Reduced Acid Deposition Leads to a New Start for Brown Trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>) in an Acidified Lake in Southern Norway

Acid deposition has led to acidification and loss of fish populations in thousands of lakes and streams in Norway. Since the peak in the late 1970s, acid deposition has been greatly reduced and acidified surface waters have shown chemical recovery. Biological recovery, in particular fish populations, however, has lagged behind. Long-term monitoring of water chemistry and fish populations in Lake Langtjern, south-eastern Norway, shows that around 2008, chemical recovery had progressed to the point at which natural reproduction of brown trout (Salmo trutta) reoccurred. The stocked brown trout reproduced in the period 2008–2014, probably for the first time since the 1960s, but reproduction and/or early life stage survival was very low. The results indicate that chemical thresholds for reproduction in this lake are approximately pH?=?5.1, Al_i?=?26 μg l^?1, ANC?=?47 μeq l^?1, and ANC_oaa?=?10 μeq l^?1 as annual mean values. These thresholds agree largely with the few other cases of documented recovery of brown trout in sites in Norway, Sweden, and the UK. Occurrence and duration of acidic episodes have decreased considerably since the 1980s but still occur and probably limit reproduction success.     

Liming of lakes,rivers and catchments in Norway

Despite a slight reduction in the level of acidic deposition in Norway, acidification of lakes and rivers continues. The Norwegian Liming Project (1979–84) demonstrated that lime treatment can be an effective measure against acidification of watercourses given appropriate adaptation to local conditions. Liming in Norway is difficult because of (1) large amounts of precipitation, (2) short retention time of lakes, and (3) episodic changes in water chemistry. In 1988 NOK 14 mill. has been allocated to operational liming and research. We report here on chemical and biological responses from lime treatment of a lake, a river and a catchment. Lake Store Hovvatn was limed in 1981 and successfully stocked with brown trout. Before reliming in 1987, fish growth had ceased, but increased post liming. The River Audna has been continuously limed since 1985. Sea trout fisheries have improved, and the stocking of Atlantic salmon smolts at the mouth of the river in 1986 has already resulted in the return of spawners. Liming of the entire terrestrial catchment to the pond Tjønnstrond in 1983 by helicopter was also successful; stocked brown trout have survived to the present.     

Acid deposition and effects in Nordic Europe. Damage to fish populations in Scandinavia continue to apace

Although the decline in fish populations due to acidicwater in Norway started as early as in the 1920's the most rapid losses appeared during the 1960–70's. Until 1978, the populations of Atlantic salmon had disappeared from the southernmost part of Norway, and in these areas, more than half of the brown trout populations had been lost. Today, in spite of no increase in acid depositions, the fishery problems seems to continue at the same speed. Data based on interviews of the local fish authority shows that lakes still holding a fish population in the late 70's, have experienced a 30% loss of brown trout populations and a 12% loss of perch in the period 1978–1983. This trend have been confirmed by testfishing in lake systems having long data series. Salmon rivers on the western coast of Norway have experienced several episodes of fish kills due to rapid changes in water quality. These fish kills have mainly affected smolts of Atlantic salmon. Spawning migrating salmon on entering their acidified home river have also been affected. In Sweden, several salmon populations along the western coast have been lost due to acidification with no positive trends so far in the 1980's. Areas in central Sweden and in some high mountain areas are still experiencing a continuous and increasing acidification with detrimental effects on invertebrates and fish. In Finland, an increase in acidic deposition during the last decades have occurred, leading to acidification in the most sensitive freshwater systems. Although some acidified freshwater lakes are reported to have lost their fish stocks, few data on fish population effects are available.     

Differences in response to liming in a lake-dwelling fish community

The Lake Fjorda water system in southern Norway consists of several lakes which exhibit a gradient in acidification. The system is inhabited by populations of brown trout, Arctic char, whitefish, perch, European minnows and Crucian carp. Populations of Arctic char, whitefish and brown trout were nearly wiped out in some of the locations, as a result of acidification, In 1985, Lake Fjorda was limed in order to improve water quality so the fish community would be recovered. Fish stock assessment by means of gill-net fishing in the epibenthic and pelagic zones was carried out before (1983) and three years after liming (1991–1993). Populations of Arctic char and whitefish have not recovered after eight years of liming. Brown trout are almost extinct and do not seem to be recovering. Perch were less affected by acidification, exhibiting good recruitment also before liming.     

