Littoral zooplankton responses to acid and aluminum stress during short-term laboratory bioassays

During autumn 1990, littoral zooplankton were collected from three alkaline lakes in Ohio, USA. Toxicity tests were performed, in which animals were placed into treatments of pH 4.5, with or without 500 microg litre(-1) Al added. Percentage survival after 24 h was determined for each test species, and compared to survival in controls (pH roughly 8.0). Three distinct responses were observed: (1) Four cladocerans, Simocephalus serrulatus, Diaphanosoma birgii, Acantholeberis curvirostris and Chydorus sphaericus, were tolerant of both acid and Al, with no significant reductions in survival in the treatments. (2) The cladoceran Eurycercus lamellatus and the capepod Acanthocyclops vernalis were sensitive to both acid and Al, and suffered 100% mortality in both treatments. (3) The cladocerans Camptocercus rectirostris, Alona costata and Pleuroxus denticulatus, and the copopod Mesocyclops edax showed decreased survival in the acid treatment, and a significantly greater decrease in the acid plus Al treatment. For nine of the ten test species, the results were consistent with previous survey and paleolimnological studies. The results indicate that direct toxic effects of H+ and Al ions largely determine the responses of these common littoral species to acidification.     

Biological responses to the reversal of acidification in surface waters of the English Lake District

Declining sulphur deposition since the late 1970s has led to increases in pH in lakes and streams of the English Lake District (Cumbria, UK). To determine whether there have been biological responses to the chemical changes, we carried out surveys of stream macroinvertebrates and lake surface sediment diatoms, and compared the results with those from earlier surveys. Macroinvertebrate taxa in five streams (current average pH range 5.1-7.0) sampled in 1999 showed clear changes from those found during 1965/1966 and 1972. For three of the streams, more taxa were present in 1999 than recorded using comparable sampling methods in the 1960s and 1970s, despite lower numbers of individuals being recovered in 1999. Values of the Margalef diversity index could be calculated for both 1965-1972 and 1999 for four of the streams; the index was significantly greater in 1999 (P < 0.001) in three streams, and unchanged in one stream (the most acid). The 1999 survey revealed the presence of acid-sensitive taxa that had been absent in the earlier surveys, notably the three stoneflies Leuctra fusca, L. moselyi and Chloroperla tripunctata. Some taxa that had been present during 1965-1972 were absent in 1999, but few of these were acid-sensitive. Diatoms from the surface sediments of six lakes were classified according to their acid sensitivities. In three of the lakes sampled in 1999 there were more diatoms characteristic of circumneutral waters, and less acidophilic species than had been found in 1983-1985. In the remaining three lakes, no noticeable changes had occurred. Overall, the observed biological changes are qualitatively as expected for the observed increases in pH, and there have been no instances of biological change in the opposite direction. The results support the expectation that changes in freshwater ecological status can be reversed by decreasing the remote emissions of acidifying pollutants.     

Recovery of crustacean zooplankton communities from acidification in Killarney Park,Ontario, 1971-2000: pH 6 as a recovery goal

Despite reductions in atmospheric SO4(2-) deposition and resultant decreases in surface water acidity, widespread biological recovery from acidification has not yet been documented. Temporal trends in crustacean zooplankton species richness (number of species) and composition were examined between 1971-2000 in 46 Killarney Park lakes, Ontario, Canada, to assess the degree of biological recovery in lakes with significant water quality improvements, i.e. pH now > 6, compared to 2 other groups: i) lakes which never acidified; and ii) lakes which are still acidified (pH < 6). Time trends in species richness could not be distinguished among the 3 groups of lakes, nor did changes in species richness indicate recovery. In contrast, the zooplankton community composition of lakes in which the pH increased to above 6, as measured by a multivariate index of species abundances, changed from a "damaged" state to one typical of neutral lakes. Some recovery in composition was also documented for the acidic lakes. While still acidic, the pH levels of these lakes have risen. The extent and pace of recovery in Killarney Provincial Park bodes well for the future of other acidified regions in North America and Europe.     

