Restoration of Zooplankton Communities in Industrially Damaged Lakes: Influences of Residual Metal Contamination and the Recovery of Fish Communities

The Sudbury, Ontario, Canada area offers a unique opportunity to develop our understanding of biotic and abiotic lake recovery processes in industrially damaged natural systems. In recent decades, lakes in the Sudbury area have shown improvements in water quality due to decreases in sulfur (S) and metal emissions from area smelters, and reduced acid deposition from more distant sources. However, biological recovery is lagging and mechanisms controlling the lag are not yet clear. Our study examines the roles of two factors, residual metal contamination and altered fish predation, on zooplankton community recovery. Data collected over three decades on six study lakes were analyzed using redundancy analysis (RDA) and partial RDA's to assess historical and present relationships of water chemistry and fish abundance with zooplankton community recovery. Continuing metal toxicity appears to be the primary cause of the absence of some zooplankton species, particularly Daphnia spp. from metal‐contaminated lakes. Conversely, once water quality is suitable and abundant planktivores reestablish, fish planktivory becomes a factor affecting Daphnia spp. establishment. The introduction of piscivores into these lakes may be necessary to facilitate the return of many Daphnia species. Further reductions in metal toxicity will also assist with the complete recovery of zooplankton communities. Studying natural systems over several decades allows us to better understand the intricate steps involved with recovery of industrially damaged lakes, and this knowledge will greatly benefit future restoration efforts in other industrially damaged systems .     

Elevated metal concentrations inhibit biological recovery of Cladocera in previously acidified boreal lakes

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Long-term trends in limnological characteristics in the Aurora trout lakes, Sudbury, Canada

Paleolimnological techniques were employed to document the limnological histories of the aurora trout lakes, located in the Sudbury region of Ontario. Two of these lakes are of special interest to fisheries managers, as they represent the only known native habitats of a rare strain of brook trout: the aurora trout. These lakes were limed as part of restoration efforts. Stratigraphic changes in diatom and chrysophyte assemblages from dated lake sediment cores indicate that all the lakes have been impacted by anthropogenic acidification, although the timing and the magnitude of acidification were different amongst the lakes. For example, Whirligig Lake was likely the most naturally acidic lake in the past, but it had further acidified since about 1960. This lake was limed in 1989 and then again in 1993. In Whitepine Lake, acidification started 1940; however, in the most recent sediments (1992), some recovery in lakewater acidity has occurred. In Little Whitepine Lake (a reference lake), acidification started earlier (1920) and the lakewater pH continued to decline until about 1990. This lake was limed in 1989. The chrysophyte paleoindicators suggest a recent recovery in this lake. The successful re-introduction of aurora trout in Whirligig and Whitepine lakes is undoubtedly related to the improved water quality through liming but, based on our paleolimnological indicators, the lakes' limnological characteristics (e.g. pH and metal concentrations) are still different from those present before atmospheric deposition of strong acids from the Sudbury smelters.     

Tracking Recovery Patterns in Acidified Lakes: A Paleolimnological Perspective

Long-term data are often lacking to effectively assess patterns of lake acidification and recovery. Fortunately, paleolimnological techniques can be used to infer past changes in lakewater acidity and related variables by means of biological indicators, such as diatom valves and chrysophyte scales, preserved in ²¹⁰Pb-dated sediment cores. We summarize paleolimnological data that we have gathered from 36 Sudbury (Ontario) and 20 Adirondack Park (New York) lakes to estimate the magnitude of lake acidification and any subsequent recovery in these lake systems. In both regions, many lakes were shown to have acidified considerably, some over two pH units, since the 1850s. Although some recovery was noted in both lake regions, Sudbury lakes generally showed larger increases in inferred lakewater pH with recent declines in sulfur emissions. Possible explanations of these differences include the greater decrease in sulfate deposition in the Sudbury area, as well as generally longer residence times of lakes in Sudbury, perhaps allowing for more in-lake alkalinity generation. In addition, Sudbury lakes generally had higher pre-industrial pH levels, suggesting that lakes with higher natural buffering capacities are more likely to recover more quickly with declines in deposition, even if they had been acidified to a great extent.     