Changes in fish populations in southernmost Norway during the last decade

Up to 1978 51% of brown trout populations and 27% of perch populations in the four southernmost counties of Norway were lost. During 1983 the fish status of lakes in the two southernmost counties which in the period 1974 to 1978 still had fish were updated. During the period 1978 to 1983, 30% of the remaining brown trout populations and 12% of the perch populations were lost. By 1983 71% of brown trout populations and 43% of perch populations in this area have been lost. In October 1983, 623 (77%) of the lakes were sampled for analyses of water quality. The status and change in fish populations during the period from 1974–78 to 1983 were highly related to water quality status.     

Monitoring fish stocks in relation to acidification in Norwegian watersheds

The Norwegian Monitoring Programme for Long-Range Transported Air Pollutants started in 1980. The biological part of this programme includes besides invertebrate studies in streams, (i) fish community status in lakes by means of interviews, test-fishing in lakes by using standard gill-net series, recruitment studies of brown trout in inland streams, and juvenile stock assess and monitoring of fish kills in salmon rivers. Damaged fish stocks are recognized within a land area of 51,500 km² in southern Norway and 30 km² in northern Norway. At least 6,000 lake-dwelling fish stocks have either been lost or are at various stages of reduction. Brown trout (Salmo trutta) is the most widespread and abundant species of fish in Norwegian watersheds, and is also most severe affected by acidification. More recently, there are some indications of an increase in the abundance of brown trout in some areas. However, analysis of age structure in lakes, and fry densities in streams in such areas revealed large annual variations in recruitment rate, which indicates unstable water chemical conditions. Atlantic salmon (Salmo salar) is virtually extinct in 25 rivers in southernmost, southwestern and western Norway.     

Population Density of Brown Trout (Salmo trutta) in Extremely Dilute Water Qualities in Mountain Lakes in Southwestern Norway

We have examined populations of brown trout in low-conductivity mountain lakes (5.0?C13.7 ??S/cm and 0.14?C0.41 mg/l Ca) in southwestern Norway during the period 2000?C2010. Inlets to the lakes were occasionally even more dilute (2007; conductivity?=?2.9?C4.8 ??S/cm and Ca?=?0.06?C0.17 mg/l). The combination of pH and conductivity was the best predictor to fish status (CPUE), indicating that availability of essential ions was the primary restricting factor to fish populations in these extremely diluted water qualities. We suggest that conductivity <5 ??S/cm is detrimental to early life stages of brown trout, and subsequently that there are lakes in these mountains having too low conductivity to support self-reproducing trout populations. Limited significance of alkalinity, Ca, Al, and color suggests that effects of ion deficit apparently overruled the effects of other parameters.     

