Use of rehabilitation experiments to understand the recovery dynamics of acid-stressed fish populations

We used rehabilitation experiments involving the stocking of 2 native sportfish, lake trout (Salvelinus namaycush) and smallmouth bass (Micropterus dolomieu), in combination with recent fish community surveys, to study the recovery dynamics of fish populations in acid-stressed lakes near Sudbury and Killarney, Ontario, Canada. Population recovery rates differed between the 2 species. Introduced lake trout did poorly in species-rich lakes and exhibited slower growth, lower survival and delayed recruitment. Smallmouth bass, in contrast, readily colonized species-rich lakes. The biomass of natural smallmouth bass recruits increased to reference lake levels within 5 years following water quality recovery and spawning by stocked fish, whereas the biomass of natural lake trout recruits remained well below reference levels 5-15 years after water quality recovery and spawning by adults occurred. We document introductions by anglers of smallmouth bass into acid-damaged lake trout lakes, including some lakes that did not contain bass prior to acidification. This range expansion of a warm-water species (bass) that can alter food-web structure and reduce the growth of a cold-water species (trout), illustrates the potential for the combination of climate warming and species introductions to greatly alter the biological recovery endpoints in acid-stressed lakes.     

Reproductive status, blood chemistry, gill histology and growth of perch (Perca fluviatilis) in three acidic lakes

Perch (Perca fluviatilis L.) were sampled soon after spawning in three small acidic lakes (pH 4.3-6.1, Al(lab) 5-106 microg litre(-1), Ca2+ 0.01-0.08 mmol litre(-1)) and in one circumneutral lake (pH 5.9-6.4, Al(lab) 4-12 microg litre(-1), Ca2+ 0.06-0.07 mmol litre(-1)) in southern Finland. Due to the delayed spawning of perch in the acidic lakes, sampling in those lakes was performed later than in the reference lake. In spite of that, the gonadosomatic index (GSI) of males in all the acidic lakes was significantly greater than in the reference lake. Of the two lakes with similar low water pH, the effects on reproduction were more prominent in the lake with higher water Al content. The plasma Ca2+ concentrations of females in the acidic lakes were significantly smaller than in the females of the reference lake. The low female:male plasma Ca2+ ratio (1.0-1.32) depicted delay of spawning. Stress in perch in acidic water was also seen in elevated blood haematocrit values, especially in females. On the other hand, a low plasma Cl- level, a common response to acidic water in salmonids, was not detected in perch in the most acidic lakes. The amount of Al accumulated in the gill epithelium was highest in the most acidified lake with high Al concentration, but was also pronounced in a lake with low pH and low Al concentration.     

Evidence of biological recovery in acid-stressed lakes near Sudbury, Canada

Reductions in the emissions of SO2 and trace metals from the Sudbury smelters have resulted in substantial improvements in water quality in many surrounding lakes. Significant biological changes have accompanied the chemical improvements. Evidence of relatively rapid recovery was found for benthic filamentous algae, phytoplankton, zooplankton, mobile species of benthic invertebrates, and some fish populations. Organisms with low dispersal ability (e.g. Hyalella azteca) have not yet recolonized these lakes. The partial recovery observed to date shows movement toward re-establishment of biological communities typical of natural Precambrian Shield lakes in this area. These findings offer strong support for further efforts to reduce industrial emissions of pollutants to the atmosphere.     

Biomass and production of amphipods in low alkalinity lakes affected by acid precipitation

Population biomass and production of the amphipod Hyalella azteca (Saussure) were found to be related to alkalinity (ranging from 0.2 to 58.1 mg liter(-1)) in 10 Canadian Shield lakes in south-central Ontario. Biomass and production of amphipods in the two lakes characterized by spring depressions of pH below 5.0 were found to be lower than those for populations inhabiting lakes that did not experience such acid pulses. The proportional biomass of amphipods in relation to the total littoral zoobenthos community was lower in lakes of low alkalinity than in circumneutral or hardwater lakes. Because production in these amphipod populations is known to depend closely on population abundance, the labour-intensive derivation of production rates yields relatively little information for biomonitoring that cannot be obtained from abundance data alone.     

