Piscivory and the development of bimodality in the size distribution of 0+ pikeperch (Stizostedion lucioperca L.)

During an eight years period the onset and development of piscivory of 0+ pikeperch was studied in the natural environment of a shallow, eutrophic Dutch lake. In some years the piscivory started as early as halfway June at a pikeperch size of circa 20 mm. The 0+ pikeperch predated mainly on the smaller 0+ smelt because these were the most slender preyfish available. The 0+ pikeperch is less and the 0+ perch is much less available because of their larger circumference. Small 0+ bream and roach became only available when their growth was retarded. The positively skewed size distribution of the 0+ pikeperch halfway June is precursory for the development of bimodality in the size distribution later in the season. A second modus of larger, piscivorous, 0+ pikeperch became relatively important in years when smelt was abundant. The survival rate of this category was appearantly higher than that of the smaller planktivorous individuals.     

Prey or predator: 0+ perch (Perca fluviatilis) in the trade-off between food and shelter

The 0+ cohort of perch can split into a slow-growing planktivorous and a fast-growing piscivorous cohort during their first months of life. Both cohorts are, however, vulnerable to predation by piscivorous fish. Laboratory experiments were performed to test the behavior of 0+ perch as a predator of cyprinids, and in the trade-off between food and shelter from the threat of predators. In the foraging trials, 0+ perch attacked bream faster than they did carp, and vegetation hampered the aggression against bream. In the second experiment, the habitat selection of two size classes of 0+ perch under the threat of predation was monitored. Overall, vegetation structures were preferred by both size classes of 0+ perch. When small fish were offered to the 0+ perch as food, the open water becomes more attractive. The results of the habitat use trials further show that the two size cohorts of 0+ perch may also differ in their behavior, in that the availability of fish as food becomes more important than the shelter of vegetation structures for the larger perch.     

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.

     

Two size classes of 0+ year perch: is phenotypic plasticity based on food resources?

Perch Perca fluviatilis of age 0+ years were caught in a gravel pit lake in June (unimodal size distribution) and in July (bimodal size distribution) to analyse morphological differences between the two growth cohorts. Independent of size, 0+ year perch developed a deeper body and perch of the large size cohort had an even deeper body than perch of the small size cohort. This might have been adaptations to either piscivory or planktivory, but might also be a hint that 0+ year perch of the small size cohort were undernourished and that they developed on a different trajectory than individuals of the large size cohort. In a second step the study was extended with a mesocosm experiment. This part of the study was designed to provide preliminary evidence for the extent to which morphological variations may be due to the consumption of different food resources when other factors such as habitat use could be neglected. Two groups of 0+ year perch in four mesocosms were fed for 40 days with the same biomass of either plankton or cyprinids. Although the experimental groups at the end of the experiment did not differ in size, they differed in morphology. The mouth of the piscivorous 0+ year perch became larger, the pectoral fins and the centre of mass of the posterior abdomen were shifted backwards. These results provide further evidence that the type of food is important and might lead to further functional adaptations in morphology.     

Feeding behaviour of major 0 + fish species in a shallow, eutrophic lake (Tjeukemeer, The Netherlands)

The forage base and the food selectivity of 0+ representatives of six abundant freshwater fish species were studied in a shallow, eutrophic Dutch lake. Most species relied on the zooplankton; the size-selective predation in early summer was directed to the smaller copepods and in late summer to larger cladocerans and copepods than concurrently present in the lake. Daphnia spp. and cyclopoid copepods were the main zooplankton taxa for smelt, perch and pikeperch. Energetically, the large cladoceran, Leptodora kindtii, was especially important for pikeperch. Bream and roach preyed upon smaller zooplankton than the other fish species. The influence of the zooplankton predation by abundant 0+ fish was clear from a small mean Daphnia size in September; this size is to be used as an indicator in fishery management. Neomysis integer, the most important macrofauna species, was consumed by perch, pikeperch and ruffe; pikeperch was most size-selective in this respect. The 0+ ruffe was à typically benthivorous fish. Only the 0+ pikeperch became piscivorous, especially in years when smelt was abundant.     

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.     

