The fish community structure in Tjeukemeer in relation to fishery and habitat utilization

The fish community in Tjeukemeer was monitored from 1971 to 1988 by trawling, and during the summer of 1988 the distribution of the more abundant species was determined in relation to size and location. Bream, pikeperch and smelt were the most dominant fishes, whereas roach, white bream, perch and ruffe comprised <10% of the total catches. After the termination in 1977 of the intensive gillnet fishery both pikeperch >50 cm and bream >30 cm increased in biomass, but roach >15 cm and perch >15 cm virtually disappeared and pikeperch <50 cm substantially decreased. Only smelt, ruffe, white bream and bream <30 cm hardly changed in biomass. Bream, pikeperch, perch and smelt were restricted to the open water zone, but roach <15 cm and 0 + bream were confined to the littoral zone. White bream and ruffe did not show a distinct habitat preference. Because the recruitment of smelt is largely dependent on immigration from the IJsselmeer, a feed-back between the smelt and pikeperch population is lacking. Because the carrying capacity of the pikeperch population is mainly determined by smelt, the other fish <15 cm are very vulnerable to predation when the smelt population is consumed, before a new year-class of smelt is recruited, or when the smelt fails to recruit. The vulnerability to predation of the different species and their feeding habits are discussed in relation to their distribution.     

Diet niche relationships among predator and prey fish species in their early life stages in Lake Võrtsjärv (Estonia)

In Lake Võrtsjärv pikeperch was observed not to shift to piscivory in their first autumn of life, although juvenile stages of a variety of fish species were abundant in the lake. It was hypothesized that the diets of predator and prey fish fry overlap and that coarse fish species are important food competitors for juvenile piscivores and thus, pikeperch and perch fry do not shift to piscivory during their first growing season. To discover the possible linkages in this pattern, in 2009 the feeding relationships of pikeperch, perch, ruffe and roach fry were analysed. The stomach content analyses showed that in the summer period, Mesocyclops leuckarti was the most frequent prey for perch and ruffe, pikeperch consumed Leptodora kindti in large quantities, and roach ate mostly plant material. Towards autumn, M. leuckarti was the most abundant prey for all percids. However, average stomach content weight and the number of prey items eaten by ruffe were considerably higher than for other fish fry. Since the feeding opportunities of fish fry are considered poor in the examined lake, the prey has the potential to restrict the recruitment to piscivory of their predators, as prey fish seem to have better abilities to persist in this ecosystem. Furthermore, supposed competition in the juvenile stage may result in a reduced top‐down effect on coarse fish.     

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.     

Formation of morphological defences in response to YOY perch and invertebrate predation in two <Emphasis Type="Italic">Daphnia</Emphasis> species coexisting in a mesotrophic lake

This study examined the formation of morphological defences by two coexisting Daphnia species, the large-sized D. pulicaria (2 mm) and the small-sized D. mendotae (1.4 mm), in response to the presence of young-of-the-year (YOY) yellow perch (Perca flavescens) and invertebrate predators (Chaoborus, Leptodora) during summer in a mesotrophic lake. We hypothesized that due to differential size-selective predation risk by YOY fish and invertebrates, the large-sized and the small-sized Daphnia species would show different morphological responses to predation threats. We followed changes in two morphological traits (relative length of the tail spine in D. pulicaria and of the helmet in D. mendotae) among different periods during summer according to YOY fish and invertebrate predation. We defined four YOY fish predation periods based on the presence of YOY perch in the pelagic zone of the lake and the relative abundance of Daphnia preys in their gut contents, and two invertebrate predation periods based on exclusive or mutual occurrence of the invertebrate predators. The large-sized (D. pulicaria) and the small-sized (D. mendotae) species showed different morphological responses to YOY fish and invertebrate predators, respectively. The tail spine ratio of the juveniles and adults of D. pulicaria did not change in response to YOY fish predation or to invertebrate predation. A gradual increase in the helmet ratio was observed in the small-sized D. mendotae over the summer period. This change was related to the co-occurrence of the invertebrate predators (Chaoborus and Leptodora) and to YOY fish predation. The warmer temperature cannot be accounted for helmet elongation since it was constant across depths, and not related with the co-occurrence of D. mendotae and YOY perch. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

Selective predation on zooplankton by pelagic Arctic charr, Salvelinus alpinus, in six subarctic lakes

