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.     

Acute effects on perch (Perca fluviatilis) and long-term effects on whitefish (Coregonus lavaretus pallasi) of liming of an acidified lake

The acute effects on perch and long‐term effects on whitefish were studied after liming one side of a lake [initial pH: 4.6–5.5; aluminium (Al): 29–54 μg L^?1; Ca²⁺: 0.02–0.07 mmol L^?1] that was divided into two parts with a plastic curtain. Some Al precipitation was observed on the gill surface of perch 1 day after liming; the concentration of plasma sodium of perch females in the limed side was also higher than in the acidic side. Nearly 6 months after liming, at the spawning time of whitefish, the plasma chloride concentration of whitefish in the limed side was higher and blood glucose concentration lower than in the acidic side. Four of five whitefish males from the limed side, but only one of five males and none of the five females from the acidic side, were ready to spawn. The growth of whitefish in the limed side was also more rapid. These changes illustrate that liming had no acute harmful effects on perch, and allowed whitefish to recover from acidity‐related physiological stress.     

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

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

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.     

Long‐term effects of liming in Norwegian softwater lakes: the rise and fall of bulbous rush (Juncus bulbosus) and decline of isoetid vegetation

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Role of magnesium in combination with liming in alleviating acid-soil stress with the aluminium-sensitive sorghum genotype CV323

An experiment to study the effects of Mg nutrition on root and shoot development of the Al-sensitive sorghum (Sorghum bicolor (L.) Moench) genotype CV323 grown in pots of sandy loam under different acid soil stress is reported. This experiment had a factorial design: four rates of liming were combined with four rates of Mg fertilization. When no Mg was added, the pH of the soil solutions (collected in ceramic cups) increased from 4.0 (unlimed) to 4.2, 4.7 and 5.9 at the increasing rates of liming. After 30 days of growth dry matter yields of the limed treatments were 40%, 115% and 199% higher than that of the unlimed treatment. Without liming and at the highest liming rate, adding Mg did not affect plant biomass significantly. At the two intermediate levels of liming, however, 11.3 mg extra Mg per kg soil increased dry matter yield to the same levels as found at the highest liming rate. Concentrations of Mg in the soil solution rose after Mg was added and fell when lime was added, but adding both Mg and lime increased Mg concentrations in the plant shoots. In plants of the limed treatments, dry matter yield was correlated closely with the Mg concentration in the shoot. This was not so in the unlimed treatment. Furthermore, in the unlimed treatments root development was inhibited, but reduced Mg uptake by the plants resulted mainly from the direct effect of Al- (or H-) ions in the soil solution rather than from impaired root development. It is concluded that Mg fertilization counteracted the interfering effects of Al- and H ions on Mg uptake.     

Lake responses to long‐term disturbances and management practices

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Odonates as Indicators of Shallow Lake Restoration by Liming: Comparing Adult and Larval Responses

Odonate assemblages were compared between replicate sets of shallow lakes that had been created and acidified by open‐cast mining across a large area (2,451 ha) of southwest France (Arjuzanx, Landes); one set of lakes (n = 5) was experimentally restored by liming with calcium carbonate, whereas another group (n = 5) was left as untreated reference lakes. Both odonate adults and exuviae were sampled bimonthly during May–August 1998. Elevated turbidity and conductivity in limed lakes were the only physicochemical measures differing between restored and reference lakes, because deacidification occurred naturally, even in reference lakes during the 17 years after the onset of restoration. Restoration by liming can apparently lead to effects on lake turbidity that might be considered adverse. Twenty‐four and 19 odonate species occurred among adults and exuviae, respectively, but there were no significant differences in richness between restored and reference sites. However, significantly, more exuviae were collected from the reference sites (588 vs. 180), where exuvial diversity and rank abundance indicated more evenly structured assemblages than those in restored lakes. Ordination showed that adult assemblages differed significantly between restored and reference lakes, and varied highly significantly with lake turbidity. This effect occurred because a small group of generally scarce adults were characteristic of reference sites (Chalcolestes viridis, Lestes virens, Cordulia aenae, Leucorrhinia albifrons, and Sympetrum sanguineum). Exuviae of these same species were less abundant at restored sites, but exuvial assemblages overall did not discriminate between restored and reference lakes. We conclude that lake restoration by liming can reduce diversity and larval numbers among odonates and subtly affects adult assemblages. In this case study, adult assemblages discriminated best between the lake types involved in the experiment, but important additional information arose from exuvial abundance and structure. This study indicates that natural recovery processes after acidification in formerly open‐cast areas––rather than chemical intervention through liming––might lead to preferable conservation outcomes.     

