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.     

Phytoplankton in 63 limed lakes in comparison with the distribution in 500 untreated lakes with varying pH

As a result of the different weathering of lime in soils and varying acid deposition, the pH of Swedish oligotrophic lake waters has a broad range from ca. 4 to 8.5. Phytoplankton species occur in all lakes, but the composition and species richness varies in relation to pH. The maximum number of taxa was recorded within the pH-range 7.0–7.6, while the number decreased at higher pH and especially at pH below 5.6. During 1975–1985, thousands of lakes in Sweden were limed with the intention to restore biotic diversity and production. In order to find out whether there are differences in quality between limed and unacidified waters, the distribution of 57 selected phytoplankton taxa in 63 limed lakes was compared with that in 500 unlimed lakes with varying pH. The study shows that most common species in unacidified lakes also occur in the limed ones. However, several species of Cyanophyceae and Conjugatophyceae were missing especially in lakes that were strongly acidified before liming. On the other hand, some taxa were more frequent in the limed lakes.     

Short-term effects of liming on perch Perca fluviatilis populations in acidified lakes in South-West Sweden

Gill net catches of perch Perca fluviatilis in eight acidified lakes immediately before liming are compared] with catches in the same lakes two years after liming. Simultaneously, six control lakes, not limed, were investigated with the same procedures. A significant increase in the average catches of perch was found in the limed lakes, but not in the control lakes. This result confirms earlier reports, although seldom supported with data from controls, on successful recovery after liming.     

How many fish populations in Finland are affected by acid precipitation?

Synopsis The number of fish populations affected or lost from small lakes in southern and central Finland due to acid precipitation is estimated. Tolerance limits (pH and labile aluminum) of common fish species were obtained from a fish status survey of 80 lakes. These tolerance values were used to estimate the proportion of affected lakes from the water chemistry data of 783 statistically selected lakes. The proportion of anthropogenically acidified lakes was estimated by calculating pre-acidification pH and aluminum concentrations of the lakes, using a steady-state model based on water chemistry. The number of fish populations for which acidification has affected growth or population structure was estimated at between 2200 and 4400. Out of these, the number of fish populations that have disappeared due to acid precipitation would be about 1000–2000. Almost 60% of the affected or lost populations are roach, Rutilus rutilus, the most sensitive of the common fish species in small lakes in southern and central Finland. Less than 15% of the damaged population is European perch, Perca fluviatilis, the most common species. This is due to the substantially higher tolerance of perch to acidified water in comparison with roach.     

Use of enclosures to detect the contribution of particular zooplankton to growth of young-of-the-year yellow perch (Perca flavescens Mitchell)

Summary Densities of the cladoceran, Holopedium gibberum, were manipulated in 18 enclosures containing juvenile (age 0+) yellow perch (Perca flavescens) and mean-lake densities of other zooplankton. In enclosures, where nearlake densities of all zooplankton species including Holopedium were maintained, young-of-the-year perch grew significantly heavier and longer than in experimental enclosures where Holopedium was excluded. Holopedium comprised between 15–45% of the diet (wet weight) of perch in the first 2 weeks of July in the control treatment (Holopedium at or near ambient lake densities) and only 3–7% of total biomass ingested in the experimental treatment (Holopedium density selectively reduced). Predation on Holopedium decreased dramatically after the 2nd week of July in the control treatment after which Chaoborus, chironomids, and Sida became dominant prey items (by weight) of juvenile perch. These findings suggest that growth and survivorship of age 0+ perch in Precambrian Shield lakes may be coupled to Holopedium abundance. Thus, utilization of Holopedium by young-of-the-year yellow perch may affect recruitment of this species since overwintering survivorship, range of accessible prey sizes or species, and vulnerability of juvenile perch to predation by larger fish depend on body size, which is reduced when Holopedium is excluded from the diet.     