Habitat-specific bioaccumulation of methylmercury in invertebrates of small mid-latitude lakes in North America

We examined habitat-specific bioaccumulation of methylmercury (MeHg) in aquatic food webs by comparing concentrations in pelagic zooplankton to those in littoral macroinvertebrates from 52 mid-latitude lakes in North America. Invertebrate MeHg concentrations were primarily correlated with water pH, and after controlling for this influence, pelagic zooplankton had significantly higher MeHg concentrations than littoral primary consumers but lower MeHg than littoral secondary consumers. Littoral primary consumers and pelagic zooplankton are two dominant prey for fish, and greater MeHg in zooplankton is likely sufficient to increase bioaccumulation in pelagic feeders. Intensive sampling of 8 lakes indicated that habitat-specific bioaccumulation in invertebrates (of similar trophic level) may result from spatial variation in aqueous MeHg concentration or from more efficient uptake of aqueous MeHg into the pelagic food web. Our findings demonstrate that littoral-pelagic differences in MeHg bioaccumulation are widespread in small mid-latitude 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.     

Challenges in assessing biological recovery from acidification in Swedish lakes

Since the 1980s, Swedish lakes have in general become less acidified. Assessment of biological recovery is, however, hampered by poor pre-acidification data, confounding effects of climate change, and few lakes with annual sampling of fish and other organisms. Only three critically acidified, but non-limed, lakes had two decades of fish monitoring. The lakes had not yet recovered to pre-industrial chemical targets. Fish had low species richness compared to other organism groups. Roach (Rutilus rutilus) and/or European perch (Perca fluviatilis) were the dominant fish species, and the acid-sensitive roach had been lost from one of the lakes. Calcium decreased, possibly approaching pre-acidification concentrations, but exceeded minimum levels needed to sustain some Daphnia species. High or increasing levels of total organic carbon, likely due to reduced acidification and climate change, might influence the biological communities in unexpected ways, for example, facilitating more frequent occurrence of the invasive algae Gonyostomum semen.     

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.     

Response of phytoplankton communities to acidification and recovery in Killarney Park and the Experimental Lakes Area,Ontario

It has been widely speculated that controls of SO2 emissions would stimulate recovery of acidified freshwater lakes in Canada, the United States and Europe. Phytoplankton communities from 22 lakes near Killarney Park Ontario, covering a pH range from 4.5-7.7, were studied from 1998-2000 and compared to data from experimentally acidified (pH decreased 6.7 to 4.5) and recovered (pH increased to 6.0) Lake 302 South at the Experimental Lakes Area (ELA), northwestern Ontario to assess recovery from acidification. Based on historical data, pH levels have rebounded to above 6.0 in several lakes in the Killarney area that were previously acidified to pH 5.0-5.5. Phytoplankton biomass was not correlated to pH, but there was a highly significant relationship between species richness and pH. Recovery trajectories were observed in a subset of 6 lakes, combining species diversity data from the present study with historical data. Correspondence analysis indicated that several of the lakes that experienced increased pH have shifted towards phytoplankton assemblages typical of circumneutral environments.     

Complex biological responses to the experimental acidification of Little Rock Lake, Wisconsin, USA

Acidification can affect aquatic organisms directly through hydrogen ion toxicity, and indirectly through disrupted food web dynamics and altered abiotic conditions. Field populations from selected taxa were studied during the Little Rock Lake whole-basin acidification experiment to illustrate patterns whose timing suggests direct (i.e. immediate) or indirect (i.e. delayed or non-uniform) responses to pH change. As the treatment basin was acidified to pH 5.6, 5.2 and 4.7, immediate changes consistent with a direct pH response were observed for species representing several trophic levels. For other taxa (e.g. littoral invertebrates associated with filamentous algal mats, several species of pelagic zooplankton), indirect mechanisms induced by food web changes were more likely explanations for abundance patterns. The results presented here suggest that the responses of aquatic ecosystems to acidification involve a complex interplay between direct pH effects and subsequent indirect interactions.     

