Piscivore-Prey Fish Interactions: Mechanisms behind Diurnal Patterns in Prey Selectivity in Brown and Clear Water

Environmental change may affect predator-prey interactions in lakes through deterioration of visual conditions affecting foraging success of visually oriented predators. Environmental change in lakes includes an increase in humic matter causing browner water and reduced visibility, affecting the behavioural performance of both piscivores and prey. We studied diurnal patterns of prey selection in piscivorous pikeperch (Sander lucioperca) in both field and laboratory investigations. In the field we estimated prey selectivity and prey availability during day and night in a clear and a brown water lake. Further, prey selectivity during day and night conditions was studied in the laboratory where we manipulated optical conditions (humic matter content) of the water. Here, we also studied the behaviours of piscivores and prey, focusing on foraging-cycle stages such as number of interests and attacks by the pikeperch as well as the escape distance of the prey fish species. Analyses of gut contents from the field study showed that pikeperch selected perch (Perca fluviatilis) over roach (Rutilus rutilus) prey in both lakes during the day, but changed selectivity towards roach in both lakes at night. These results were corroborated in the selectivity experiments along a brown-water gradient in day and night light conditions. However, a change in selectivity from perch to roach was observed when the optical condition was heavily degraded, from either brown-stained water or light intensity. At longer visual ranges, roach initiated escape at distances greater than pikeperch attack distances, whereas perch stayed inactive making pikeperch approach and attack at the closest range possible. Roach anti-predatory behaviour decreased in deteriorated visual conditions, altering selectivity patterns. Our results highlight the importance of investigating both predator and prey responses to visibility conditions in order to understand the effects of degrading optical conditions on piscivore-prey interaction strength and thereby ecosystem responses to brownification of waters.     

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%).     

Feeding efficiency of planktivorous perch Perca fluviatilis and roach Rutilus rutilus in varying turbidity: an individual‐based approach

The feeding rate of perch Perca fluviatilis showed high individual variation at low and moderate turbidities, when one individual had consumed more Daphnia pulex than any other fish, whereas no such variation in feeding efficiency was observed with roach Rutilus rutilus. There was a significant decrease in total consumption of D. pulex by P. fluviatilis with increasing turbidity, but no correlation was observed in the case of R. rutilus. The results suggest that the difference in the ontogeny of P. fluviatilis and R. rutilus may be detectable as behaviour‐related species‐specific trait differences in the early planktivorous feeding stage of the two common species.     

Trophic dynamics in a whole lake experiment: size-structured interactions and recruitment variation

Horizontal and vertical heterogeneity as a result of size‐structured processes are important factors influencing indirect effects in food webs. In a whole‐lake experiment covering 5 years, we added the intermediate consumer roach (Rutilus rutilus) to two out of four lakes previously inhabited by the omnivorous top predator perch (Perca fluviatilis). We focused our study on the direct consumption effect of roach presence on zooplankton (and indirectly phytoplankton) versus the indirect effect of roach on zooplankton (and phytoplankton) mediated via effects on perch reproductive performance. The patterns in zooplankton and phytoplankton abundances were examined in relation to population density of roach and perch including young‐of‐the‐year (YOY) perch in the light of non‐equilibrium dynamics. The presence of roach resulted in changed seasonal dynamics of zooplankton with generally lower biomasses in May–June and higher biomasses in July–August in roach lakes compared to control lakes. Roach presence affected perch recruitment negatively and densities of YOY perch were on average higher in control lakes than in treatment lakes. In years when perch recruitment did not differ between lakes as a result of experimental addition of perch eggs, total zooplankton biomass was lower in treatment lakes than in control lakes. Phytoplankton biomass showed a tendency to increase in roach lakes compared to control lakes. Within treatment variation in response variables was related to differences in lake morphometry in treatment lakes. Analyses of the trophic dynamics of each lake separately showed strong cascading effects of both roach and YOY perch abundance on zooplankton and phytoplankton dynamics. Consideration of the long transients in the dynamics of top predators (fish) in aquatic systems that are related to their long life span involving ontogenetic niche shifts is essential for making relevant interpretations of experimental perturbations. This conclusion is further reinforced by the circumstance that the intrinsic dynamics of fish populations may in many cases involve high amplitude dynamics with long time lags.     