Whitefish stocking in acidified lakes: ecological and physiological responses

In autumn 1986, six small lakes at different stages of acidification were stocked with one-summer-old whitefish, Coregonus pallasi Valenciennes 1848, in order to see whether whitefish stocking would be a suitable method for the mitigation of acidification effects. In two of the lakes the introduction was a complete failure: the whitefish did not survive, evidently due to high acidity and high aluminium concentrations of the lake waters. In one of the most acidified lakes (pH 4.3–4.8, Al_lab 29–125 g 1^–1) and in two less acidic lakes (pH 5.0–5.2 and 5.4–6.4), introduction was successful. Three years after the introduction, the mean weights of the fish in those three lakes were 580, 250 and 360 g respectively, with the weight and also the condition factor of stocked whitefish being highest in the most acidified lake. In that lake there were few or no fish present during the introduction, whereas in the less acid lakes there were dense populations of perch and therefore a potential interspecific competition for food. Different availability of food in the lakes was presumed to be the main reason for the growth differences. Plasma Na⁺ and Cl^– concentrations of whitefish were lower in the acidic lakes than in the lake with pH around 6 three years after stocking. This suggests that, despite the good growth and highest condition factor of whitefish in the most acid lake, the fish still experienced some acid stress.     

Acidification affects the perch,Perca fluviatilis,populations in small lakes,of southern Finland

Synopsis Characteristics of perch were studied in 4 recently acidified lakes in southern Finland. The densities of the populations were 0–250 ha^–1 compared with 1400–3300 ha^–1 in two circumneutral reference lakes. The perch were close to extinction in one acidic lake and dominated by 7 year old fish in two other acidic lakes, suggesting decreased reproduction. Individual fecundity was not reduced by the acidity, but high mortality (close to 100%) of eggs in two acidic lakes indicated recruitment failure. In one of the acidified lakes reproduction had continued resulting in a population dominated by younger fish. However, increased egg mortality, low density, and the resulting high growth rate of perch in this lake were considered to be consequences of the acidity. These findings are the first documentation on the possible decrease of fish populations in Finland due to acid precipitation.     

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.     

Fish species distribution in relation to water quality gradients in the North Branch of the Moose River Basin

The distribution of fish species in the North Branch of the Moose River (Lake Rondaxe to headwaters of Big Moose Lake) was determined by intensive netting and electrofishing surveys of lakes and streams in the watershed during 1982–83. A chronology of changes in fish species occurrence in the drainage system was reconstructed from earlier published surveys conducted in 1882 and 1931 and unpublished survey data obtained by the NYSDEC during the period 1948–1975. Native species present in 1882 were also collected in 1931. Smallmouth bass (Micropterus dolomieu) were introduced in the early 1900's and were present in collections made in 1931. Major changes in the fish community have taken place since 1931. The smallmouth bass and many of the native species found in the earlier surveys were either absent or restricted in occurrence to downstream sites (eg. L. Rondaxe and Moss L. sub-drainage) in 1982. Non-native species introduced after 1931 (yellow perch,Perca flavescens; central mudminnow,Umbra limi; banded killifish,Fundulus diaphanus) are currently widely distributed throughout the drainage system. In particular, the yellow perch is now a dominant species in the larger lakes of the basin. Comparisons of survival rates for caged fish transferred from high to low pH sites in the Big Moose drainage system demonstrated relatively greater acid tolerane of non-native species (yellow perch, mudminnow, killifish) than native cyprinids. Watershed acidity gradients (pH and aluminum concentrations) and relative physiological acid tolerance are major determinants of currently observed fish species distribution patterns in the North Branch of the Moose River. Differences in age and size structure of fish populations inhabiting acidic and non-acidic lakes of the drainage system were apparent, but difficult to interpret without additional information onpopulation size and potential density dependent parameters such as age specific growth and survival rates. Differential hatching success was observed for yellow perch eggs reciprocally transferred between acid (Big Moose) and neutral (Moss L.) lakes. Eggs transferred from Moss L. to Big Moose L. exhibited poor hatching success as a result of alterations in egg membrane structure that inhibited normal egg expansion and the hatching process. This effect was not evident in eggs from the same parents reared in Moss Lake nor in eggs from the Big Moose parents reared in both lakes. These experimental observations suggested possible genetic adaptation to acid stress by the yellow perch population inhabiting Big Moose Lake.     

Littoral microcrustaceans (Cladocera and Copepoda) as indices of recovery of a limed water system

A 7 year study (1992–1998) of littoral microcrustaceans (Cladocera and Copepoda) in the watercourse of the River Rore, South Norway, illustrates that qualitative data on cladocerans and copepods are well suited to indicate the recovery of lakes following liming. Eight limed, two acid and two circum neutral reference lakes, were sampled twice a year (June/July and September/October). In the limed lakes, species associated with neutral lakes have become more common, whereas apparently acid-tolerant species have become rarer. In Lake Rore and Lake Syndle, the two largest limed lakes which exhibited a gradual increase in pH, the changes in species composition indicated that these lakes were about to recover. Species composition in Lake Røynelandsvatn, which has reacidified after liming, first reflected improved water quality, then reverted to dominance by acid-tolerant species. In the remaining lakes, the species composition reflects a fauna which has recovered compared with the preliming situation. There is strong evidence, however, that temporary fluctuations in pH have a negative influence on the speed of recovery, confirming the importance of keeping pH stable.     

