its implication for biomanipulation

Stocking 0+ pike in shallow eutrophic lakes as an indirect way to facilitate clear water by increased phytoplankton grazing by zooplankton, has been used in several studies with varying success. Lack of piscivory could be one reason for poor impact of the pike stockings, on reducing the 0+ cyprinid fish population. In relation to this, we present data on food and feeding habits of 936 0+ pike (Esox lucius L.) (2–18 cm) sampled during five years in a small eutrophic lake undergoing biomanipulation. Overall, the food of 0+ pike followed a sequence of diet shifts including microcrustacea, insects, macrocrustacea and finally vertebrates (fish) with increase in pike length. Despite a high abundance of potential prey fish in the lake, in general less than 50% of the pike were piscivorous until a length of 10.1–13.7 cm. As piscivory increased, the principal prey fish (0+ roach) apparently displayed antipredatorial behaviour which decreased their availability for 0+ pike. We conclude that lack of piscivory caused by e.g. anti predator tactics by prey fish, may reduce the impact of 0+ pike stockings in lake restoration projects.

     

Substitution of top predators: effects of pike invasion in a subarctic lake

1. Invasions of top predators may have strong cascading effects in ecosystems affecting both prey species abundance and lower trophic levels. A recently discussed factor that may enhance species invasion is climate change and in this context, we studied the effects of an invasion of northern pike into a subarctic lake ecosystem formerly inhabited by the native top predator Arctic char and its prey fish, ninespined stickleback. 2. Our study demonstrated a strong change in fish community composition from a system with Arctic char as top predator and high densities of sticklebacks to a system with northern pike as top predator and very low densities of sticklebacks. A combination of both predation and competition from pike is the likely cause of the extinction of char. 3. The change in top predator species also cascaded down to primary consumers as both zooplankton and predator‐sensitive macroinvertebrates increased in abundance. 4. Although the pike invasion coincided with increasing summer temperatures in the study area we have no conclusive evidence that the temperature increase is the causal mechanism behind the pike invasion. But still, our study provides possible effects of future pike invasions in mountain lakes related to climate change. We suggest that future pike invasions will have strong effects in lake ecosystems, both by replacing native top consumers and through cascading effects on lower trophic levels.     

Prey vulnerability to a gape-size limited predator: behavioural and morphological impacts on northern pike piscivory

The northern pike ( Esox lucius ) is a selective and important predator in lake ecosystems. Prey size in pike is limited by pike gape size, which is a linear function of pike body length. Here we show that the absolute gape-size limit in pike is greater than previously considered, and that maximum ingestible prey size is limited by prey body depth. Further, we experimentally show that pike prefer shallow-bodied roach before deeper-bodied common bream, and small prey sizes within each prey species. Handling time in pike increases with prey body depth, and since common bream are deeper-bodied than roach, handling time is longer for bream than for roach of the same length, but equal considering body depth. Prey handling time is suggested to be a major cost to the pike, since it increases the risk of losing the prey, as well as exposure to predation, kleptoparasitism and cannibalism. Consequently, prey vulnerability is determined by risk of predation and intraspecific interactions, and behavioural preferences in the pike, and not by pike gape-size limits. The consequences for natural populations is evaluated by analysing size structures of predator and prey fish populations in a eutrophic lake.     

Prey naivety in the behavioural responses of juvenile Chinook salmon (Oncorhynchus tshawytscha) to an invasive predator