Selective predation by planktivore fish appears to be an important regulatory factor of zooplankton communities, potentially causing large changes in species composition and size distributions within populations. In this study, prey preferences and size-selective predation on zooplankton by Arctic charr were examined in six subarctic lakes with Arctic charr as the dominant pelagic fish species. Most of the lakes had a zooplankton community dominated by copepods (Cyclops scutifer and Eudiaptomus graciloides), but the pelagic charr evidently selected cladoceran species (Bythotrephes longimanus, Daphnia sp. and Bosmina sp.), likely because the copepods have a higher mobility and evasiveness than the cladocerans. Furthermore, a strong size selection was also revealed for both Bosmina sp. and Daphnia sp., as individual prey from Arctic charr stomachs were exclusively larger than individuals sampled in the environment. Additionally, visibility due to size, morphology and pigmentation (egg-carrying females) was also a major factor for the selection of zooplankton prey. In conclusion, Arctic charr was found to be highly selective on zooplankton both in respect to species composition and individual size of Bosmina sp. and Daphnia sp.     

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.     

P-load,phytoplankton, zooplankton and fish stock in Loosdrecht Lake and Tjeukemeer: confounding effects of predation and food availability

The fish community in the Loosdrecht lakes is dominated by bream, pikeperch and smelt and is characteristic of shallow eutrophic lakes in The Netherlands. The biomasses of the respective fish species amount to ca. 250, 25 and 10 kg ha^–1 and correspond to those in Tjeukemeer, another lake in The Netherlands. The average size of bream, however, is much smaller in the Loosdrecht lakes as a consequence of poorer feeding conditions. The zooplankton community in the Loosdrecht lakes is predominantly composed of relatively small species such as Daphnia cucullata, Bosmina coregoni and cyclopoid copepods, whereas in Tjeukemeer, Daphnia hyalina is permanently present in relatively high densities and the other species show a larger mean length. In the Loosdrecht lakes, the absence of D. hyalina and the smaller sizes of the other zooplankton species could be the consequence of a higher predation pressure, in combination with unfavourable feeding conditions for the zooplankton including the low density of green algae and the high density of filamentous cyanobacteria. A biomanipulation experiment in Lake Breukeleveen, one of the Loosdrecht lakes, indicated that feeding conditions were too unfavourable for large zooplankton to develop in spring, when the reduced fish biomass was not yet supplemented by natural recruitment and immigration.     

Responses of cyanobacteria to herbivorous zooplankton across predator regimes: who mows the bloom?

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The ecology of ruffe,Gymnocephalus cernuus (Pisces: Percidae) introduced to Mildevatn,western Norway

Synopsis Diet, habitat use, diel and seasonal activity and a number of population parameters were studied on ruffe,Gymnocephalus cernuus, introduced to Mildevatn, western Norway. This lake is sited outside the natural range of the ruffe and has a lower fish diversity and a different fish species composition than within its native range. From June through September the ruffe was planktivorous and mainly caught at 4 to 6 m depth in the benthic zone. At other times of year ruffe was feeding on zoobenthos and caught deeper in the benthic zone. Ruffe was mainly day active. Zooplankton feeding during summer is the clearest difference compared to ruffe populations living within its natural range. Presence of large zooplankton organisms available for ruffe is suggested as the main reason for the difference found in food choice. The availability of large zooplankton is probably due to community structure caused by a predator and lack of interspecific competition for zooplankton in the deeper parts of the lake. Piscivorous brown trout.Salmo trutta, restrict the habitat of threespined stickleback,Gasterosteus aculeatus, to the zone of littoral vegetation, allowing high densities of larger zooplankton species likeBythotrephes longimanus to be present in the lake. Brown trout is present only in the upper light and well oxygenated parts of the lake, leaving a refuge for the ruffe, where they can feed on the rich zooplankton community.     

Food and habitat partitioning among juveniles of three fish species in the pelagial of a mesotrophic lake

An analysis of the diet of 0⁺ perch, smelt and roach caught at night of the pelagial of a mesotrophic lake showed that their food was composed mainly of herbivorous and predatory cladocerans, copepods and Chaoborus larvae during summer, and of herbivorous cladocerans and copepods during October. An analysis of habitat use by juveniles revealed separation among the species: roach occurred in the upper, perch and smelt in the deeper water layers. Food and habitat were alternatively partitioned among the juveniles. In early summer the lowest food overlap between perch and smelt coincided with high habitat overlap. In October high food overlap between perch and smelt corresponded with low habitat overlap. Smelt and roach fry used common food resources throughout the season, but they were segregated in habitat.     

