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.     

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.     

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.     

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.     

Acidity status and phytoplankton species richness,standing crop,and community composition in Adirondack,New York,U.S.A. lakes

The mid-summer phytoplankton communities of more than 100 Adirondack lakes ranging in pH from 4.0 to 7.2 were characterized in relation to 25 physical-chemical parameters. Phytoplankton species richness declined significantly with increasing acidity. Acidic lakes (pH < 5.0) averaged fewer than 20 species while more circumneutral waters (pH > 6.5) averaged more than 33 species. Phytoplankton abundance was not significantly correlated with any of the measured physical-chemical parameters, but standing crop parameters, i.e., chlorophyll a and phytoplankton biovolume, did correlate significantly with several parameters. Midsummer standing crop correlated best with total phosphorus concentration but acidity status affected the standing crop-phosphorus relationship. Circumneutral waters of low phosphorus content, i.e. < 10 µg·1^–1 TP, averaged 3.62 µg·1^–1 chlorophyll a whereas acidic lakes of the same phosphorus content averaged only 1.96 µg·1^–1 chlorophyll a. The midsummer chlorophyll content of lakes of high phosphorus content, i.e. > 10 µg·1^–1 TP, was not significantly affected by acidity status.Adirondack phytoplankton community composition changes with increasing acidity. The numbers of species in midsummer collections within all major taxonomic groups of algae are reduced with increasing acidity. The midsummer phytoplankton communities of acidic Adirondack lakes can generally be characterized into four broad types; 1) the depauperate clear water acid lake assemblage dominated by dinoflagellates, 2) the more diverse oligotrophic acid lake community dominated by cryptomonads, green algae, and chrysophytes, 3) the productive acid lake assemblage dominated by green algae, and 4) the chrysophyte dominated community. The major phytoplankton community types of acid lakes are associated with different levels of nutrients, aluminum concentrations, and humic influences.     

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.     

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.     

Population dynamics,production and angling catch of brown trout,Salmo trutta,in a mature upland reservoir in mid-Wales

Synopsis The brown trout in Llyn Frongoch, a mature upland reservoir, and its nursery stream was sampled during 1983. The stream stock consisted largely of the 1983 and 1982 year classes, with fish reaching mean lengths of 7.0 and 11.6 cm at one and two years of age. The size and biomass of the stream stock at the beginning of 1983 and 1984 were estimated to be 120 and 125 (1.20 and 1.25 fish m^–2) and 1.41 and 0.69 kg (14.1 g m^–2 and 6.9 g m^–2) respectively. Annual stream production ranged from an estimated minimum of 2.49 kg (24.9 g m^–2) to an estimated maximum of 4.59 kg (45.9 g m^–2). Both downstream and upstream movements of 0+ juveniles were recorded. The adult spawning stock was estimated at 79 males and 32 females, a sex ratio of 2.5:1, with most spawners belonging to the 1980 yearclass. The average size of the lake stock over the year was estimated to be 1 650 (229 fish ha^–1) or 250.8 kg (34.8 kg ha^–1). The 1980 yearclass was predominant; there were few fish older than five years. Seasonal variations in netting catches suggested movements to and from the littoral region. Growth in the lake was moderately fast, with fish reaching mean lengths of 21.7 and 27.2 cm by three and four years of age. Fish entering the lake after one year appeared to grow faster than fish which remained in the stream for two years. Annual production in the lake was estimated at 136.7 kg (19.0 kg ha^–1). The total angling catch for the season was estimated to be 62.6 kg (8.7 kg ha^–1).     

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.     

Effects of liming on the occurrence and abundance of fish populations in acidified Swedish lakes

Test fishing with multimesh gillnets was performed in 103 acidified Swedish lakes before and 2–4 years after liming. In a subset of 39 lakes additional test fishing was carried out 5–9 years after liming. Taking into account the sampling biases due to differences between water temperature and number of nets used on different sampling occasions, an increase of the total CPUE, the number of species caught and recruitment of dominating species was evident after liming. Decreased recruitment was found for perch (Perca fluviatilis) at a pH below 5.2 and for roach (Rutilus rutilus) at a pH below 6.1. During the acidification phase and 2–4 years after liming acid-tolerant perch dominated the lakes, but when a longer time had elapsed after liming species dominance shifted and acid-sensitive planktivorous species increased in CPUE while perch decreased.     