1. Non‐native predators might inflict proportionally higher mortality on prey that have no previous experience of them, compared to species that have coexisted with the predator for some time. 2. We tested whether juvenile Chinook salmon (Oncorhynchus tshawytscha) were less able to recognise a non‐native than a native predator, by investigating behavioural responses to the chemical cues of the invasive smallmouth bass (Micropterus dolomieu) and the native northern pikeminnow (Ptychocheilus oregonensis) in both laboratory and field experiments. 3. Laboratory results demonstrated strong innate antipredator responses of individual juvenile Chinook salmon to northern pikeminnow; fish spent 70% of time motionless and exhibited 100% greater panic response than in controls. By contrast, antipredator responses to the chemical cues of smallmouth bass did not differ from controls. 4. These results were supported by similar differences in recognition of these predator odours by groups of juvenile Chinook salmon in fully natural conditions, though responses reflected a greater range of antipredator behaviours by individuals. In field trials, responses to northern pikeminnow odour resulted in increased flight or absence, reductions in swimming and foraging, and increased time spent near the substratum, compared to smallmouth bass odour. 5. Given that survival of juvenile fish is facilitated by predator recognition, our results support the hypothesis that naivety may be an important factor determining the effect of non‐native predators on prey populations. Efforts to manage the effect of native and non‐native predators may benefit by considering complex behavioural interactions, such as these at the individual and group levels.     

Fish–duck interactions in boreal lakes in Finland as reflected by abundance correlations

We studied the hypothesis that fish play an important role in lake use by ducks (pairs and broods) in boreal lakes. The study was based on densities of different duck and fish species in 28 boreal lakes in southern Finland. We focused on the three most common duck species (mallard Anas platyrhynchos, green-winged teal A. crecca and common goldeneye Bucephala clangula) and on the three most common fish species (perch Perca fluviatilis, roach Rutilus rutilus and pike Esox lucius) in the region. We considered both competitive and predatory interactions between ducks and fish, the perch and roach being potential competitors with ducks and the pike a potential predator of ducks. We found a negative association between green-winged teal brood density and total fish density, the other duck species having only a weak association with total fish density. When the three fish species were considered separately, a negative association, suggesting food competition, was found between perch, green-winged teal and goldeneye, whereas the role of roach as a food competitor seemed to be of minor importance. We did not find any clear signs of predatory effects of pike on ducks. Our results suggest that food competition is a more important factor than pike predation in affecting lake use by ducks in oligotrophic boreal environments in southern Finland.     

Restoration by biomanipulation in a small hypertrophic lake: first-year results

Biomanipulation was carried out in order to improve the water quality of the small hypertrophic Lake Zwemlust (1.5 ha; mean depth 1.5 m). In March 1987 the lake was drained to facilitate the elimination of fish. Fish populations were dominated by planktivorous and benthivorous species (total stock c. 1500 kg) and were collected by seine- and electro-fishing. The lake was subsequently re-stocked with 1500 northern pike fingerlings (Esox lucius L.) and a low density of adult rudd (Scardinius erythrophthalmus). The offspring of the rudd served as food for the predator pike. Stacks of Salix twigs, roots of Nuphar lutea and plantlets of Chara globularis were brought in as refuge and spawning grounds for the pike, as well as shelter for the zooplankton.The impact of this biomanipulation on the light penetration, phytoplankton density, macrophytes, zooplankton and fish communities and on nutrient concentrations was monitored from March 1987 onwards. This paper presents the results in the first year after biomanipulation.The abundance of phytoplankton in the first summer (1987) after this biomanipulation was very low, and consequently accompanied by increase of Secchi-disc transparency and drastic decline of chlorophyll a concentration.The submerged vegetation remained scarce, with only 5 % of the bottom covered by macrophytes at the end of the season.Zooplankters became more abundant and there was a shift from rotifers to cladocerans, comprised mainly of Daphnia and Bosmina species, the former including at least 3 species.The offspring of the stocked rudd was present in the lake from the end of August 1987. Only 19% of the stocked pike survived the first year.Bioassays and experiments with zooplankton community grazing showed that the grazing pressure imposed by the zooplankton community was able to keep chlorophyll a concentrations and algal abundance to low levels, even in the presence of very high concentrations of inorganic N and P. The total nutrient level increased after biomanipulation, probably due to increased release from the sediment by bioturbation, the biomass of chironomids being high.At the end of 1987 Lake Zwemlust was still in an unstable stage. A new fish population dominated by piscivores, intended to control the planktivorous and benthivorous fish, and the submerged macrophytes did not yet stabilize.     