Changes in the density, population structure, growth and reproduction of perch, Perca fluviatilis L. in two acidic forest lakes in southern Finland

The density of the perch populations decreased rapidly during the 1980s in two acidified lakes in southern Finland. In an acid-humic lake the density of the perch population decreased from 240 to 60 ha(-1) and in an acid clearwater lake from 150 to 20 ha(-1). In both lakes the rapid decrease was mainly due to failures in reproduction, although post-spawning mortality of adult fish was also recorded. The perch spawned annually in both lakes but according to the field exposures of egg strands, most of the eggs died before hatching. Nevertheless, some new recruits appeared in both lakes. These perch, born in 1983-1986, grew faster than those of the more abundant year-classes of the late 1970s, indicating a clear density-dependent growth response. According to the mortality estimate and the age distribution of perch, the latest year-classes that produced more than 100 adult perch ha(-1) were born in 1978 or 1979 in both lakes. It was calculated that the density of both populations was >500 ha(-1) in the early 1980s.     

Effects of acidic water on freshwater snails: results from a study of 1000 lakes throughout Norway

This paper summarizes some results from a monograph which focused on ecological and biogeographical aspects of the biota in Norwegian lakes and rivers with particular reference to the snail fauna. Field studies were made in the course of 20 summers within the period of 1953-1987. Acidic water is the main reason why snails are absent from some 4000 lakes in a low-calcium area in southern Norway ('Acid south'). This area is heavily influenced by acidic precipitation. Snails were not detected in lakes with pH below 5.2. In one lake, three snail species disappeared in parallel with acidification from pH 5.2 to 4.2. The calcium level modified the effect of low pH.     

Towards a model of acidification effects on waterfowl in Eastern Canada

Data from 212 lakes in central Ontario were used to examine the relationship between presence of breeding waterfowl and loons and the following lake characteristics: pH, presence of fish, lake area, dissolved organic carbon (DOC) and total phosphorus (TP) concentration. In univariate analyses, the two fish-eating species preferred large, high pH lakes with fish, while insectivorous species showed little consistent pattern of lake association. Logistic regression analyses confirmed that large lake size and presence of fish were important determinants of presence of piscivores, though the residual effect of pH differed between the two species. Broods of three of the four insectivorous species avoided lakes with fish, and independently showed a positive response to pH. By calculating the probability that fish will be present on a lake of given area and pH, it is possible to estimate the net effect of pH change on these waterfowl. Relationships such as those presented here can, with some assumptions, be linked to models of lake acidification to estimate response of waterfowl to predicted changes in acidic deposition.     

Critical loads for Finnish lakes: comparison of three steady-state models

Three simple steady-state water-chemistry models are used to calculate critical loads of sulfur for lakes in Finland. Because of the high concentrations of organic matter in Finnish lakes, the influence of organic anions on the calculation of critical loads has been given special attention. The first two methods are well known ion-balance methods which have been used in many previous lake-acidification studies. The third method, developed for this study, includes the numerical solution of equilibrium equations for organic anions, inorganic carbon species and inorganic monomeric aluminum. The original pH and aluminum concentration of the lakes are estimated with this model, and a method to estimate the original acid neutralizing capacity (ANC) by simulating a Gran-titration is also tested on the lake data. Uncertainty in the predictions is estimated by varying the most critical model parameters.     

Temporal and spatial variation of RNA content in nymphs of the mayfly, Stenonema femoratum (Say)

Temporal variation of RNA: weight, RNA: protein, RNA: DNA, and protein: DNA ratios in Stenonema femoratum (Say) was assessed by monitoring at the same site in a small midwestern stream over a two-year period. Variation components were estimated using a two-way mixed model analysis of variance with weight class and collection date as main effects. About 30% of the variation in RNA concentration was within-cell variation associated with differences among individuals and analytical error. Temporal variation accounted for about 50% of the total variability in RNA concentration data. RNA: weight had 11% and RNA: protein had 17% variation attributed to weight class effects indicating that these ratios were dependent on the size of the mayfly. RNA: DNA ratios were not influenced by mayfly weight, suggesting that this ratio would be the most appropriate for comparing populations based on means of mixed individual sizes. Comparison of mean RNA: DNA ratios among collection dates yielded significant increases in the ratio during late fall in two consecutive years. Spatial variations in RNA: DNA ratios of S. femoratum collected from sites of similar habitats were not significantly different. These data encourage the future development of RNA concentration as an in situ biochemical indicator of growth rate and possibly of stress related effects on growth rate.     