Paleolimnological reconstruction of the effects of atmospheric deposition of acids and heavy metals on the chemistry and biology of lakes in New England and Norway

Sediment cores from nine lakes in southern Norway (N) and six in northern New England (NE) were dated by ¹³⁷Cs, ²¹⁰Pb and in NE also by pollen, and were analyzed geochemically and for diatoms. Cores from two N and three NE lakes were analyzed for cladocerans. ¹³⁷Cs dating is unreliable in these lakes, probably due to mobility of Cs in the sediment. In Holmvatn sediment, an up-core increase in Fe, starting ca. 1900, correlates with geochemical indications of decreasing mechanical erosion of soils. Diatoms indicate a lake acidification starting in the 1920's. We propose that soil Fe was mobilized and runoff acidified by acidic precipitation and/or by soil acidification resulting from vegetational succession following reduced grazing. Even minor land use changes or disturbances in lake watersheds introduce ambiguity to the sedimentary evidence relating to atmospheric influences. Diatom counts from surface sediments in 36 N and 31 NE lakes were regressed against contemporary water pH to obtain coefficients for computing past pH from subsurface counts. Computed decreases of 0.3–0.8 pH units start between I890 and I930 in N lakes already acidic (pH 5.0–5.5) before the decrease. These and lesser decreases in other lakes start decades to over a century after the first sedimentary indications of atmospheric heavy metal pollution. It is proposed that the acidification of precipitation accompanied the metal pollution. The delays in lake acidification may be due to buffering by the lakes and watersheds. The magnitude of acidification and heavy metal loading of the lakes parallels air pollution gradients. Shift in cladoceran remains are contemporary with acidification, preceding elimination of fishes.     

Planktonic rotifer community structure in Adirondack,New York,U.S.A. lakes in relation to acidity,trophic status and related water quality characteristics

The structure of the mid-summer planktonic rotifer communities of 101 Adirondack lakes ranging in pH from 4.0 to 7.3 were characterized in relation to acidity and selected water quality parameters. More than 70 rotifer species were identified from collections in 1982 and 1984. None of the species collected could be considered acidobiontic or alkalibiontic. Keratella taurocephala was the most commonly collected rotifer, occurring in 94 of the study lakes. It was abundant throughout the range of pH investigated but was particularly dominant in acidic waters, averaging > 85 % of the rotifers collected from waters of pH < 5.0.Rotifer community structure can be related to three groups of water quality parameters. Community parameters (richness and diversity) are most highly correlated with parameters indicative of acidity status. Rotifer abundance correlates with trophic state indicators, i.e. chlorophyll a and total phosphorus, over the full range of pH investigated. However, in acidic lakes, rotifer abundance is related to true color and DOC, indicators of humic influences.The rotifer communities of the Adirondacks can be classified into four broad types: 1) A diverse, productive community of the more alkaline lakes, generally with 13 species, and dominated by Conochilus unicornis, Kellicottia bostoniensis, Kellicottia longispina, and Polyarthra major; 2) Relatively diverse communities of productive, highly colored acid lakes, with 8 species, and often with very large populations (> 200 · 1^–1) dominated by K. bostoniensis and K. taurocephala; 3) Depauperate (< 4 species) communities of clear water acid lakes with generally low density populations dominated by K. taurocephala (> 90 % of rotifers in each sample); and 4) Extremely depauperate (2–3 species) acid lake communities associated with small lakes with high flushing rates dominated by C. unicornis.     

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.     

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.     

Species-specific population-density thresholds in cladocerans?

The population density of a Daphnia species seems more dependent on properties specific to the species than on those specific to the point in the season, location within a lake basin or the given lake itself. In spite of week-to-week fluctuations, the population density for each of two common European Daphniaspecies was found to be remarkably similar within single lakes (from station to station on a single date, and from date to date at a single station) as well as from lake to lake, regardless of trophic state. All lakes on all dates revealed densities in the range 10–50 ind l^–1 for the smaller-bodied D. cucullata and 1–5 ind l^–1 for the larger-bodied D. hyalina, in spite of different intensity of reproduction resulting from different food levels (chlorophyll a between 0.2 and 4.2 g/l). It can be asserted that the population density of each species remains far below the carrying capacity of the habitat K, and does not depend on food levels, which merely set the rate of population increase, while the population density reflects the species' vulnerability to predation by planktivorous fish. The reactive distance (the distance over which a foraging fish can see its prey) in 1+ roach, a dominant planktivore in the lakes studied, has been found to be twice as great for D. hyalina as for D. cucullata, whatever the light intensity. The relative reactive field volume was therefore an order of magnitude greater for the former than for the latter, showing that the densities of the two prey species would be assessed by a foraging roach as equal when, in reality, they differed by an order of magnitude, as they do in various lakes and in various seasons.The first of the two conclusions is that whatever the growth and reproduction in a population of a cladoceran such as Daphnia, its density would be fixed by mortality induced by fish predation. The second would be, that the difference between the bottom-up and top-down effects in freshwater is more than merely the upward or downward direction along the food web, since the bottom-up effects are about the flow control (the rate of net production, individual growth rate, rate of reproduction) and the top-down about the standing-crop control (biomass, individual body size, population density level).     

