Diet shifts and food selection of perch Perca fluviatilis and roach Rutilus rutilus in humic lakes of varying water colour

The field data from four humic lakes suggested that water colour may have both direct and indirect effects on inter‐ and intra‐specific interactions of perch Perca fluviatilis and roach Rutilus rutilus. The results agree with suggestions that, compared with R. rutilus, P. fluviatilis may be an inferior forager on zooplankton in highly coloured water. As an indirect effect, water colour decreases the coverage of macrophytes and limits suitable littoral habitats, benefiting R. rutilus over P. fluviatilis. Perca fluviatilis benefiting from complex habitats does not have the advantage in macrophyte‐poor highly coloured water.     

Behavioral differences of various ecological groups of roach Rutilus rutilus and perch Perca fluviatilis

The intrapopulation morphological and behavioral polymorphism of roach (Rutilus rutilus) and perch Perca fluviatilis) was studied. It was shown that marker indices for distinguishing intrapopulation groups are the size of the mouth opening and the index of the body shape in roach and the coefficient of pigmentation in perch. Based on analysis of the data of indices, it was found that there are two ecological groups—coastal and pelagic in populations of roach and perch. Several behavioral differences in individuals of various ecological groups were revealed. Comparison of the defense reaction of perch individuals belonging to various ecological groups was performed. The assessment of the effects of hydrodynamic activity on the survival of perch individuals belonging to various ecological groups was made. Differences in the swimming capacity of juvenile roach developing under conditions of biotopes of various types were established. Suggestions were made on the mechanisms of formation of intrapopulation groups of roach and river perch.     

Ergasilid copepods as parasites of perch Perca fluviatilis and roach Rutilus rutilus in Central Finland: seasonality, maturity and environmental influence

A total of 1255 roach Rutilus rutilus (L.) and 866 perch Perca fluviatilis (L.) from four interconnected lakes in Central Finland differing in trophic status and pollution level were studied for parasitic ergasilid copepods between August 1985 and December 1988. In addition, 109 whitefish ( Coregonus sp.) were studied from one of the lakes.
Four ergasilid species were found: (the prevalence and intensity/fish, respectively, for the whole material are given in parentheses) Ergasilus briani (16·9%, 0·5), Neoergasilus japonicus (15·6%, 0·4) and Paraergasilus longidigitus (2·1%, 0·02) on the roach and Ergasilus sieboldi (9·9%, 0·1) and P. longidigitus (4·9%, 0·05) on the perch. Logit analysis was used to assess the significance of the year, the lake and the season for the prevalence of infection. Generally, the prevalence of E. briani on roach depended on the season only, with higher prevalences in summer. That of E. sieboldi on perch was mainly dependent on the lake, being most frequent in the oligotrophic lake. Both lake, year and season affected the prevalence of N. japonicus in roach. The more complicated system for N. japonicus was explained by the long duration of its free-living stage, its loose attachment and different location on the host, being found on the fins whereas E. briani and E. sieboldi were on the gills.
All species were found throughout the year, and age had no clear influence on infection. The occurrence of egg-sacs and length distributions of the copepods suggest that there are two generations a year, e.g. an overwintering E. briani generation which produces offspring in June to attack the fish in July, and a second generation which attacks the fish from August onwards. Whitefish were not infected by these parasites, only one fish being found which carried a single individual of E. sieboldi.     