Selenium Health Benefit Values as Seafood Safety Criteria

Selenium (Se) is absolutely required for activity of 25–30 genetically unique enzymes (selenoenzymes). All forms of life that have nervous systems possess selenoenzymes to protect their brains from oxidative damage. Homeostatic mechanisms normally maintain optimal selenoenzyme activities in brain tissues, but high methylmercury (MeHg) exposures sequester Se and irreversibly inhibit selenoenzyme activities. However, nutritionally relevant amounts of Se can replace the Se sequestered by MeHg and maintain normal selenoenzyme activities, thus preventing oxidative brain damage and other adverse consequences of MeHg toxicity. Findings of studies that seem contradictory from MeHg exposure perspectives are entirely consistent from MeHg:Se molar ratio perspectives. Studies that have reported dose-dependent consequences of maternal MeHg exposures on child development uniformly involved seafoods that contained much more Hg than Se. Meanwhile more typical varieties of ocean fish contain much more Se than Hg. This may explain why maternal MeHg exposure from eating ocean fish is associated with major IQ benefits in children instead of harm. Therefore, instead of being avoided, ocean fish consumption should be encouraged during pregnancy. However, the safety of freshwater fish consumption is less certain. In freshwater fish, MeHg bioaccumulation and toxicity are both inversely related to Se bioavailability. Their Se can be far lower than their MeHg contents, potentially making them more dangerous than pilot whale meats. Therefore, to provide accurate and appropriate regulatory advice regarding maternal consumption of seafoods and freshwater fish, Hg:Se molar ratios need to be incorporated in food safety criteria.     

Acquired Recognition of Chemical Stimuli from Pike,Esox lucius,by Brook Sticklebacks,Culaea inconstans (Osteichthyes,Gasterosteidae)

In this study we test whether brook sticklebacks (Culaea inconstans) can acquire predator recognition through releaser-induced recognition learning, i.e. simultaneous exposure to aversive ('releasing') stimuli and neutral stimuli causing learned aversion to the neutral stimuli. We exposed wild-caught pike-naive brook sticklebacks (collected from a creek containing fathead minnows, Pimephales promelas, but not pike, Esox lucius) to chemical stimuli from pike that were mixed with brook stickleback skin extract, fathead minnow skin extract, or a control of distilled water. In subsequent tests 2 d later, when only pike stimuli were presented, sticklebacks conditioned with stickleback skin extract and fathead minnow skin extract exhibited antipredator behaviour (i.e. increased schooling and movement toward the substrate), while those conditioned with distilled water did not. Sticklebacks conditioned with stickleback skin extract responded to pike with a more intense response, in terms of movement toward the substrate, than those conditioned with fathead minnow skin extract, suggesting that conspecific skin extract may be a stronger stimulus than heterospecific skin extract for learning recognition of predators. To our knowledge this is the first study to demonstrate that an acanthopterygian fish can acquire predator recognition through the pairing of conspecific alarm pheromone with the cue of a predator. Furthermore, our results are the first to demonstrate that fish can acquire predator recognition through the pairing of a heterospecific alarm pheromone with the cue of a predator. These results suggest that brook sticklebacks will benefit by being in close proximity to fathead minnows. Acquired predator recognition has long-term consequences in mediating predator-prey interactions.     

Dynamics of a Boreal Lake Ecosystem during a Long-Term Manipulation of Top Predators

We assessed the long-term (16 years) effects of introducing piscivores (northern pike) into a small, boreal lake (Lake 221, Experimental Lakes Area) containing abundant populations of two planktivorous fish species. After the introduction, pearl dace were extirpated and yellow perch abundance was greatly reduced. Daphnia species shifted from D. galeata mendota to larger bodied Daphnia catawba, but the total zooplankton biomass did not increase, nor did the biomass of large grazers such as Daphnia. Phytoplankton biomass decreased after the northern pike introduction, but increased when northern pike were partially removed from the lake. Phosphorus (P) excretion by fish was ∼0.18 mg P m⁻² d⁻¹ before pike addition, declined rapidly to approximately 0.03–0.10 as planktivorous perch and dace populations were reduced by pike, and increased back to premanipulation levels after the pike were partially removed and the perch population recovered. When perch were abundant, P excretion by fish supported about 30% of the P demand by primary producers, decreasing to 6–14% when pike were abundant. Changes in phytoplankton abundance in Lake 221 appear to be driven by changes in P cycling by yellow perch, whose abundance was controlled by the addition and removal of pike. These results confirm the role of nutrient cycling in mediating trophic cascades and are consistent with previous enclosure experiments conducted in the same lake.     