Distribution and Diet of 0+ Fish within a Canyon‐Shaped European Reservoir in Late Summer

The distribution and diet of age 0+ fish were studied in the deep canyon‐shaped Římov Reservoir (Czech Republic), which is characterized by a longitudinal trophic gradient. During late summer of two years, 0+ fish were sampled from inshore and offshore habitats along the longitudinal reservoir axis. Offshore catches of 0+ fish from the surface layer were dominated by roach (Rutilus rutilus ), bream (Abramis brama ) and perch (Perca fluviatilis ), whereas in the deeper open water perch predominated. Inshore catches of 0+ fish were constituted mainly by perch and roach. The proportions of roach in the inshore catches were highest at the upper and most eutrophic part of the reservoir, whereas the proportions of perch in the inshore catches were higher at the downstream areas. Total catches of both inshore and offshore 0+ fish increased upstream in the reservoir. Offshore 0+ perch were of consistently smaller size than inshore 0+ perch. Inshore 0+ perch had significantly smaller size at the upstream reservoir part than at the downstream, more lacustrine regions. The diet of both inshore and offshore 0+ fish consisted predominantly of crustacean zooplankton. Perch diet was generally dominated both by cladocerans and copepods, whereas roach diet consisted chiefly of cladocerans. (© 2006 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

The fish community of the Loosdrecht Lakes and the perspective for biomanipulation

In a two-year-study, the fish community of the Loosdrecht Lakes area was characterized with regard to the professional fishery and the forage base. The lakes are shallow (mean depth 2 m) and eutrophic to hypertrophic. The bream,Abramis brama, dominates the fish biomass. The pikeperch,Stizostedion lucioperca, is the main predator.Bream up to 30 cm have a slow growth rate and are in a bad condition. A faster growth rate and a better condition are shown by bream of 30 cm and more. The small bream feeds on chironomid larvae, benthic cladocerans and zooplankton. The better condition of bream over 30 cm is explained by the more efficient feeding of larger bream onChironomus plumosus larvae. Pikeperch show a fast growth rate and a good condition. Recruitment is limited by the low densities of smelt,Osmerus eperlanus, leaving cannibalism as the most important way for the 0+ pikeperch to become piscivorous and to manifest a fast growth.The impact of the professional gillnet fishery on bream and pikeperch is small because the mesh sizes in use are as large as 75–100 mm bar mesh.The planktivorous 0+ pikeperch consumes mainly the carnivorous zooplanktersLeptodora kindtii and cyclopoids. The zooplankton community lacks large herbivorous species likeDaphnia hyalina, capable of consuming bluegreens. A possible experiment in biomanipulation with a view to find out whether the development ofD.hyalina is depressed by the small planktivorous cyprinids, is predator enhancement with the aid of a stocking programme for indoor-raised 0+ pikeperch in early summer.     

Variation in abundance and survival of fish larvae in shallow eutrophic lake Tjeukemeer

Synopsis Variations in abundance and survival of larvae of the major fish species, bream, roach, pikeperch, perch and smelt were studied in shallow eutrophic lake Tjeukemeer (1986–1988). Except for smelt, the size of the spawning stock and the abundance of larvae were not related. Both the timing of appearance of larvae and their growth rate were determined by water temperature as was shown by a growth model developed earlier for Tjeukemeer fishes. In cyprinids and percids the survival of the larvae was hypothesized to be negatively related to the duration of the earliest stages. Except in cyprinids, the abundance of 0+ juvenile fish could be explained by the number of larvae. In cyprinids 0+ juvenile abundance at the end of the year was not related to the number of larvae in May and June. The cyprinid juvenile survival rates are hypothesized to be related to the size of the main cyprinid predator, 0+ pikeperch.     

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.     

Food web manipulation in a small,eutrophic Lake Wirbel,Poland: effect of stocking with juvenile pike on planktivorous fish

In a four year experiment (1988–1991) carried out in a shallow, eutrophic lake, an increase in young-of-the-year pike (Esox lucius) density was used to control planktivorous fish assemblages consisting mainly of small-sized cyprinids: roach (Rutilus rutilus), Leucaspius delineatus, and white bream (Blicca bjoerkna). Stocking with pike fry of ca 30 mm in three successive springs resulted in large-scale mortality among prey of vulnerable sizes. Rotenone treatment at the end of the fourth year showed that roach and white bream populations contained few 0+ to 2+ fish, but very high numbers of 3+ to 6+ fish. By this time, short-lived, slow-growing L. delineatus, with no size refuge from predators, had been driven almost to extinction.     

