Overwinter changes in mass and lipid content of Perca fluviatilis and Gymnocephalus cernuus

The body condition, lipid reserves and mortality of 0 and 1 year‐old perch Perca fluviatilis and ruffe Gymnocephalus cernuus , sampled during the winter in Lake Constance, Germany, were compared. Length‐frequency analyses did not provide evidence for overwinter mortality in either species. The fresh and dry mass of perch as well as their lipid contents decreased during winter, while ruffe were heavier and contained more lipid at the end of the winter. The superior performance of ruffe was mainly attributed to its sensory capabilities, which allowed it to ingest zoobenthos throughout the winter, while the zooplankton feeding of perch was constrained by low light levels. In lakes that undergo a process of re‐oligotrophication, this advantage of ruffe over perch may be even more pronounced, since lower food supply during the growth season and thus lower fish lipid content at the start of winter is probably better tolerated by ruffe than by perch.     

The Effects of Zebra Mussels (Dreissena polymorpha) on the Foraging Success of Eurasian Perch (Perca fluviatilis) and Ruffe (Gymnocephalus cernuus)

Complex habitat structures can influence the foraging success of fish. Competition for food between fish species can therefore depend on the competitors' abilities to cope with structural complexity. In laboratory experiments, we comparatively assessed effects of zebra mussels (Dreissena polymorpha Pall .) on the foraging success of Eurasian perch (Perca fluviatilis L.) and ruffe (Gymnocephalus cernuus (L.)). In single‐species and mixed‐species experiments, the fish were fed caddisfly larvae (Tinodes waeneri (L.)) over complex (mussel‐covered stones) and less‐complex (bare stones) substrates. With intraspecific competition, food consumption by perch and ruffe decreased significantly when the complex substrate was used. With interspecific competition, food consumption by perch and ruffe did not change with substrate complexity, but perch clearly out‐competed ruffe on both substrates. Zebra mussel beds provide a refuge for macrozoobenthos against predation by ruffe and probably also by perch. (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Temperature-dependent differences in foraging ability of two percids,Perca fluviatilis and Gymnocephalus cernuus

Synopsis The effects of temperature on capture rate, handling time, capture probability and routine swimming performance were studied in two fish species, perch Perca fluviatilis and ruffe Gymnocephalus cernuus. In addition, the reaction distance (RD) of both species was measured. Handling time decreased and capture rate increased with increasing temperature for both perch and ruffe. The response of handling time and capture rate to temperature was more pronounced for perch than for ruffe. Routine swimming performance increased with increasing temperature for perch but not significantly so for ruffe. The laboratory study showed that the foraging ability of ruffe was less sensitive to temperature than it was for perch. These results were consistent with the vertical distributions of both species in thermally stratified lakes; ruffe, which occurred at all depths, utilized a wider temperature range than perch.     

The ecology of ruffe,Gymnocephalus cernuus (Pisces: Percidae) introduced to Mildevatn,western Norway

Synopsis Diet, habitat use, diel and seasonal activity and a number of population parameters were studied on ruffe,Gymnocephalus cernuus, introduced to Mildevatn, western Norway. This lake is sited outside the natural range of the ruffe and has a lower fish diversity and a different fish species composition than within its native range. From June through September the ruffe was planktivorous and mainly caught at 4 to 6 m depth in the benthic zone. At other times of year ruffe was feeding on zoobenthos and caught deeper in the benthic zone. Ruffe was mainly day active. Zooplankton feeding during summer is the clearest difference compared to ruffe populations living within its natural range. Presence of large zooplankton organisms available for ruffe is suggested as the main reason for the difference found in food choice. The availability of large zooplankton is probably due to community structure caused by a predator and lack of interspecific competition for zooplankton in the deeper parts of the lake. Piscivorous brown trout.Salmo trutta, restrict the habitat of threespined stickleback,Gasterosteus aculeatus, to the zone of littoral vegetation, allowing high densities of larger zooplankton species likeBythotrephes longimanus to be present in the lake. Brown trout is present only in the upper light and well oxygenated parts of the lake, leaving a refuge for the ruffe, where they can feed on the rich zooplankton community.     

