Population structure,growth and fecundity of perch (Perca fluviatilis L.) in an acidified river system in Southern Norway

Population structure, density and reproductive potential of perch (Perca fluviatilis L.) from 4 acidified lakes in a river system in Southern Norway were investigated. The upper 3 lakes were most affected by acidification and the number of perch caught per unit effort indicate three-fold increase in density from the upper Gjerstadvann to the lower Brøbørvann. Low density of perch in the upper lakes is explained by; 1) Abnormal mortality occurring in episodes due to acidification, probably caused by aluminium toxicity at pH = 5.2–5.4. 2). Acid water (pH ≤ 5.0) during spawning and development of eggs and larvae, resulting in recruitment failure. Juvenile mortality of perch may depend on the density of the parental stock. In the 3 upper lakes, the density of adult perch probably was too low to produce significant juvenile mortality, and in these lakes the recruitment probably depended more on the May water quality. Improved water quality increases egg hatching and survival and also benefits zooplankton production, the main food for perch during the first summer. Low population density has reduced competition for food and thus improved growth of perch in the 3 upper lakes compared to the lower lake. The perch in Brøbørvann mature at higher age and have lower individual fecundity than perch in Gjerstadvann, an effect of both slower growth and lower length specific fecundity in Brøbørvann. In Gjerstadvann, the perch therefore have a higher reproductive potential relative to the stock density, and may, when the water quality is good enough, give rise to strong year-classes and more frequent year-class fluctuations than perch in the less acid Brøbørvann.     

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.     

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.     

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.     

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.     

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.     

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.     

Biological recovery of the Bohemian Forest lakes from acidification

A limnological survey of eight small, atmospherically acidified, forested glacial lakes in the Bohemian Forest (?umava, Böhmerwald) was performed in September 2003. Water chemistry of the tributaries and surface layer of each lake was determined, as well as species composition and biomass of the plankton along the water column, and littoral macrozoobenthos to assess the present status of the lakes. The progress in chemical reversal and biological recovery from acid stress was evaluated by comparing the current status of the lakes with results of a survey four years ago (1999) and former acidification data since the early 1990s. Both the current chemical lake status and the pelagic food web structure reflected the acidity of the tributaries and their aluminium (Al) and phosphorus (P) concentrations. One mesotrophic (Ple?né jezero) and three oligotrophic lakes (?erné jezero, ?ertovo jezero, and Rachelsee) are still chronically acidified, while four other oligotrophic lakes (Kleiner Arbersee, Prá?ilské jezero, Grosser Arbersee, and Laka) have recovered their carbonate buffering system. Total plankton biomass was very low and largely dominated by filamentous bacteria in the acidified oligotrophic lakes, while the mesotrophic lake had a higher biomass and was dominated by phytoplankton, which apparently profited from the higher P input. In contrast, both phytoplankton and crustacean zooplankton accounted for the majority of plankton biomass in the recovering lakes. This study has shown further progress in the reversal of lake water chemistry as well as further evidence of biological recovery compared to the 1999 survey. While no changes occurred in species composition of phytoplankton, a new ciliate species was found in one lake. In several lakes, this survey documented a return of zooplankton (e.g., Cladocera: Ceriodaphnia quadrangula and Rotifera: three Keratella species) and macrozoobenthos species (e.g., Ephemeroptera and Plecoptera). The beginning of biological recovery has been delayed for ～20 years after chemical reversal of the lakes.     

Effects of acidic pH and phytoplankton on survival and condition of Bosmina longirostris and Daphnia pulex

The effects of pH, algal composition and algal biomass on abundance, size, reproduction and condition of Daphnia pulex and Bosmina longirostris were tested in a field experiment using water and natural phytoplankton assemblages from a circumneutral (pH 6.43) and a moderately acidic (pH 5.75) lake in south-central Ontario. Both species were affected by pH and phytoplankton composition, with decreased egg production, lipid reserves, body size or abundance in treatments containing algae and/or water from the more acidic lake compared to treatments containing water and phytoplankton from the circumneutral lake. This result was unexpected for Bosmina, which often increases in relative and/or absolute abundance in acidified lakes. The negative effect of acidic conditions on Bosmina suggests that the population increase observed in most acidified lakes is not due to a positive response to low pH or ambient phytoplankton, but to altered biotic interactions possibly involving reduced competition.     