Poor nutritional quality of primary producers and zooplankton driven by eutrophication is mitigated at upper trophic levels

Eutrophication and rising water temperature in freshwaters may increase the total production of a lake while simultaneously reducing the nutritional quality of food web components. We evaluated how cyanobacteria blooms, driven by agricultural eutrophication (in eutrophic Lake Köyliöjärvi) or global warming (in mesotrophic Lake Pyhäjärvi), influence the biomass and structure of phytoplankton, zooplankton, and fish communities. In terms of the nutritional value of food web components, we evaluated changes in the ω‐3 and ω‐6 polyunsaturated fatty acids (PUFA) of phytoplankton and consumers at different trophic levels. Meanwhile, the lakes did not differ in their biomasses of phytoplankton, zooplankton, and fish communities, lake trophic status greatly influenced the community structures. The eutrophic lake, with agricultural eutrophication, had cyanobacteria bloom throughout the summer months whereas cyanobacteria were abundant only occasionally in the mesotrophic lake, mainly in early summer. Phytoplankton community differences at genus level resulted in higher arachidonic acid, eicosapentaenoic acid (EPA), and docosahexaenoic acid (DHA) content of seston in the mesotrophic than in the eutrophic lake. This was also reflected in the EPA and DHA content of herbivorous zooplankton (Daphnia and Bosmina) despite more efficient trophic retention of these biomolecules in a eutrophic lake than in the mesotrophic lake zooplankton. Planktivorous juvenile fish (perch and roach) in a eutrophic lake overcame the lower availability of DHA in their prey by more efficient trophic retention and biosynthesis from the precursors. However, the most efficient trophic retention of DHA was found with benthivorous perch which prey contained only a low amount of DHA. Long‐term cyanobacterial blooming decreased the nutritional quality of piscivorous perch; however, the difference was much less than previously anticipated. Our result shows that long‐term cyanobacteria blooming impacts the structure of plankton and fish communities and lowers the nutritional quality of seston and zooplankton, which, however, is mitigated at upper trophic levels.     

Management of a pikeperch stock in a long-term biomanipulated stratified lake: efficient predation vs. low recruitment

Pikeperch is an efficient predator in the pelagic zone of eutrophic and hypertrophic lakes and has, therefore, been used in several biomanipulation projects. Pikeperch stocking has also been part of the manipulation strategy in Feldberger Haussee, Germany. From 1996 to 1999, population development, growth and diet composition of pikeperch in Feldberger Haussee were studied. From 1996 to 1998, the population was dominated by two cohorts. Only in years with warm summers, recruitment from 0+ fish to older age-classes was successful. It appears likely that due to the relatively high depth of the lake the spawning period is delayed as a result of a slower temperature increase in spring. Therefore, 0+ pikeperch cannot benefit from the spring maximum of the zooplankton. This may be the reason for the slow growth and consequently, hampers the switch to piscivory for the majority of 0+ pikeperch. However, after becoming piscivorous, the pikeperch showed a reasonably good growth. Predation was mainly directed towards roach and 0+ perch, the major planktivores. The results of the present study confirm the role of pikeperch as an efficient predator of zooplanktivorous fish in open-water dominated lakes. Yet, it is suggested that the probability of successful recruitment of pikeperch in deeper, stratified lakes is reduced. Therefore, stable self-sustaining pikeperch populations cannot be expected in such lakes and regular stocking measures may be necessary even if a potential spawning stock exists.     

The food, growth and abundance of five co-existing cyprinids in lake basins of different morphometry and water quality

The food, growth and abundance of five co-existing cyprinid fish species in the eutrophic Lake Hiidenvesi were studied. The diet overlaps within the cyprinid community in shallow lake basins were compared with those in a deep basin, where littoral resources are less available. Roach, bleak and white bream inhabited both the shallow and the deep parts of the lake. Their growth rate was slow, probably due to the low availability of animal food, indicated by the increasing proportion of detritus and plant material in the diets towards the end of the summer. In the deep basin, roach and bleak, contrary to white bream, did not forage on the very abundant invertebrate Chaoborus flavicans, explained by the migration behaviour of C. flavicans. Blue bream, unlike other cyprinids, utilized copepods and had a relatively fast growth rate, but was mostly restricted to the shallow areas. The condition of the bream stock was weak both in terms of growth and abundance. The availability of zoobenthos was low and bream was not able to compete for zooplankton with the more specialized planktivores.     