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

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

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

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

Long-term effects of catchment liming on invertebrates in upland streams

1. Catchment liming to mitigate acidification causes major chemical change in freshwaters but longer‐term effects are poorly understood. Using a replicated basin‐scale experiment with a multiple BACI design (= before‐after‐control‐impact), we assessed chemical and biological effects for 10 years after the catchments of three acidified Welsh streams at Llyn Brianne were limed in 1987/88. 2. Stream chemistry was measured weekly to monthly, and macroinvertebrates monitored annually, between 1985 and 1998. Biological change through time was assessed from the abundance and taxon richness of invertebrates. We paid particular attention to 18 species known to be acid‐sensitive. The effects of liming were assessed by comparing chemical and biological trends among the three replicate limed streams, three acid reference streams and two naturally circumneutral streams. 3. Following single lime applications, acid‐base chemistry in treated streams changed significantly. High mean pH (> 6), increased calcium (> 2.5 mg L^?1) and low aluminium (< 0.1 mg L^?1) persisted throughout the 10 years following liming. 4. The effects of liming on invertebrates were modest. Acid sensitive taxa increased significantly in abundance in limed streams, but only during 2 years following treatment. Significant effects on richness were more sustained, but on average added only 2–3 acid‐sensitive species to the treated streams, roughly one‐third of their average richness in adjacent circumneutral streams. Only the mayfly Baetis rhodani and the stonefly Brachyptera risi occurred significantly more often in limed streams after treatment than before it. 5. Despite these modest long‐term effects on invertebrates, nearly 80% of the total pool of acid‐sensitive species has occurred at least once in the limed streams in the 10 years since treatment. This pattern of occurrence suggests that the colonization of limed streams by acid‐sensitive taxa reflects limited persistence rather than restricted dispersal. We present evidence to show that episodes of low pH continued to affect acid‐sensitive taxa even after liming. We highlight the importance of extending the time‐periods over which the effects of large‐scale ecological experiments are assessed.     

Influence of dolomitic lime on DOC and DON leaching in a forest soil

The influence of liming on leaching and distribution of dissolved organic carbon (DOC) and nitrogen (DON) in mineral soil was investigated in a leaching experiment with soil columns. Soil samples from separate horizons (O, A and B horizons) were collected from control and limed plots in a field liming experiment in a spruce forest in southern Sweden. The field liming (0.88 kg m^-2) had been carried out 8 years before sampling. To minimize the variation among replicates, soil profiles were reconstructed in the laboratory so that the dry weight was the same for each individual soil horizon regardless of treatment. Two soil column types were used with either the O+A horizons or the O+A+B horizons. One Norway spruce seedling (Picea abies (L.) Karst) was planted in each soil column. Average pH in the leachate water was greater in the limed treatment than in the control treatment (5.0 versus 4.0 for O+A columns and 4.3 versus 3.8 for O+A+B columns). After reaching an approximate steady state, the leaching of DOC was 3--4 times greater from the limed O+A and O+A+B columns than from the corresponding control columns but the leaching of DON increased (3.5 times) only in the limed O+A columns. There was a significant correlation between DOC and DON in the leachates from all columns except for the control O+A+B columns, which indicated a decoupling of DOC and DON retention in the B horizon in the control treatment. This might be explained by a selective adsorption of nitrogen poor hydrophobic compounds (C/N ratio: 32--77) while there was a lower retention of nitrogen rich hydrophilic compounds (C/N ratio: 14--20). Proportionally more hydrophobic compounds were leached from the limed soil compared to the unlimed soil. These hydrophobic compounds also became more enriched in nitrogen after liming so in the limed treatment nitrogen might be adsorbed at nearly the same proportion as carbon, which might explain the fact that there was no decoupling of leached DOC and DON from the B horizon after liming.     

Advances in the positional cloning of nodulation genes in soybean

The effect of liming on the decomposition of Norway spruce needle litter was studied in 40–60-year-old Norway spruce stands. Finely-ground limestone had been spread about 30 years ago at a dose of 2 t ha^–1 and reliming was carried out about 20 yr later at a dose of 4 t ha^–1. Needle litter was collected from both control and limed plots, and it was placed in litter bags in the middle of the humus layer of the plot from which they originated, and similarly to the other plot in May. Litter bags were sampled after 4, 12 and 16 months. The site of origin of the needle litter, whether from control plot or from limed plot, affected mainly the early stages of decomposition. Initially the effect of liming was seen as decreased concentration of water soluble material and then, during decomposition, as decreased mass loss and decreased degradation of lignin, and increased C/N ratio. The incubation site, whether the control or the limed plot, did not affect decomposition significantly.Decomposition of Scots pine needles in a young Scots pine plantation was also studied. The treatments were: 2 t ha^–1 of finely-ground limestone and 2.5 t ha^–1 of bark ash spread 8 months before this study. The treatments did not affect decomposition much, but some stimulation of the treatments on decomposition was observed. Compared to spruce needles, the C/N ratio of pine seedles was lower, they contained less lignin and more water soluble material, and decomposed faster in the first summer.     