The interactions of abiotic and biotic factors influencing perch Perca fluviatilis and roach Rutilus rutilus populations in small acidified boreal lakes

Four small, acidified boreal lakes, all sustaining populations of perch Perca fluviatilis, roach Rutilus rutilus and pike Esox lucius, were studied in four successive years. Three lakes were moderately acidified (mean pH of 5·61-5·83), while the fourth was more acidic (mean pH of 5·16) and had a sparse population of R. rutilus. Perca fluviatilis density was higher in this lake (1004 ha(-1)) than in the other three (355-717 ha(-1)), where R. rutilus dominated in terms of numbers (981-2185 ha(-1)). Large, potentially predatory, P. fluviatilis were most abundant in the lake with clearest water, and these seemed to have a negative effect on P. fluviatilis density. Perca fluviatilis mean mass was negatively correlated with R. rutilus biomass and was highest in the most acidic lake with the sparse R. rutilus and the highest P. fluviatilis density. Perca fluviatilis mass correlated positively with pH in two lakes (with the highest fish biomass), suggesting that low pH affected P. fluviatilis mass negatively. Perca fluviatilis growth correlated positively with summer (July to August) air temperature in the lake with sparse R. rutilus, thus differing from P. fluviatilis and R. rutilus growth in the other three lakes. The mean age of P. fluviatilis was generally lower than that of R. rutilus and was lowest in the two lakes with the highest fish biomass, indicating that adult mortality was affected by density-induced factors.     

Effect of temperature and roach Rutilus rutilus group size on swimming speed and prey capture rate of perch Perca fluviatilis and R. rutilus

Effects of temperature and group size of roach Rutilus rutilus on foraging behaviour of perch Perca fluviatilis and R. rutilus were tested in two laboratory experiments. A temperature experiment with P. fluviatilis and R. rutilus in aquaria (with either one P. fluviatilis or two R. rutilus) was tested at five temperatures: 4, 8, 12, 16 and 20° C, and showed that P. fluviatilis had a lower swimming speed and capture rate than R. rutilus, especially at 4 and 8° C. The effect of group size was tested at four R. rutilus abundances: 0, 2, 4 and 6, all at 16° C, and revealed that swimming speed and capture rate of P. fluviatilis were lowest at the highest R. rutilus abundance, whereas R. rutilus was relatively unaffected. Perca fluviatilis occupied positions closer to the bottom than R. rutilus, especially when feeding, and this tendency was reinforced at the highest roach abundance.     

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).     

Population responses of perch (Perca fluviatilis) and roach (Rutilus rutilus) to recovery from acidification in small Finnish lakes

Decreasing trends in atmospheric emissions and acidic deposition during the 1990s have resulted in chemical recovery from acidification in the sensitive surface water systems of southern Finland. Responses of perch and roach populations to the improved water quality were studied in 30 small lakes with the aid of water chemistry monitoring data gathered in 1987–2002 and the data collected from two consecutive periods of gillnet test fishing, 1985–1988 and 2001–2002. In the most acidified lakes, alkalinity and ANC have increased and sulphate and labile aluminium concentrations decreasæed markedly. The response of perch populations to the improved water quality is seen in improved reproduction success, indicated by a higher CPUE in numbers and a lower mean weight. The growth rate of perch has declined as the population density has increased. Roach populations have not recovered in the same way as perch, there being no major changes in NPUE or mean weight. Lower growth rates were, however, observed in the roach populations of all study lakes. The increased perch population density as an obstacle to the recovery of roach populations is discussed. Despite the chemical and biological recovery of the study lakes, the buffer capacity of many headwater lakes is low and the lakes will be sensitive to any increases in acidic deposition in the future.     

Environmental factors regulate the effects of roach Rutilus rutilus and pike Esox lucius on perch Perca fluviatilis populations in small boreal forest lakes

In this study of 18 small boreal forest lakes, the effects of abiotic and biotic factors (roach Rutilus rutilus and pike Esox lucius) on various population variables of perch Perca fluviatilis were examined. As a single variable, the gillnet catch per unit effort (CPUE) of R. rutilus was negatively related to the mean mass of small (< 200 mm) and the growth rate of young (1–2 years) P. fluviatilis. The mean mass of large (≥ 200 mm) P. fluviatilis was the highest at intermediate CPUE of R. rutilus. Redundancy analysis including environmental factors and P. fluviatilis population variables suggested that ‘predation–productivity–humus' gradient affected P. fluviatilis populations by decreasing the CPUE and mean mass of small individuals but increasing these variables of large individuals. The CPUE of R. rutilus and lake area had a negative effect on small and a positive effect on large P. fluviatilis growth rate. In small boreal forest lakes, P. fluviatilis populations are affected by the partially opposite forces of competition by R. rutilus and predation by E. lucius, and the intensity of these interactions is regulated by several environmental factors.     