WHOLE-CATCHMENT LIMING AT TJØNNSTROND,NORWAY: AN 11-YEAR RECORD

In June 1983 a whole-catchment liming experiment was conducted at Tjønnstrond, southernmost Norway, to test the utility of terrestrial liming as a technique to restore fish populations in remote lakes with short water-retention times. Tjønnstrond consists of 2 small ponds of 3.0 and 1.5 ha in area which drain a 25-ha catchment. The area is located at about 650–700 meters above sea-level in sparse and unproductive forests of spruce, pine and birch with abundant peatlands. A dose of 3 ton/ha of powdered limestone were spread by helicopter to the terrestrial area. No limestone was added to the ponds themselves. The ponds were subsequently stocked with brown and brook trout. Liming caused large and immediate changes in surface water chemistry; pH increased from 4.5 to 7.0, Ca increased from 40 to 200 μeq/L, ANC increased from –30 to +70 μeq/L, and reactive-Al decreased from about 10 to 3 μmol/L. During the subsequent 11 years the chemical composition of runoff has decreased gradually back towards the acidic pre-treatment situation. The major trends in concentrations of runoff Ca, ANC, pH, Al and NO₃ in runoff are all well simulated by the acidification model MAGIC. Neither the measured data nor the MAGIC simulations indicate significant changes in any other major ion as a result of liming. The soils at Tjønnstrond in 1992 contained significantly higher amounts of exchangeable Ca relative to those at the untreated reference catchment Storgama. In 1992 about 75% of the added Ca remains in the soil as exchangeable Ca, 15% has been lost in runoff, and 10% is unaccounted for. The whole-catchment liming experiment at Tjønnstrond clearly demonstrates that this liming technique produces a long-term stable and favourable water quality for fish. Brown trout in both ponds in 1994 have good condition factors, which indicate that the fish are not stressed by marginal water quality due to re-acidification. The water quality is still adequate after 11 years and >20 water renewals. Concentrations of H⁺ and inorganic Al have gradually increased and approach levels toxic to trout, but the toxicity of these are offset by the continued elevated Ca concentrations. Reduced sulphate deposition during the last 4 years (1990–94) has also helped to slow and even reverse the rate of reacidification. The experiment at Tjønnstrond demonstrates that for this type of upland, remote terrain typical of large areas of southern Norway, terrestrial liming offers a suitable mitigation technique for treating acidified surface waters with short retention times.     

The effects of liming on juvenile stocks of Atlantic salmon (Salmo salar) and brown trout (Salmo trutta) in a Norwegian river

The effects of liming on juvenile stocks of Atlantic salmon (Salmo salar) and brown trout (Salmo trutta) in the river Vikedalselva in southwestern Norway were assessed. From 1987 to 1989, the river was limed only during the spring snow melt, and pH varied in the range between 5.5 and 7.0. In 1990 to 1993, the river was limed to pH 6.2 from 15 February to 1 June and to pH 5.7 during the rest of the year. Since 1994, the pH during late winter and spring was maintained above 6.5. Prior to liming fish kills were evident during spring snow melt, but these have not occurred since liming. Electrofishing in the autumn between 1981 and 1994 showed no significant change in densities of juvenile salmon and brown trout after liming, mean densities ranged between 19–50 and 9–32 individuals 100 m^–2 respectively. A significant linear correlation between production and biomass of both species was found, indicating that factors directly controlling density affect juvenile production and cause production to remain below carrying capacity. In spite of a clear increase in pH and a reduction in the concentration of labile aluminium after liming, the conditions still do not seem to be optimal for juvenile salmonids. We suggest that a complexity of different factors impose limitations on fish production in the river: inadequate egg deposition, environmental factors such as water temperature and flow, osmoregulatory failure in mixing zones between limed and acidic water and gill damage through deposition of aluminium and iron. However, there are several indications of a reduction in toxic effects after the pH was raised to 6.5 during spring snow melt.     

Relationship between fish populations and pH for lakes in Southermost Norway

Fish status in terms of ‘good, ‘sparse’, ‘lost’, ‘never had fish’, has previously been reported for several thousands of lakes in southermost Norway. In more than a thousand of these lakes pH and conductivity have also been measured. These data have been used to establish a relationship between pH and fish status (brown trout). It is estimated that a uniform pH increase of 0.2 units will result in status changes from category ‘lost’ to ‘sparse’ in 12% of the lakes (27% of the lakes which have lost the fish population). Altogether 21 % of the lakes are predicted to change to a better category. We have used a fixed pH shift in order to make the approach applicable. This is a rather drastic simplification since the lakes will respond very differently to a reduction in S deposition depending on the original acidity and a number of other factors. The limitations of the approach and an alternative method used by Chester (1982) are discussed in detail.     

Recovery of Young Brown Trout in Some Acidified Streams in Southwestern and Western Norway

Temporal changes in densities of young brown trout (Salmo trutta), mainly of age 0+, and in water quality (pH and alkalinity) were assessed by means of electrofishing in lake tributaries in three acidic, software watercourses in western and southwestern Norway; Gaular and Vikedal (1987–1999) and Bjerkreim (1988–1999). Approximately 74 sites were sampled each year. Most of the streams were acidic with mean annual pH levels between 5.1–5.9. Alkalinity and pH increased significantly in all three areas during the study period. Brown trout fry densities increased significantly during the period in Vikedal and Bjerkreim. Also in Gaular, the density of young brown trout has exhibited a positive trend in recent years. We suggest that the increase in the density of young brown trout is because the study areas have became less acidified during recent years due to reduction in sulphate deposition.     