Nutrient responses to the liming of lake Gårdsjön

The acidified lakes Lake Gårdsjön and Lake Stora Hästevatten the reference lake have been monitored since 1979 and 1980 respectively. The lakes are situated in SW Sweden; in an area severly affected by acid deposition. Lake Gårdsjön was limed in spring 1982. This paper analyses changes in nutrient concentrations upon liming of Lake Gårdsjön. The liming of Lake Gårdsjön was followed by a slight increase in ammonium, nitrate, and dissolved organic nitrogen concentrations. A drastic decrease occurred in particulate nitrogen and particulate carbon, whereas dissolved organic carbon increased. Total phosphorus and particulate phosphorus concentrations were similar to pre-limed conditions. The long-term decrease in phosphorus concentration, exhibited by the reference lake, was not identified in Lake Gårdsjön after liming, but total phosphorus concentration was still less than half compared to Lake Gårdsjön in the early 1970's. Additional measures such as phosphorus fertilization, should in certain cases be considered in addition to liming if the goal is to restore lakes to their pre-acidic conditions.     

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

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

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.     

Temporal distribution of perch (Perca fluviatilis L.) in a lake-reservoir (Moselle,France): analysis of catches with vertical gill nets

Four fishing periods within a yearly cycle between 1990 and 1991 were carried out in the lake-reservoir of Mirgenbach, lake of the nuclear power station in Cattenom (Moselle, France). The catches of perch (Perca fluviatilis L.) were analyzed.In all, about 630 perch were caught in 8 sampling stations. The analysis of seasonal catches of this carnivorous fish was made according to size and mesh; a simple correspondence factor analysis enabled us to establish that this species lived in different areas depending on the season.     

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

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

Dissolved organic carbon, seston and macroinvertebrate drift in an acidified and limed humic stream

SUMMARY. 1. Total seston, and invertebrate drift were studied before and after lime addition to Fyllean River, a stream-iake system in Halland county, southwest Sweden, with poorly buffered waters undergoing acidification. 2. The largest effect of liming was on the chemistry of the water. Following liming with 23 mg CaCO₃ l^?1 the pH of the water changed from 5.8 to 6.8 and alkalinity from 0.04 to 0.13 meq l^?1.Turbidity increased from 3.4 to 4.7 JTU with no change in colour. 3. Dissolved organic carbon (DOC) concentration of all samples was in the range 10.7–13.3 mg C l^?1 with no significant change occurring due to liming. 4. Total seston increased from 4.35 mg DM 1^?1 in unlimed conditions to 6.25 mg DM l^?1 after lime addition. All significant changes in seston occurred in the smaller size fraction (0.45–25 μm). 5. Liming reduced the organic content of the partieulate material from an average of 61% to 39% immediately downstream of a lime silo (within 1 km) but had little effect when the river course was interrupted by lakes and impoundments. 6. The lakes in the river system had a larger effect on seston concentration than any effect of the lime addition by itself. Particle concentrations were reduced by 50–55% and DOC by about 1 mg C l^?1as the water passed through the lakes. 7. Macroinvertebrate drift density was low in all samples before and after liming and typical of oligotrophic streams. Drift was significantly lower at limed (0.024 ind. m^?3) than at unlimed (0.083 ind. m^?3) locations. The decrease was only in total drift density with no significant change in the relative abundance of functional groups or in densities of single taxa, except for a reduction in drift of predators in the limed condition.     

Birch leaf processing and associated macroinvertebrates in an acidified lake subjected to liming

In connection with the liming of an acid lake in southern Norway, a series of litter bags was placed in a pH-gradient in the limed lake and a nearby unlimed lake. During the experiments, which lasted two years, no significant differences in decomposition rates between the various localities at the same depth were noticed. The chironomid collector Tanytarsus pallidicornis was the dominant invertebrate species in the benthic samples. This demonstrates the importance of fine particulate organic matter as a food source in the lake. Collector dominance was lower in the leaf packs, which consisted mostly of coarse particulate organic matter. A poor fauna of detritus grazers probably contributed to the unchanged rates of decomposition of the leaves after liming.