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.     

Delayed spawning of perch, Perca fluviatilis L., in acidified lakes

The timing of spawning of perch was examined in four acidified lakes (pH 4.4–4.8) and in one circum-neutral lake (pH 6.3) in southern Finland in spring 1987. In three of the lakes, perch started to spawn soon after the ice melt (4–14 degree days > 5° C) and had spawned by the end of May at about 100 degree days > 5° C. In the two most acidified lakes, fish started to spawn later, at 35 and 60 degree days > 5° C, and had spawned in early June, at about 200 degree days > 5° C. The maturing of gonad products was delayed in both males and females.     

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.     

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.     

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.     

Long-term studies on the Windermere populations of perch (Perca fluviatilis), pike (Esox lucius) and Arctic charr (Salvelinus alpinus)

SUMMARY. 1, Windermere is a glacial valley lake with two basins separated by shallows and a total area of 14.8 km². Perch, pike and charr are the dominant fish species and have been studied on a continuous basis for over 45 years.
2. The initial slow-growing stocks of numerous small perch and moderate numbers of large pike were subjected to a heavy initial fishing pressure that was subsequently relaxed for perch but not for larger (>550 mm) pike. The number of charr in the lake has increased 6–8-fold during the study, probably due to the sustained reduction in the abundance of the larger pike that heavily predate the autumn spawning race of charr when it enters the shallow littoral areas to spawn.
3. The biomass of perch did not recover from the initial fishery until the late 1950s when strong year-classes occurred in warm summers. However, it was now composed of larger and faster growing individuals. In 1976, disease killed over 98% of all adult perch in the lake and the subsequent recovery has again been slow.
4. Existing models have explained the variation in perch year-class strength for the years 1959–74 in terms of adult perch biomass, pike cohort strength and temperature in the year of hatch. A new. more generally applicable model, incorporating a stock-recruitment expression, gives a better fit to the earlier data, particularly from the North Basin, where there have been periods of extremely low adult biomass.
5. The importance of continuing to extend these long-term data sets and the scope for future modelling work are discussed.     

Temperature effects on spawning and egg development in Eurasian perch

Spawning, egg strand development and embryo survival was studied during 1995–1996 in perch Perca fluviatilis exposed to heated effluent from two coastal nuclear power plants on the Baltic. Spawning was earlier and more prolonged in the heated areas. Although the fertilization rate was high, most egg strands sampled in the heated areas did not develop to hatching. The egg strands disintegrated within a few days of spawning, and egg mortality was very high. All samples collected from reference areas developed up to hatching. Exposure to high temperatures during the final maturation of the gonad influenced the follicular mechanism producing the jelly membrane which constitutes the matrix of the egg strand. Effects were observed in fish in open effluent areas as well as in an enclosed research basin. These results suggest that there is a conflict between optimal reproduction and temperature preference in temperate area fish. Fish are attracted to cooling water effluent, at the expense of reduced reproductive performance.     

The population dynamics of pike,Esox lucius,and perch,Perca fluviatilis,in a simple predator-prey system

Synopsis The population dynamics and predator-prey relationship of pike, Esox lucius, and perch, Perca fluviatilis, were examined in simple fish communities in two adjacent shallow lakes, Lochs Kinord and Davan, Deeside, Scotland. Few perch survive to age 3 but Z is low for fish > 3 years and perch live up to 17 years. Population fecundity remained relatively high and constant in perch because of the multi-age spawning stock and the presence of older more fecund perch. Growth rates of perch in both lochs are relatively high as a consequence of low stock abundance. The N, B, and P of adult perch were unusually low. The age range of pike, and N, B, P, and growth were in the range of values reported elsewhere. There was little variation in the strength of pike year classes and the importance of cannibalism and low occurrence of alternative prey in the lochs suggest that the populations were self-regulating. Cannibalism by adults was responsible for most of mortality in perch larvae, and predation by pike and adult perch was responsible for the majority of juvenile losses. This conclusion is supported by the high biomass ratios of pike:juvenile perch of 1.0–30.8. While the number of adult fish was almost equal, the biomass of adult pike was 2–3 × that of perch in Kinord and 6 × in Davan. In L. Kinord, where year class strength was stable, high predation pressure from perch and pike reduced perch abundance rather than eliminated year classes. Perch year classes fluctuated in abundance in L. Davan and were eliminated in the first summer in two sampling years. The pike, and particularly the perch populations, have features characteristic of fish communities in unperturbed ecosystems: namely, a wide range of age classes, stability in biomass with variation dampened by longevity, and low production.     

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.     