Projected shifts in fish species dominance in Wisconsin lakes under climate change

Temperate lakes may contain both coolwater fish species such as walleye (Sander vitreus) and warmwater fish species such as largemouth bass (Micropterus salmoides). Recent declining walleye and increasing largemouth bass populations have raised questions regarding the future trajectories and management actions for these species. We developed a thermodynamic model of water temperatures driven by downscaled climate data and lake‐specific characteristics to estimate daily water temperature profiles for 2148 lakes in Wisconsin, US, under contemporary (1989–2014) and future (2040–2064 and 2065–2089) conditions. We correlated contemporary walleye recruitment and largemouth bass relative abundance to modeled water temperature, lake morphometry, and lake productivity, and projected lake‐specific changes in each species under future climate conditions. Walleye recruitment success was negatively related and largemouth bass abundance was positively related to water temperature degree days. Both species exhibited a threshold response at the same degree day value, albeit in opposite directions. Degree days were predicted to increase in the future, although the magnitude of increase varied among lakes, time periods, and global circulation models (GCMs). Under future conditions, we predicted a loss of walleye recruitment in 33–75% of lakes where recruitment is currently supported and a 27–60% increase in the number of lakes suitable for high largemouth bass abundance. The percentage of lakes capable of supporting abundant largemouth bass but failed walleye recruitment was predicted to increase from 58% in contemporary conditions to 86% by mid‐century and to 91% of lakes by late century, based on median projections across GCMs. Conversely, the percentage of lakes with successful walleye recruitment and low largemouth bass abundance was predicted to decline from 9% of lakes in contemporary conditions to only 1% of lakes in both future periods. Importantly, we identify up to 85 resilient lakes predicted to continue to support natural walleye recruitment. Management resources could target preserving these resilient walleye populations.     

Development of fish communities in lakes after biomanipulation

Biomanipulation measures in the Netherlands are usually a combination of a drastic fish stock reduction and an introduction of pike fingerlings. In three small shallow lakes (Noorddiep, Bleiswijkse Zoom and Zwemlust) these measures resulted in a clear water state and the development of macrophytes. After the measures the fish community developed differently because of the new physical and biological conditions. Results of lake Noorddiep and lake Bleiswijkse Zoom showed that the fish community became more divers. Bream and carp became less dominant and were partly replaced by roach and perch. The importance of the main predator pike-perch was strongly reduced and replaced by pike and perch. The share of piscivorous fish in the total fish stock increased at all sites. The recruitment of young-of-the-year was similar or even higher in the clear overgrown areas than in the turbid water before the measures, but the recruitment of young-of-the-year to older fish differed between the species. Predation by pike and perch could not control the young-of-the-year cyprinids, but their predation may have contributed to the shift from bream to roach, because of selective predation on bream in the open water, while roach was hiding in the vegetation. The macrophytes provide new refugia and feeding conditions that favour roach and perch, but offer relatively poor survival conditions for bream and carp.     