Photoreceptors and eyes of pikeperch Sander lucioperca,pike Esox lucius,perch Perca fluviatilis and roach Rutilus rutilus from a clear and a brown lake

The photoreceptors and eyes of four fish species commonly cohabiting Fennoscandian lakes with different light transmission properties were compared: pikeperch Sander lucioperca, pike Esox lucius, perch Perca fluviatilis and roach Rutilus rutilus. Each species was represented by individuals from a clear (greenish) and a humic (dark brown) lake in southern Finland: Lake Vesijärvi (LV; peak transmission around 570 nm) and Lake Tuusulanjärvi (LT; peak transmission around 630 nm). In the autumn, all species had almost purely A2-based visual pigments. Rod absorption spectra peaked at c.526 nm (S. lucioperca), c. 533 nm (E. lucius) and c. 540 nm (P. fluviatilis and R. rutilus), with no differences between the lakes. Esox lucius rods had remarkably long outer segments, 1.5–2.8-fold longer than those of the other species. All species possessed middle-wavelength-sensitive (MWS) and long-wavelength-sensitive (LWS) cone pigments in single, twin or double cones. Rutilus rutilus also had two types of short-wavelength sensitive (SWS) cones: UV-sensitive [SWS1] and blue-sensitive (SWS2) cones, although in the samples from LT no UV cones were found. No other within-species differences in photoreceptor cell complements, absorption spectra or morphologies were found between the lakes. However, E. lucius eyes had a significantly lower focal ratio in LT compared with LV, enhancing sensitivity at the expense of acuity in the dark-brown lake. Comparing species, S. lucioperca was estimated to have the highest visual sensitivity, at least two times higher than similar-sized E. lucius, thanks to the large relative size of the eye (pupil) and the presence of a reflecting tapetum behind the retina. High absolute sensitivity will give a competitive edge also in terms of short reaction times and long visual range.     

A survey of the spawning of perch (Perca fluviatilis), pike (Esox lucius), and roach (Rutilus rutilus), using artificial spawning substrates in lakes

The development of spawning in perch, pike and roach in Lake Geneva has been studied by means of artificial spawning substrates, laid at different depths, from 1984 to 1993. In Lake Geneva, perch spawned in May. A rise of surface water temperature up to 14 °C stimulated spawning activity while bad weather (surface temperature at 10 °C) induced a spread of the spawning period over more than one month. The spawning period was delayed in years when the mean width of perch egg-ribbons was the largest; this corresponded to the biggest females. At the beginning of the spawning period (early in May), perch preferentially chose a depth of 4 m to spawn. In contrast, at the end of the spawning period, maximum spawning intensity was observed at a depth of 12 m. This phenomenon was more pronounced when water temperature rose above 14 °C in the top 4 m while it remained below 12 °C at a depth of 12 m. Pike spawned at the end of April and at the beginning of May in Lake Geneva. They preferred spruce branches among the different spawning substrates that we tested in Lake Vouglans. When the water temperature increased at the surface of Lake Geneva, pike preferred to lay their eggs on substrates set at 3 m depth where the temperature was cooler than in the surface layer (10.5 °C vs 14 °C). Roach spawned during the last two weeks of May or during the first two weeks of June in Lake Geneva, depending on water temperature. Spawning had generally been spread over a week, but a sudden decrease of water temperature could slow spawning intensity. Roach were able to lay their eggs on natural or synthetic substrates, located 0.5 m below the surface near the shore as well as, at a distance of several hundred metres from the shore. The survival of eggs was always above 90% for perch and roach and generally above 70% for pike.     