Acid deposition from the Russian Kola Peninsula: Are sensitive fish populations in north-eastern Finnish Lapland affected?

Sulphur emissions from Russian Kola Peninsula smelters are known to cause surface water acidification in the border areas between Norway and Russia. The sulphur deposition is also high in the eastern part of Finnish Lapland. In 1990, a monitoring programme was started to survey the effects of acid deposition on sensitive fish populations in north-eastern Finnish Lapland. Altogether 103 sites in three areas were electrofished and autumn water samples were taken. Besides the brown trout (Salmo trutta), special attention was paid to the occurrence of minnow (Phoxinus phoxinus) since it is a common species in small waters and is highly sensitive to acidification. During the first three years of monitoring no signs of acidification were recorded. The alkalinity values of brooks generally exceeded 0.1 mmol/1. Brown trout, minnow and burbot (Lota lota) were caught frequently in the study sites. Later the study was focused on the uninhabited Vätsäri area which is receiving the highest sulphur deposition in Finnish Lapland. The alkalinity values of the sampled brooks were in most cases below 0.05 mmol/1, indicating a decreased buffer capacity. However, the electrofishing of the brooks showed no acid-induced damage. The lowest alkalinity values were detected from a group of small upland ponds. In four such ponds the alkalinity was zero or negative. No minnows were caught from these four ponds apart from one, where the minnows were exceptionally large. The results show that the waters near the eastern border of northern Finnish Lapland are threatened by acidification. No damage to fish populations subject to fishing was observed. The absence of minnows in some small waters is possibly the first sign of acid-induced fish population damage.     

Effects of acidity on fish communities in southwestern Québec (Canada)

Surveys of fish populations were conducted on 74 lakes of the Outaouais hydrographic region during the summers of 1985 and 1986 to assess the potential impact of acidity on ichtyologic fauna. Results show that species diversity declined with the increasing acidity. The color of water does not seem to mitigate the adverse effects of acidity. On the contrary, the number of species decreased similarly in both brown lakess (>30 Hazen) and clear lakes (≤30 Hazen) with the increase of acidity. The species tolerance threshold levels to acidity show that 72% of fish species are no longer captured when pH reaches 5.0, compared to 32% at pH ≤5.5. The pH range 5.0 to 5.5 can possibly be regarded as the break point for the occurrence of most fish species in this area. Analysis of the size frequency distributions show that recruitment failures have occurred in acidic waters for walleye and lake trout. We have estimated that anthropogenic acidification is responsible for the loss of more than 10000 fish populations in the Outaouais area.     

Impact of acid precipitation on freshwater ecosystems in Norway

Extensive studies of precipitation chemistry during the last 20 yr have clearly shown that highly polluted precipitation falls over large areas of Scandinavia, and that this pollution is increasing in severity and geographical extent. Precipitation in southern Norway, Sweden, and Finland contains large amounts of H⁺, SO⁼ ₄, and NO^? ₃ ions, along with heavy metals such as Cu, Zn, Cd, and Pb, that originate as air pollutants in the highly industrialized areas of Great Britain and central Europe and are transported over long distances to Scandinavia, where they are deposited in precipitation and dry-fallout. In Norway the acidification of fresh waters and accompanying decline and disappearance of fish populations were first reported in the 1920s, and since then in Sørlandet (southernmost Norway) the salmon have been eliminated from several rivers and hundreds of lakes have lost their fisheries. Justifiably, acid precipitation has become Norway's number-one environmental problem, and in 1972 the government launched a major research project entitled ‘Acid precipitation — effects on forest and fish’, (the SNSF-project). Studies of freshwater ecosystems conducted by the SNSF-project include intensive research at 10 gauged watersheds and lake basins in critical acid-areas of southern Norway, extensive surveys of the geographical extent and severity of the problem over all of Norway, and field and laboratory experiments on the effect of acid waters on the growth and physiology of a variety of organisms. Large areas of western, southern, and eastern Norway have been adversely affected by acid precipitation. The pH of many lakes is below 5.0, and sulfate, rather than bicarbonate, is the major anion. Lakes in these areas are particularly vulnerable to acid precipitation because their watersheds are underlain by highly resistant bedrock with low Ca and Mg contents. Apart from the well-documented decline in fish populations, relatively little is known about the effects of acid precipitation on the biology of these aquatic ecosystems. Biological surveys indicate that low pH-values inhibit the decomposition of allochthonous organic matter, decrease the species number of phyto-and zooplankton and benthic invertebrates, and promote the growth of benthic mosses. Acid precipitation is affecting larger and larger areas of Norway. The source of the pollutants is industrial Europe, and the prognosis is a continued increase in fossil-fuel consumption. The short-term effects of the increasing acidity of freshwater ecosystems involve interference at every trophic level. The long-term impact may be quite drastic indeed.     