Assessment of Littoral Benthic Invertebrate Communities at the Land–Water Interface in Lakes Recovering from Severe Acid- and Metal-Damage

Benthic invertebrate communities within confluence sites, or areas of sediment deposition, are shaped by the input of catchment products including coarse woody debris, organic and inorganic particulates, and contaminants, but these sites also appear to be potential “hotspots” where recolonization of severely damaged ecosystems begins. Two species of leaf packs and a sweep netting technique were used to assess benthic invertebrate communities across a gradient of 14 confluence sites in 3 recovering lakes near the copper and nickel smelters in Sudbury, Canada. Environmental variables including delta habitat composition, delta area and length, and composition of deposited materials were used to detect spatial patterns in littoral benthic invertebrate communities. Benthic invertebrate community relationships with water chemistry were also assessed. Partial redundancy analysis (pRDA) showed that all sampling methods detected similar gradients of increasing invertebrate community richness and diversity as area and length of the sediment delta and the surface organic matter abundance increased. Two-way nested ANOVAs showed significant differences (p < .05) in taxa richness and diversity metrics among sites. Of the three methods, the benthic invertebrate community measurements from the birch leaf packs provided the strongest correlations with measures of organic matter inputs or habitat characteristics of the confluence zones. These correlations suggest that tree planting in riparian areas, or organic matter or macrophyte additions to littoral zones, may enhance littoral benthic invertebrate richness and diversity in acid and metal damaged lakes.     

Abiotic and biotic processes in lakes recovering from acidification: the relative roles of metal toxicity and fish predation as barriers to zooplankton re‐establishment

1. Recovery of acidified aquatic systems may be affected by both abiotic and biotic processes. However, the relative roles of these factors in regulating recovery may be difficult to determine. Lakes around the smelting complexes near Sudbury, Ontario, Canada, formerly affected by acidification and metal exploration, provide an excellent opportunity to examine the factors regulating the recovery of aquatic communities. 2. Substantial recovery of zooplankton communities has occurred in these lakes following declines in acidity and metal concentrations, although toxicity by residual metals still appears to limit survival for many species. Metal bioavailability, not simply total metal concentrations, was very important in determining effects on zooplankton and was associated with a decrease in the relative abundance of cyclopoids and Daphnia spp., resulting in communities dominated by Holopedium gibberum. 3. As chemical habitat quality has improved and fish, initially yellow perch and later piscivores (e.g. smallmouth bass, walleye), have invaded, biotic effects on the zooplankton are also becoming apparent. Simple fish assemblages dominated by perch appear to limit the survival of some zooplankton species, particularly Daphnia mendotae. 4. Both abiotic (residual metal contamination) and biotic (predation from planktivorous fish) processes have very important effects on zooplankton recovery. The re‐establishment of the zooplankton in lakes recovering from stress will require both improvements in habitat quality and the restoration of aquatic food webs.     

Recovery of copepod, but not cladoceran, zooplankton from severe and chronic effects of multiple stressors

In the mid‐twentieth century, many lakes near Sudbury, Canada, were severely contaminated by acid and metal emissions from local smelters. For example, in the early 1970s, Middle Lake had pH of 4.2, and Cu and Ni levels both >0.5 mg L^?1 . To determine if crustacean zooplankton could recover from such severe and chronic damage, Middle Lake was neutralized in 1973. A comparison of its zooplankton with that of 22 reference (pH > 6) lakes indicates that the planktonic Copepoda completely recovered by 2001. In contrast, the cladoceran assemblage improved but did not recover. Colonist sources existed – Cladocera and Copepoda occurred with equal frequency in area lakes – but six separate colonizations by cladoceran species failed. We argue that local factors, metal toxicity and predation by yellow perch, have, to date, prevented cladoceran recovery. Nonetheless, the complete copepod recovery is encouraging, given the severity and duration of pre‐neutralization stress.     

Acid rain — perspectives on lake recovery

During the 1970‘s and 1980‘s, the acidification of surface waters by atmospherically deposited sulphur became a major international concern. Large sulphur emission control programs were implemented in Europe and North America with the expectation that many affected aquatic ecosystems would recover. Because of a variety of factors, these positive expectations have been slow to be realized. Only limited evidence of the chemical recovery of acid lakes has emerged from areas other than the Sudbury, Canada region, where sulphur emission reductions were particularly large. Lake response models indicate that when current sulphur emission control strategies in Europe and North America are fully implemented, many lakes will still be acid-damaged even though substantial overall improvements in lake chemistry are expected. An increasing body of evidence indicates that substantial biological recovery, among many groups of organisms,can be expected when chemical conditions improve in lakes. Not all species, however, are capable of unassisted recovery and some lakes can pose biological or physical barriers to colonizers. Thus, stocking may be an important element in management strategies for the restoration of some recovering lakes. Communities in recovered lakes may not achieve pre disturbance conditions, but establishment of typical communities appears to be a reasonable recovery target. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effects of Copper,Nickel, and Sulfate from the Smelters at Sudbury,Ontario (Canada) on Microbial Communities in Lakes

