Food selection in juvenile three-spined sticklebacks studied in relation to size,abundance and biomass of prey

The food selection and diet of juvenile three-spined sticklebacks (Gasterosteus aculeatus L.) was studied in the littoral of a brackish-water sea area. The sampling stations were located in a eutrophicated area, where the plankton was characterized by rotifers, in a noneutrophicated zone, where copepods predominated, and in the intermediate zone, characterized by cladocerans. In all areas the fish preferred microcrustaceans to rotifers, but they did not feed strictly according to the size-selection hypothesis. Nor did they take the most abundant prey species. With some exceptions the species chiefly eaten by and also selected by fish, were those which had the largest biomass in the plankton.     

Differences in feeding selectivity and efficiency between young-of-the-year European perch (Perca fluviatilis) and roach (Rutilus rutilus) — field observations and laboratory experiments on the importance of prey movement apparency vs. evasiveness

Feeding selectivity and efficiency of young-of-the-year European perch and roach were compared under field and laboratory conditions. In laboratory experiments, the importance of prey evasiveness versus prey movement conspicuousness for fish selectivity was evaluated with respect to changing Cladocera/Copepoda prey ratio. Feeding efficiency was additionally investigated in relation to feeding time (5, 10, 20 min) and prey density (approx. 50, 200, 700 ind. L⁻¹). In Říov Reservoir, the diet of both fish species was nearly exclusively composed of crustacean zooplankton. In roach, diet shifted from rotifers and bosminids in May, towards Daphnia sp. and Leptodora kindtii in June and July. Daphnia contributed almost exclusively to the roach diet since June, composing on average more than 94% of total prey. Cyclopoid copepods, occurred in the roach’s diet only on the first sampling date; later on both cyclopoid and calanoid copepods were completely absent. On the other hand, copepods played an important role in the diet of perch. In early and mid-June when their share in the zooplankton was particularly high, copepods contributed by more than 50% to the diet of perch. Although their contribution dropped with their decline in zooplankton in June/July, by the end of July they again comprised about one third of perch’s diet. In both fish species, the increase in numbers of cladocerans in their diet was related to increase in SL. In roach, the numbers of consumed prey were doubled every twenty days during the investigated period. In perch the increase was not so consistent, but significantly higher efficiency of perch was reported on three out of six sampling dates. In laboratory experiments, roach showed a distinct avoidance for copepods and a preference for cladocerans. Both prey categories were only fed non-selectively when they dominated the prey mixture. Perch selectivity was more diversified. Contrary to roach, perch were fed copepods non-selectively on a balanced prey ratio. Further, with an increasing share of Cladocera, a situation resembling that of roach and Copepoda was avoided. However, when the share of copepods in the prey mixture dropped below ten percent, they were consumed non-selectively and with their ongoing decline in the prey mixture their preference even increased. Feeding efficiency differed significantly between perch and roach when foraging on copepods exclusively or on a prey mixture where copepods predominated. In the short time feeding experiment (5 min) with copepods, perch consumed on average 5.9 times more prey than roach. Although roach increased their success with increasing time it was still 1.7 times greater than for perch in the long time feeding experiment (20 min). Total numbers of prey consumed were positively affected by prey density and feeding time. With increasing feeding time, the consumption rate generally declined. With a fourfold increase in feeding time, the numbers of consumed prey increased on average only two times. Only in roach feeding on copepods did the numbers of prey consumed per minute of feeding increase with increasing feeding time. However, the overall numbers were low. Differences in feeding selectivity and efficiency between perch and roach juveniles were found to be significant both in the field and laboratory experiments. In roach, selectivity was determined solely by prey evasiveness. By contrast, perch’s selectivity was influenced by prey movement conspicuousness; prey escape abilities did not play an important role. Perch were more efficient foragers on evasive prey, but its feeding efficiency for non-evasive prey was not lower than that of roach. According to our observations, we suggest feeding behaviour to be responsible for the roach’s inefficiency in capturing evasive copepods.     