Environmental factors as forces structuring the fish community of the Elbe Estuary

Smelt, flounder, eel, three-spined stickleback, twaite shad, ruffe and herring were the major species among 62 forming the fish community of the Elbe estuary. Species richness, species diversity, evenness and total fish biomass decreased in the upstream direction. Total abundance was greatest during summer. Salinity was the most significant physico-chemical factor affecting species richness and total fish biomass, whereas water temperature was the best predictor of total abundance. Marine fish species decreased with decreasing salinity, while the freshwater species roach and ide were absent at salinities > 15‰. Bream, blue bream, white bream, perch and pikeperch occurred more frequently in shallow marginal regions with lower current velocities, while herring and other marine species occurred closer to the deep central regions with high current velocities. Annual and seasonal variations of the community structure mainly reflected in population dynamics of smelt, flounder, twaite shad, three-spined stickleback and eel, especially those of 0-age smelt. Populations of smelt and ruffe and their importance in the fish community increased between 1989 and 1992, while those of twaite shad and eel decreased. At oxygen concentrations <3 mg I¹, smelt and flounder were rare, but eels were at their maximum at 3.5 mg I⁻¹. Changes of species composition and fish abundance were closely related to daytime and tidal cycle.     

0+ perch predation on 0+ bream: a case study in a eutrophic gravel pit lake

SUMMARY 1. We studied the population dynamics of 0+ fish in a eutrophic gravel pit lake in which predation of 0+ Eurasian perch on other 0+ fish was suspected to influence perch growth and the structure of the fish community, with effects on the lake food web. 2. The adult fish community was dominated by piscivorous species, especially perch, and the 0+ fish community was dominated by perch and bream. Bream grew to a total length of 80 mm between May and the end of August, gradually decreased in numbers during the summer, and completely disappeared in autumn. Stomach analysis revealed that 0+ perch ≥28 mm fed on 0+ bream. 3. The initially unimodal cohort of 0+ perch gradually broadened and became bimodal by the end of July. Fish of the larger, piscivorous cohort grew faster (1.4 mm day^?1) than the smaller, zooplankton‐consuming fish (0.6 mm day^?1). Although individuals of both cohorts later consumed zooplankton and grew at similar rates (0.5 mm day^?1), only perch of the large cohort (mean TL 125 mm) were found by mid‐October. Intraspecific competition for food or cannibalism of older perch may have contributed to the disappearance of the smaller perch. 4. Early piscivory of the fast‐growing 0+ perch apparently resulted in the disappearance of 0+ bream by the end of the growing season and precocious maturation of male perch. In contrast to findings in other studies, these large 0+ perch thus avoided the juvenile bottleneck by switching to piscivory early during their ontogenetic development. 5. The observations of this study suggest that early piscivory of 0+ perch can have a long‐lasting impact on fish communities in eutrophic lakes, particularly if prey fish are abundant and the structural complexity of the lake is low. Furthermore, early piscivory of 0+ perch may help prevent the expected increase in 0+ cyprinids following reduction of adult cyprinids, which is considered important to ensure the long‐term success of biomanipulation experiments.     

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.     

The relations between 0+ fish density,zooplankton size and the vulnerability of pikeperch,Stizostedion lucioperca,to angling in the Frisian lakes

During a five year study, the size and abundance of zooplankton and 0+ fish in September were registered in a number of Frisian lakes. The mean size of Daphnia was related to the abundance of planktivorous 0+ fish in September. Immigration of fish larvae into the Frisian Lake District from the IJsselmeer in June was observed in 1979 and 1980. The abundance of 0+ fish was negatively related and the size of Daphnia was positively related to the vulnerability to angling of predatory pikeperch.     

Cladoceran Densities,Day-to-Day Variability in Food Selection by Smelt,and the Birth-Rate-Compensation Hypothesis

Late-evening gut inspection of a dominant planktivore (smelt) and evaluation of densities, fecundities, and body-size distributions in dominant zooplankton prey (cladocerans) were made in day-to-day sequences in June–July (24 days in 1999 and 24 days in 2000). This was conducted as a field test of the hypothesis that species-specific population densities in cladocerans result from size-selective predation by a dominant fish assumed to be a general predator, switching from one prey to another as relative abundance changes. Little of the expected coincidence has been revealed between population density declines and increased numbers of a given prey in smelt diet. However, the data were consistent with the notion that fish would switch from one prey to another depending on the prey relative abundance (the number of prey a fish would see in its reaction field volume). Each cladoceran population fluctuated around its species-specific density level, lower or higher, depending on individual susceptibility to smelt predation, from 0.2 ind. l^–1 in large-bodied Daphnia hyalina and Leptodora kindtii, to 30.0 ind. l^–1 in small-bodied Daphnia cucullata andBosmina thersites. In spite of high fish-to-fish and day-to-day variability in both smelt diet and smelt selectivity for different prey, all cladocerans (also copepods and midge larvae) were equally persistent in smelt diet, and smelt selectivity was similar for small- and large-bodied prey categories, but lower for elongated-(Daphnia, Diaphanosoma) than for compact-body (Bosmina, Chydorus) species, when integrated for the entire sampling time. Closer examination of D. cucullata and B. thersites revealed strong smelt selection for later instars and females with greater clutches, showing that size distribution in a cladoceran population might be structured by fish predation in a similar way to that a cladoceran community (species relative abundance) is structured in a lake habitat. The birth-rate-compensation hypothesis is offered to explain why the value of food selectivity index in a planktivorous fish would remain the same for alternate prey categories with similar life-history traits, unless they differ in susceptibility to predation before the time of first reproduction.     