Analysis of water and sediments from deep and shallow environments in lakes located 6–154 km east or southeast of the smelters at Sudbury, Ontario (Canada) revealed variable, interactive effects of copper, nickel, and sulfate from smelter fallout on lacustrine microfloras. Metal species in sediments were differentiated by sequential extractions, and the nature, abundances, and activities of microbial populations were represented by chlorophyll-a in water and by CO₂ production, fatty acids, phospholipids, dehydrogenase, alkaline phosphatase, and spectral properties of humic matter in sediments. Smelter fallout declined logarithmically with distance from the smelters, and its effects on microfloras depended on the type of microorganism or microbial process and on environmental factors and the abundances of metal species and detoxifying agents. Extractable copper and nickel had toxic effects, which were not attributable solely to the exchangeable fractions, but in certain cases nickel counteracted copper toxicity. Sedimentary sulfide as a whole or sulfide produced by bacterial sulfate reduction, or low oxidation–reduction potential regardless of sulfide concentration, ameliorated metal toxicity by making the metals less bioavailable; and toxicity showed a “quantum jump” when detoxifying agents fell below certain critical concentrations, implying the existence of threshold levels of bioavailable metals above which toxicity increased abruptly. In some cases metal toxicity was lowest in the lakes closest to the smelters (because sulfate concentrations were highest) as well as in the lakes furthest away, and was highest at intermediate distances. The results also suggest that nickel pollution led to ecological succession whereby nickel-tolerant microbial populations replaced nickel-sensitive ones.     

Acid rain recovery may help to mitigate the impacts of climate change on thermally sensitive fish in lakes across eastern North America

From the 1970s to 1990s, more stringent air quality regulations were implemented across North America and Europe to reduce chemical emissions that contribute to acid rain. Surface water pH slowly increased during the following decades, but biological recovery lagged behind chemical recovery. Fortunately, this situation is changing. In the past few years, northeastern US fish populations have begun to recover in lakes that were historically incapable of sustaining wild fish due to acidic conditions. As lake ecosystems across the eastern United States recover from acid deposition, the stress to the most susceptible populations of native coldwater fish appears to be shifting from acidification effects to thermal impacts associated with changing climate. Extreme summer temperature events – which are expected to occur with increasing frequency in the coming century – can stress and ultimately kill native coldwater fish in lakes where thermal stratification is absent or highly limited. Based on data from northeastern North America, we argue that recovery from acid deposition has the potential to improve the resilience of coldwater fish populations in some lakes to impacts of climate change. This will occur as the amount of dissolved organic carbon (DOC) in the water increases with increasing lake pH. Increased DOC will reduce water clarity and lead to shallower and more persistent lake thermoclines that can provide larger areas of coldwater thermal refuge habitat. Recovery from acidification will not eliminate the threat of climate change to coldwater fish, but secondary effects of acid recovery may improve the resistance of coldwater fish populations in lakes to the effects of elevated summer temperatures in historically acidified ecosystems. This analysis highlights the importance of considering the legacy of past ecosystem impacts and how recovery or persistence of those effects may interact with climate change impacts on biota in the coming decades.     

Modelling the hydro-geochemistry of acid-sensitive catchments in Finland under atmospheric deposition and biomass harvesting scenarios

The dynamic hydro-geochemical Model of Acidification of Groundwater in Catchments (MAGIC) was used to predict the response of 163 Finnish lake catchments to historic and future atmospheric deposition (1880–2100) and future tree harvesting practices. Deposition was assumed to follow current legislated European emission reduction policies (CLe) and a scenario based on maximum (technically) feasible reductions (MFR). Future harvesting was assumed to shift from stem-only harvesting (SOH) to whole-tree harvesting (WTH) owing to the potential increased utilisation of biofuels. Despite the modest changes in atmospheric deposition under CLe (compared to current day), these reductions are predicted to halt the decline in soil base saturation; however, further reductions are required to improve soil and lake water chemistry. The MFR scenario predicted a significant long-term improvement in soil base saturation leading to continued long-term recovery in surface waters (all lakes with ANC > 0 by 2100). However under the WTH scenario, significant long-term impacts (re-acidification) were predicted for soil and surface water chemistry (14 lakes with ANC < 0 by 2100). To some extent the long-term negative impacts were reduced under MFR, indicating that increased utilisation of biofuels will necessitate ‘trading emissions for timber’, or soil amendment, to maintain ecosystem quality and sustainable forest growth. The current practice of SOH is close to the sustainable maximum harvesting under current (legislated) atmospheric deposition in Finland.     

