The diet and growth of roach (Rutilus rutilus (L.)) in Lake Vesijärvi and possible changes in the course of biomanipulation

Previous studies have suggested that the roach Rutilus rutilus (L.) stock of Lake Vesijärvi is one of the main factors delaying the recovery of the lake after sewage diversion. This study is concerned with the documentation of the diet of roach in the lake. In total, 531 roach were examined. Both in the pelagial and in the littoral the roach had mixed diets in May and in September—October. The importance of zooplankton decreased and the importance of benthos and plants increased with increasing size of roach. In July, in the pelagic zone all sizes of roach fed exclusively on zooplankton (Bosmina spp.), while in the littoral zooplankton had the highest volume proportions only in the smallest (<130 mm) roach. The frequent use of plant food and slow growth rate of large roach indicate a low availability of animal prey. As the fish densities decrease due to the mass removal taking place in the lake, the percentage of plant food in the diets of roach will probably decrease and the growth of roach will increase. Additionally, the tendency of the roach to migrate into the pelagic zone in early summer may be reduced, which would decrease their predation on the zooplankton.     

Intraspecific competition drives multiple species resource polymorphism in fish communities

It has been hypothesized that inter-specific competition will reduce species niche utilization and drive morphological evolution in character displacement. In the absence of a competitor, intra-specific competition may favor an expansion of the species niche and drive morphological evolution in character release. Despite of this theoretical framework, we sometimes find potential competitor species using the same niche range without any partitioning in niche. We used a database on test fishing in Sweden to evaluate the factors (inter- and intraspecific competition, predation, and abiotic factors) that could influence habitat choice of two competitor species. The pattern from the database shows that the occurrence of perch and roach occupying both littoral and pelagic habitats of lakes in Sweden is a general phenomenon. Furthermore, the results from the database suggest that this pattern is due to intra-specific competition rather than inter-specific competition or predation. In a field study, we estimated the morphological variation in perch and roach and found that, individuals of both species caught in the littoral zone were more deeper bodied compared to individuals caught in the pelagic zone. Pelagic perch fed more on zooplankton compared to littoral perch, independent of size, whereas the littoral perch had more macroinvertebrates and fish in their diet. Pelagic roach fed more on zooplankton compared to littoral roach, whereas littoral individuals fed more on plant material. Furthermore, we sampled littoral and pelagic fish from another lake to evaluate the generality of our first results and found the same habitat associated morphology in both perch and roach. The results show a consistent multi-species morphological separation in the littoral and pelagic habitats. This study suggests that intra-specific competition is possibly more important than inter-specific competition for the morphological pattern in the perch-roach system.     

Determinants of the distribution of juvenile fish in the littoral area of a shallow lake

1. Research has often focused on the pelagic areas of lakes; the littoral zone has received less attention. The few studies concerning fish distribution in littoral habitats have concentrated on stands of submersed macrophytes, whereas other littoral habitat types have seldom been investigated. 2. This study aimed to predict the occurrence of juvenile fish in several littoral habitats of a shallow lake as a function of food availability, complexity of habitat structure, water depth and substrate. Habitats comprising reed, woody structures, and two open water areas differing in depth were sampled for fish and invertebrate biomasses on two shores, over 6 months and during both daylight and at night. 3. The juvenile fish community consisted almost exclusively of 0+ and 1+ roach and perch. There was a strong diel component in habitat use, with a predominant occurrence of fish in complex habitats (mainly woody structures) during the day, and a partial migration towards the open habitats at night, more strongly expressed in roach than in perch. 4. The diet of all fish groups was relatively constant over the seasonal cycle, and was independent of habitat. There was a higher degree of planktivory in roach than in perch, but both species fed on benthic macroinvertebrates to a substantial extent. 5. According to a logistic regression model, the biomass of potential food organisms in the different habitats had little predictive effect on the spatial distribution of the fish, whereas the structural complexity of the habitats combined with the diel cycle explained about 28% of the occurrence patterns in 0+ and 1+ perch and 1+ roach.     