Competition,predation, cannibalism: the development of young‐of‐the‐year perch populations in ponds with bream or roach

Piscivory of perch can occur within a few weeks after perch hatch, leading to the development of two size‐cohorts, with the larger perch becoming cannibals. However, the possibility of early piscivory is assumed to depend on the hatch timing of the prey and predator. Two species, bream (2006) and roach (2007), were tested as the prey fish. The bream (hatching 12 days after perch) were preyed upon by the perch, leading to the predicted development of two sizes of cohorts as well as to cannibalism. With roach (hatching simultaneously with perch), however, no piscivory or cannibalism occurred and the perch population was unimodally distributed. The results of this experimental pond study underpin recent theoretical findings that size differences between predator and prey, determined through differences in the timing of hatching as well as differences in juvenile growth rates, foster the occurrence of early piscivory in YOY perch that may lead to bimodality and finally to intra‐cohort cannibalism.     

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.     

The distribution and growth of coarse fish in gravel-pit lakes in south-east England

SUMMARY. Age and growth rates of the coarse fish of thirty-nine gravel-pit lakes in south-east England are compared. Species composition was very variable, the commonest species being roach ( Rutilus rutilus ), bream ( Abramis brama ), tench ( Tinca tinca ), perch ( Perca fluviatilis ) and pike ( Esox lucius ). Two main kinds of fish community were identifiable from the survey: (1) those consisting of a mixture of cyprinids, usually roach and bream; and (2) those in which cyprinids were rare or absent and the fish community was composed of perch and pike. The majority of the lakes fell into the first category. Comparison of growth curves derived by back-calculation showed that, in all species, variation occurred from lake to lake and from year to year. The majority of gravel-pit lake fishes had 'medium' growth rates when compared with published data by other workers on different waters. Seasonal growth of roach infected with plerocercoids of Ligula intestinalis was observed at monthly intervals. Annulus formation on the scales of these fish occurred at the onset of growth in May. The heavy Ligula burden in 89% of these fish had not caused the formation of any additional or false rings.     

Distribution patterns of fishes in a canyon-shaped reservoir

In August 2004 and 2005, an extensive study of the fish community was carried out in the largest water supply reservoir in the Czech Republic and Central Europe, the canyon‐shaped ?elivka Reservoir, using a fleet of Nordic multimesh gillnets. Fishes were sampled at eight locations along the longitudinal profile of the reservoir and at five benthic depth layers covering depths from the surface down to 18 m (benthic gillnet 1·5 m high), and at three pelagic depth layers down to the depth of 5 m above the bottom (pelagic gillnets 4·5 m high). Catches of both juvenile (age 0+ year) and adult (fishes >1 year) fishes were highest in the upper layers of the water column (i.e. in the epilimnion down to 5 m, and down to 10 m in the benthic habitats). Along the tributary–dam axis in the pelagic habitats, both juvenile and adult fishes preferred the upper part of the reservoir, where the maximum number of species and also the greatest abundance of zooplankton were found. In the benthic habitats, fishes selected location according to factors other than trophic status. More juvenile fishes were recorded in the benthic habitats than in the pelagic habitats. Depth had the largest explanatory power for predicting fish community composition, followed by the affiliation with benthic and pelagic habitats, and location on the longitudinal axis of the reservoir. The fish community was represented mainly by cyprinids and consisted of two distinct groups of species, with bleak Alburnus alburnus, rudd Scardinius erythrophthalmus and asp Aspius aspius dominating the offshore group while perch Perca fluviatilis and ruffe Gymnocephalus cernuus were affiliated with the inshore group of the adult fish community. Roach Rutilus rutilus, bream Abramis brama and pikeperch Sander lucioperca occurred in important proportions in both the inshore and the offshore zones. All species, with the exceptions of adult perch (1+ year and older), 0+ year perch and 0+ year roach, preferred the most eutrophic tributary part of the reservoir. The fish community was relatively stable between the 2 years sampled.     

Seasonal variation in the interactions between piscivorous fish,planktivorous fish and zooplankton in a shallow eutrophic lake

The interactions between the higher trophic levels in a shallow eutrophic lake were studied during the course of a year. Three fish species determined the main pathways of organic matter flow within the system: the predominantly planktivorous bream (Abramis brama), the obligate planktivorous smelt (Osmerus eperlanus), and the piscivorous pikeperch (Stizostedion lucioperca). Of the thirteen common zooplankton taxa Daphnia hyalina and cyclopoid copepods were utilized most by the planktivorous fish, while the large production of small cladocerans is almost left unutilized.The seasonal variations of production and consumption are large. This is mainly affected by seasonal variation of the water temperature. The production of O + smelt is efficiently utilized by the pikeperch. Being the most important zooplankton consumer, as well as the most important prey group, O + fish plays a key role in the Tjeukemeer food web.