Experience affects performance of ten-spined sticklebacks foraging on zooplankton

To study the effects of short-term experience on prey size-selection ten-spined sticklebacks (Pungitius pungitius) were fed 7–13 days with five differing diets of novel prey, Daphnia magna. The diets consisted either of a mixture of two prey size classes (1.7 and 2.2 mm) or of single-sized (1.7 and 2.2 mm) prey. Before and after the diets, the sticklebacks' prey size selection was tested with a 1:1 ratio of 1.7 and 2.2 mm D. magna. Sticklebacks made more attempts to capture large than small prey, but their foraging success was better for small than for large prey. Sticklebacks fed with a diet of both prey sizes chose significantly more large prey on the 13th day than on the 7th day or at the beginning of the experiment. Handling times for both Daphnia size classes decreased slightly with increasing foraging experience. Inexperienced sticklebacks made more unsuccessful strikes on large prey than did experienced fish. Foraging success on large prey improved somewhat with increasing experience in all but one diet group. The results indicate that experience affected ten-spined sticklebacks' prey selection.     

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.     

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.     

Food and feeding behaviour of a planktivorous cyprinid, Pelecus cultratus (L.), in a shallow eutrophic lake, Neusiedler See (Austria)

Pelecus cultratus (razor fish), a cyprinid fish, has become a dominant species in Neusiedler See. Gut content analyses of 400 specimens collected in 1989 and 1990 showed that Pelecus fed mainly on large zooplankton (Diaphanosoma, Leptodora and Arctodiaptomus), although their diet also included Insecta (larvae, pupae and adults) and Arachnida (spiders), occasionally small fishes. Comparison of the relative abundance of the zooplankton species in the stomach to the lake indicated that Pelecus strongly selected cladocerans over copepods, and fed mostly on large-sized individuals of Diaphanosoma, Leptodora and Arctodiaptomus. The fish showed a significant positive selectivity only for individuals of Diaphanosoma > 1.0 mm and Leptodora > 4.0 mm. In contrast, selectivity increased continuously in relation to the diameter of the compound eye of both prey species. This suggested that prey visibility was a key factor in determining the prey selectivity by Pelecus. It also seems likely that the persistence of the Leptodora population in Neusiedler See can be attributed to negligible predation pressure on the smaller sized individuals of this species.     

Production,consumption and prey availability of northern pike (Esox lucius), pikeperch (Stizostedion lucioperca) and European catfish (Silurus glanis): a bioenergetics approach

Bioenergetics models were applied for the assessment of food consumption of northern pike, pikeperch and European catfish in drainable ponds of 0.4 ha. The ponds were stocked with known numbers of the 0⁺ predators. The prey fish consisted of naturally recruited 0⁺ cyprinids (rudd, roach and bream). The study shows that the impact of the three 0⁺ piscivores on the aquatic system differs, which is related to the feeding behaviour and habitat use of the fish. All piscivores realised part of their production on a non-fish diet.     

Impacts of a submerged plant (Elodea canadensis) on interactions between roach (Rutilus rutilus) and its invertebrate prey communities in a lake littoral zone