Degradation of littoral habitats by residential development: woody debris in lakes of the Pacific Northwest and Midwest, United States

One of the least understood aspects of aquatic ecology is the role of riparian zones of lakes, and how these habitats and their functions are impacted by human development of lakeshores. We investigated the effects of residential lakeshore development on littoral coarse woody debris (CWD) distribution and on riparian forest characteristics by comparing 18 lakes in the U.S. Pacific Northwest with 16 previously surveyed lakes in the U.S. Upper Midwest. Residential development had a strong negative effect on CWD and riparian forest characteristics at both local and whole-lake scales. There was a strong positive correlation between riparian forest density and littoral CWD abundance in both regions. We found regional variation in CWD and riparian forest characteristics, mostly owing to differences in native forests. Our results suggest the role of local processes in determining CWD distribution and point to potential regional differences in littoral habitat structure associated with forest composition and lakeshore development that may have consequences for littoral-pelagic coupling in lakes.     

Acid and aluminum effects on the survival of littoral macro-invertebrates during acute bioassays

Six common macro-invertebrates were exposed to soft water at pH 4.5, with or without 200 microg liter(-1) Al added. Survivals were determined at 6, 12, 24 and 48 h and compared with neutral pH, Al-free controls. The order of acid-sensitivity among the test animals, from greatest to least (with mean 24/48 h survivals in the pH 4.5, low Al treatment in parentheses), was: Caenis sp. (2%) > Hyalella azteca (12%) > Enallagma sp. (20%) > Gyraulus sp. (55%) > Chironomidae (94%) > Hydracarina (99%). Aluminum significantly reduced the survivals of Gyraulus, Hyalella and Chironomidae. The latter group experienced no significant mortality at pH 4.5 except when Al was present. In contrast, the Hydracarina were unaffected by both acid and acid plus Al exposure, and the survivals of Enallagma and Caenis at low pH were enhanced by Al. These differential responses to the treatments indicate that both acid and Al stress may control the structure of the littoral macroinvertebrate community in acid lakes.     

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.     

Comparison of episodic acidification in Canada, Europe and the United States

Episodic acidification is practically a ubiquitous process in streams and drainage lakes in Canada, Europe and the United States. Depressions of pH are often smaller in systems with low pre-episode pH levels. Studies on European surface waters have reported episodes most frequently with minimum pH levels below 4.5. In Canada and the United States, studies have also reported a number of systems that have had minimum pH levels below 4.5. In all areas, change in water flowpath during hydrological events is a major determinant of episode characteristics. Episodic acidification is also controlled by a combination of other natural and anthropogenic factors. Base cation decreases are an important contributor to episodes in circumneutral streams and lakes. Sulphate pulses are generally important contributors to episodic acidification in Europe and Canada. Nitrate pulses are generally more important to episodic acidification in the Northeast United States. Increases in organic acids contribute to episodes in some streams in all areas. The sea-salt effect is important in near-coastal streams and lakes. In Canada, Europe and the United States, acidic deposition has increased the severity (minimum pH reached) of episodes in some streams and lakes.     

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.     

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.     

A palaeolimnological assessment of diatom production responses to lake acidification

To quantify diatom palaeoproduction responses to lake acidification, a whole-basin diatom accumulation rate was calculated for the small acidified lake Gaffeln, in the G?rdsj?n catchment in south-west Sweden. Changes in the relative frequency of diatom assemblages to acidification were typical of other lakes in the area, notably a decline in planktonic diatoms after approximately 1900 and increase in acid-tolerant benthic species (e.g. Eunotia spp. and Tabellaria binalis). Single deep-water cores could be used to infer past changes in diatom production because of changed sediment-microfossil deposition patterns, probably due to the development of a benthic algal mat in the littoral zone. The basin mean total diatom accumulation rate (based on eight cores) was approximately constant over the last 150 years, while planktonic diatoms decreased from 0.3 x 10(6) frustules cm(-2) year(-1) prior to 1900 to trace levels in the 1970s. There was, however, a corresponding increase in the accumulation of benthic diatoms over the same period, from 0.5 to 1 x 10(6) frustules cm(-2) year(-1) between 1950 and the present, together with changed community structure. The increase in benthic species probably reflects an expansion of the littoral zone as light transparency increased.     

