Chernobyl-derived radiocaesium in fish as dependent on water quality and lake morphometry

¹³⁷Cs concentrations in perch Perca fluviatilis , pike Esox lucius and roach Rutilus rutilus obtained from lakes of different size and water quality in an area which received about 10–67 kBq m⁻²¹³⁷Cs, were compared with environmental data. Radiocaesium concentrations were highest in pike, and were about two to three times higher in the pike and perch than in the roach. The largest perch had about four times more ¹³⁷Cs than the smallest ones, but the activities in the pike and roach were independent of fish size. All of the water quality parameters examined correlated with ¹³⁷Cs concentrations in the fish, but the concentrations of ¹³⁷Cs in the bottom sediment did not. ^l37Cs fallout and water retention time in lakes showed a positive correlation with fish ¹³⁷Cs, while the size of the catchment area, phosphorus content of the water, water colour, pH and electrical conductivity showed a negative one. Thus there was more ¹³⁷Cs in the fish from the oligotrophic lakes than from the eutrophic ones, from the less humic lakes than from the more humic ones, from the more acid than from the less acid ones and from the lakes containing less potassium than from those containing more potassium. The depth, area and volume of the lake did not affect radiocaesium in the fish. Regression equations are presented for predicting ¹³⁷Cs in fish on the basis of environmental data. Fallout, total phosphorus and pH together had the highest influence in multiple regression models but were less significant in the roach than in the perch and pike.     

Environmental correlates of population variables of perch (<Emphasis Type="Italic">Perca fluviatilis</Emphasis>) in boreal lakes

We examined relationships among perch population variable parameters in two types of lakes, lakes with perch (P-lakes, n?=?15) and lakes with perch and roach (PR-lakes, n?=?10) using redundancy analysis (RDA) to relate population variables to environmental factors. Effects from environmental factors were tested for significance by means of permutation tests (Monte Carlo). Three factors, pH, altitude and fraction of roach (by number) in the gill net catches, explaining 47.9% of the variation, had significant effects on perch population variables. The significance of pH was improved by partialing out the effect of conductivity and roach. Similarly, the significance of altitude was improved by partialing out the effect of pH and roach, and the significance of roach was improved by partialing out the effect of pH and altitude. When the fraction of pike in the catch was included in the analysis, the effect of roach was not significant and vice-versa, as roach and pike fractions were correlated with each other. The effect of pike was significant when roach was not included, but the effect was not as strong as the effect of roach. A biplot was constructed by plotting population variables on the first and second RDA axis, with arrows showing five selected environmental factors. Growth of 3+ to 5+ perch was positively related to pH and altitude, perch catch per unit effort was negatively related to pH and altitude, and age specific perch weight was negatively related to fraction of roach. The relationship between growth of 2+ perch and pH was not as strong as the relationship between pH and the growth of older perch. Moreover, the growth of 2+ perch was negatively related to the fraction of roach, probably due to competition between young zooplankton feeding perch and roach.     

Stunted growth of pikeperch Sander lucioperca in Lake Sahajärvi, Finland

Growth of pikeperch Sander lucioperca in the eutrophic and clay-turbid Lake Sahajärvi, Southern Finland, was extremely slow in comparison with other lakes at similar latitudes. The most important food item in July was phantom midge larvae Chaoborus flavicans for all sizes of S. lucioperca (239–423 mm total length L _T), while later, in August and September, the diet of S. lucioperca (149–407 mm L _T) consisted of small (30–100 mm L _T) perch Perca fluviatilis , ruffe Gymnocephalus cernuus and roach Rutilus rutilus .     

Consumption of cyanobacteria by roach (Rutilus rutilus): useful or harmful to the fish?