Rapid changes in fish community structure and habitat distribution following the precipitation of lake phosphorus with aluminium

1. Fish community structure and habitat distribution of the abundant species roach, perch and ruffe were studied in Lake Nordborg (Denmark) before (August 2006) and after (August 2007) aluminium treatment to reduce internal phosphorus loading. 2. Rapid changes in fish community structure, abundance and habitat distribution occurred following a decline in in‐lake phosphorus concentrations from 280 to 37 μg P L^?1 and an increase in Secchi depth transparency from 1.1 to 1.9 m (August). The proportion of perch in overnight gill net catches increased, whilst roach decreased, and the average weight of all key species increased. 3. The habitat distribution of perch and roach changed from a high proportion in the upper pelagic and littoral zones in 2006, towards enhanced proportions in the deeper pelagic and profundal zone in 2007. The abundance of large‐bodied zooplankton increased and the abundance of benthic invertebrates decreased in the same period, suggesting that the habitat shift was not induced by food limitation. 4. Ruffe shifted from the littoral and upper profundal zones towards the deep profundal zone, likely reflecting an increased predation risk in the littoral zone and better oxygen conditions in the deep profundal. 5. Our results indicate that enhanced risk of predation in the upper pelagic and the littoral zones and perhaps improved oxygen concentrations in the deeper profundal zone at decreasing turbidity are responsible for the observed habitat shift. The results indicate that fish respond rapidly to changes in nutrient state, both in terms of community structure and habitat use.     

Resource limitation during early ontogeny: constraints induced by growth capacity in larval and juvenile fish

The presence of and mechanisms behind density-dependent growth and resource limitation in larval and juvenile stages of organisms with high mortality such as fish are much debated. We compare observed consumption and growth rates with maximum consumption and growth rates to study the extent of resource limitation in young-of-the-year (YOY) roach (Rutilus rutilus) and perch (Perca fluviatilis). Diet, habitat use, consumption rate and growth rate were measured under varying YOY fish densities over 2 years in four lakes. In the first year, YOY roach and perch were studied under allopatric conditions. Experimental addition of perch roe in the second year also allowed study of YOY of the two species under sympatric conditions in two of the lakes. The diet of YOY roach was dominated by cladoceran zooplankton and YOY roach habitat use was restricted to the shore region in both years. This restricted habitat use did not involve any cost in foraging gain in the first year as consumption and growth rates were very close to maximum rates. During the second year, when the two species coexisted, resources were limited in late season, more so in the littoral than in the pelagic habitat in one lake while the reverse was the case in the other lake. The diet of YOY perch was also dominated by zooplankton, and with increasing perch size the proportion of macroinvertebrate prey in the diet increased. After hatching, YOY perch first utilized the pelagic habitat restricting their habitat use to the shore after 1 to several weeks in the pelagic zone. During the larval period, perch were not resource limited whereas juvenile perch were resource limited in both years. The fact that YOY perch were more resource limited than YOY roach was related to the higher handling capacity and lower attack rate of perch relative to roach, rendering perch more prone to resource limitation. Estimates of resource limitation based on consumption rates and growth rates yielded similar results. This supports the adequacy of our approach to measure resource limitation and suggests that this method is useful for studying resource limitation in organisms with indeterminate growth. Our results support the view that density-dependent growth is rare in larval stages. We suggest that density-dependent growth was absent because larval perch and roach were feeding at maximum levels over a wide range of larvae densities. Received: 14 June 1999 / Accepted: 29 October 1999     