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.     

Can species‐specific prey responses to chemical cues explain prey susceptibility to predation?

The perception of danger represents an essential ability of prey for gaining an informational advantage over their natural enemies. Especially in complex environments or at night, animals strongly rely on chemoreception to avoid predators. The ability to recognize danger by chemical cues and subsequent adaptive responses to predation threats should generally increase prey survival. Recent findings suggest that European catfish (Silurus glanis) introduction induce changes in fish community and we tested whether the direction of change can be attributed to differences in chemical cue perception. We tested behavioral response to chemical cues using three species of freshwater fish common in European water: rudd (Scardinius erythrophthalmus), roach (Rutilus rutilus), and perch (Perca fluviatilis). Further, we conducted a prey selectivity experiment to evaluate the prey preferences of the European catfish. Roach exhibited the strongest reaction to chemical cues, rudd decreased use of refuge and perch did not alter any behavior in the experiment. These findings suggest that chemical cue perception might be behind community data change and we encourage collecting more community data of tested prey species before and after European catfish introduction to test the hypothesis. We conclude that used prey species can be used as a model species to verify whether chemical cue perception enhances prey survival.     

Effects of liming on the occurrence and abundance of fish populations in acidified Swedish lakes

Test fishing with multimesh gillnets was performed in 103 acidified Swedish lakes before and 2–4 years after liming. In a subset of 39 lakes additional test fishing was carried out 5–9 years after liming. Taking into account the sampling biases due to differences between water temperature and number of nets used on different sampling occasions, an increase of the total CPUE, the number of species caught and recruitment of dominating species was evident after liming. Decreased recruitment was found for perch (Perca fluviatilis) at a pH below 5.2 and for roach (Rutilus rutilus) at a pH below 6.1. During the acidification phase and 2–4 years after liming acid-tolerant perch dominated the lakes, but when a longer time had elapsed after liming species dominance shifted and acid-sensitive planktivorous species increased in CPUE while perch decreased.     

Structural equation models suggest that bottom‐up processes override top‐down processes in boreal pikeperch (Sander lucioperca) lakes

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Fish–duck interactions in boreal lakes in Finland as reflected by abundance correlations

We studied the hypothesis that fish play an important role in lake use by ducks (pairs and broods) in boreal lakes. The study was based on densities of different duck and fish species in 28 boreal lakes in southern Finland. We focused on the three most common duck species (mallard Anas platyrhynchos, green-winged teal A. crecca and common goldeneye Bucephala clangula) and on the three most common fish species (perch Perca fluviatilis, roach Rutilus rutilus and pike Esox lucius) in the region. We considered both competitive and predatory interactions between ducks and fish, the perch and roach being potential competitors with ducks and the pike a potential predator of ducks. We found a negative association between green-winged teal brood density and total fish density, the other duck species having only a weak association with total fish density. When the three fish species were considered separately, a negative association, suggesting food competition, was found between perch, green-winged teal and goldeneye, whereas the role of roach as a food competitor seemed to be of minor importance. We did not find any clear signs of predatory effects of pike on ducks. Our results suggest that food competition is a more important factor than pike predation in affecting lake use by ducks in oligotrophic boreal environments in southern Finland.     

Morphological variation of perch Perca fluviatilis in humic lakes: the effect of predator density,competition and prey abundance

Between and within‐lake variations in morphology of perch Perca fluviatilis were studied in four humic lakes in eastern Finland. Perca fluviatilis were more streamlined and smaller headed in a lake with the highest abundance of cyprinids, but lowest abundance of predators (Lake Tuopanjärvi), indicating adaptation to planktivorous feeding and low predator density. Highest bodied fish were found from a lake with the lowest cyprinid but highest predator abundance (Lake Koppelojärvi), which conversely indicates adaptation to more effective predator avoidance. Furthermore, the length of the paired fins was longest in Lake Kinnasjärvi and Lake Tuopanjärvi, where the abundance of benthic macroinvertebrates was lowest, suggesting selection for more effective benthivory. Clear morphological differences of P. fluviatilis between habitats were found only in Lake Kinnasjärvi, whereas in Lake Koppelojärvi and Lake Tuopanjärvi only the length of the paired fins differed and in Lake Harkkojärvi no differences were found. Taken together, these results suggest that inter and intrapopulation morphological differences are probably highly dependent on different biotic factors (i.e. predation risk, resource availability and competition). Spatial and temporal variations in these factors may have a great effect on body morphology of P. fluviatilis.     