1. The ability of roach to use cyanobacterial food is generally believed to be one reason for the dominance of roach over perch in eutrophic European lakes. The aim of this study was to test whether cyanobacteria really are a suitable food for juvenile roach. Special attention was paid to differences between the two cyanobacteria species Aphanizomenon and Microcystis which are common in eutrophic lakes and are ingested by roach there.
2. We performed growth and behaviour experiments with juvenile roach fed with zooplankton and the different cyanobacteria. Growth rate with Aphanizomenon was lower than with Daphnia but significantly higher than without food, whereas growth rate with Microcystis was as low as without food.
3. In cultivation experiments of roach faeces, Microcystis was found not to have been digested and grew exponentially after passing through the gut whereas Aphanizomenon stayed at low biomass. Differences in growth were not related to the toxin content of cyanobacteria. Investigations of roach motility showed no differences whether fed with Aphanizomenon or Microcystis .
4. In contrast to Microcystis , Aphanizomenon can be regarded as a suitable food source for juvenile roach probably because of its better digestability. We conclude that the ability to feed on cyanobacteria is not a general competitive advantage for roach, but the outcome depends on the species composition of the cyanobacteria.     

Shift in pike, Esox lucius L., predation pressure following the introduction of ruffe, Gymnocephalus cernuus (L.) to Loch Lomond

Rapid and dramatic change in the fish community of Loch Lomond has resulted from a series of fish introductions in recent years. A comparison of the diet of pike, Esox lucius L., in 1989–1990 with data from 1955–1967, prior to recent introductions, demonstrates a shift in prey choice. In 1955–1967 powan, Coregonus lavaretus (L.), dominated in the diet (57% of prey by number) by 1989–1990 the introduced ruffe, Gymnocephalus cernuus (L.), was the commonest prey species (44% by number). This shift in pike predation to an abundant population of introduced ruffe has consequences for native species. Assuming that the pike population has not increased in response to increased food availability due to the introduction of ruffe, pike predation pressure on native species will be relieved. This is likely to have the greatest effect on powan. Comparison of the predation rate in 1955–1967 with 1989–1990 supports the hypothesis that the rate of predation on powan has declined, although the effect that this may have on the powan population is unclear, as the role of predation in the regulation of population size is unknown for this species.     

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.     

Associations between water chemistry and fish community composition: a comparison between isolated and connected lakes in northern Sweden

1. The correlation between water chemistry, physical variables and fish community composition was examined in 40 small (≤30 ha) coastal lakes in northern Sweden. Twenty of the 40 lakes were isolated from other water bodies and 20 were connected to the Baltic Sea. Lakes were fished in summer, using three different methods. Water chemistry was sampled in late winter prior to ice‐out and pH was measured additionally in summer. 2. Our central question was whether water chemistry plays a greater role in the composition of fish communities in isolated lakes than in connected lakes, as isolated lakes cannot be recolonised once a species has become extinct. 3. Results indicate that winter anoxia affects community composition only in isolated lakes, whereas acidity is of importance in both connected and isolated lakes. Methane (indicating anoxia), was significantly correlated with variation in fish community composition in isolated lakes, and a group of variables that indicate anoxia (CH₄, pCO₂, inorganic carbon and dissolved oxygen) explained 24–34% of the variation. pH alone explained 12% of the variation in community composition for connected lakes and a group of variables indicating acidity (summer and winter pH and ANC) explained 10–20% of the variation in isolated lakes. Lake area was the most important physical variable, being significantly correlated with the variation in fish community composition in connected lakes. 4. In isolated lakes, the presence of pike (Esox lucius), perch (Perca fluviatilis) and roach (Rutilus rutilus) was associated with low CH₄. The occurrence of crucian carp (Carassius carassius) and roach was positively correlated with pH, and the crucian carp was apparently also affected by predation by pike and perch. In connected lakes the effect of anoxia was low, probably due to the possibility of recolonisation and pockets of oxygenated water, allowing pike and perch to persist and thereby limiting the distribution of crucian carp.     