Human exposure to mercury may decrease as acidic deposition increases

It has been hypothesized that human mercury (Hg) exposure via fish consumption will increase with increasing acidic deposition. Specifically, acidic deposition leads to reduced lake pH and alkalinity, and increased sulphate ion concentration ([SO₄ ^2?]), which in turn should cause increased Hg levels in fish, ultimately resulting in increased human Hg exposure via fish consumption. Our empirical test of this hypothesis found it to be false. We specifically examined Hg levels in the hair of Ontario Amerindians, who are known consumers of fish from lakes across the province, and observed a weak negative association with increasing sulphate deposition. An examination of Hg levels in lake trout, northern pike and walleye, three freshwater fish species commonly consumed by Ontario Amerindians, found a similar weak negative association with increasing sulphate deposition. Further analysis of these fish data found that fish [Hg] was most significantly (positively) associated with lake water concentrations of dissolved organic carbon (DOC), not pH, alkalinity or [SO₄ ^2?]. Lake DOC levels are lower in regions of greater acidic deposition. We propose an alternate hypothesis whereby human Hg exposure declines with increasing acidic deposition. In particular, we propose that increasing sulphate deposition leads to reduced lake DOC levels, which in turn leads to lower Hg in fish, ultimately reducing human Hg exposure via fish consumption.     

The REFISH (restoring endangered fish in stressed habitats) project, 1988–1994

The REFISH (Restoring Endangered Fish In Stressed Habitats) Project was established in 1988 to assess acid-tolerance among indigenous Norwegian strains of brown trout. The work, comprising both laboratory and field studies, has involved the restocking and subsequent test-fishing of thirteen lakes with five brown trout strains. There was considerable variation in the ability of individual lakes to support adult trout. This did not appear related to ANC (acid neutralising capacity) or any single chemical factor. One strain, Bygland, was found to be relatively acid-tolerant, accounting for more than 60% of all fish recaptured by test-fishing over 1990–1994. This is consistent with better survival of young life-stages of the Bygland strain, compared with that of the other strains, in laboratory experiments employing acidic conditions. Strainspecific differences in calcium metabolism may be the physiological basis for acid tolerance.     

Liming of acidified lakes: Induced long-term changes

This study presents data concerning long-term trends after neutralization of four acidified lakes in two regions on the Swedish west coast. Neutralization was achieved by a di-Ca-silicate with 52% CaO and about 11.5% MgO. Between 61 and 74% of the spread lime product dissolved during a 5 to 7 yr period. The liming increased pH, from a range of 4.5 to 5.2 to near neutral and restored alkalinity in the range of 0.2 to 0.3 meq l^?1 and the Ca-content became 3 to 4 times higher than before liming. In two lakes transparency decreased significantly presumably due to changed phytoplankton composition. These changes successively declined due to dilution and continuous acid loading. The changes in water chemistry and development of stocked brown trout (Salmo trutta) populations initiated biotic changes. Phyto- and zooplankton communities reacted both instantly and later with successions in species composition. Changes of benthic macroinvertebrate species occured over several years, but some pelagic species, e.g. corixids were rapidly reduced due to predation of fish. Observed changes were predominantly due to expanding populations of species present at very low abundances even during acid state of the lakes. Some organisms found during preacid state of the lakes did not establish new populations and this process may need a prolonged time with favorable conditions. Reacidification towards the end of the study period significantly stressed the brown trout population and also favored expansion of the filamentous algaMougeotia sp. andSphagnum sp. that almost vanished during the first year after liming. Decreasing concentration of total P was not influenced by neutralization and may be mostly dependent on negative changes in the soils surrounding the lakes. If generally valid, this process may be an important factor for the oligotrophication of lakes in areas where acid deposition is high.     