Heterogeneity in levels of serum neutralizing antibodies against viral hemorrhagic septicemia virus genotype IVB among fish species in Lake St. Clair, Michigan, USA

The presence of neutralizing antibodies against viral hemorrhagic septicemia virus (VHSV-IVb) was investigated in sera of 13 fish species collected from Lake St. Clair, Michigan, USA, a VHSV-endemic water body. We tested 297 sera collected May 2004-June of 2010, using a complement-dependent 50% plaque neutralization test (50% PNT). Neutralizing antibodies were detected in 23% (67/297) of the samples. The highest overall antibody prevalence (85%, 34/40) and mean positive antibody titer (12,113 ± 11,699 SD) were detected in muskellunge (Esox masquinongy). Antibodies were also detected in 50% (15/30) of sampled northern pike (E. lucius), 25% (15/61) of freshwater drum (Aplodinotus grunniens), and 7% (3/41) of smallmouth bass (Micropterus dolomieu). All sera from channel catfish (Ictalurus punctatus), lake sturgeon (Acipenser fulvescens), quillback (Carpiodes cyprinus), rock bass (Ambloplites rupestris), shorthead redhorse (Moxostoma macrolepidotum), silver redhorse (M. anisurum), walleye (Sander vitreus), white perch (Morone americana), and yellow perch (Perca flavescens) were negative. Antibodies in one or more fish species were detected in all sampling years (2004, 2006, 2007, 2009, and 2010), whereas in parallel sampling periods, VHS virus was detected only in 2006 and 2009. Our results suggest the continued presence of VHSV-IVb in the Lake St. Clair ecosystem, and underscore the importance of assessing immune responses of fish populations to determine prior virus exposure.     

A Probabilistic Risk Assessment of the Effects of Methylmercury on Great Egrets and Bald Eagles Foraging at a Constructed Wetland in South Florida Relative to the Everglades

This case study summarizes an assessment of risk of methylmercury (MeHg) exposure to fish–eating birds foraging at Stormwater Treatment Area–2 (STA–2). This assessment was required as a special condition for a permit modification authorizing flow–through operation of STA–2 Cell 1 without it first satisfying formal mercury start–up criteria. The assessment estimates the risks posed by MeHg to the great egret (Ardea albus) and the bald eagle (Haliaeetus leucocephalus). Exposure models were based on literature–derived life history parameters combined with site–specific MeHg concentrations in water, sediment, and fish. To assess risk, daily MeHg intake by females and cumulative MeHg consumed by nestlings were compared to literature–derived effects thresholds. The results indicated the likelihood was low that MeHg exposures to birds foraging throughout STA–2 would exceed effects thresholds at the time of this assessment. Birds foraging exclusively from Cell 1 or the small discharge canal were predicted to experience greater exposures and could be at potential risk. However, this worst–case risk was comparable or lower than risk levels encountered in nearby water conservation areas or the Everglades National Park.     

Changes in the fish community and water quality during seven years of stocking piscivorous fish in a shallow lake

SUMMARY 1. Piscivores (annual stocking of 1000 individuals ha^?1 of 0+ pike and a single stocking of 30 kg ha^?1 of large 20–30 cm perch) were stocked in seven consecutive years in a shallow eutrophic lake in Denmark. The stocking programme aimed at changing food‐web structure by reducing zooplanktivorous and benthivorous fish, with resultant effects on lower trophic levels and ultimately water quality. 2. The fish community and water quality parameters (Secchi depth, concentrations of total phosphorus, chlorophyll a and suspended solids) were monitored between 1996 and 2000 and relationships were evaluated between predatory fish and potential prey and between zooplanktivorous or benthivorous fish and water quality parameters. In addition, potential consumption of piscivorous fishes was calculated. 3. The density of fish feeding on larger zooplankton or benthos (roach >15 cm, crucian carp >15 cm) declined distinctly during the study period. This effect was attributed to predation by large (>50 cm) pike. Based on scale readings, we cautiously suggest that the stocking of 0+ pike boosted the adult pike population to produce an unexpected impact in later years. Conversely, no direct impact of stocked 0+ pike was detected on 0+ roach. 4. A major decline in the recruitment strength of 0+ roach was observed in 2000. A combination of (i) the indirect effect of large pike preying on adult roach, with negative effect on roach reproduction and (ii) the direct predation effect of 0+ pike and/or 1+ and 2+ perch recruited in the lake, provides the most likely explanation of this phenomenon. 5. A marked increase in Secchi depth in 2000 and declining trends in suspended solids, chlorophyll‐a and total phosphorus concentrations were observed. These changes may also be attributable to changes in the fish community, although the relationships were not straightforward. 6. This 7‐year study indicates that piscivorous fish may be a significant structuring force in shallow eutrophic lakes, suggesting that stocking piscivores can increase predation pressure on cyprinids. However, the general lack of impact of 0+ pike points to the need of refining current stocking practices in several countries across Europe.     