The biology of pike, Esox lucius L., in a southern eutrophic lake

The biology of pike, Esox lucius L., in a southern productive lowland lake is described. Scales and opercular bones were difficult to read and interpret, and growth estimated had perforce to be based on scale readings. Growth of Slapton pike is average and intermediate between recorded extremes. Both male and female pike reached maximum condition in February, just prior to spawning. The population of pike of over 450 mm fork length in the lake was estimated at 870·6 ± 389·3 in 1975, and 950·4 ± 143.3 in 1976/77. This leads to the conclusion that the pike population has one of the highest biomasses per surface area of water ever recorded. Number of pike per unit area of surface was also high, despite the fact that a significant proportion of the population (fish under 450 mm) could not be adequately sampled. Mean instantaneous mortality rate was 0·53, and mean instantaneous survival rate was 0·59. The roach provided the main item of diet of the pike, with perch taken less readily. Immature pike ate a significant proportion of invertebrates, but roach was again a common feature of the diet of even small fish. Two spawning migrations were identified; at other times of the year, pike were non territorial. The majority of pike spawned in March. Fecundity of a sample of females was assessed. Overall sex ratio was 1:1.     

Invasive fish species in the largest lakes of Scotland,Northern Ireland,Wales and England: the collective UK experience

An invasive species is defined as an alien (or introduced or non-native) species whose establishment and spread threaten ecosystems, habitats or species with harm. Such threats to UK lake fish communities have long been appreciated and this review assembles case histories, including new data, from the largest lakes of Scotland, Northern Ireland, Wales and England to examine the hypothesis that at least some of these introductions have become invasive. Loch Lomond in Scotland has experienced six introductions [chub (Leuciscus cephalus), common bream (Abramis brama), crucian carp (Carassius carassius), dace (Leuciscus leuciscus), gudgeon (Gobio gobio) and ruffe (Gymnocephalus cernuus)], of which the most significant has been that of the percid ruffe, which has been implicated in a recent decline of the native coregonid whitefish (Coregonus lavaretus). In Northern Ireland, the introduction of the cyprinid roach (Rutilus rutilus) to Lough Neagh has apparently had a negative impact on some overwintering waterfowl, although the native coregonid pollan (Coregonus autumnalis) remains abundant. Llyn Tegid in Wales has received three introductions [rudd (Scardinius erythrophthalmus), ruffe and silver bream (Blicca bjoerkna)], although no impacts on the native whitefish or other fish populations have been observed. In England, individuals of at least 12 native and non-native fish species have been brought to Windermere for the purpose of live-baiting, although only those of the cyprinids roach and common bream have established abundant populations. At the same time, the native salmonid Arctic charr (Salvelinus alpinus) has declined markedly while the native esocid pike (Esox lucius) has shown changes in abundance, distribution and individual condition, although these developments have not been shown to be causally linked. None of these introductions were sanctioned by appropriate fisheries or other regulatory bodies and almost all of them probably arose from the release or escape of live-bait used by pike anglers. Of the 10 species introductions documented here, four (common bream, gudgeon, roach and ruffe) have established abundant populations and two of these (roach and ruffe) have apparently caused or currently threaten harm, supporting the hypothesis that at least some of these introductions have become invasive.     

Tests of artificial substrata as nursery habitat for young fish

Recruitment areas for freshwater fish are often negatively affected by eutrophication and physical disturbances. Vegetated areas, which are important nursery habitats, are reduced and water turbidity increased. As a method of compensation, we tested artificial substrata for young-of-the-year fish. The structures were made of spruce bundles, with and without surrounding nets, and placed in a hyper-eutrophic very turbid environment and in an undisturbed area with clear water. Both habitats were devoid of submerged vegetation. Young fish abundance in treated areas was compared with adjacent reference sites. In the clear water area, the abundance of all investigated species – perch, pike, bream, silver-bream and roach – was higher in areas with artificial refuges. A similar response was evident for cyprinids in the turbid environment. High abundance of pikeperch and ruffe appeared in the hyper-eutrophic test area. Neither of these species, nor perch, was attracted to the artificial refuges. The lack of response in perch and pikeperch suggests that the importance of structural refuge decreases in very turbid water for these species. Of the two methods tested, spruce bundles with surrounding nets generally attracted most young fish, implying that the nets further increased the refuge capacity by reducing predation risk. The conclusion is that artificial habitats could improve recruitment habitats and that protective devices can increase refuge capacity.     