High methane emissions from a littoral zone on the Qinghai-Tibetan Plateau

The littoral zones of lakes have been regarded as hotspots of methane (CH₄) fluxes through several studies. In the present study, we measured CH₄ fluxes in six kinds of littoral zones of Huahu Lake on the Qinghai-Tibetan Plateau in the peak growing season of 2006 and 2007. We found that CH₄ efflux (ranging from −0.1 to 90 mg CH₄ m⁻² h⁻¹) from the littoral zones of this lake was relatively high among those of boreal and temperate lakes. Our results also showed that emergent plant zones (Hippuris vulgaris and Glyceria maxima stands) recorded the highest CH₄ flux rate. The CH₄ flux in the floating mat zone of Carex muliensis was significantly lower than those of the emergent plant zones. CH₄ fluxes in the floating-leaved zone of Polygonum amphibium and bare lakeshore showed no significant difference and ranked last but one, only higher than that of the littoral meadow (Kobresia tibetica). Plant biomass and standing water depths were important factors to explain such spatial variations in CH₄ fluxes. No significant temporal variations in CH₄ fluxes were found due to the insignificant variations of physical factors in the peak growing season. These results may help in our understanding of the importance of the littoral zone of lakes, especially the emergent plant zone, as a hotspot of CH₄ emission.     

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.     

Recolonization of acid-damaged lakes by the benthic invertebrates Stenacron interpunctatum,Stenonema femoratum and Hyalella azteca

The recolonization of acid-damaged lakes in Killarney Park, Canada is described for 3 species of benthic invertebrates; 2 mayflies (Stenonema femoratum, Stenacron interpunctatum) and an amphipod (Hyalella azteca). Synoptic surveys of 119 lakes for amphipods and 77 lakes for mayflies were conducted between 1995 and 1997 and defined pH thresholds of 5.6 for S. femoratum and H. azteca and pH 5.3 for S. interpunctatum. In an intensive study of 2 acid-damaged lakes and 2 reference lakes from 1997 to 2002, reestablishment of S. interpunctatum, S. femoratum and H. azteca occurred, when timing of the events could be estimated, less than 4-8 years after pH thresholds for specific taxa were reached. Dispersal of S. interpunctatum to all habitat patches within a lake was completed 3 years after recolonization was detected in the smallest lake (11 ha). It is anticipated that dispersal throughout the largest lake (189 ha) will take much longer. The time lag from estimated pH recovery to reestablishment and subsequent dispersal of mayflies to all suitable habitats within a lake was as much as 11 to 22+ years. The density of S. interpunctatum increased in the recovering lakes to levels higher than in reference lakes, but stable endpoints have not yet been reached during 6 years of monitoring.     

The variations of aluminium species in mountainous forest soils and its implications to soil acidification

Aluminium (Al) speciation is a characteristic that can be used as a tool for describing the soil acidification process. The question that was answered is how tree species (beech vs spruce) and type of soil horizon affect Al speciation. Our hypotesis is that spruce and beech forest vegetation are able to modify the chemical characteristics of organic horizon, hence the content of Al species. Moreover, these characteristics are seasonally dependent. To answer these questions, a detailed chromatographic speciation of Al in forest soils under contrasting tree species was performed. The Jizera Mountains area (Czech Republic) was chosen as a representative mountainous soil ecosystem. A basic forestry survey was performed on the investigated area. Soil and precipitation samples (throughfall, stemflow) were collected under both beech and spruce stands at monthly intervals from April to November during the years 2008–2011. Total aluminium content and Al speciation, pH, and dissolved organic carbon were determined in aqueous soil extracts and in precipitation samples. We found that the most important factors affecting the chemistry of soils, hence content of the Al species, are soil horizons and vegetation cover. pH strongly affects the amount of Al species under both forests. Fermentation (F) and humified (H) organic horizons contain a higher content of water extractable Al and Al³⁺ compared to organo-mineral (A) and mineral horizons (B). With increasing soil profile depth, the amount of water extractable Al, Al³⁺ and moisture decreases. The prevailing water-extractable species of Al in all studied soils and profiles under both spruce and beech forests were organically bound monovalent Al species. Distinct seasonal variations in organic and mineral soil horizons were found under both spruce and beech forests. Maximum concentrations of water-extractable Al and Al³⁺ were determined in the summer, and the lowest in spring.