Lake acidity and mercury content of fish in Darwin National Reserve, Russia

Darwin National Reserve is a protected natural area on the north-west shore of the Rybinsk Reservoir, 350 km north of Moscow. In June 1989, six lakes in the Reserve and the reservoir were surveyed to assess lake acidity and the mercury content of perch, Perca fluviatilis. Five were seepage lakes with no permanent inlets or outlets and one was a drainage lake with both an inlet and an outlet. The seepage lakes were acidic (mean pH 4.6-4.8) and varied in colour from 20 to 200 Hazen units. The drainage lake and reservoir were alkaline (mean pH 8.0-8.1) and colour spanned a similar range. The mean mercury content of perch dorsal epaxial muscle ranged from 0.5 to 1.1 microg g(-1) wet weight in the five acidic lakes and from 0.1 to 0.2 microg g(-1) in the alkaline lakes. Fish mercury content was negatively correlated with lake pH (r=-0.93, P=0.002) if all waters were considered together, and positively correlated with apparent colour (r=0.91, P=0.03) in the seepage lakes.     

Resilience of epilithic algal assemblages in atmospherically and experimentally acidified boreal lakes

Algal assemblages can be highly responsive to environmental changes in recovering acidified lakes. We compared epilithic algal assemblages in boreal lakes during chemical recovery from atmospheric (Killarney Park, Ontario) and experimental (Lake 302S, Experimental Lakes Area, Ontario) acidification to assess the impact of spatial and temporal scale of severe acidification on taxonomic resilience (i.e. recovery rate). Resilience was measured as the distance traveled by lakes in ordination space during pH recovery based on canonical correspondence analysis. Resilience was relatively negligible in the Killarney lakes, suggesting that eight years of experimental acidification in Lake 302S had less impact on biological recovery than did decades of regional acidification. Increases in dissolved organic carbon, dissolved inorganic carbon, and calcium best explained temporal variance of epilithic species abundances in the recovering acidified lakes. In Lake 302S, contrasting trajectories of taxonomic resilience and resistance, i.e. displacement from reference conditions following a perturbation, indicated that ecological factors affecting epilithon differed at corresponding pH levels during recovery and acidification. Our findings reveal that modeling of ecosystem recovery from severe acidification must account for the spatial and temporal scale of the perturbation, and biological delay responses that result in differences between recovery and acidification trajectories.     

Food chain enrichment of organochlorine compounds and mercury in clean and polluted lakes of Finland

Lakes polluted by pulp mill and urban wastes including chlorobleaching of pulp, semipolluted lakes and reference lakes in nearly natural condition in Central Finland were studied for contents of mercury, methyl mercury and organochlorine compounds in sediment, plankton, roach and pike. Chlorobleaching had caused a 30-fold concentration of Hg in bottom sediment related to that of the purest reference lake. This was not reflected to the mercury levels in fish which were highest at one natural condition (humic) lake and rather high also at semipolluted lake Päijänne. Mercury in fish was shown to be mostly methylated but not completely and its time trends could be estimated. Chloroform showed no but carbon tetrachloride, tetrachloroethylene and chlorinated cymenes significant bioaccumulation in fish. Levels of chlorophenols from bleaching had strongly decreased but pentachlorophenol and 2,3,4,6-tetrachlorphenol levels remained related to earlier results. Using fat basis attenuated the power of estimation of food chain enrichment by a three throphic level model for lipohilic biocides and a strong proof was obtained of the enrichment of hexachlorobenzene. The time trends at Päijänne were decreasing for mercury and DDE but increasing for PCB.     

Evaluation of aquatic environmental fate of 2,4,6-trichlorophenol with a mathematical model

A mathematical model, Simplified Approach for Fate Evaluation of Chemicals in Aquatic Systems (SAFECAS), was applied to evaluate fate of 2,4,6-trichlorophenol(TCP) in three types of aquatic environments; a river, an oligotrophic lake, and an eutrophic lake.Since phenols are dissociative substances, equilibrium constants and rate constants for transfer and transformation processes were measured or estimated as a function of pH.Results suggest that most of TCP is distributed to the water phase in aquatic environments, and is almost removed by the advective outflow in the river and by the photodegradation in lakes.     