Effects of brown and turbid water on piscivore–prey fish interactions along a visibility gradient

1. Environmental changes such as eutrophication and increasing inputs of humic matter (brownification) may have strong effects on predator–prey interactions in lakes through a reduction in the visual conditions affecting foraging behaviour of visually oriented predators. 2. In this experiment, we studied the effects of visual range (25–200 cm) in combination with optically deteriorating treatments (algae, clay or brown humic water) on predator–prey interactions between pike (Esox lucius) and roach (Rutilus rutilus). We measured effects on reaction distance and strike distance for pike and escape distance for roach, when pike individuals were exposed to free‐swimming roach as well as to roach held in a glass cylinder. 3. We found that reaction distance decreased with decreasing visual range caused by increasing levels of algae, clay or humic matter. The effect of reaction distance was stronger in turbid water (clay, algae) than in the brown water treatment. 4. Strike distance was neither affected by visual range nor by optical treatment, but we found shorter strike distances when pike attacked roach using visual cues only (roach held in a cylinder) compared to when pike could use multiple senses (free‐swimming roach). Escape distance for roach was longer in turbid than in brown water treatments. 5. Changes in environmental drivers, such as eutrophication and brownification, affecting the optical climate should thus have consequences for the strength of predator–prey interactions through changes in piscivore foraging efficiency and prey escape behaviour. This in turn may affect lake ecosystems through higher‐order interactions.     

Antipredator Behavior of American Bullfrogs (Lithobates catesbeianus) in a Novel Environment

Invasive species capable of recognizing potential predators may have increased establishment rates in novel environments. Individuals may retain historical predator recognition and invoke innate responses in the presence of taxonomically or ecologically similar predators, generalize antipredator responses, or learn to avoid risky species in novel environments. Invasive amphibians in aquatic environments often use chemical cues to assess predation risk and learn to avoid novel predators via direct experience and/or associated chemical cues. Ontogeny may also influence recognition; experience with predators may need to occur at certain developmental stages for individuals to respond correctly. We tested predator recognition in invasive American bullfrog ( Lithobates catesbeianus) tadpoles that varied in experience with fish predators at the population and individual scale. We found that bullfrog tadpoles responded to a historical predator, largemouth bass ( Micropterus salmoides), only if the population was locally sympatric with largemouth bass. Individuals from a population that did not co‐occur with largemouth bass did not increase refuge use in response to either largemouth bass chemical cues alone or chemical cues with diet cues (largemouth bass fed bullfrog tadpoles). To test whether this behavioral response was generalized across fish predators, we exposed tadpoles to rainbow trout ( Oncorhynchus mykiss) and found that tadpoles could not recognize this novel predator regardless of co‐occurrence with other fish species. These results suggest that environment may be more important for predator recognition than evolutionary history for this invasive species, and individuals do not retain predator recognition or generalize across fish predators.     

Consumption of crucian carp (Carassius carassius L., 1758) by restocked pike (Esox lucius L., 1758) in a lake with frequent winter hypoxia