Diet-dependent body morphology and ontogenetic reaction norms in Eurasian perch

Individual morphology and performance are directly or indirectly under the influence of variation in resource levels. To study the effects of different resource conditions and their effects on morphology and ontogenetic reaction norms in young‐of‐the‐year (YOY) perch (Perca fluviatilis), we used three different approaches. First, we examined the morphological trajectories over early ontogeny in relation to lake‐specific resources in a field study. Second, one lake that lacked perch recruitment was stocked with perch eggs from a control lake in a whole‐lake experiment to study ontogenetic reaction norms. Third, we compared the development of YOY perch in the three lakes that mainly inhabited the littoral zone with YOY perch experimentally confined to enclosures in the pelagic zone of the lakes.
Overall body morphology of the YOY perch changed both as a function of size and as a function of diet. As perch increased in size they developed a deeper body morphology corresponding to an increased proportion of benthic macroinvertebrates in their diet. In pelagic enclosures where perch were constrained to feed mainly on zooplankton they had a more fusiform body morphology than perch in the lakes that fed on a mixture of zooplankton and macroinvertebrates. Similarly, the ontogenetic reaction norm of perch was related to the diet and lake‐specific zooplankton levels in the whole‐lake experiment.
In the pelagic enclosures, perch with high growth rates had a more fusiform body morphology than slow‐growing perch, whereas the opposite was found in the lakes, where perch included more macroinvertebrates in their diets. Perch in lakes with a higher proportion of macroinvertebrates in the diet also had deeper body morphology. The opposite morphology – growth rate relationship found between perch in the pelagic versus those using the whole lake suggest a morphological trade‐off between foraging on zooplankton and foraging on macroinvertebrates. Our results suggest that YOY perch show different ontogenetic reaction norms as a function of lake‐specific resource levels, which may allow YOY recruitment to later stages. Our results further suggest that diet‐related changes in morphology are a rapid process.     

Habitat shift in roach (Rutilus rutilus) induced by pikeperch (Stizostedion lucioperca) introduction: predation risk versus pelagic behaviour

Changes in the fish community structure and habitat use were followed after the introduction of pikeperch (Stizostedion lucioperca) to the roach-dominated Lake Gjersjøen. Quantitative echosounding showed that the density of juvenile roach (Rutilus rutilus) was dramatically reduced in pelagic areas, from 12 000–15 000 fish/ha to 250 fish/ha, while total fish density remained unchanged in littoral areas. At the same time, the habitat segregation between different size groups of roach was altered as larger roach utilized the pelagic zone after pikeperch introduction. The loss of the pelagic refuge for juvenile roach increased the availability of juvenile roach to littoral predators, notably perch. In littoral areas, the fish community changed from one dominated by roach (> 95%) to one dominated by perch (> 50%).     

The effect of core compression on the measurement of zinc concentrations and anthropogenic burdens in lake sediments

The whitefish (Coregonus lavaretus) in Lake Tyrifjorden, southeastern Norway, is monomorphic with regard to gill raker number: (x=32), and exhibits a bimodal length distribution. The small whitefish (<27 cm) were distributed mainly in the littoral zone and the deeper layers of the pelagic zone. The larger whitefish (>27 cm) were mainly distributed in the upper layers of the pelagic zone. Both small and large whitefish fed on zooplankton in the pelagic zone. In the littoral zone the diet of the two size groups was significantly different; the small whitefish fed mainly on zooplankton whereas the large whitefish fed mainly on zoobenthos.     

Feeding of fish in Lake Glubokoe

Lake Glubokoe is inhabited by 13 species of fish. The commonest are roach, bream, perch and ruff. Comparison of seine catches for 1950 and 1984 revealed a considerable decrease in the share of perch (from 31% to 2%) and ruff (from 18.5 to less than 0.1%) in 1984. The richest food resource in the lake is crustacean plankton. In summer, it is mostly consumed by roach. Estimation of the electivity shows that, unlike perch, roach selectively consumes small crustaceans (Bosmina and Polyphemus). Poor growth rate of all common species of fish points to their food limitation. In summer the diets of common species of fish are differentiated to a considerable extent. Of 36 pairs of combinations a high food overlap was found only in 2 cases: 1) O + perch and Leucaspius delineatus, 2) yearlings and adult roach. Low diet overlap in fish during intensive feeding in summer probably reflects a feedback between the volume and intensity of food competition. The populations of bream and ruff probably compete for chironomid larvae which inhabit the sublittoral, as perch, roach and Leucaspius delineatus probably do for large littoral insects. However, intraspecific competition appears to be more important than interspecific for the roach and bream populations (the two most numerous species in the lake).     