Individually assessed boldness predicts Perca fluviatilis behaviour in shoals,but is not associated with the capture order or angling method

Selectivity of recreational angling on fish behaviour was studied by examining whether capture order or lure type (natural v. artificial bait) in ice‐fishing could explain behavioural variation among perch Perca fluviatilis individuals. It was also tested if individually assessed personality predicts fish behaviour in groups, in the presence of natural predators. Perca fluviatilis showed individually repeatable behaviour both in individual and in group tests. Capture order, capture method, condition factor or past growth rate did not explain variation in individual behaviour. Individually determined boldness as well as fish size, however, were positively associated with first entrance to the predator zone (i.e. initial risk taking) in group behaviour tests. Individually determined boldness also explained long‐term activity and total time spent in the vicinity of predators in the group. These findings suggest that individual and laboratory‐based boldness tests predict boldness of P. fluviatilis in also ecologically relevant conditions, i.e. in shoals and in the presence of natural predators. The present results, however, also indicate that the above‐mentioned two angling methods may not be selective for certain behavioural types in comparison to each other.     

Predictions of establishment risk highlight biosurveillance priorities for invasive fish in New Zealand lakes

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Temporal and spatial macrofaunal community changes along a salinity gradient in seagrass meadows of Akkeshi-ko estuary and Akkeshi Bay,northern Japan

Changes in water clarity (secchi disc transparency) in relation to the presence/absence of introduced, exotic fish, including rudd (Scardinius erythrophthalmus), tench (Tinca tinca), perch (Perca fluviatilis), brown bullhead catfish (Ameiurus nebulosus), goldfish (Carassius auratus), and koi carp (Cyprinus carpio) were determined for 49 small, North Island, New Zealand lakes. There was a negative association between water clarity and the presence of exotic fish independent of lake depth. Moreover, a ‘before-and-after’ comparison and examination of case-studies indicated that introductions of exotic fish reduce water clarity. The number of species introduced affected the relationship between lake depth and water clarity but the specific role of each species could not be distinguished because most of the lakes (83%) contained more than one exotic fish species. A model incorporating the known mechanisms by which planktivorous, benthivorous and herbivorous fish can influence water clarity in lakes showed that control over just one species or feeding guild may not result in an improvement in water clarity because of the additive and synergistic effects of different species on lake trophic processes. Handling editor J. Cambray     

Predator-induced respiratory responses in juveniles of vendace Coregonus albula, whitefish C. lavaretus, perch Perca fluviatilis and roach Rutilus rutilus

Predator-induced respiratory responses of juvenile vendace, whitefish, perch and roach were studied in an intermittent-flow respirometer by creating visual contact between test fishes and predators (juvenile northern pike). Vendace and whitefish always responded to the presence of pike with increased oxygen consumption. The response of perch was the opposite: oxygen consumption decreased to near the standard respiratory level. Roach responded more variably: some individuals increased their oxygen uptake while others decreased it. Changes in the rate of respiration during exposure to predation are supposed to be caused by the differences in locomotory activity due to induced antipredator behaviour. According to their responses, these species could be grouped into escapers (coregonids), hiders (perch) or indeterminates (roach). The species-specific differences in the antipredator behaviour originate in morphological and physiological characteristics which together determine the effectiveness of a particular antipredator style (i.e. either hiding or escaping).     

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.     

Are behavioural traits in prey sensitive to the risk imposed by predatory fish?