Reconstruction of centuries-old Daphnia communities in a lake recovering from acidification and metal contamination

Paleolimnological and molecular genetic techniques were combined to reconstruct the long-term patterns in Daphnia community composition in Hannah Lake--a lake recovering from industrial acidification, metal contamination and faunal extirpation. Like many zooplankters, Daphnia produce diapausing eggs that can remain viable for decades and even over a century. Yet, the appearance of D. mendotae in Hannah Lake during the last two decades is likely the outcome of dispersal from other nearby lakes, not by colonization from the sediment egg-bank. Our genetic tests using PCR, SSCP and sequencing indicate that D. mendotae diapausing eggs are absent within the sediment record of the previous 250 years and that, prior to metal smelting operations in the region, the community was dominated by D. pulicaria. This species shift following the lake's chemical restoration is consistent with earlier historical changes in lake acidity. Environmental fluctuations may have governed community composition throughout Hannah Lake's more ancient past. Extending this molecular-paleolimnological approach to other lakes should help develop more accurate formulations of the biological recovery process.     

A survey of lakes in the Republic of Ireland: hydrochemical characteristics and acid sensitivity

In 1997, as part of a national program to determine and map critical loads, a lake survey was carried out in the Republic of Ireland. In total 200 lakes were sampled, which represents approximately 3.3% of the total lake population. The majority of lakes were situated in remote, high-altitude, acid-sensitive areas along the coastal margins of the country. Lake chemistry was dominated by marine inputs. Approximately 50% of the lakes had DOC > 5 mg L-1 due to the presence of organic soils in a large proportion of the catchments. Nonmarine sulfate concentrations were at background levels (< 20 mu eq L-1) in 50% of the lakes. Exceedance of critical load was observed in 7% of the sampled lakes (13 lakes). However, there are uncertainties in the critical load calculations due to the interference of sea salts and organic acids; accurate estimation under such conditions requires long-term lake and deposition chemistry.     

A test of fluctuating asymmetry in Bluegills (Lepomis macrochirus Rafinesque) as a measure of pH-related stress

Fluctuating asymmetry of bilateral anatomical characters, previously proposed as an indicator of stress induced by agents affecting the calcium metabolism of fishes, was measured in bluegills (Lepomis macrochirus) from eleven Wisconsin lakes. The fluctuating asymmetries of three meristic characters-numbers of pored lateral line scales, pectoral fin rays and gill rakers on the outer arch-were either unrelated or only marginally related to either lake pH or waterborne calcium concentration. The asymmetry variance of gill raker counts was negatively correlated with both lake pH and waterborne calcium concentration; however, the overall asymmetry mean of gill raker counts in bluegills from five low pH lakes only slightly exceeded that in bluegills from six lakes with circumneutral pH. Fluctuating asymmetry in adult fishes was judged to be insensitive as a potential measure of pH-related stress.     

Changes in trace metal concentrations in lake water and biota during experimental acidification of Little Rock Lake, Wisconsin, USA

Little Rock Lake, a small (18 ha), low-alkalinity (25 microeq litre(-1), pH 6.1) seepage lake in northern Wisconsin, was divided into two basins by a flexible, inert barrier and, beginning in spring 1985, the north basin was acidified in three 2-year steps to pH 5.6, 5.1 and 4.7. The annual average pH of the reference basin remained near 6.1. As part of a comprehensive programme to determine the chemical and biological responses to acidification, minor metals (Al, Fe, Mn) and trace metals (Cd, Cu, Pb, Zn) in lake water (0.4 microm pore filtered samples), periphyton, zooplankton, and yellow perch (Perca flavescens) were measured. At pH 5.6, dissolved Mn and Fe increased in the acidified basin. At pH 5.1 and 4.7, dissolved Al, Fe, Mn, Cd and Zn were elevated in the acidified basin. At pH 4.7, dissolved Pb in the acidified basin became elevated over reference basin levels. Dissolved Cu remained similar in both basins down to pH 4.7. Cd burdens in periphyton collected on artificial substrates were lower in the treatment basin at pH 5.1 (1.8 microg g(-1) dry wt.) than in the reference basin at pH 6.1 (7.5 microg g(-1) dry wt.), but Al and Fe burdens in periphyton were similar in both basins. Likewise, Cd levels in muscle tissue of perch from the treatment basin at pH 4.7 were lower (26 ng g(-1) dry wt.) than in the reference basin at pH 6.1 (36 ng g(-1) dry wt.); Al and Fe burdens were similar in perch muscle tissue from both basins. Levels of Cd and Fe in zooplankton from the acidified basin at pH 4.7 were approximately equal to 2x higher than in animals from the reference basin. In both basins of the lake, Al and Cd levels in lake biota decreased with increasing trophic level, demonstrating that food chain biomagnification does not occur for these metals.     