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.     

Restructuring of Fish Assemblages in Swedish Lakes Following Amelioration of Acid Stress through Liming

A large-scale liming program was initiated in Sweden in the mid-1970s to ameliorate the effects of acidification and to restore acidified lakes and rivers to conditions present before acidification. I compared the fish fauna composition and biomass among limed, acidified, and consistently circumneutral lakes. The study focused on (1) recolonization and fish assemblage development and (2) fish biomass and assemblage composition. Time-series data from 14 limed, 5 acid, and 10 circumneutral lakes, and a data set of 323 limed and 346 nonlimed circumneutral lakes, are analyzed. Long-term monitoring revealed that fish species richness increased after liming and was of the same magnitude in limed lakes as in nonlimed circumneutral lakes after 10–20 years. Species proportions differed between limed and nonlimed lakes, and the relative biomass of Perca fluviatilis (Eurasian perch) was in general higher, whereas cyprinid and pelagic species were less important in the limed lakes than in the nonlimed circumneutral lakes. Recolonization of fish species was found to be a crucial factor for the development of the fish assemblages after liming, and in several limed lakes the reintroduction of fish species has accelerated the recovery of pre-acidification fish assemblages.     

Intensive fish predation: an obstacle to biological recovery following liming of acidified lakes?

Several thousand lakes have been limed inNorway and Sweden during the last threedecades. Most fishless lakes are stocked withsalmonid game fish after liming, predominantlybrown trout (Salmo trutta). Thecombination of intensive anthropogenic stockingand improved natural reproduction possibilitiesin brooks and rivers often leads to denselypopulated lakes and excess fish planktivory.Another major fish species in southern Norway,Eurasian perch (Perca fluviatilis),exhibits strong numerical increases afterliming, and within a short period of time thelakes harbor dense and often stuntedpopulations. The cladoceran zooplanktoncommunity was used as a diagnostic tool forstudying fish status, patterns and processes oflakes in the present area. Systems that werelimed before all fish species were lost, andbefore major dominance shifts had taken placewithin the invertebrates, recovered morequickly. Recovery periods are severelyprolonged in strongly disturbed, chronicallyacidified ecosystems. The control of fishabundance in optimal human resource managementof limed lakes is a delicate balance betweenthe necessity to suppress potentialinvertebrate predators and fish competitorssuch as corixids and chaoborids, and to avoidover-exploitation of the planktonic and benthiccommunities. If the combined watershed and lakesystem does not naturally produce sub-optimalspawning and recruitment possibilities, eitherfor brown trout or Eurasian perch, resulting insmaller populations, chances are high that thesite will lose its attraction as a fishingobject due to overpopulation. This should be amajor field of concern both for the ecology ofthese lakes and their resource management bythe fishery and environmental officials.     

Benthic fauna in a series of lakes displaying a gradient of pH

The benthic fauna was examined in a series of four isolated headwater lakes, displaying a pH gradient of 5.4–7.0. A slight reduction in number of taxa present occurred below pH 6, with the fauna below 3 m dominated by the Diptera. Although epibenthic gastropods were rare, in contrast with European studies, Amphipoda, Ephemeroptera and Pisidium were common in the least buffered lake, which experiences spring pH values as low as 4.7. There were little relation between the pH or alkalinity, and the abundance, and biomass of the fauna at depths greater than 3 m. However, biomass of the littoral fauna increased significantly in the lakes with lower pH, as a result of an increase in large littoral species normally susceptible to fish predation.     