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.     

Genetic and morphological divergence along the littoral–pelagic axis in two common and sympatric fishes: perch,Perca fluviatilis (Percidae) and roach,Rutilus rutilus (Cyprinidae)

Individuals are constantly in competition with one another and, on both ecological and evolutionary timescales, processes act to reduce this competition and promote the gain of fitness advantages via diversification. Here we have investigated the genetic (AFLP) and morphological (geometric morphometrics) aspects of the littoral–pelagic axis, a commonly observed resource polymorphism in freshwater fishes of postglacial lakes. We found a large degree of variation in the genetic and morphological divergence between littoral and pelagic perch and roach across Swedish lakes. Although there was evidence of assortative mating (elevated kinship values) in both species, we could not find any significant coupling of morphology and genetic divergence. Instead, there was evidence that the extent of resource polymorphism may be largely caused by phenotypic plasticity. These results suggest that assortative mating, which can lead to genetically determined adaptive divergence, does occur in these species, particularly perch, but not according to genetically fixed morphological traits. The behavioural mechanisms facilitating associative mating need to be investigated to explore the interaction between phenotypic plasticity and adaptive genetic divergence and their roles in diversification. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 929–940.     

Antipredator behaviour of 0+ year Perca fluviatilis: effect of vegetation density and turbidity

In this study, the combined influence of vegetation density and water turbidity on habitat utilization of a prey fish, 0+ year perch Perca fluviatilis , under predation risk (pike, Esox lucius ) was investigated. The vegetated habitat was overall preferred over the open habitat in the presence of a predator. The level of turbidity, and to a lesser extent vegetation density, however, influenced the response of 0+ year perch. The use of the vegetated habitat was lower in very turbid than in clear and turbid conditions, suggesting reduced antipredator behaviour in very turbid water. The effect of vegetation density on antipredator behaviour was only present in clear water, where the use of a structural refuge decreased with increasing vegetation density. No such effect was observed in turbid and very turbid water. The results showed that the structuring role of vegetation or habitat complexity may diminish with increased turbidity. The observed masking effect of turbidity suggests that predator‐prey interactions in vegetated habitats are more complex than what has generally been thought.     

Consumption, growth and respiration of bleak, Alburnus alburnus (L.), and roach, Rutilus rutilus (L.), during early ontogeny

Three components of the energy budget, consumption ( C ), production ( P ) and respiration ( R ) in juvenile roach and bleak kept under controlled food ( Anemia salina , 2400 ind. l⁻¹) and temperature (20° C) conditions were measured in a study aimed at defining differences between the two species and elucidating the patterns of energy partitioning during ontogeny.
Daily food consumption rates (J day⁻¹ fish⁻¹) increased allometrically ( C =a W ^b) with body size ( W , mg dry weight) in both species. Covariance analysis indicated no differences in slope or intercept for the two regression lines ( P ≤ 0.05, n = 82). However, the two species grew at significantly different rates, roach faster than bleak.
The dependence of the respiration rate (μmol h⁻¹ fish ⁻¹) on body weight ( W ) can be described by an allometric function: R = a W ^b, where a ± 95% C.L. = 0.17 ± 0.15 for roach and 0.18 ± 0.20 for bleak. The slope for roach (b ± 95% C.L. = 0.78 ± 0.01) is slightly higher than that for bleak (0.69 ± 0.03).
Assimilation efficiency [AE = ( P + R ) C ⁻¹] was significantly higher in roach than in bleak. Different levels of AE correlated with differences in relative gut length (gut length as percentage of body length). Due to the shorter relative gut length above a weight of 5 mg, bleak has lower powers of digestion, which may explain lower production rates. These differences in energetic performance between the two species indicate mechanisms leading to niche differentiation in the early life history of the fishes.     