Improved growth in stunted brown trout (Salmo trutta L.) after reliming of Lake Hovvatn,Southern Norway

The chronically acidic Lake Store Hovvatn and the adjoining pond Pollen in southernmost Norway were limed in March 1981. The two locations were stocked with brown trout (Salmo trutta L.) at low and high densities in Hovvatn and Pollen, respectively. After 6 yr of reacidification, the locations were relimed in July 1987. Growth depression during the reacidification process in spite of low fish densities and superabundance of food was observed in Lake Store Hovvatn. Three months after reliming, a substantial growth response was found in trout from Lake Store Hovvatn; Mean annual length increment was 68% higher than that of the preceding year. In Pollen, reliming had no apparent effect on growth. In both populations reliming caused increased swimming activity measured as an increase in CPUE-values. These results show that the growth response to liming depends on population density and food availability. Moreover, the results indicate that the food conversion rate of the trout is negatively affected in acid waters.     

Evidence of fish population responses to acidification in the Eastern United States

The hypothesis that acidification has reduced or eliminated fish populations in certain areas of the eastern United States was investigated by examining present and historical fishery survey records. The causes of acidification (e.g., atmospheric deposition) were not specifically considered, although instances of obvious alternative explanations (e.g., acid mine drainage, organic acids) were avoided. The number of usable data sets located was small. Trend analyses are severely limited by the lack of high quality historical data. The strongest evidence for fisheries declines associated with acidification is provided by data for the Adirondack Mountain region of New York. In some lakes, fish populations have declined or disappeared; lakes experiencing fishery declines are now acidic. Alternative explanations for changes in fish communities over time were examined. In 49 lakes, some or all fish populations have apparently been lost with no available explanation other than acidification. Extrapolation of these data to the entire Adirondack Mountain region suggests that perhaps 200 to 400 lakes may have lost fish populations as a result of acidification. Streams in Pennsylvania and Massachusetts also had documented declines in fish populations that were associated with acidity; however, the data are fewer and less complete than those for New York. Acidification effects on fishery resources in other regions of the eastern United States are apparently minimal. The extent of the damage to date appears small relative to the total resource.     

Liming the acid lake Hovvatn,Norway: A whole-ecosystem study

Hovvatn, a 1-km² chronically-acidified lake in southernmost Norway, was treated with 200 tonne of powdered limestone in March 1981. An additional 40 tonne were added to a 0.046 km² pond (Pollen) draining into Hovvatn. The lakes were stocked with brown trout in June 1981 and in each subsequent year. At ice-out pH rose from 4.4 to 6.3 (Hovvatn) and 7.5 (Pollen), Ca and alkalinity increased, and total Al decreased by 120 μg L^?1. None of the other major ions exhibited significant changes in concentration. Total organic Cand Pincreased after liming. The phytoplankton community was dominated by chrysophytes and did not change significantly following liming. The zooplankton community was typical of acid lakes prior to liming. There was a clear succession in species dominance following treatment, although no new species immigrated to the lakes. Zoobenthos changed from a community characterized by low abundance and reduced number of species to increased abundances of oligochaetes, mayflies and chironomids. Hovvatn and Pollen were barren of fish prior to stocking. The stocked fish showed remarkably high growth rate during the first years. Liming apparently improved conditions for zoobenthos, enhancing the processing of fine detritus which in turn resulted in elevated levels of TOC and P in the lakewaters during the first year after liming. The “oligotrophication” process typical of acid lakes was temporarily reversed by liming. The interactions between groups of organisms in Hovvatn and Pollen indicates that many years are required before a new steady-state can be attained following liming.