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.     

State-dependent invasion windows for prey in size-structured predator-prey systems: whole lake experiments

1. In size-structured communities where individuals grow in size over their life cycle, interactions between species will shift between competitive and predatory interactions depending on size relationships. The outcome of interactions will subsequently depend on the strength of competitive and predatory interactions, respectively. 2. In a whole lake experiment including four experimental lakes, it was tested under which conditions the competing prey, roach Rutilus rutilus, could successfully recruit into systems previously occupied by the predator, perch Perca fluviatilis. Two replicated introduction experiments were carried out 3 years apart. 3. Roach were able to successfully recruit into three of the four experimental lakes of which two were also inhabited by the top predator pike Esox lucius. Resource levels were unrelated to whether roach could successfully recruit into the systems as recruiting roach in all years were feeding close to their maximum rate. 4. High population fecundity of roach and low predation pressure by perch combined were necessary ingredients for successful recruitment and the presence of only one of these conditions did not result in successful recruitment. 5. It is hypothesized that, although roach were able to successfully recruit into one lake with only perch present in addition to the two lakes that also inhabited pike, long-term coexistence of roach and perch depends on the presence of another top predator (e.g. pike) selectively preying on perch. This hypothesis was supported by data on co-occurrence of perch and roach in different lakes. 6. Overall, the results are in accordance with expectation of size-structured life-history omnivory theory suggesting that coexistence between top predator and intermediate consumer is fragile.     

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.     

Age, growth and length-weight relationship of brown trout Salmo trutta (L.) in the Loch of Strathbeg, Aberdeenshire

Most age and growth studies on brown trout in British lakes have been restricted to highland or moorland areas. In the unusual environment of a coastal dune lake (Loch of Strath-beg, Aberdeenshire) the trout population was short-lived with few fish surviving longer than three years. Growth was rapid compared with other naturally-stocked lakes. Growth in length and weight differed between 1–and 2–migrants. Faster growth in the burn was the factor causing early migration by 1–migrants (71% of population), but both groups grew at the same rate on entering the loch. The effect of gill-net selection on the estimation of the length-weight relationship was demonstrated. A small difference was found in the length-weight relationship among the mature stages. Factors responsible for rapid growth and small age-range of the population are discussed.     

The interactions of abiotic and biotic factors influencing perch Perca fluviatilis and roach Rutilus rutilus populations in small acidified boreal lakes

Four small, acidified boreal lakes, all sustaining populations of perch Perca fluviatilis, roach Rutilus rutilus and pike Esox lucius, were studied in four successive years. Three lakes were moderately acidified (mean pH of 5·61-5·83), while the fourth was more acidic (mean pH of 5·16) and had a sparse population of R. rutilus. Perca fluviatilis density was higher in this lake (1004 ha(-1)) than in the other three (355-717 ha(-1)), where R. rutilus dominated in terms of numbers (981-2185 ha(-1)). Large, potentially predatory, P. fluviatilis were most abundant in the lake with clearest water, and these seemed to have a negative effect on P. fluviatilis density. Perca fluviatilis mean mass was negatively correlated with R. rutilus biomass and was highest in the most acidic lake with the sparse R. rutilus and the highest P. fluviatilis density. Perca fluviatilis mass correlated positively with pH in two lakes (with the highest fish biomass), suggesting that low pH affected P. fluviatilis mass negatively. Perca fluviatilis growth correlated positively with summer (July to August) air temperature in the lake with sparse R. rutilus, thus differing from P. fluviatilis and R. rutilus growth in the other three lakes. The mean age of P. fluviatilis was generally lower than that of R. rutilus and was lowest in the two lakes with the highest fish biomass, indicating that adult mortality was affected by density-induced factors.     

