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.     

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.     

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.     

Impact of black bullhead (Ameiurus melas) on turbidity in a diked wetland

Benthivorous fish in shallow, aquatic systems have been correlated with increased turbidity and declines in macrophyte production and wildlife use. Bullheads have been credited with increasing turbidity, but this has been seldom tested and has not been studied in a diked marsh. To assess the relationships of black bullhead (Ameiurus melas) and turbidity, we assembled mesocosmsin the Show Pool Management Unit of The Ottawa National Wildlife Refuge, OH, U.S.A. We stocked treatment enclosures with different biomasses of black bullhead at weekly intervals. Mean turbidity within treatment enclosures was significantly higher than within controls but remained lower than that of the open marsh. Both surface and bottom turbidity increased with adult and juvenile black bullhead biomass. Turbidity increased with fine sand concentration only in the presence of juvenile fish. Wind speed and direction were significant influences on the turbidity of the open marsh, but not within control enclosures. That treatment turbidity – even at extreme biomasses – remained significantly lower than the turbidity of the open marsh implicates fetch in having a greater influence on a marsh's turbidity than the presence of black bullhead. The greater impact of benthivorous fish on turbidity within shallow systems may be an indirect one through the destruction of macrophytes and subsequent destabilization of unconsolidated substrates.     

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.     

Environmental Conditions and Intraspecific Interference: Unexpected Effects of Turbidity on Pike (Esox lucius) Foraging

Interference among predators decreases per capita foraging rates and has implications for both community dynamics and top-down trophic processes. Interference originates from behavioural interactions among foragers, and these behaviours could be affected by environmental conditions. In experiments on pike foraging alone or among conspecifics in different levels of water turbidity, we expected high turbidity to decrease the perceived risk of intraspecific interactions among pike, and thereby decrease the strength of interference, as turbidity would decrease the visual contact between individuals and act as a refuge from behavioural interactions. The results show that this is not the case, but suggest that interference is induced instead of reduced in high turbidity. Per capita foraging rates do not differ between pike foraging alone or in groups in our clear and moderately turbid treatments, indicating no effect of interference. As high turbidity enhances prey consumption for pike individuals foraging alone, but does not have this effect for pike in groups, high turbidity induces the relative interference effect. We suggest that future evaluations of the stabilizing effects of interference on community dynamics and its reduction of predation impact on top-down trophic cascades should consider potential unexpected effects of environmental conditions.     

The effects of thermal effluent exposure on the gametogenesis of female fish

High temperature in Swedish and Lithuanian thermal effluent areas influenced gametogenesis of female perch Perca fluviatilis , roach Rutilus rutilus and pike Esox lucius negatively, indicating reduced reproductive capacity. Oocyte atresia started during vitellogenesis in autumn, and was often followed by asynchronous egg cell development. Among other anomalies, multi-nucleus oocytes and hermaphroditism were observed. No significant impact was seen in silver bream Blicca bjoerkna . Ruffe Gymnocephalus cernuus reacted by a tendency to produce an additional mature oocyte generation during the spawning period. Ovaries in roach from coastal areas were often infected by a microsporidian parasite Pleistophora mirandellae , causing severe damage to the gonad. Parasites were also detected in pike, but neither in perch nor in lake populations of roach. Fish living in open coastal environments did not avoid impact by moving out of the heated areas. There seems to be a conflict in some temperate fish between temperature preference behaviour and safeguarding normal reproduction.     

Growth of pike larvae: effects of prey, turbidity and food quality

We studied experimentally the effects of turbidity and prey composition on pike larval growth and hypothesized that pike larval growth varies with turbidity and food quality. We reared the first-feeding pike larvae (Esox lucius) in laboratory tanks with (1) clear or (2) turbid water provided with zooplankton rations from (3) an inner and (4) an outer archipelago site. The sites differ in physical features, salinity, eutrophication status, zooplankton community structure and density. Pike larvae showed the highest weight increase in clear water with zooplankton from the outer site and the poorest weight increase in turbid water with zooplankton as prey from the inner site. Our fatty acid analysis revealed that unsaturated fatty acid levels were highest in the outer site. The relative percentage of copepods was also higher in the outer site. This study supports the hypothesis that turbidity weakens the ability of pike larvae to capture certain prey. Further, zooplankton community composition matters in turbid water, but is not a primary factor in clear water.     

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.     

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.     

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.     