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.     

A trophic pathway from biogenic methane supports fish biomass in a temperate lake ecosystem

Although some primary consumers such as chironomid larvae are known to exploit methane‐derived carbon via microbial consortia within aquatic food webs, few studies have traced the onward transfer of such carbon to their predators. The ruffe Gymnocephalus cernuus is a widespread benthivorous fish which feeds predominantly on chironomid larvae and is well adapted for foraging at lower depths than other percids. Therefore, any transfer of methanogenic carbon to higher trophic levels might be particularly evident in ruffe. We sampled ruffe and chironomid larvae from the littoral, sub‐littoral and profundal areas of Jyväsjärvi, Finland, a lake which has previously been shown to contain chironomid larvae exhibiting the very low stable carbon isotope ratios indicative of methane exploitation. A combination of fish gut content examination and stable isotope analysis was used to determine trophic linkages between fish and their putative prey. Irrespective of the depth from which the ruffe were caught, their diet was dominated by chironomids and pupae although the proportions of taxa changed. Zooplankton made a negligible contribution to ruffe diet. A progressive decrease in δ¹³C and δ¹⁵N values with increasing water column depth was observed for both chironomid larvae and ruffe, but not for other species of benthivorous fish. Furthermore, ruffe feeding at greater depths were significantly larger than those feeding in the littoral, suggesting an ontogenetic shift in habitat use, rather than diet, as chironomids remained the predominant prey item. The outputs from isotope mixing models suggested that the incorporation of methane‐derived carbon to larval chironomid biomass through feeding on methanotrophic bacteria increased at greater depth, varying from 0% in the littoral to 28% in the profundal. Using these outputs and the proportions of littoral, sub‐littoral or profundal chironomids contributing to ruffe biomass, we estimated that 17% of ruffe biomass in this lake was ultimately derived from chemoautotrophic sources. Methanogenic carbon thus supports considerable production of higher trophic levels in lakes.     

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

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

Some like it deep: Intraspecific niche segregation in ruffe (Gymnocephalus cernua)

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Changes in reproductive strategy in the ruffe during a period of establishment in a new habitat

The pattern of maturation, body size and fecundity was examined in a population of ruffe ( Gymnocephalus cernuus L.) three times during a period of rapid growth, and eventual stabilization, following its introduction to a new habitat. When the ruffe were less common, maturing ruffe were relatively large and immature ruffe relatively small, compared with when the ruffe were abundant. Intermediate ruffe population size showed a maturation pattern intermediate between these two extremes. It is suggested that this pattern of maturation is a response of the ruffe population to changing growth opportunity induced by changing intraspecific competition. This fluctuating maturation pattern is interpreted in terms of a threshold-dependent maturation trigger, operating on the rate of accumulation of energy and a trade-off between somatic growth and gonad development. When the ruffe population was large, high intraspecific competition resulted in low opportunity for growth; only fish with the highest rate of food acquisition were able to mature in a given year–the investment in gonadal tissue reducing somatic growth. When the ruffe population was low, the high rate of energy acquisition in the population resulted in the triggering of maturation, even at small size, only the very smallest fish remaining immature. High growth opportunity allowed maturing fish to develop gonad and maintain somatic growth. The pattern of size related fecundity also changed over the three periods. When growth opportunity was low, size related fecundity was greater than when opportunity for growth was high. This suggests that maturing females faced with poor growth conditions compensated by increasing egg number for a given body size either by decreasing egg size or by increasing total investment in ovarian tissue.     