The growth of whitefish in relation to water quality and fish species composition

Water quality in 16 Finnish lakes did not affect directly densely gill-rakered whitefish growth, except possibly in an acid (pH 4·9) lake, Iso Lehmaälampi, where acidity may have retarded the growth of whitefish. The density of roach affected whitefish growth in the second year of life: highest growth rates were in lakes without a roach population and lowest growth rates in lakes having strong roach populations. Competition by vendace also retarded the growth of young whitefish. The efficient mass removal of roach from a eutrophic lake was considered to have increased the growth rate of young whitefish. It is suggested that an examination of the fish species composition and relative abundance, as well as the growth of whitefish, can be used as an aid in predicting the success of stocking with whitefish.     

Radioactive strontium (85Sr) in marking newly hatched pike and success of stocking

Immersion of newly hatched pike fry in an aqueous solution of 1·85 MBq ml⁻¹⁸⁵Sr did not affect their growth or survival over 3 weeks. Cannibal pike fry retained only l.2–3·3% of the ⁸⁵Sr from their labelled prey. Newly hatched ⁸⁵Sr-marked fry stocked into two lakes in 3 years were distinguishable from wild fish 4–6 weeks later. Stocked and wild fry grew equally well, and stocked pike survived best where density of wild pike was highest. There was no evidence of increase in density as a result of Stocking.     

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.     

Pike predators induce morphological changes in young perch and roach

The morphological response of two 1 year-old prey species, perch Perca fluviatilis and roach Rutilus rutilus to the presence of predatory pike Esox lucius , was tested in a 6 week aquarium experiment. The growth of perch was higher than that of roach but there was no difference in growth of prey between predator and control treatments. The presence of pike caused morphological changes in both perch and roach. Perch showed an increase in body depth during the experiment whereas the morphological response of roach to pike was more related to a displacement of the dorsal and pelvic fins and width of the anal fin. Differences in morphological responses of perch and roach to pike can be interpreted by their overall differences in antipredator strategies.     

The population density, growth rate and production of roach Rutilus mtilus (L.) in Tjeukemeer, The Netherlands

A method of estimating the population density of roach in Tjeukemeer (21.3 km²) using 20 529 introduced fin-clipped fish is described. Fyke nets proved to be an effective method of sampling the population for marked fish during the spawning season. A total of 20 277 roach were processed during the recapturing period. The population density of roach (⋝ 14 cm) was estimated to be 1 246 458. The growth rate of roach in the lake although relatively poor (von Bertalanffy's L ∞ for males and females, 22 and 26 cm respectively) for the species was similar to that recorded in three other Friesian lakes. The mortality rate of males was higher than that of females. Logarithmic length-weight regression analyses showed that the value of the coefficient varied both within and between the sexes, that of females being higher (range 3.03–3.375) than that of males (range 2.76–3.254). Seasonal changes in the size of the coefficient were due to a disproportionate loss of gonad weight in larger fish. The fecundity of the population was comparatively high for the species.
The total production of the population was estimated to be 95 hg ha⁻¹ of which 39 kg ha⁻¹ was contributed by fry. In older (⋝ IV) fish the production of females (12 kg ha⁻¹) exceeded that (2 kg ha⁻¹) of males, due to differences in their growth and mortality rates. The relatively poor performance of roach in Tjeukemeer, in terms of biomass and production was due to a scarcity of zoobenthos and competition from other species offish. There is no evidence either from this study or the literature that the productive potential of roach in lakes is high, even though macrophytes and detritus can be consumed in significant amounts.