Biomass dependent interactions in pond ecosystems: responses of lower trophic levels to fish manipulations

The effects of mature benthivorous cyprinid fish and theirrecruitment on sediment resuspension, turbidity, phyto- andzooplankton, and benthic macroinvertebrates were studied in fourexperimental ponds. The ponds were stocked with bream (Abramisbrama L.), white bream (Blicca bjoerkna L.), roach (Rutilus rutilus L.) and wild carp (Cyprinus carpio L.) of3+–5+ age classes at standing crop biomass varying from 0 to500 kg ha-1. Cyprinids caused an increase in sedimentresuspension and in turbidity, in proportion to their biomass. Meancrustacean biomass did not significantly affect phytoplanktonbiomass due to intense grazing by fish during spring. Ponds withhigh fish stocks showed reduced midge biomass and vegetation coverand increased biomass of predatory invertebrates.     

Diversity of microbial communities of the intestinal mucosa and intestinal contents of fish from Lake Chany (Western Siberia)

The diversity of microbial communities associated with the intestinal mucosa and the intestinal contents of fish with different nutritional profiles have been investigated. Differences in the intestinal microbiota of fish with different dietary preferences have been revealed using group-specific primers for the main large taxa of bacteria. Representatives of the phyla Planctomycetes, Vernicomicrobia, and Cyanobacteria have been detected in the intestinal contents of all investigated nonpredatory (crucian carp, roach, and dace) and predatory (pikeperch, pike, and perch) fish. Representatives of the phylum Firmicutes were detected in the intestinal mucosa and intestinal contents of predatory fish species, but not in those of the nonpredatory fish.     

Foraging Trade-offs in Fathead Minnows (Pimephales promelas,Osteichthyes, Cyprinidae): Acquired Predator Recognition in the Absence of an Alarm Response

Pike-naive fathead minnows (Pimephales promelas) were fed ad libitum or deprived of food for 12, 24, or 48 h and then exposed to either conspecific alarm pheromone or distilled water and the odour of a predatory northern pike (Esox lucius). Minnows fed ad libitum or deprived for 12 h showed a stereotypic alarm response to the alarm pheromone (increased time under cover objects and increased occurrence of dashing and freezing behaviour); those deprived of food for 24 h showed a significantly reduced alarm response, while those deprived of food for 48 h did not differ significantly from the minnows exposed to a distilled water control. Upon subsequent testing in an Opto-Varimex activity meter, all groups initially exposed to alarm pheromone and pike odour exhibited an alarm response when exposed to pike odour alone. Those initially conditioned with distilled water and pike odour did nor show an alarm response to pike odour alone. These results demonstrate that there exists a significant trade-off between hunger level and predator-avoidance behaviour in fathead minnows and that minnows can learn the chemical cues of a predatory northern pike through association with alarm pheromone even in the absence of an observable alarm response.     

Comparison of losses of planktivorous fish by predation and seine-fishing in a lake undergoing long-term biomanipulation

SUMMARY 1. Piscivore stocking at artificially high densities and fishing are the two common approaches to reduce the amount of planktivorous and benthivorous fish in lake biomanipulation programmes. Both measures have advantages and disadvantages, but their relative efficacy has not previously been directly compared.
2. We calculated the average annual catch of roach and bream in a lake undergoing long-term biomanipulation (Feldberger Haussee, Germany) by seining each year between 1992 and 1998. We compared this value with a bioenergetics estimate of annual consumption rates of the dominant cohorts of piscivores, pikeperch and pike, in 1997 and 1998. We also determined species composition and length distribution of prey fish in stomachs of the piscivores.
3. Roach was the dominant prey species of both pikeperch and pike, whereas bream was rarely taken by either piscivorous species. Seining removed on average larger specimens of roach than were found in the stomachs of the piscivores.
4. Based on stocking densities of the piscivores, published mortality rates, and individual consumption rates, feeding of pikeperch and pike on roach exceeded the manual removal of roach by seining by a factor of 4–15 (biomass) in 1997 and 1998.
5. Based on these results, a combination of fishing and piscivore enhancement is recommended. Whereas the stocks of adult roach and bream have to be reduced mainly by fishing, the predation of piscivores should be directed predominantly towards the juvenile zooplanktivorous fish. Therefore, small size-classes of piscivorous fish should be promoted by fisheries management, including stocking and harvest regulations.     