Temperature dependency of cupular mechanics and hair cell frequency selectivity in the fish canal lateral line organ

The mechanical frequency selectivity of the cupula located in the supraorbital lateral line canal and the frequency selectivity of the hair cells driven by the cupula were measured simultaneously in vivo. Laser interferometry was used to measure cupular mechanics and extracellular receptor potentials were recorded to determine hair cell frequency selectivity. Results were obtained from two teleost fish species, the ruffe (Acerina cernua L.), a European temperate zone freshwater fish, and the tropical African knife fish (Xenomiystus nigri). In both species cupular displacement grows with increasing frequency of canal fluid displacement, reaching a maximum at 115 Hz in the ruffe and at 460 Hz in the African knife fish. Cupular best frequencies were independent of temperature. Cut-off frequencies of hair cell frequency selectivity were found to depend on temperature with a Q10 of 1.75, ranging from 116 Hz (4 degrees C) to 290 Hz (20 degrees C), as established in the ruffe. At normal habitat temperatures of the two fish species (ruffe, 4 degrees C; African knife fish, 28 degrees C), this results in hair cell cut-off frequencies that match the two different cupular best frequencies remarkably well. This match suggests adjusted signal transfer in these two peripheral stages of canal lateral line transduction.     

Collapse of the native ruffe (<Emphasis Type="Italic">Gymnocephalus cernua</Emphasis>) population in the Biesbosch lakes (the Netherlands) owing to round goby (<Emphasis Type="Italic">Neogobius melanostomus</Emphasis>) invasion

We investigated the change in benthic fish communities in three artificial lakes of the Biesbosch area in the Netherlands between two time periods: before and after the invasion of round goby (Neogobius melanostomus). Native ruffe (Gymnocephalus cernua), the dominant species in benthic gillnet and littoral beach seining catches before the invasion, almost completely disappeared in all lakes only 2 years after the invasion. We found a significant increase in 0 + perch (Perca fluviatilis) and, in some lakes, pikeperch (Sander lucioperca) abundance in gillnet catches after invasion. In the post-invasion period, the 0 + fish community was dominated by perch, and the older fish community was dominated by round goby. The species richness of 0 + fish increased in the post-invasion period owing to the invasion of gobiids. However, it did not change for older fish between periods. Our results clearly show that, owing to a similar benthic lifestyle and high niche overlap, ruffe was the only species negatively influenced by the round goby invasion. The competitive superiority of round goby over ruffe is so strong that the once-dominant species of the overall benthic fish community collapsed after only a few years of coexistence.     

Predicting habitat use and trophic interactions of Eurasian ruffe,round gobies,and zebra mussels in nearshore areas of the Great Lakes

The Laurentian Great Lakes have been subject to numerous introductions of nonindigenous species, including two recent benthic fish invaders, Eurasian ruffe (Gymnocephalus cernuus) and round gobies (Neogobius melanostomus), as well as the benthic bivalve, zebra mussel (Dreissena polymorpha). These three exotic species, or “exotic triad,” may impact nearshore benthic communities due to their locally high abundances and expanding distributions. Laboratory experiments were conducted to determine (1) whether ruffe and gobies may compete for habitat and invertebrate food in benthic environments, and (2) if zebra mussels can alter those competitive relationships by serving as an alternate food source for gobies. In laboratory mesocosms, both gobies and ruffe preferred cobble and macrophyte areas to open sand either when alone or in sympatry. In a 9-week goby–ruffe competition experiment simulating an invasion scenario with a limited food base, gobies grew faster than did ruffe, suggesting that gobies may be competitively superior at low resource levels. When zebra mussels were added in a short-term experiment, the presence or absence of mussels did not affect goby or ruffe growth, as few zebra mussels were consumed. This finding, along with other laboratory evidence, suggests that gobies may prefer soft-bodied invertebrate prey over zebra mussels. Studies of interactions among the “exotic triad”, combined with continued surveillance, may help Great Lakes fisheries managers to predict future population sizes and distributions of these invasive fish, evaluate their impacts on native food webs, and direct possible control measures to appropriate species.     