Diel diet shift of roach and its implications for the estimation of daily rations

In a small, 12 ha, mesotropic lake, roach Rutilus rutilus performed diel habitat shifts that clearly influenced the composition of their diet. During daytime, roach stayed in the littoral zone and concentrated on littoral prey. At night they were found in the pelagic zone, and pelagic prey items such as Daphnia spp. or Chaoborus flaricans dominated their food. On a seasonal scale, there were shifts in the importance of different food items and in the diel pattern of feeding intensity. Bioenergetics modelling in combination with an evacuation rate method for estimating daily rations allowed for changes in feeding modes to be taken into account, and so food item specific daily rations over the season could be determined. With the evacuation rate method applied on selected days, diel changes in diet compositions and feeding intensities could be quantitatively accounted for. When the 24 h integrated diet proportions were then used as an input parameter for bioenergetics modelling, food item specific consumption could be determined over the entire sampling season. The consideration of the diel diet shifts proved to be essential for the model output. If only the daytime or the night-time diet composition (derived from one single daily sampling) was taken into account for bioenergetics modelling, severe under- or overestimations of daily rations for specific food items resulted.     

Effects of piscivore-mediated habitat use on growth, diet and zooplankton consumption of roach: an individual-based modelling approach

SUMMARY 1. We used an individual based modelling approach for roach to (i) simulate observed diel habitat shifts between the pelagic and littoral zone of a mesotrophic lake; (ii) analyse the relevance of these habitat shifts for the diet, activity costs and growth of roach; and (iii) quantify the effects of a hypothetical piscivore-mediated (presence of pikeperch) confinement of roach to the littoral zone on roach diet, activity costs and growth.
2. The model suggests that in the presence of pikeperch, roach shifts from zooplankton as the primary diet to increased consumption of less nutritious food items such as macrophytes, filamentous algae and detritus.
3. The growth of roach between May and October was predicted to be significantly higher in the absence of pikeperch, although the net activity costs were about 60% higher compared with the scenario where pikeperch were present.
4. These modelling results provide quantitative information for interpreting diel horizontal migrations of roach as a result from a trade-off between food availability and predation risk in different habitats of a lake.
5. Altering the habitat selection mode of planktivorous roach by piscivore stocking has the potential to reduce zooplankton consumption by fish substantially, and could therefore be used as a biomanipulation technique complementing the reduction of zooplanktivorous fish.     

Morphology dependent foraging efficiency in perch: a trade‐off for ecological specialization?

Trade-offs in foraging efficiency leading to divergent natural selection between and within populations exploiting different resources are thought to be a primary cause of trophic polymorphism. In this study we focused on the trade-offs in foraging efficiency and growth in a polymorphic perch population. Specifically, we related habitat-specific growth and diet of perch to perch morphology. In a subsequent laboratory study we experimentally tested the trade-off by testing the efficiency of perch with different morphology feeding on pelagic ( Daphnia sp., Chaoborus sp.) and littoral (mayfly larvae) food resources. The feeding performance was tested in different physical environments to see if we could predict growth patterns in the field based on foraging rate and behavior of perch.
In the field study, we found that the perch from the littoral and the pelagic zones differed in both morphology and diet. Within the littoral zone the deeper-bodied individuals grew faster compared to the more streamlined individuals, whereas the opposite pattern was found in the pelagic zone. In the aquarium experiments, perch from the littoral zone had higher capture rates on the pelagic prey types in vegetation trials and on mayfly larvae in both open water and vegetation trials. The pelagic perch had higher capture rates on the pelagic prey types in open water trials. The littoral perch had lower search velocity than the pelagic perch in open water trials whereas the opposite pattern was found in vegetation trials. The attack velocity of the pelagic perch was also higher than that of the littoral perch independent of vegetation structure. Our results suggest that there is a functional trade-off between performance in alternate habitats and general body form in perch. Such trade-offs may promote divergent natural selection and could be the mechanism that give rise to and upholds the pattern in the field.     