Allopatric and sympatric diversification within roach (Rutilus rutilus) of large pre‐alpine lakes

Intraspecific differentiation in response to divergent natural selection between environments is a common phenomenon in some northern freshwater fishes, especially salmonids and stickleback. Understanding why these taxa diversify and undergo adaptive radiations while most other fish species in the same environments do not, remains an open question. The possibility for intraspecific diversification has rarely been evaluated for most northern freshwater fish species. Here, we assess the potential for intraspecific differentiation between and within lake populations of roach (Rutilus rutilus)—a widespread and abundant cyprinid species—in lakes in which salmonids have evolved endemic adaptive radiations. Based on more than 3,000 polymorphic RADseq markers, we detected low but significant genetic differentiation between roach populations of two ultraoligotrophic lakes and between these and populations from other lakes. This, together with differentiation in head morphology and stable isotope signatures, suggests evolutionary and ecological differentiation among some of our studied populations. Next, we tested for intralacustrine diversification of roach within Lake Brienz, the most pristine lake surveyed in this study. We found significant phenotypic evidence for ecological intralacustrine differentiation between roach caught over a muddy substrate and those caught over a rocky substrate. However, evidence for intralacustrine genetic differentiation is at best subtle and phenotypic changes may therefore be mostly plastic. Overall, our findings suggest roach can differ between ecologically distinct lakes, but the extent of intralacustrine ecological differentiation is weak, which contrasts with the strong differentiation among endemic species of whitefish in the same lakes.     

Shifts in the microhabitat of larval and juvenile roach, Rutilus rutilus (L.), in a floodplain channel

The microhabitat of roach larvae (stages L1–L6) and 0+ juveniles was examined in a braided channel (Ilon) of the Upper Rhone River, France, from weekly samples (spring-autumn 1985) using Point Abundance Sampling by electrofishing. During early development, microhabitat exploitation was characterized by two transitions: the first at larval step L3, when morphological and physiological attributes permit the active choice of microhabitat; the second at the end of the larval and start of the juvenile period. At larval step L3, and up to L5, roach were strongly associated with lentic waters of medium depth (0.5–1.0m) and with either macrophytes or ligneous debris. The L6 and 0+ juveniles were found more often than expected in shallower waters (< 0.2m, 0.2–0.5 m) and were no longer associated with dense vegetation or ligneous debris. However, both larval (L3–L6) and 0+ juvenile roach preferred silty sand and/or silted gravel, and a lack of current. This shift, from moderately deep waters with vegetation and/or branches as larvae to open shallow waters as juveniles, demonstrates the young roach's perception and response to environmental change, with shallows exploited as a refuge from predation when protective structures become scarce or unavailable.     

Diet composition and prey preference of tench, Tinca tinca (L.), common bream, Abramis brama (L.), perch, Perca fluviatilis L. and roach, Rutilus rutilus (L.), in two contrasting gravel pit lakes: potential trophic overlap with wildfowl

The abundance of benthic macroinvertebrate taxa and the relative abundance of zooplankton taxa were compared with the diet of bream, tench. perch and roach from two gravel pit lakes during May-July 1986 and 1987. Significant food preferences were demonstrated between species and between lakes. Chironomid pupae dominated the macroinvertebrate diet of perch, roach, ate predominantly either Spirogyra sp. (St Peters Lake) or Daphniu hyalina (Main Lake).
The Main Lake, bream fed largely upon chironomid larvae and the bivalve Sphuerium but in St Peters Lake they positively selected a variety of less abundant benthic invertebrate taxa. Bream switched from benthos to zooplankton in the Main Lake in 1986. Tench ate large numbers of Aselhs and showed positive selection of various macroinvertebrate prey in St Peters but ate D. hjulim in Main Lake. Tropic overlap for chironomids and other macroinvertebrate prey was demonstrated between perch, bream and tench, and potentially with wildfowl which used the gravel pits for breeding and wintering.     