Influence of fish assemblage and shallow lake productivity on waterfowl communities in the Boreal Transition Zone of western Canada

1. Shallow lakes in the Boreal Transition Zone (BTZ) in Alberta, Canada are naturally productive systems that provide important breeding and moulting habitat for many waterfowl (Anseriformes). To examine the relative importance of biotic and abiotic factors on waterfowl population densities, species richness and community composition, we surveyed 30 shallow lakes and evaluated the relationships among fish communities, lake characteristics and waterfowl in both breeding and moulting habitat. Shallow lakes were either fishless (n = 15), contained only small‐bodied fishes (n = 10) or contained large‐bodied, mostly predatory, fish in addition to small‐bodied fish (n = 5). 2. Environmental factors, including water colour, submerged aquatic vegetation, lake area and potassium, explained 24.3% of the variation in breeding waterfowl communities. Fish assemblage contributed independently to a small but significant proportion (13.4%) of the variation, while 13.8% of the explained variation was shared between environmental factors and fish assemblage. In total, 51.5% of the variation in breeding waterfowl communities was explained. 3. Overall, 55.5% of the total variation in moulting waterfowl communities was explained. Environment alone [especially total phosphorus, lake area, maximum depth and dissolved organic carbon (DOC)] and variation shared by fish and environment similarly accounted for most of the explained variation in moulting waterfowl communities (21.7% and 25.7% respectively), while fish assemblage was only one‐third as important (8.1%). 4. Both breeding and moulting waterfowl densities increased with lake productivity, even in eutrophic and hypereutrophic lakes. Breeding waterfowl density was also twice as great in fishless lakes than in lakes with fish, after accounting for lake area. 5. Certain waterfowl taxa were linked to fishless lakes, especially in the moulting season. Canvasback and moulting ring‐necked ducks were linked to small‐bodied fish lakes, whereas moulting common goldeneye were indicators of large‐bodied fish lakes. Knowledge of fish presence and species composition can therefore help guide conservation and management of waterfowl habitat in western Canada. Our results suggest that management efforts to maintain the most productive waterfowl habitat in the BTZ should focus on smaller, shallow, fishless lakes, particularly given that larger fish‐bearing systems have greater regulatory protection.     

The recovery of crustacean zooplankton from acidification depends on lake type

Acidification has harmed freshwater ecosystems in Northern Europe since the early 1900s. Stricter regulations aimed at decreasing acidic emissions have improved surface-water chemistry since the late 1980s but the recovery of biotic communities has not been consistent. Generally, the recovery of flora and fauna has been documented only for a few lakes or regions and large-scale assessments of long-term dynamics of biotic communities due to improved water quality are still lacking. This study investigates a large biomonitoring dataset of pelagic and littoral crustacean zooplankton (Cladocera and Copepoda) from 142 acid-sensitive lakes in Norway spanning 24 years (1997–2020). The aims were to assess the changes in zooplankton communities through time, compare patterns of changes across lake types (defined based on calcium and humic content), and identify correlations between abiotic and biological variables. Our results indicate chemical and biological recovery after acidification, as shown by a general increase in pH, acid neutralizing capacity, changes in community composition and increases in the total number of species, number of acid-sensitive species and functional richness through time. However, the zooplankton responses differ across lake types. This indicates that the concentration of calcium (or alkalinity) and total organic carbon (or humic substances) are important factors for the recovery. Therefore, assessment methods and management tools should be adapted to the diverse lake types. Long-term monitoring of freshwater ecosystems is needed to fully comprehend the recovery dynamics of biotic communities from acidification.     