REGULAR PAPERS Habitat distribution and size structure in freshwater fish communities: effects of vendace on interactions between perch and roach

In a study of habitat selection and size structure in three fish species in 115 Swedish lakes, roach Rutilus rutilus used the pelagic habitat to a lesser extent and were lower in pelagic biomass in the presence of vendace Coregonus albula. Size distributions of roach were skewed toward larger sizes in lakes with vendace, although a similar pattern of size-dependent habitat use was found regardless of lake group. In disagreement with predictions, vendace did not affect relative perch Perca fluviatilis biomass, size structure, or the proportion of large perch. However, a higher growth for large perch indicated higher per capita energy intake in lakes with vendace.     

Effects of habitat and food resources on morphology and ontogenetic growth trajectories in perch

Studies on resource polymorphism have mainly been considered at the end stage of ontogeny, whereas many species undergo diet changes as they grow. We conducted a field survey to analyze the role of adaptive variation during ontogeny in Eurasian perch (Perca fluviatilis). We caught perch from the littoral and pelagic zones of a lake to investigate whether perch differ in morphology and diet between these habitats. We also investigated whether there were any differences in morphological trajectories during the ontogeny of perch collected from the two habitats. We found that perch caught in the littoral habitat, independently of size, had a deeper body, larger head and mouth and longer fins than perch caught in the pelagic zone. Macroinvertebrates and fish dominated the diet of littoral perch, whereas the diet of the pelagic perch consisted mainly of zooplankton and to some extent fish. Independently of size, the more streamlined individuals had a larger proportion of zooplankton and a smaller proportion of macroinvertebrates in their diet than the deeper-bodied individuals, indicating a relation between diet and morphology. Some morphological characters followed different ontogenetic trajectories in the two habitats; e.g. the changes to a deeper body and a larger head were faster in the littoral than in the pelagic perch. The relationship between the length of perch and the size of the mouth and fins also differed between perch from the two habitats, where the increase in the length of the pelvic fin and the area of the mouth increased faster with size in the littoral perch. Our findings show that variation in morphology between habitats differs during ontogeny in a way that corresponds to functional expectations for fish species that occupy these habitats.     

Effect of vertebrate predation on the spatio-temporal distribution of cladocerans in a temperature eutrophic lake

We analysed the spatio-temporal distribution of zooplankton along a profile of 10 stations from the shore to the pelagic zone from April to September 1988, the period when the larvae and juveniles Rutilus rutilus, the most abundant species in the Lake, are in the littoral zone. The digestive tracts of the young roach were analysed. They fed essentially on rotifers and on cladocerans. For comparison, zooplankton was also analysed at one littoral area without fish fry. There was an increase of cladoceran density from the vegetated nearshore zone to the offshore zone. Considering the density of Bosmina longirostris, Daphnia longispina, Chydorus sphaericus and Ceriodaphnia quadrangula, we observed a different distribution pattern in the course of the year. In the nearshore zone, the relative abundance of small species, Bosmina and Chydorus, was much higher than that of the larger Daphnia. From April to September, predation pressure mainly affected the smallest species: in contrast to the inshore station without fish fry, the density of Bosmina decreased in May in the littoral with fish. Chydorus was concentrated in the littoral between February and April, then grew into the pelagic zone, where predation pressure obviously was low during the warm season. The number of Daphnia, which was eaten by the fish fry at any time, remained low in the nearshore zone, which suggests that the presence of fish may cause Daphnia to avoid this zone. Ceriodaphnia which was not affected by this predation, was scarce in the nearshore zone during mid-summer. The low density of the cladocerans in the nearshore zone is likely associated with vertebrate predation by roach fry and juveniles, the result of such a process being either a depletion in density of the prey, or an avoidance behaviour.     