Spatial and seasonal variation of the zooplankton in the coastal zone and main khors of Lake Nasser (Egypt)

A prominent feature of Lake Nasser is the presence of khors (dendritic side extensions). We studied the zooplankton of the larger khors and coastal zone of the main lake in 1996 and 1997, and found an assemblage of rotifers, cladocerans and copepods that was partly tropical, partly temperate, at relatively high biomasses. Spatial differences were weak, but the upstream khors (Toushka and Korosko) were consistently richer than the downstream khors (El-Ramla and Kalabsha), with a rather sudden transition around km 150 at El Madiq. Summer standing crops were higher than those in winter by a factor 2–3. The zooplankton of the littoral of the main channel showed the same spatial pattern as that in the khors, being more abundant in spring (average 82300 ind m^–3) than in autumn (average 72700 ind m^–3). Zooplankton dry weight increased from 4 g m^–2 at khor El-Ramla to 7 g m^–2 at khor Toushka. These rather high values had low variation. The number of species, diversity and evenness all showed a high degree of similarity among the khors and in the littoral of the main lake. The lake fish fauna is poor, lacking a pelagic planktivore. The predominance of medium-sized Copepoda (one calanoid, two cyclopoids) in the zooplankton suggests that fish predation is moderate. This is confirmed by the persistence of two Daphnia species at low abundance, although rather strong variations in time suggest that Cladocera suffer from summer predation by invertebrates (copepods) as well as vertebrates (mostly larval fish). Because the zooplankton is underutilised by higher trophic levels, we suggest to assess the benefits of introducing a pelagic zooplanktivorous fish.     

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.     

The effects of selective fish predation on the horizontal distribution of pelagic Cladocera in a southern French reservoir

A four-year study showed a clear seasonal succession of species within the cladoceran community of the large, oligotrophic Sainte-Croix reservoir (S.E. France). Diaphanosoma brachyurum and Ceriodaphnia pulchella were strictly limited to the warm stratified period (July to October), whereas Bosmina longirostris and Bosmina coregoni were dominant during spring and autumn. Daphnia longispina was the only species to occur throughout the year with higher densities in spring.In spring and late autumn, the discharge of the inflowing river Verdon was high and the abundance of all species showed a gradient over the whole lake with lower densities close to the inflow. During the stratified period, water inflow was very low and species showed different patterns. Densities of the small form Ceriodaphnia pulchella were similar all along the long axis of the lake, whereas Daphnia densities were significantly higher near the dam. The distribution pattern of Diaphanosoma, an intermediate-sized species, showed similar trends to that of Daphnia. The only planktivorous fish in the pelagic zone, the bleak (Alburnus alburnus), fed mostly on large-bodied species (> 1.0 mm) and was more abundant close to the inflow current. A comparison between the length frequency distributions of cladocera upstream and downstream provided a clear demonstration of the effects of size-selective predation on prey populations. Finally, the interactions between spatial heterogeneity and long-term development of the zooplankton community and the indirect effects of predation are discussed.     

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.     

Fish impingement at power stations situated along the rivers Rhine and Meuse in The Netherlands

Summary Substantial numbers of predominantly O⁺ fish were impinged on the cooling water screens of Harculo, Gelderland, Merwedehaven and Amer power stations. Low numbers were impinged at the two stations located on the river Meuse (Maas and Claus). Two possible reasons for this difference are suggested,viz. a low fish density in the river Meuse and the deep position of the cooling water intakes. In comparison with Bergum power station situated on a lake the impingement rate at the river stations was more than ten times as low.Most fish were impinged during the summer months when large numbers of juvenile fish are present in the rivers. Dominating species impinged were roach, pikeperch, bream and perch. The abundance of pikeperch in the screensamples, points to a recent increase in the population of pikeperch in the rivers. The growth pattern of O⁺ pikeperch and O⁺ roach during summer seemed very similar for power stations located at different rivers. Thirty fish species and four crustacean species were collected, among whichOrconectus limosus was quite abundant at Maas and Claus power stations.