The annual reproductive cycle of pike, Esox lucius L., in two Scottish lakes

Histological changes in the ovaries of pike from two shallow, adjacent, Scottish lakes are described and related to a macroscopic scale of maturity stages and to changes in gonad weight. There was no evidence of pre-ovulatory degeneration although occasional residual oocytes were present in spent ovaries. The gonadosomatic index (GSI) was high in males from October to March, and in females it increased through the winter months to a maximum immediately prior to spawning. Maximum ovarian weight was 8.3–9.0% of total body weight compared with only 0.9–1.1% for testes. These GSIs were low compared with other reports for the British Isles, suggesting lower reproductive output. Spawning occurred in the first two weeks of April at a water temperature of 6.0–7.5° C. Fecundity was described from a regression of egg number (F) on fork length (f.l.): log₁₀F= -1.51 + 3.33 log₁₀ f.l. (cm). Mean egg diameter after preservation varied from 1.1 to 2.4 mm over the size range examined. Males were mature at age 2 at a length of ≥ 27 cm, while females first spawned at age 3 at a length of ≤ 28 cm, although a minority were mature at age 2. The sex ratio was 1.2 males: 1 female at Kinord and 1.4 : 1 at Davan. Greater egg production at Davan, 50 678 ha^?1, compared with Kinord, 33 557 ha^?1, was due to higher individual fecundity-at-age resulting from better growth. Fecundity (19.29 × 10³ calculated for a pike of 55 cm F.L.), relative fecundity of only 10–24 eggs per gram wet weight of adult, and GSI values were low compared with pike in North America and Eurasia. Food supply may restrict reproductive output of pike in Lochs Kinord and Davan. The abundance ratio of predator: prey is high with the main prey species, perch, present in low numbers, and the only alternative species is eel in these simple fish communities.     

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.     

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.     

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).     

Loss of genetic variability in reintroduced roach (Rutilus rutilus) populations

Genetic variation at five microsatellite loci and in mtDNA was surveyed in reintroduced and 'control' populations of roach ( Rutilus rutilus ) in Sweden. Microsatellite allelic richness and allele size ranges were significantly reduced in reintroduced populations, and mtDNA diversity was nearly significantly reduced in reintroduced populations. These measures of genetic variability were strongly correlated with lake characteristics that influence population size and food availability.     

Latitudinal gradients in onset date, onset temperature and duration of spawning of roach

Onset date, onset temperature and duration of spawning of roach Rutilus rutilus were related to latitude (40–66° N) and altitude (−1–825 m) based on data from the literature over the whole geographic distribution of roach. Onset date was positively and duration was negatively correlated with latitude, while no correlation was found between onset temperature and latitude. No significant correlations were found between any of the analysed variables of spawning and altitude, probably because most of the analysed populations were below 300 m altitude.     

Differences in growth of perch (Perca fluviatilis L.) in two small forest lakes

Growth patterns and food composition of perch, Perca fluviatilis L., was studied in two small forest lake populations in southern Finland. Size and morphometry of the lakes and physical and chemical properties of water are similar. There is a clear difference in the growth rates of perch between the two lakes. The difference in growth is highly significant in all age groups. In the first lake there is a perch population of 2 000 (1750 ind · ha^–1) adult fishes. In the second lake there is a small population of pike, that keeps the perch population down: 200 adult perch (530 ind · ha^-1). The main food items of perch are crustacean zooplankton, Asellus aquaticus L. and Trichoptera larvae in the first lake and zooplankton, Odonata larvae, Ephemeroptera larvae and Heteroptera in the second.It is concluded that the main reason for the growth difference of studied perch populations is the different population density. There are also differences in species composition of bottom fauna of the lakes, maybe owing to the floating Sphagnum peat moss vegetation in the second lake. This can also affect the growth difference between the two populations of perch.     

Reproduction effort versus the environment; case histories of Windermere perch, Perca fluviatilis L., and pike, Esox lucius L.

The theoretical limits of net reproduction rates for perch, Perca fluviatilis L. and pike, Esox lucius L, have been estimated from the upper and lower limits of fecundity, growth, mortality, age of maturity and biomass of parental stock, observed in Windermere over 40 years. Differences in egg numbers due to changes in fecundity were for perch × 1.5, for pike × 2. Assuming the same mortality, a fast growing cohort would produce more eggs during its lifespan than a slow growing, for perch × 2, for pike × 9. Changes in mortality from high to low resulted in more eggs, for perch × 7, and for pike (where high mortality includes natural and fishing mortality) × 10. Change in age at first maturity from 3 to 2 years was unimportant. Biomass of parental stock varied by × 6 for perch and × 3 for pike, and number of eggs laid varied by x2 for perch and × 5 for pike. The number of recruits at age 2 years varied by × 300 for perch and × 7 for pike. It was concluded that temperature in the first summer of life and predation were of major importance in regulating the number of recruits. Ricker recruitment curves and a simple model showed that perch, but not pike, could produce maximum recruitment from minimum parental stock. For perch, but not pike, compensatory density dependent mortality increased with increased parental stock. It was concluded that responses by the perch and pike populations to a changing environment can not be of sufficient magnitude to prevent wide fluctuations in recruitment.