Sensitivity of fitness to macroparameter changes: an analysis of survivorship and individual growth in sea urchin life histories

Summary Feeding relationships between roach and ide from two sites in a mesotrophic lake SE Norway are presented and discussed. When animal food supply was scarce, both fish species increased their consumption of macrophytes; roach sevenfold, and ide threefold. Along a typical littoral vegetation gradient, ide fed among helophytes, while roach fed in the zone outside. This different habitat selection was reflected in the most important plants consumed (ide: Equisetum fluviatile, roach: Characeae), and confirmed by gill net catches. During the vegetative season, roach avoided areas with dense vegetation. In shallow areas beyond the littoral, the most important food plant for both fish species was Potamogeton perfoliatus, which constituted 80% of the total food consumed (dry weight basis) in the roach and 35% in the ide. The diet shift to plants seemed to be strongly influenced by the supply of animal food and the intensity of intra-and interspecific competition.     

Biomass of planktonic crustaceans and the food of young cyprinids in the littoral zone of Lake Balaton

The littoral zone of Lake Balaton and its periphyton-zooplankton-fish communities have been investigated intensively in recent years. Total average number of crustacean plankton varied from 36 to 126 ind l^–1, their biomass from 0.49 to 1.86 mg ww l^–1 month^–1 at different areas of the littoral zone. In general, these values for the above parameters were higher in hypertrophic areas. 23 fish species occurred in the littoral zone with cyprinids dominating. The seasonal food spectra of Y-O-Y roach (Rutilus rutilus), white bream (Blicca bjoerkna) and bream (Abramis brema) were based mainly on planktonic crustaceans and benthic/periphytic invertebrates. According to the frequency of occurrence of crustaceans and other invertebrates, the food composition of young cyprinids differed significantly in the NE and SW-basins of the lake.     

Chaos and stability of age-0 fish assemblages in a temperate deep reservoir: unpredictable success and stable habitat use

Large year-to-year variability in different fish species recruitment has been confirmed by previous studies while diurnal patterns of occupation in two basic reservoir habitats (pelagic and littoral) by different age-0 fish species in late summer are still unclear. Data collected over an 11-year period regarding late-summer age-0 fish assemblages in pelagic and littoral habitats of a reservoir were used to test the recruitment instability and to investigate diurnal habitat use. Trawling was conducted in the pelagic habitat at night while beach seining was conducted in the littoral habitat during day and night. Fluctuations in age-0 fish abundance and species composition were observed with both sampling methods; however, the following spatio-temporal patterns were relatively stable in most investigated years: (1) pelagic species (pikeperch; Sander lucioperca, small perch; Perca fluviatilis, bream; Abramis brama at night), (2) littoral species (large perch, asp; Leuciscus aspius, dace; Leuciscus leuciscus), (3) migratory species likely performing diel horizontal migrations (bleak; Alburnus alburnus), (4) species abundant in the littoral habitat both during day and night and also in pelagic habitat at night (roach; Rutilus rutilus) and (5) species detected in both habitats exclusively at night (ruffe; Gymnocephalus cernuus).     

Spatial and temporal separation of littoral zone fishes of Lake Geneva (Switzerland-France)

The spatial (distance from the shore) and seasonal (summer-winter) distributions of the ichtyofauna in the littoral zone of Lake Geneva were studied between October 1991 and December 1992. Gill nets were set simultaneously at three different sites of a chosen littoral zone: shore, middle and far (50, 300 and 550 m from the shore, respectively).The results show that in summer more species and individuals are present in the littoral zone than in winter. Throughout the year, species richness is greater at the shore site than at the middle or far site. In summer, total abundance is greater at the far site, whereas there are more fish at the shore site during winter. Perch (juveniles and adults), roach and gudgeon (adults only), have maximum abundance during the summer, while lake trout, grayling and whitefish (adults) have maximum abundance during the winter. For five species (adults only) the site comparison shows spatial segregation of the habitats: greater numbers of perch are present at the far site while roach, tench, lake trout and grayling prefer the shore site.     

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.