Competition-induced switch in young of the year perch,Perca fluviatilis: an experimental test of resource limitation

Synopis Reproductively developed male fathead minnows, Pimephales promelas, exhibited courtship behaviour in the presence of female conspecifies under laboratory conditions. Male courtship consisted of several distinctive and visually conspicuous behaviours directed toward females, including approach, display, and two contact behaviours, as well as leading behaviour from the female to a suitable spawning site. An ovulated condition in females was not necessary to generate male courtship behaviour; in fact, the amount of courtship exhibited by males may depend inversely on the readiness of females to spawn.     

Delayed consequences of extremely low‐frequency magnetic fields and the influence of adverse environmental conditions on roach Rutilus rutilus embryos

This study presents data collected over a 6 year period on the effects of extremely low‐frequency magnetic fields (MFs) (1·4–1·6 µT, 500 Hz and 1·4–1·6 µT, 72·5 Hz) and MFs in combination with other environmental stressors (elevated temperature, 0·01 mg l^?1 trichlorfon, 0·01 mg l^?1 copper sulphate pentahydrate) on roach Rutilus rutilus embryos. Effects were studied during different stages of early development. Rutilus rutilus were raised in ponds for 4 months after exposure to MFs. The mass, standard length (L_S) and morphological characteristics of underyearlings which were exposed as embryos were recorded. An increase in embryo mortality and a decrease in L_S and mass indices in underyearlings were noted after they had been exposed to a combination of MFs and different adverse environmental factors. In addition, exposure to MFs led to changes in the total number of vertebrae and the number of seismosensory system openings in the mandibular bones of underyearlings. MFs of different frequency caused both increases (500 Hz) and decreases (72·5 Hz) in morphological diversity. The stressors used in this study, however, did not increase the fluctuating asymmetry of bilateral morphological characteristics. The possible microevolutionary effects of exposure to MFs alone and in combination with other adverse environmental factors upon natural fish populations are discussed.     

Interactive effects of season and temperature on enzyme activities,tissue and whole animal respiration in roach,Rutilus rutilus

Synopsis This paper reviews investigations on the ecophysiology of a population of roach, Rutilus rutilus, from a subalpine oligotrophic lake in the Austrian Tirol. Metabolic responses to season and temperature were studied in whole animals, tissues and selected enzymes. The exponent of the relationship between body mass and three levels of the metabolic rate of acclimated fish was 0.82 ± 0.02, 0.60 ± 0.15, and 0.75 ± 0.01 at 4, 12, and 20° C respectively. Various combinations of long-term acclimation to constant or seasonally fluctuating temperatures and long-term (up to 14 days) monitoring of O₂ at the acclimation temperature led to the conclusion that the aerobic power of fish swimming in the routine mode does not show any sign of being temperature compensated. On the other hand, there are several indications that the energy expenditure of spontaneously swimming fish is adjusted to the seasonal pattern of environmental change and that these responses of metabolism and behaviour are controlled by both endogenous and exogenous factors. The rate of oxygen consumption of gill and muscle tissue brei from fish caught during a seasonal cycle and measured at 15° C appears to follow closely the reproductive and gonadal cycle of the living fish. The same holds for the activities of phosphofructokinase, acetoacetyl-CoA thiolase, and cytochrome oxidase. On the other hand, the Na⁺, K⁺-ATPase of the kidney shows near perfect temperature compensation when fish acclimated to 5 and 25° C are compared, whereas an equally pronounced case of inverse temperature acclimation has been reported for the activity of digestive enzymes in the gut. Summarizing these data it is pointed out that the temperature relationship of a poikilothermic organism is the sum of often very diverse temperature relationships of specific metabolic and behavioural functions. In the case of the roach, strong effects of acclimation temperature on the molecular level, sometimes in the opposite direction, combine with seasonal effects on enzyme activities and tissue respiration. However, on the whole animal level the fish behave as strictly non-compensating poikilotherms, the reproductive cycle being the only detectable influence capable of modulating the basic temperature relationship of energy expenditure.