Lake restoration with and without dredging of phosphorus-enriched upper sediment layers

Human activity has been the cause of continuing decline of water quality in most Dutch lakes. Development of lake restoration programmes must take into account the lake functions. Major reduction of the nutrient and pollutant loading is the primary step in lake restoration. Still, the recovery of eutrophic lakes is retarded frequently because of internal phosphorus loading by the lakes' sediments. Sediment dredging, as an additional tool for water quality management to accelerate accomplishing the desired water quality, is studied. In this paper we evaluate the preliminary results of eight lake restoration projects in the Netherlands. The lakes are compared in order to estimate the magnitude of the internal phosphorus loading. Dredging as an additional measure was carried out twice in the peatlake Geerplas. In the Nieuwkoop Lakes only the external phosphorus loading was substantially reduced from 0.9 to 0.2 g P m^–2 y^–1. Provisional results of these two shallow peatlake restoration projects focussed on eutrophication abatement with and without dredging, are presented. Both show a decrease in phosphorus concentration in the lakes. The necessity to dredge the lakes is discussed.     

Direct and Indirect (Foodweb Mediated) Effects of Metal Exposure on the Growth of Yellow Perch (Perca flavescens): Implications for Ecological Risk Assessment

Yellow perch (YP) are metal tolerant fish that form large populations in many metal-impacted regions across Canada. While they are able to survive and reproduce successfully in environments with water and sediment metal concentrations that are toxic to many invertebrates, perch experience a suite of direct and indirect impacts. YP were studied in a series of Canadian Precambrian Shield lakes near Sudbury, Ontario, along a gradient of metal exposure downwind from metal smelters. In lakes at the high end of our exposure gradient, concentrations of metals (Cu, Ni, and Cd) in YP liver and kidney were well above levels seen in fish from reference lakes. Direct effects linked to metal-exposure were observed, ranging from effects at the cellular level, to effects in organs and tissues, individuals, and populations. In addition to direct or physiological effects, we also documented indirect, foodweb-mediated effects of metals on YP in the most contaminated lakes resulting from the elimination of metal-sensitive large benthic invertebrates and their replacement by small metal-tolerant taxa. The most common indication of such indirect effects on YP is severely stunted growth coupled with a high degree of zooplankton dependence throughout their life. Such indirect effects have important implications for ecological risk assessment (ERA) because they indicate that higher trophic levels may be functionally altered even though the functional groups that they depend on are present and abundant. Although the functional groups important to yellow perch can be abundant in metal-impacted lakes, their benthic communities are impoverished and this is strongly reflected in their size structure upon which energy transfer to higher trophic levels depends. Thus indirect (foodweb-mediated) effects can be important in situations where invertebrate size structure is impacted in such a way as to reduce the efficiency of energy transfer to higher trophic levels, and therefore measures should be taken to protect and/or restore large sensitive benthic species.     

Histopathological characteristics of Coregonus fishes under the impact of metal industry

This article provides results from monitoring studies of the state of common whitefish (Coregonus lavaretus L.) inhabiting the border Pasvik River and a number of lakes of Northern Fennoskandia. The water catchment area is subject to a significant contamination by coper, nickel, and sulfur dioxide due to the airborne transport of emissions from the Pechenganikel smelter. The ecological situation near the smelter is evaluated as very unfavorable and requires constant control, including the monitoring of aquatic systems. We used histopathologies of gills, liver, and kidneys as biomarkers to diagnose the state of sparsely rakered whitefish. Changes in their cellular structure were diagnosed and classified. The presence of the revealed abnormalities is associated with long-term aerotechnogenic stress on the investigated water bodies. The possibility of using the obtained data for assessing water quality has been considered.     

The recovery of acid‐damaged zooplankton communities in Canadian Lakes: the relative importance of abiotic,biotic and spatial variables