1. Using 5‐m² field enclosures, we examined the effects of Elodea canadensis on zooplankton communities and on the trophic cascade caused by 4–5 year old (approximately 16 cm) roach. We also tested the hypothesis that roach in Elodea beds use variable food resources as their diet, mainly benthic and epiphytic macroinvertebrates, and feed less efficiently on zooplankton. Switching of the prey preference stabilises the zooplankton community and, in turn, also the fluctuation of algal biomass. The factorial design of the experiment included three levels of Elodea (no‐, sparse‐ and dense‐Elodea) and two levels of fish (present and absent). 2. During the 4‐week experiment, the total biomass of euplanktonic zooplankton, especially that of the dominant cladoceran Daphnia longispina, decreased with increase in Elodea density. The Daphnia biomass was also reduced by roach in all the Elodea treatments. Thus, Elodea provided neither a favourable habitat nor a good refuge for Daphnia against predation by roach. 3. The electivity of roach for cladocerans was high in all the Elodea treatments. Roach were able to prey on cladocerans in Elodea beds, even when the abundance and size of these prey animals were low. In addition to cladocerans, the diet of roach consisted of macroinvertebrates and detrital/plant material. Although the biomass of macroinvertebrates increased during the experiment in all Elodea treatments, they were relatively unimportant in roach diets regardless of the density of Elodea beds. 4. Euplanktonic zooplankton species other than Daphnia were not affected by Elodea or fish and the treatments had no effects on the total clearance rate of euplanktonic zooplankton. However, the chlorophyll a concentration increased with fish in all the Elodea treatments, suggesting that fish enhanced algal growth through regeneration of nutrients. Thus, our results did not unequivocally show that Elodea hampered the trophic cascade of fish via lowered predation on grazing zooplankton. 5. In treatments with dense Elodea beds (750 g FW m^?2), chlorophyll a concentration was always low suggesting that phytoplankton production was controlled by Elodea. Apparently, the top‐down control of phytoplankton biomass by zooplankton was facilitated by the macrophytes and operated simultaneously with control of phytoplankton production by Elodea.     

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.

     

Ontogenetic niche shifts and resource partitioning in a subarctic piscivore fish guild

The feeding ecology of three piscivorous fish species (perch (Perca fluviatilis), pike (Esox lucius) and burbot (Lota lota)), was studied in the subarctic Pasvik watercourse (69 °N), northern Norway and Russia. All three species primarily occupied the benthic habitats in the watercourse. Perch and burbot exhibited distinct ontogenetic niche shifts in food resource use, perch changing from a dominance of zooplankton to zoobenthos to fish, and burbot from zoobenthos to fish. Fish prey dominated the diet of all the investigated size-classes of pike, but small-sized pike (<20 cm) were not represented in the sample. Fish prey size was positively related to predator size in all three species. Whitefish (Coregonus lavaretus) was the dominant prey of pike and large-sized burbot and perch. Nine-spined sticklebacks (Pungitius pungitius) was also an important prey and appeared to be a dietary stepping-stone enhancing the transition from invertebrate feeding to consumption of large-sized whitefish prey for all three predators. A cluster analysis separated the different size groups of the three predator species into five functional feeding groups, most of them containing two or all three species. Within these feeding groups, and especially among the piscivorous size-classes, there was a strong and significant interspecific overlap in prey selection, and the dietary similarities between the species were in general much larger than the intraspecific similarities between ontogenetic stages. All three piscivorous species are important top predators in the aquatic food web of the watercourse, and their ontogenetic diet shifts and resource partitioning patterns generate a substantial food web complexity in this subarctic ecosystem.     

Changing handling times during feeding and consequences for prey size selection of 0+ zooplanktivorous fish

The interrelationship of fish size, prey size and handling time within a 15-min feeding period was studied in three size groups of 0 + roach, Rutilus rutilus, and bleak, Alburnus alburnus. Four size classes of cladoceran prey were used to measure changes in feeding rate and handling time from initial rapid feeding to sustained feeding. Observed differences in increase of handling time between prey size classes led to a change in the prey profitability ranking of those size classes within the first 2 min of the experiments. A 1-min feeding period is interpreted as reflecting an intermediate motivational status between extreme hunger and satiation. The use of average handling times for this period revealed a substantial change in prey profitability estimates compared to previous studies which used handling times based on short-term (a few seconds up to 1 min) feeding. It is not the largest prey items a fish can handle and swallow that are most profitable, but prey of intermediate size. By this approach a closer fit between expectations derived from optimal foraging theory and empirical data on prey size selection of 0 + zooplanktivorous fish is qualitatively achieved. Optimal prey size was found to be close the mouth gape width in small fish of 15 mm standard length, decreasing to 50% of mouth gape width in fish of 40 mm standard length.     