Fish status survey of Nordic lakes: effects of acidification,eutrophication and stocking activity on present fish species composition

The status of fish populations in 3821 lakes in Norway, Sweden and Finland was assessed in 1995-1997. The survey lakes were chosen by stratified random sampling from all (126 482) Fennoscandian lakes > or = 0.04 km2. The water chemistry of the lakes was analyzed and information on fish status was obtained by a postal inquiry. Fish population losses were most frequent in the most highly acidified region of southern Norway and least common in eastern Fennoscandia. According to the inquiry results, the number of lost stocks of brown trout (Salmo trutta), roach (Rutilus rutilus), Arctic char (Salvelinus alpinus) and perch (Perca fluviatilis) was estimated to exceed 10000. The number of stocks of these species potentially affected by the low alkalinity of lake water was estimated to exceed 11000. About 3300 lakes showed high total phosphorus (> 25 microg L(-1)) and cyprinid dominance in eastern Fennoscandia, notably southwestern Finland. This survey did not reveal any extinction of fish species due to eutrophication. One-third of the lakes had been artificially stocked with at least one new species, most often brown trout, whitefish (Coregonus lavaretus s.l.), Arctic char, rainbow trout (Oncorhynchus mykiss), pike-perch (Stizostedion lucioperca), grayling (Thymallus thymallus), pike (Esox lucius), bream (Abramis brama), tench (Tinca tinca) and European minnow (Phoxinus phoxinus). The number of artificially manipulated stocks of these species in Fennoscandian lakes was estimated to exceed 52000. Hence, the number of fish species occurring in Nordic lakes has recently been changed more by stockings than by losses of fish species through environmental changes such as acidification.     

Mercury content in roach (Rutilus rutilus L.) in circumneutral lakes--effects of catchment area and water chemistry

The environmental influence on the mercury content in roach (Rutilus rutilus L.) is investigated using partial least square regression on 46 environmental variables describing the land use in the catchment area, various catchment area and lake characteristics, lake water chemistry, and fish stock. The Hg content in the fish from the 78 investigated circumneutral lakes is heavily influenced by the land use in their surroundings. The boreal forest lakes possessed the highest Hg levels in roach, whereas fish from lakes surrounded by arable land had lower levels. The Hg levels also showed a negative relationship to the amount of dissolved ions and the total amount of nutrients in lake water. Lake pH did not have any significant influence on the Hg content in roach in these non-acidified lakes. The Hg levels in lakes influenced by large amounts of wetland were less well explained by the presently investigated environmental variables, which implies that the Hg burden in fish from this kind of lake is governed by other factors.     

Decreased acid deposition and the chemical recovery of Killarney,Ontario, lakes

Lakes in Killarney Park near Sudbury, Ontario, Canada, have shown dramatic water quality changes including general increases in pH and alkalinity, and decreases in SO4(2-), base cations and metals. While some lakes have recovered to pH > 6.0, many are still highly acidic despite decades of improvement. Very high historical S deposition related to emissions from the Sudbury metal smelters dominated the acidification process in this region. However, since the implementation of substantial S emission controls (90%) at the smelters, the Sudbury emissions are no longer the major source of S deposition in the Sudbury area. Wet deposition of SO4(2-) and SO4(2-) concentrations in lakewaters at Killarney now approach values in the Dorset, Ontario, area, about 200 km from Sudbury. This suggests that the S deposition to the Killarney area is now primarily from long-range transport, not from local sources. Studies of Killarney lakes are revealing the complex nature of the chemical recovery process. As lake acidity decreases, other changes including decreased Ca2+ concentrations, increased transparency, and altered thermal regimes may potentially affect some of these ecosystems. It is clear that continuing assessments of the recovery of Killarney lakes, within a multiple-stressor framework, are needed.