Stock status and current tendencies in the population dynamics of commercial fishes of Pskov-Chudskoe water body

Dynamics of eight commercial populations of fish from Pskov-Chudskoe Lake forming the main bulk (up to 80%) of annual catch was analyzed using an ADAPT-VPA model making it possible to reconstruct the dynamics of the total stock biomass and fishing mortality. The populations of five species (European smelt Coregonus albula, whitefish C. lavaretus maraenoides, perch Perca fluviatilis, roach Rutilus rutilus, ruffe Gymnocephalus cernuus) in the present period demonstrate a tendency for a decrease in biomass while the stocks of two species (bream Abramis brama and zander Sander lucioperca) beginning from the 1980s considerably increased. A particularly drastic (multifold) increase in biomass is recorded in zander population. The causes of the observed population changes are related mainly to ecosystemic transformations taking place in the water body; only with respect to two species-perch and roach-can one state that the decrease in stocks is determined in part by the fishery impact. The increase in zander population is, in all probability, the result of the combined effects of eutrophication forming favorable conditions for the feeding and survival of its juveniles and fishery that led to the decrease in the biomass of one of its main food competitors-perch. The drastic increase in zander numbers in turn promoted the decrease in the biomass of species being objects of its feeding.     

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.     

Dispersal of the flea Ctenophyllus hirticrus and spreading of plague epizooties in Gorny Altai

Gradual dispersion of an abundant flea species Ctenophyllus hirticrus specific to the Pallas's pika (the main plague carrier), is revealed in the Gorno-Altai natural plague focus on the territory, occupied by two populations of this lagomorph. Spreading of Yersinia pestis in these areas took place a short time later the rise of this ectoparasite's abundance. It is supposed that the colonization of these areas by C. hirticrus was one of the factors determined epizooties spreading within the focus and formation of new sites of stable Y. pestis preservation.     

Predator odor recognition and antipredatory response in fish: does the prey know the predator diel rhythm?

We studied in a laboratory experiment using stream tanks if two percid prey fish, the perch (Perca fluviatilis) and the ruffe (Gymnocephalus cernuus), can recognize and respond to increased predation risk using odors of two piscivores, the pike (Esox lucius) and the burbot (Lota lota). Burbot is night-active most of the year but pike hunts predominantly visually whenever there is enough light. Perch is a common day-active prey of pike and dark-active ruffe that of burbot. We predicted that besides recognizing the predator odors, the prey species would respond more strongly to odors of the predator which share the same activity pattern. Both perch and ruffe clearly responded to both predator fish odors. They decreased movements and erected the spiny dorsal fins. Fin erection showed clearly the black warning ornamentation in the fin and thus erected fin may function besides as mechanical defense also as warning ornament for an approaching predator. No rapid escape movements were generally observed. Both perch and ruffe responded more strongly to pike odor than to burbot. There were no clear differences in response between day and night. In conclusion, we were able to verify clear predator odor recognition by both prey fish. Both perch and ruffe responded to both predator odors and it seemed that pike forms a stronger threat for both prey species. Despite of diel activity differences both perch and ruffe used the same antipredatory strategies, but the day-active perch seemed to have a more flexible antipredatory behavior by responding more strongly to burbot threat during the night when burbot is active.     

The fish community structure in Tjeukemeer in relation to fishery and habitat utilization

The fish community in Tjeukemeer was monitored from 1971 to 1988 by trawling, and during the summer of 1988 the distribution of the more abundant species was determined in relation to size and location. Bream, pikeperch and smelt were the most dominant fishes, whereas roach, white bream, perch and ruffe comprised <10% of the total catches. After the termination in 1977 of the intensive gillnet fishery both pikeperch >50 cm and bream >30 cm increased in biomass, but roach >15 cm and perch >15 cm virtually disappeared and pikeperch <50 cm substantially decreased. Only smelt, ruffe, white bream and bream <30 cm hardly changed in biomass. Bream, pikeperch, perch and smelt were restricted to the open water zone, but roach <15 cm and 0 + bream were confined to the littoral zone. White bream and ruffe did not show a distinct habitat preference. Because the recruitment of smelt is largely dependent on immigration from the IJsselmeer, a feed-back between the smelt and pikeperch population is lacking. Because the carrying capacity of the pikeperch population is mainly determined by smelt, the other fish <15 cm are very vulnerable to predation when the smelt population is consumed, before a new year-class of smelt is recruited, or when the smelt fails to recruit. The vulnerability to predation of the different species and their feeding habits are discussed in relation to their distribution.     