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.     

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.     

Rapid and unexpected effects of piscivore introduction on trophic position and diet of perch (Perca flavescens) in lakes recovering from acidification and metal contamination

1. Yellow perch (Perca flavescens) are often the only surviving fish species in acidified lakes. We studied four lakes along a gradient of recovery from acidification and that had different food web complexities. All had abundant yellow perch, two had low piscivore abundance, one had a well‐established piscivore population and one was manipulated by introducing piscivorous smallmouth bass (Micropterus dolomieu). We hypothesised that there would be strong effects on perch abundance, behaviour and diet induced by the presence of piscivores. 2. In the manipulated lake, the bass reduced yellow perch abundance by 75% over a 2‐year period. Concomitantly, perch use of the pelagic habitat fell from 48 to 40%. 3. In contrast to findings from less disturbed systems, yellow perch in the littoral zone of the manipulated lake did not strongly shift from zooplankton to benthic food sources after the arrival of piscivores. Diet analysis using stable carbon isotopes revealed a strong continued reliance on zooplankton in all lakes, independent of the degree of piscivory. The failure to switch to benthos in the refuge area of the littoral zone is most likely related to the depauperate benthos communities in these formerly acidified lakes. 4. Yellow perch in lakes recovering from acidification face a considerable ecological challenge as the necessary switch to benthic diet is hindered by a low abundance of benthos. The arrival of piscivores in these recovering lakes imposes further restrictions on perch access to food items. We infer that future recovery of perch populations (and higher trophic levels) will have to be preceded by the re‐establishment of diverse benthic macroinvertebrate communities in these lakes.     

Food quality for Daphnia in humic and clear water lakes

1. Growth and reproduction of Daphnia fed lake seston were measured in two categories of meso‐ to eutrophic lakes differing with respect to terrestrial organic matter influence (humic and clear water lakes). The content of highly unsaturated fatty acids (HUFA), P and N, as well as the taxonomical composition of seston were analysed. 2. Seston HUFA and C : P ratios were similar between lake categories, whereas C : N ratios were lower in the clear water lakes in both spring and summer. Despite the similarity in HUFA and P content of seston, Daphnia growth rate, clutch size and the proportion of gravid females were, respectively, about 1.5, 3 and 6 times higher in the clear water lakes. 3. Differences in growth and reproduction were related to a combination of higher N content and good fatty acid quality of the seston in the clear water lakes. Relatively high biomass of edible algae, such as Rhodomonas sp. and Cryptomonas sp., in the clear water lakes, and differences in water pH likely contributed to the observed differences in Daphnia growth and reproduction between lake categories. Additionally, it is possible that Daphnia was energy limited in the humic lakes despite high particulate organic carbon (POC) concentrations, as the contribution of non‐algal and detrital C to the POC pool was high. 4. Our results suggest that dietary HUFA content has the potential to improve herbivore growth and reproduction if N and P are not limiting. N merits more attention in studies of zooplankton nutrition.     

Phosphorus in acidified lakes: The example of Lake Gårdsjön,Sweden

Supply and turnover of phosphorus was studied in an acidified lake ecosystem, Lake Gårdsjön, located in southwestern Sweden. This study included transport and budget calculations combined with field and laboratory experimental work on abiotic phosphorus chemistry and biological utilization of phosphorus. The main conclusions presented in this paper are:

- The acidification process in inland waters resulting from acid atmospheric deposition is accompanied by ‘oligotrophication’ because of reduced input of phosphorus from the drainage area, possibly due to efficient fixation of phosphorus to aluminium complexes in the B-horizon of podzol soils

- Primary productivity in acidified lakes is limited mainly by low phosphorus supply

- Algal utilization of phosphorus in acidified lakes is impaired, yielding lower biomass than could be expected from ambient phosphorus concentrations. One possible reason for this could be that enzymatic recycling of organic phosphates is prevented by high levels of aluminium in lake water.

     

The effect of anthropogenic acidification on the hydrofauna of the lakes of the West Tatra Mountains (Slovakia)

Fourteen West Tatra lakes were studied, of which one could be considered to be recently anthropogenically acidified and eight others classified as acidification-endangered. In the anthropogenically acidified lake, the zooplankton assemblage has been substantially altered (three mountain-lake crustacean species have been eliminated). Several littoral macrobenthic species sensitive to acidification have either been eliminated from the acidified and acidification-endangered lakes or occur only sporadically. The effect of acidification has so far not been observed on the benthic fauna of the lakes medial which is probably due to the higher pH below the surface of sediment. In comparison with the High Tatra, acid depositions have had a less pronounced effect on the lakes of the West Tatra.     