1. Acidification has damaged biota in thousands of lakes and streams throughout eastern North America. Fortunately, reduced emissions of sulphur dioxide and nitrogen oxides beginning in the 1960s have allowed pH levels in many affected systems to increase. Determining the extent of biological and pH recovery in these systems is necessary to assess the success of emissions reductions programmes. 2. Although there have been promising signs of biological recovery in many systems, recovery has occurred more slowly than expected for some taxa. Past studies with crustacean zooplankton indicate that a mixture of local abiotic variables, biotic variables and dispersal processes may influence the structure of recovering communities. However, most studies have been unable to determine the relative importance of these three groups of variables. 3. We assessed chemical and biological recovery of acid‐damaged lakes in Killarney Park, Ontario. In addition, we assessed the relative importance of local abiotic variables, biotic variables and dispersal processes for structuring recovering communities. We collected zooplankton community data, abiotic and biotic data from 45 Killarney Park lakes. To assess the recovery of zooplankton communities, we compared zooplankton data collected in 2005 to a survey conducted for the same lakes in 1972–73 using several univariate measures of community structure, as well as multivariate methods based on relative species abundances. To determine the factors influencing the structure of recovering zooplankton communities, we used hierarchical partitioning for univariate measures and spatial modelling and variation partitioning techniques for multivariate analyses. 4. Our survey revealed significant pH increases for the majority of sampled lakes but univariate measures of community structure, such as species richness and diversity, indicated that only minor changes have occurred in many acid‐damaged lakes. Hierarchical partitioning identified several variables that may influence our univariate measures of recovery, including pH, dissolved organic carbon (DOC) levels, fish presence/absence, lake surface area and lake elevation. 5. Multivariate methods revealed a shift in communities through time towards a structure more typical of neutral lakes, providing some evidence for recovery. Variation partitioning suggested that the structure of recovering copepod communities was influenced most by dispersal processes and abiotic variables, while biotic (Chaoborus densities, fish presence/absence) and abiotic variables were more important for cladoceran zooplankton. 6. Our results indicate that the recovery of zooplankton communities in Killarney Park is not yet complete, despite decades of emission reductions. The importance of variables related to acidification, such as pH and DOC, indicates that further chemical recovery may be necessary. The differing importance of abiotic, biotic and dispersal processes for structuring copepod versus cladoceran zooplankton might indicate that different management approaches and expectations for recovery are needed for these groups.     

Climate drivers of diatom distribution in shallow subarctic lakes

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Effects of Changes in Acidity on Aquatic Insects in Rocky Littoral Habitats of Lakes Near Sudbur y, Ontario

Benthic aquatic insects were collected from rocky nearshore areas (< 1 m deep) of 17 lakes near Sudbury, Ontario, with a pH range of 4.7–7.3 and a size range of less than 10 ha to over 10,000 ha. These insect communities were composed of taxa common to lake soft-sediments and streams. Direct and indirect effects of lake acidity appeared to be major controls on the structure of these communities, implying that several factors may be involved in restructuring during acidification or recovery. Declines in abundances of several taxa of Ephemeroptera at pH below 5.5 were attributable to acid toxicity, while increases in the abundances of Odonata and Diptera at pH below 5.5 were associated with the absence of fish predators and other indirect effects of acidity. The communities of two experimentally neutralized lakes restructured rapidly within 5 years, approaching but not achieving community structures typical of our near-neutral survey lakes. Neutralization led to recolonization or increased abundance of the acid-sensitive mayfly, Stenacron interpunctatum , and the dragonfly, Boyeria grafiana ; however, recolonization by other taxa expected to be present in near-neutral lakes ( Stenonema femoratum, Eurylophella, and Basiaeshna janata ) was not observed. Consistent with results for the survey lakes, declines in the abundances of the dragonflies Aeshna interrupta, Aeshna eremita , and Leucorrhinia glacialis in the neutralized lakes were associated with reintroductions of Salvelinus fontinalis (aurora trout) and increased fish predation pressure, while reduced abundances of the dipterans Ceratopogonidae, Psectrocladius , and Stackelbergina may be related to indirect effects of acidity other than fish predation. Although community composition varied greatly across the acidity gradient, total species richness and abundance were not correlated with lake chemistry or number of fish species.     

Invertebrate food sources for waterbirds provided by the reconstructed wetland of Nyirkai-Hany,northwestern Hungary

Detailed studies of the macroinvertebrate benthos and zooplankton communities in Loch Leven, the largest shallow lowland lake in Scotland, UK, were carried out from 1966 to 1973 as part of the International Biological Programme (IBP). The results revealed a reduction in species diversity that was attributed to increasing eutrophication. This work provides a baseline against which the response of the invertebrate communities to subsequent changes in management can be assessed. This article compares macroinvertebrate benthos and zooplankton data from the IBP study with the post-IBP era during which changes at Loch Leven included a 60% reduction in the phosphorus input from external sources and variations in fish stocking rates. Only in recent years has there been evidence of ecological recovery by the invertebrate communities: the number of macroinvertebrate and zooplankton taxa has increased (including taxa considered to be sensitive to nutrient enrichment) and invertebrate abundances have declined. These changes appear to reflect the improvements in water quality and habitat conditions at Loch Leven that have occurred as a result of the recent reduction in nutrient loads, albeit with a substantial delay before any ecological response could be detected. This time lag in recovery has important implications for assessing improvements in the ecological status of other lake systems, as is required by the EU Water Framework Directive.