Growth,Reproduction and Condition of Roach (Rutilus rutilus (L.)), Perch (Perca fluviatilis,L.) and Ruffe (Gymnocephalus cernuus (L.)) in Eutrophic Lake Aydat (France)

The growth, reproduction and condition of adults of the three dominant fish species (roach, Rutilus rutilus, (L.); perch, Perca fluviatilis, L. and ruffe, Gymnocephalus cernuus, (L.)) in the eutrophic Lake Aydat were studied over one year cycle. Compared to published data, the growth of R. rutilus and G. cernuus was about average, while that of P. fluviatilis was below average. Comparing to literature, the fecundity of R. rutilus and G. cernuus was low but their oocytes were large. In contrast, P. fluviatilis had a high fecundity but small oocytes. At the end of summer, an abrupt decrease in the condition was recorded only for perch, probably due to stress as a result of environmental conditions. The sex-ratio was in favour of females for the three studied species but, in contrast to G. cernuus, the sex-ratio of R. rutilus and P. fluviatilis increased significantly also with age. It is concluded that Lake Aydat is a more favourable environment for R. rutilus and G. cernuus than for P. fluviatilis.     

Competition between perch (Perca fluviatilis) and ruffe (Gymnocephalus cernuus): the advantage of turning night into day

1. The outcome of interspecific competition for food resources depends both on the competitors’ sensory abilities and on environmental conditions. In laboratory experiments we tested the influence of daylight and darkness on feeding behaviour and specific growth rate (SGR) of two species with different sensory abilities. 2. We used perch (Perca fluviatilis) as a visually orientated, and ruffe (Gymnocephalus cernuus) as a mechano‐sensory oriented predator and tested their growth rates and behaviour under conditions of interspecific and intraspecific competition. Three different foraging conditions were used: food supplied (i) only during the day, (ii) only during the night or (iii) during both day and night. 3. In perch neither SGR nor feeding behaviour were influenced substantially by interspecific competition during daylight. During darkness their foraging behaviour changed markedly and their access to the food source as well as their SGR were negatively affected by the presence of ruffe. 4. Ruffe's foraging behaviour did not change during either day or night with interspecific competition. During the night ruffe's SGR was higher with interspecific competition, probably because of a release from intraspecific competition and the competitive inferiority of perch during the night. 5. Because of its seonsory abilities ruffe feeds predominantly at night, thereby reducing competitive interference from perch.     

Nutrient release and resuspension generated by ruffe (Gymnocephalus cernuus) and chironomids

1. Benthivorous fish may play an important role in internal nutrient loading. Ruffe are highly specialised, feeding exclusively on bottom animals; thus all nutrients released via their feeding are derived from the bottom and are new to the water column. The fish can also release nutrients from the sediment through resuspension while searching for food. 2. The aim of this study was to estimate experimentally in the laboratory the effect on water quality of resuspension and nutrient release by ruffe and bottom animals (chironomids). 3. Ruffe released nutrients during 8 h experiments as follows: total P 1.4, dissolved PO₄ 0.6, total N 24.0 and NH₄‐N 15.9 μg g^?1 WW h^?1. A decreasing trend in mass‐specific release was observed over time, probably because of starvation. The mass‐specific release of total N and NH₄‐N decreased as the mean weight of fish increased. The mean ratio of excreted N : P was 32. 4. In 26 h experiments with sediment and both ruffe and chironomids, ruffe increased nutrient concentrations and turbidity values significantly but chironomids had an effect only on turbidity. Neither ruffe nor chironomids affected the ratio of inorganic N : P concentrations. An interaction between ruffe and chironomids was found for turbidity. 5. According to these results, benthivorous fish may increase nutrient concentrations in the water column and need to be taken into account when estimating internal loading.