Comparison of Plankton-Water Chemistry Relationships in Three Acid Stressed Lakes in Nova Scotia,Canada

A field program was conducted in 1980 and 1981 on three lakes in Kejimkujik National Park, Nova Scotia, to study plankton-water chemistry relationships in a region of potential acid precipitation stress. Among lake and between year comparisons were made. The three study sites represent two different types of lakes. Kejimkujik Lake (pH 4.8) and Pebbleloggitch Lake (pH 4.5) are both high-colored, dystrophic acid lakes. Beaverskin Lake is a clear, oligotrophic lake which is not as acidic (pH 5.4). The phytoplankton of Kejimkujik Lake was dominated by diatoms, while Pebbleloggitch Lake was dominated by chlorophytes and chrysophytes, and Beaverskin Lake was dominated by cyanophytes. Kejimkujik Lake had the highest total algal cell volume per liter, and Pebbleloggitch Lake the lowest. Rotifer populations composed the majority of the zooplankton communities, while the crustacean zooplankton were dominated by the acid-tolerant copepod Diaptomus minutus. Kejimkujik Lake had the lowest zooplankton biomass and Pebbleloggitch Lake the highest zooplankton biomass per m³. A pattern of highest algal density in Beaverskin Lake, but highest zooplankton density in Pebbleloggitch Lake emerged in both years. This is likely an effect of abundant, but unsuitable food (blue-green algae) in Beaverskin Lake and important detrital food resources in Pebbleloggitch Lake in both years. Multiple regression analysis of water chemistry variables with plankton species produced many significant effects, but failed to show clear patterns. Cause and effect relationships in aquatic ecosystems are poorly delineated by such techniques.     

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.     

Growth of larval and juvenile perch: the importance of diet and fish density

In August, growth rate of young–of–the–year (YOY) Perch In lake enclosures could be explained by both YOY density and mean cladoceran biomass, suggesting that in a lake where YOY perch are dominant, growth may be density dependent in late summer and mediated through top–down control on daphnid biomass. In June, growth rate of YOY perch could not be fully explained by YOY density or by mean cladoceran biomass, suggesting that growth and survival during the first part of the summer is negatively affected by a diet of Bosmina and cyclopoid copepods only. The experiments also suggest why YOY perch have a slow growth and a low abundance in eutrophic lakes where small zooplankton dominate. The June experiment also indicated that growth of late larval or early juvenile perch improved when a larger cladoceran became available and was included in the diet.     

Periphyton-macroinvertebrate interactions in light and fish manipulated enclosures in a clear and a turbid shallow lake

In a clear and a turbid freshwater lake the biomasses of phytoplankton, periphytic algae and periphytonassociated macrograzers were followed in enclosures with and without fish (Rutilus rutilus) and four light levels (100%, 55%, 7% and < 1% of incoming light), respectively. Fish and light affected the biomass of primary producers and the benthic grazers in both lakes. The biomass of primary producers was generally higher in the turbid than the clear lake, and in both lakes fish positively affected the biomass, while shading reduced it. Total biomass of benthic grazing invertebrates was higher in the clear than in the turbid lake and the lakes were dominated by snails and chironomids + ostracods, respectively. While light had no effect on the biomass of grazers in the clear lake, snail breeding was delayed in the most shaded enclosures and presence of fish reduced the number of snails and the total biomass of grazers. In the turbid lake ostracod abundance was not influenced by light, but was higher in fish-free enclosures. Density of chironomids correlated positively with periphyton biomass in summer, while fish had no effect. Generally, light-mediated regulation of primary producers was stronger in the turbid than in the clear lake, but the regulation did not nambiguously influence the primary consumers. However, regulation by fish of the benthic grazer community was stronger in the clear than in the turbid lake, and in both lakes strong top-down effects on periphyton were seen. The results indicate that if present-day climate in Denmark in the future is found in coastal areas at higher latitudes, the effect of lower light during winter in such areas will be highest in clear lakes, with typically lower fish biomass and higher invertebrate grazer density.     

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.