Ontogenetic dietary changes of whitefish larvae: insights from field and experimental observations

Ontogenetic changes in resource use are widespread in many fish species. This study investigated the feeding habits of whitefish (C. lavaretus L.) larvae in Lake Annecy (France) coupled with experimental behavioral studies in order to identify the underlying mechanisms of the ontogenetic shifts in the diet. The predatory behavior of wild larvae, and the escape responses of their zooplankton prey were both videorecorded in experimental tanks under controlled laboratory conditions. Ontogenetic diet patterns showed that young whitefish larvae have a preference for small cyclops, while older larvae selectively predate cladocerans. Our experimental observations showed that the capture success rate also varied in relation to ontogenetic development in fish. Young larvae were more successful in capturing small copepods, whereas old larvae were more successful in capturing Daphnia. In addition, the larvae were able to adjust their predatory behavior (speed, pursuit) according to the swimming pattern of the prey. These observations suggest that the selective predation on cladocerans observed in old larvae is the outcome of both active and passive choices depending on the escape swimming behavior of the prey, and handling time of the predator.     

Rudd (Scardinius erythrophthalmus) introduced to the Iberian peninsula: feeding ecology in Lake Banyoles

The first data on the ecology of rudd (Scardinius erythrophthalmus) introduced to the Iberian peninsula are presented. The habitat and diet variation of rudd were studied in Lake Banyoles (Spain), an oligotrophic karstic lake dominated by exotic fish species. Rudd were strictly littoral and the diet was based on detritus and plant material. The most important animal prey were the cladocerans Daphnia longispinaand Scapholeberis rammneri, amphipods and several late stages of nematoceran dipterans. Rudd were more zooplanktivorous in spring and autumn and less in summer. There was also a size-dependent diet shift, from microcrustaceans to macroinvertebrates. The diet of rudd was also distinguished by the importance of plant material and various small neustonic invertebrates, particularly S. rammneriand late stages of nematocerans, showing a strong resource partitioning with other fish species. The degree of herbivory in Lake Banyoles was lower than usual.     

Possible competitive interactions between overwintering tufted duck (Aythya fuligula (L.)) and fish populations of Lough Neagh,Northern Ireland: evidence from diet studies

The overwintering population of tufted duck (Aythya fuligula) on Lough Neagh, Northern Ireland, has varied over the last three decades, with numbers declining in the early 1980s but then increasing to former levels in the late 1980s. Population fluctuations of recently introduced roach (Rutilus rutilus) mirrored these trends. The present study explores the possibility that competition for benthic food resources is responsible for these changes by examining the diets of tufted duck, roach and other major fish species of the lake. Diet overlaps were generally high due to the common consumption of chironomid larvae. The diet of tufted duck overlapped most with that of roach because these two species were the only significant consumers of molluscs. This evidence provides further support for a tufted duck — roach competition hypothesis.     

Direct and indirect impacts of predation by fish on the zooplankton community: an experimental analysis using tanks

The impact of Pseudorasbora parva, a common zooplanktivorous fish species in Japan, on a zooplankton community was analyzed in experimental tanks, half of which were stocked with the fish. Different zooplankton species showed different responses to the introduction of the fish. In the presence of the fish, the populations of the large cladoceran Ceriodaphnia and the predatory copepod Mesocyclops were reduced, but the population of the herbivorous copepod Eodiaptomus and the small cladocerans Bosmina fatalis and Bosminopsis deitersi increased relative to the controls. The increase of Mesocyclops seen in the control tanks might have suppressed the populations of the small cladocerans, which are vulnerable to invertebrate predation. The results suggest that the population densities of the large prey items preferred by the fish, Ceriodaphnia and Mesocyclops, were controlled directly by fish predation, but the population densities of the smaller and less preferred zooplankton were controlled indirectly through the food-web cascade.     