Preferences for familiar fish do not vary with predation risk in the European minnow

In the near-natural conditions of a fluvarium in the River Frome, U.K., European minnows Phoxinus phoxinus displayed schooling preferences for familiar conspecifics. Schools were composed of 75% familiar fish. The minnows in this river co-occur with many piscivores, the most notable being the pike Esox Iucius . The preference for familiar shoalmates was not increased by the threat of predatory attack from a model pike, despite the antipredator benefits afforded to individual members of schools of familiar fish.     

Mercury content in muscles of fish of the Selenga River and lakes of its basin (Russia)

Differences between the muscle mercury contents in fish from lakes Gusinoye and Karasinoye and the Selenga River may indicate an inconsiderable income of mercury in aquatic ecosystems with atmospheric precipitation, along with a more intensive migration of mercury with the riverine flux. The minimal concentration of mercury (less than 0.15 mg/kg, dry weight) has been registered in the muscles of Amur sleeper from Lake Karasinoye and of Baikal omul from Lake Baikal, while the maximal concentration (1.0–2.3 mg/kg, dry weight) has been revealed in the fish caught in the Selenga River delta (in predatory pike, Amur catfish, and perch, as well as in omnivorous roach and ide). The muscle metal content in dace decreased along the downstream direction.     

Prey vulnerability to a gape-size limited predator: behavioural and morphological impacts on northern pike piscivory

The northern pike ( Esox lucius ) is a selective and important predator in lake ecosystems. Prey size in pike is limited by pike gape size, which is a linear function of pike body length. Here we show that the absolute gape-size limit in pike is greater than previously considered, and that maximum ingestible prey size is limited by prey body depth. Further, we experimentally show that pike prefer shallow-bodied roach before deeper-bodied common bream, and small prey sizes within each prey species. Handling time in pike increases with prey body depth, and since common bream are deeper-bodied than roach, handling time is longer for bream than for roach of the same length, but equal considering body depth. Prey handling time is suggested to be a major cost to the pike, since it increases the risk of losing the prey, as well as exposure to predation, kleptoparasitism and cannibalism. Consequently, prey vulnerability is determined by risk of predation and intraspecific interactions, and behavioural preferences in the pike, and not by pike gape-size limits. The consequences for natural populations is evaluated by analysing size structures of predator and prey fish populations in a eutrophic lake.     

Molecular and phenotypic evidence of a new species of genus Esox (Esocidae, Esociformes, Actinopterygii): the southern pike, Esox flaviae

We address the taxonomic position of the southern European individuals of pike, performing a series of tests and comparisons from morphology, DNA taxonomy and population genetics parameters, in order to support the hypothesis that two species of pike, and not only one, exist in Europe. A strong relationship emerged between a northern genotype supported by COI, Cytb, AFLP and specific fragments, and a phenotype with round spot skin colour pattern and a large number of scales in the lateral line, clearly separated from a southern genotype with other skin colour pattern and a low number of scales in the lateral line. DNA taxonomy, based on a coalescent approach (GMYC) from phylogenetic reconstructions on COI and Cytb together with AFLP admixture analysis, supported the existence of two independently evolving entities. Such differences are not simply due to geographic distances, as northern European samples are more similar to Canadian and Chinese samples than the southern Europe ones. Thus, given that the differences between the two groups of European pike are significant at the phenotypic, genotypic and geographical levels, we propose the identification of two pike species: the already known northern pike (Esox lucius) and the southern pike (E. flaviae n.sp.). The correct identification of these two lineages as independent species should give rise to a ban on the introduction of northern pikes in southern Europe for recreational fishing, due to potential problems of hybridisation.     

Turbidity decreases anti-predator behaviour in pike larvae, <Emphasis Type="Italic">Esox lucius</Emphasis>

Synopsis We tested how algal turbidity and light conditions influence anti-predator behaviour of first-feeding pike. Results showed that pike larvae were able to detect the predator by both chemical and visual signals in turbid water. However, the anti-predator behaviour was reduced in turbid water compared with clear water. Larvae hid in the vegetation in the presence of predator cues more in clear water than in turbid water. The attack rate on zooplankton in clear water was lower in the presence of predator cues, whereas no such difference was detected in turbid water. Both of these results indicate that turbidity acted as a refuge for larvae. Light proved to be an important regulating factor for feeding pike in the absence of predators, demonstrated as lowered attack rates in 50 light level in both clear and turbid water. This indicates that long-term turbidity may be critical for small larvae, which need to feed continuously to survive.