Factors influencing changes in mercury concentrations in lake water and yellow perch (<Emphasis Type="Italic">Perca flavescens</Emphasis>) in Adirondack lakes

Over the past 20+ years, fish with elevated concentrations of mercury (Hg) have been observed in remote lake districts, including the Adirondack region of New York. Across eastern North America studies have also reported a negative correlation between fish Hg concentration and lake pH. Recent controls in emissions of sulfur dioxide (SO₂) have resulted in some improvement in the acid–base status of acid-impacted surface waters including Adirondack lakes. In addition, there has been an apparent decrease in atmospheric Hg deposition. A synoptic survey of 25 lakes in the Adirondacks was conducted in 1992–1993 to analyze spatial patterns of Hg in the water column and yellow perch (Perca flavescens). The same cluster of 25 lakes was resurveyed in 2005–2006 to evaluate if changes in lake concentrations of Hg species or fish Hg have occurred. We observed a varied response of changes in water chemistry and fish Hg concentrations. In twelve of the resurveyed lakes the yellow perch had lower Hg concentrations, six lakes had yellow perch with higher Hg concentrations, and in seven lakes yellow perch Hg concentrations did not change significantly (α = 0.05). Four variables appear to influence the change in yellow perch Hg concentrations in the Adirondacks: watershed area, elevation, change in pH, and change in fish body condition. We hypothesize that as the acidity in lakes is attenuated, the lakes may become more productive and/or water quality conditions less stressful to fish leading to increasing fish body condition. As fish body condition improves, fish exhibit “growth dilution” of tissue contaminants leading to lower fish Hg concentrations.     

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

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

Environmental factors influencing zooplankton species composition of lakes in north-central Ontario,Canada

To assess the relative importance of lake chemistry, morphometry and zoogeography on limnetic zooplankton, we collected zooplankton, water, and morphometric data from 132 headwater Canadian Shield lakes in 6 regions across north-central Ontario. A subset of these lakes (n = 52) were fished with gill nets. We clustered lakes based on their zooplankton species composition (presence/absence). Discriminant analysis was employed to determine how well lake characteristics could predict zooplankton community types. Correct classification of zooplankton communities for three models ranged from 72 to 91%. Lake size, lake location, and buffering capacity were ranked as the most important factors separating lake groups. Fish abundance (CPUE) was not significant in distinguishing between zooplankton communities. Though the range of lake sizes was limited (1–110 ha), larger lakes tended to support more species. Lake location (zoogeography) also influenced species composition patterns. Although Algoma lakes tended to be larger (\-x = 18.0 ha, other lakes \-x = 2.5 ha), they supported relatively depauperate zooplankton communities. Buffering capacity was ranked third in the discriminant analysis models, but pH and alkalinity were not significantly different between lake groups.     

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.     

The destruction of an endemic species flock: quantitative data on the decline of the haplochromine cichlids of Lake Victoria

Synopsis The Lake Victoria fish fauna included an endemic cichlid flock of more than 300 species. To boost fisheries, Nile perch (Lates sp.) was introduced into the lake in the 1950s. In the early 1980s an explosive increase of this predator was observed. Simultaneously, catches of haplochromines decreased. This paper describes the species composition of haplochromines in a research area in the Mwanza Gulf of Lake Victoria prior to the Nile perch upsurge. The decline of the haplochromines as a group and the decline of the number of species in various habitats in the Mwanza Gulf was monitored between 1979 and 1990. Of the 123+ species originally caught at a series of sampling stations ca. 80 had disappeared from the catches after 1986. In deepwater regions and in sub-littoral regions haplochromine catches decreased to virtually zero after the Nile perch boom. Haplochromines were still caught in the littoral regions where Nile perch densities were lower. However, a considerable decrease of species occurred in these regions too. It is expected that a remnant of the original haplochromine fauna will survive in the littoral region of the lake. Extrapolation of the data of the Mwanza Gulf to the entire lake would imply that approximately 200 of the 300+ endemic haplochromine species have already disappeared, or are threatened with extinction. Although fishing had an impact on the haplochromine stocks, the main cause of their decline was predation by Nile perch. The speed of decline differed between species and appeared to depend on their abundance and size, and on the degree of habitat overlap with Nile perch. Since the Nile perch upsurge, the food web of Lake Victoria has changed considerably and the total yield of the fishery has increased three to four times. Dramatic declines of native species have also been observed in other lakes as a result of the introduction of alien predators. However, such data concern less speciose communities and, in most cases, the actual process of extinction has not been monitored.     

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The numbers of perch Perca fluviatilis in the Cikola backwater system of the river Danube were estimated by mark-recapture techniques using multiple fishings. Fish were caught by fyke nets and electrical fishing. The latter method was unselective for sex and the catch data could be used to correct the sex bias in the fyke net catches. A population estimate in April of 6059 ind ha^–1 (95% C.L. 5135–7386) for perch > 5 cm was obtained for a 0.6 ha area of backwater and 1665 ind ha^–1 (95% C.L. 1204–2692) in May for a 0.4 ha area of a small bay.