Perch,Perca fluviatilis L., in Small Lakes: Relations between Population Characteristics and Lake Acidity

Perch population characteristics of small lakes were examined with a special reference to lakes which have acidified recently due to atmospheric deposition of air pollutants. The population density and biomass of perch were higher in an acid (pH 4.5) clear water lake with low aluminium concentration than in recently acidified (pH 4.3–4.7) clear water lakes with higher aluminium levels. The structure of perch population in an acid (pH 4.4) humic lake was similar to recently acidified clear water lakes. The population density and the biomass of perch were significantly higher in 6 lakes with pH>5.0 than in 6 lakes of pH<5.0 whereas the mean age and length at a given age were higher in the more acid lakes.     

Abundance, Annual Survival, and Recruitment of Unexploited and Exploited Lake Charr, Salvelinus Namaycush, Populations at the Experimental Lakes Area, Northwestern Ontario

We studied abundance, annual survival, and recruitment of nine lake charr, Salvelinus namaycush, populations at the Experimental Lakes Area, Ontario, for periods of 9–24 years. We used the Jolly–Seber mark-recapture method to evaluate abundance and annual survival in all populations, and total catches of individual year classes to evaluate recruitment. Seven populations were unexploited and unaffected by whole-lake experiments. One population was exploited prior to mark-recapture study and another was affected by experimental acidification. Abundance ranged from 8 to 24 fish ha^–1 in the unaffected populations. Annual survival ranged from 78% to 93%yr^–1 in all populations except the exploited population. This population may have been additionally affected by northern pike, Esox lucius, predation. Yearly recruitment was relatively constant in all populations and related to lake charr abundance. Lake charr abundance did not recover in populations affected by exploitation or acidification. Unexploited lake charr populations were characterized by relatively constant abundance, high annual survival, and relatively constant annual recruitment.     

Disruption of sulfur cycling and acid neutralization in lakes at low pH

Experimental acidification of a softwater lake to below pH 5 fundamentally changed the sulfur cycle and lowered internal alkalinity generation (IAG). Prior to reaching pH 4.5, the balance of sulfur reduction and oxidation reactions within the lake was in favour of reduction, and the lake was a net sink for sulfate. In the four years at pH 4.5 the balance of reduction and oxidation reactions was in favour of oxidation, and there was a net production of sulfate (SO₄ ^2–) within the lake. Evidence indicating a decrease in net SO₄ ^2– reduction at pH 4.5 was also obtained in an anthropogenically acidified lake that had been acidified for many decades. In both lakes, the decrease in net SO₄ ^2– reduction appeared to be linked not to a simple inhibition of SO₄ ^2– reduction but rather to changes in benthic ecosystem structure, especially the development of metaphytic filamentous green algae, which altered the balance between SO₄ ^2– reduction and sulfur oxidation.At pH's above 4.5, net SO₄ ^2– reduction was the major contributor to IAG in the experimental lake, as it is in many previously studied lakes at pH 5 and above. At pH 4.5, the change in net annual SO₄ ^2– reduction (a decrease of 110%) resulted in a 38% decrease in total IAG. Because of the important role of net SO₄ ^2– reduction in acid neutralization in softwater lakes, models for predicting acidification and recovery of lakes may need to be modified for lakes acidified to pH <5.     

Recovery of the fish community and changes in the lower trophic levels in a eutrophic lake after a winter kill of fish

A severe oxygen deficit induced a fish kill in the eutrophicated two-basin Lake Äimäjärvi in southern Finland during winter 2002–2003, resulting in cascading effects on the lower trophic levels of the lake. Pikeperch disappeared from the lake and bleak and white bream decreased to low numbers. The recovery of the populations of other species started immediately when strong year-classes of roach and perch appeared in summer 2003 and onwards. A sharp increase in the growth of perch and roach was recorded, and perch became the dominant fish species during 2004–2006. Consequent responses after the fish kill included increased Secchi depth, expansion of submerged macrophytes, decreased nutrient concentrations and reduction of Cyanophyta from the more eutrophic northern basin of the lake, and a temporary increase in the size of Daphnia in the early summer 2003. However, the ecosystem of the lake returned gradually to the earlier structure and level of eutrophication. Bluegreen algae have returned since 2005, the water has become more turbid, macrophytes declined in 2007 and the fish community was again dominated by small cyprinid fishes in 2008.