Zooplankton community structure driven by vertebrate and invertebrate predators

Summary A zooplankton community was established in outdoor experimental ponds, into which a vertebrate predator (topmouth gudgeon: Pseudorasbora parva) and/or an invertebrate predator (phantom midge larva: Chaoborus flavicans) were introduced and their predation effects on the zooplankton community structure were evaluated. In the ponds which had Chaoborus but not fish, small- and medium-sized cladocerans and calanoid copepods were eliminated while rotifers became abundant. A large-sized cladoceran Daphnia longispina, whose juveniles had high helmets and long tailspines as anti-predator devices, escaped from Chaoborus predation and increased. In the ponds which had fish but not Chaoborus, the large-sized Daphnia was selectively predated by the fish while small-and medium-sized cladocerans and calanoid copepods predominated. In the ponds containing both Chaoborus and fish, the fish reduced the late instar larvae (III and IV) of Chaoborus but increased the early instar larvae (I and II). Small- and large-sized cladocerans were scarcely found. The former might have been eliminated by predation of the early instar larvae of Chaoborus, while the latter was probably predated by fish. Consequently, the medium-sized cladocerans, which may have succeeded in escaping from both types of predator, appeared abundantly. The results suggest that various combinations of vertebrate and invertebrate predators are able to drive various kinds of zooplankton community structure.     

Piscivorous fishes diet dominated by the Asian cyprinid invader,topmouth gudgeon (<Emphasis Type="Italic">Pseudorasbora parva</Emphasis>)

Predation on the non-native cyprinid, Pseudorasbora parva, was reported during an annual study in a small connecting canal about 40 km south of Brno, Czech Republic. Two cyprinid fish species (Pseudorasbora parva, Rutilus rutilus) were identified in the diet of 23 piscivores belonging to three native fishes (Perca fluviatilis, Sander lucioperca, Esox lucius). The relative abundance of prey fishes changed seasonally and was dominated by the topmouth gudgeon in spring. Thereafter roach incidence continuously increased. In contrast, topmouth gudgeon was found to be a more sized-available prey and was the most dominant prey species in the diet of all piscivorous fishes during the whole season. Strong piscivory towards this alien might be beneficial for its eradication or control to support ecological integrity in natural and modified seminatural habitats.     

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.     

The relationship between cladoceran body size and the growth of underyearling roach (Rutilus rutilus) (L.) in two shallow lowland lakes: a mechanism for density-dependent reductions in growth

Large planktonic Cladocera are typically the most important components of the diet of underyearling roach. Selection for large species and individuals by fish can result in a shift in the species composition of the cladoceran community as well as a reduction in the mean size of the individuals of large species and in the assemblage as a whole. Laboratory experiments demonstrated that underyearlings feeding on Daphnia hyalina smaller than 1 mm in length has a significantly lower intake of prey volume per unit time than when feeding on prey greater than 1.5 mm. A decrease in the nutritional quality of zooplankton prey, brought about by increasing predation pressure is suggested as the mechanism for density-dependent reductions in the growth of underyearling roach in eutrophic water bodies.     

Diet spectra and resource partitioning in the larvae and juveniles of three species and six cohorts of cyprinids from a subalpine lake

Summary Diet composition based on gut analyses was studied in larvae and juveniles belonging to six (out of eight) age groups (cohorts) of three species of cyprinids (Rutilus rutilus L., Leuciscus cephalus L., Scardinius erythrophthalmus L.) from a small meso-oligotrophic lake in Tyrol, Austria.A basic pattern of ontogenetic shifts of resource use is postulated for the first weeks after hatching, consisting of the sequence: phytoplankton-rotifers-crustaceans-chironomid larvae. However, there are several variations to this general theme.Diet overlap is of about the same magnitude between representatives of different species or different cohorts, and between members of schools belonging to one cohort. This points to the importance of random food selection in all larvae and juveniles during this phase of life.Prey size is a very poor predictor of food choice by young cyprinids, but there is greater similarity in diet between the larger juveniles than between the smaller larvae, irrespective of whether the fish compared represent different species, different cohorts or are members of homogeneous groups.The lack of correlation between prey size and predator size may be explained by assuming that out of a limited range of available prey size the fish always try to include in their diet also the largest items they are able to swallow. This would be a good strategy considering that growth rates are positively correlated with food size.One clearcut interspecific difference in resource use may be noted: The larvae of L. cephalus are distinguished from those of the other two species by the absence of rotifers and nauplii in their diet, and by their greater ability to handle both adult copepods and chironomid larvae.