To assess the effects of stocked pike (Esox lucius L.) on crucian carp (Carassius carassius L.) biomasses, the annual consumption of pike was estimated and compared with removal fishery catches. The studied lake, Lake Savijärvi, is a small (40 ha), shallow and eutrophicated lake in southern Finland with frequent algal blooms during summers and fish kills during winters. Until the 1980s, the fish fauna consisted of pike, perch (Perca fluviatilis L.), roach [Rutilus rutilus (L.)], crucian carp, and tench (Tinca tinca L.). Since 2003, when crucian carp were abundant in the extreme and with only a few roach and tench individuals, the lake has been biomanipulated by removal seining. To enhance the effects of seining, piscivorous pike were restocked in the spring of 2008. During the 3‐year study, the catch of crucian carp decreased from 243 to 136 kg while the catch of pike increased from 0.1 to 5.7 kg per seining hectare due to their spawning in the lake as of 2009. The total crucian carp consumption by captured pike was 588 kg during the 2008–2010 study period. When applying the number of pike estimated with the mark‐recapture method, consumption estimate increased to 917 kg, or 22.9 kg per lake hectare in 2010. Thus, pike consumption of crucian carp was about 17% of the removal catch in 2010.     

The consequences of a drastic fish stock reduction in the large and shallow Lake Wolderwijd,The Netherlands. Can we understand what happened?

In 1990 an experiment started in the large and shallow lake Wolderwijd (2700 ha, mean depth 1.5 m) to improve the water quality. About 75% of the fish stock was removed (425 000 kg fish). The fish was mainly composed of bream and roach. In May 600000 young pikes (3–4 cm) were introduced.In May 1991 the water became very clear (Secchi depth 1.8 m) during a spring bloom of large Daphnia. Then the grazing by zooplankton was eight times higher than the primary production of algae and the total suspended matter concentration became very low. Compared to the situation before the fish reduction, the grazing had increased only slightly, while the primary production had decreased significantly in early spring. The fish stock reduction might have contributed to the reduction in primary production by a reduced internal nutrient load. The clear water period lasted six weeks. Daphnia disappeared in July due to food limitation, the algal biomass increased and the Secchi depth became 50 cm. Daphnia did not recover during summer, due to predation that was not caused by 0 + fish but by the mysid shrimp Neomysis integer. Neomysis could develop abundantly, because of the reduced biomass of the predator perch. The production of young fish had been low because of the cold spring weather. The cold weather was probably also responsible for the slow increase in density of macrophytes. After 1991, perch probably can control Neomysis. Due to lack of spawning places and shelter for 0 + pike, pike was probably not able to control the production of 0 + fish. In a lake of this scale, it will not be easy to get more than 50% coverage of macrophytes, which seems necessary to keep the algal biomass low by nutrient competition. Therefore, we expect also in the future a decrease in transparency in the summer. Locally, especially near Characeae, the water might stay clear.     

Information Transfer across Fish Shoals under Predator Threat

The hypothesis that shoaling fish can obtain information about a predator's approach from changes in the behaviour of other shoal members was tested in an experiment in which receiver minnows (Phoxinus phoxinus) behind a one-way-mirror could observe the reaction of transmitter minnows threatened by the stalk of a pike (Esox lucius) model. Although the receiver minnows were out of visual contact with the pike model they decreased their foraging behaviour and started hiding when the pike model came towards the feeding patch of the transmitter fish. The presence of skittering behaviour and inspection behaviour in the transmitter but not in the receiver fish suggests that individual confirmation of the predator does play a role in determining the nature of the anti-predator response. Nevertheless there is clear evidence that receiver fish modify their own behaviour on the basis of information obtained from individuals which have seen a predator.     

Behaviour in a standardized assay,but not metabolic or growth rate,predicts behavioural variation in an adult aquatic top predator Esox lucius in the wild

This study tested for links among behaviour, state and life‐history variables as predicted by the pace‐of‐life hypothesis in adult pike Esox lucius. First, a standardized open‐field behavioural assay was developed to assess individual behaviour of wild‐captured adult E. lucius. Behaviour within the standardized assay predicted swimming behaviour in the lake, providing an ecological validation of the assay. There was no relationship between standardized behaviour and any of the life‐history and state variables, including metabolism, body condition, juvenile growth rate and adult growth rate in contrast to predictions from the pace‐of‐life hypothesis. This study demonstrates that it is possible to assess ecologically relevant behavioural variation in a large‐bodied top predator using a standard open‐field assay, but it is noteworthy that this standardized behaviour is not systematically related to standard metabolism or growth.