1. Behavioural differences among prey species may result from evolutionary adaptations that facilitate coexistence with different predators and influence vulnerability to predators. It has been hypothesised that prey species modify their behaviour in relation to the risk posed by particular predators. 2. We examined the relationship between anti‐predator behaviour and predation risk in five species of larval odonates in combination with three predatory fish species (perch, gudgeon and rudd) that differ in foraging behaviour. The odonates, Platycnemis pennipes, Coenagrion puella, Lestes sponsa, Sympetrum striolatum and Libellula depressa, differ with regard to their life cycle and habitat, including water depth, occurrence in temporary ponds and co‐existence with fish. 3. The odonate species differed in their response to fish: (i) Two species showed a flexible response. Larval C. puella reduced activity in the presence of fish, regardless of species, whereas L. depressa altered their activity only in the presence of gudgeon. (ii) Independent of fish species, all odonates except L. depressa exhibited spatial avoidance of fish. This was interpreted as a more general anti‐predator response. (iii) In some cases the odonates showed no response to predators and their behaviour was thus independent of predation risk. 4. Our results confirm that all odonates responded to the presence of at least some predatory fish, and that some odonate species discriminated between fish species. However, we found no significant correlation between behavioural modifications and predation risk, indicating that anti‐predator responses and predation risk depend on the particular predator and the species being preyed on.     

Direct Estimation of Food Consumption of Juvenile Fish in a Shallow Inlet of the Southern Baltic

The species composition and feeding behaviour of fish larvae and juveniles was investigated in a shallow inlet of the Southern Baltic. The aim of the study was to examine if there was a predatory impact of young fish on the observed annual collapse of the copepod population. Dominant fish species were herring (Clupea harengus), perch (Perca fluviatilis) and roach (Rutilus rutilus). Roach larvae preferred shallower areas, whereas young herring and perch were found in open water. Highest consumption values were always observed in the evening. By computing complete evacuation of the digestive tract in approximately 3.5 hours, daily consumption in percent of fish fresh weight was 1.3, 4.7 and 5.5% for young herring, roach and perch, respectively. The impact of young fish on the dynamics of the population of the copepod species Eurytemora affinis is negligible.     

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

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

Correlated geographic variation in predation risk and antipredator behaviour within a wide‐ranging snake species (Notechis scutatus,Elapidae)

Geographic variation in antipredator behaviour within wide‐ranging species may be driven by both genetic and environmental influences. We quantified antipredator responses in neonatal (laboratory born, n = 555) and adult (field caught, n = 346) tiger snakes (Notechis scutatus) from 11 mainland and island sites in southern Australia. We used these data to test predictions from Bonnet et al.'s hypothesis that the vigour of antipredator responses in this species reflects behavioural plasticity (in turn, driven by an individual snake's exposure to predators during its lifetime) rather than by genetic variation in this trait. We used the number of predator taxa in each area as an index of predator risk. As predicted, adult snakes from predator‐rich areas had more vigorous defensive responses when handled, whereas neonatal behaviour (although also variable among populations) was unrelated to predator species richness. Adult males bit more readily than adult females (as expected from the greater predation exposure of males during mate searching) but no such sex difference was evident in neonates. Although alternative models remain possible, our data are most consistent with the hypothesis that geographic divergence in antipredator tactics within this species primarily reflects developmentally plastic responses to local predation risk.     

The influence of simulated aquatic macrophytes on the zooplankton consumption rate of juvenile roach, Rutilus rutilus, rudd, Scardinius erythrophthalmus, and perch, Perca fluviatilis

Laboratory experiments showed that, of zooplanktivorous roach, rudd and perch, in the absence of any environmental structure roach were the most efficient feeders, but high densities of simulated submerged marcrophytes elevated perch to this position; rudd matched the performances of perch and roach only at intermediate structure densities. These changes in efficiency rankings with increased structure were the result of overall decreases in the performances of roach and rudd. Simulated water lilies and emergent reeds had their own influences on consumption rate. These effects were largely mediated through relationships between structure density and swimming speed; roach and rudd, not perch, showed reduced swimming speeds in the more structured environments. Observations of the routine behaviour of the fish suggest that their assessment of predation risk may be an important factor in determining their activity level and hence foraging strategy.