Biomass and Production of Lake Charr During the Acidification and PH Recovery of A Small Ontario Lake

The biomass and production of lake charr, Salvelinus namaycush, was studied in Lake 223, a lake that was intentionally acidified to pH 5.0 and then allowed to gradually recover, and in Lake 224, a natural lake of neutral pH. During the first 6 years pH decreased, biomass of Lake 223 lake charr increased, primarily due to high recruitment during the initial years of acidification. Biomass then decreased the final year of acidification. Biomass of Lake 223 lake charr remained low during the subsequent 10 years of pH recovery and 5 additional years after lake pH had returned to neutral pH. Production of lake charr decreased during acidification and increased during pH recovery. Production was still at least an order of magnitude lower at the end of this study than 22 years earlier before lake pH was lowered. The continued low production was caused primarily by low abundance during the years of pH recovery and following years. Production per individual fish in Lake 223 decreased during acidification and quickly increased during pH recovery to values similar to that prior to acidification. Production, biomass, and production per individual varied little between years for lake charr in Lake 224.     

Growth and population size of crayfish in headwater streams: individual- and higher-level consequences of acidification

1. Environmental stress may have indirect positive effects on population size through modification of food‐web interactions, despite having negative effects on individuals. Here we evaluate the individual‐ and population‐level effects of acidification on crayfish (Cambarus bartonii) in headwater streams of the Allegheny Plateau (PA, U.S.A.) with field experiments and survey data. Median baseflow pH of 24 study reaches in nine streams varied from 4.4 to 7.4, with substantial variation found both among and within streams. 2. Two bioassays were conducted to evaluate the relationship between stream pH and crayfish growth rates. Growth rates were always higher in circumneutral reaches than in acidic reaches. Crayfish originating in acidic water grew less when transplanted into neutral water than did crayfish originating in neutral water, providing some evidence for a cost of acclimation to acidity. 3. Stream surveys showed that fish were less abundant and crayfish more abundant in acidified streams than in circumneutral streams. Crayfish density was sixfold higher in reaches with the lowest pH relative to circumneutral reaches. Large crayfish made up a higher proportion of crayfish populations at sites with high fish biomass, consistent with the hypothesis that fish predation on small individuals may be limiting crayfish population size at these sites. 4. Although individual crayfish suffered lower growth in acidified streams, increased acidity appeared to cause an increase in crayfish population size and shifts in size structure, possibly by relieving predation pressure by fish.     

Evolutionary shifts in copepod acid tolerance in an acid-recovering lake indicated by resurrected resting eggs

We employed zooplankton resting eggs to track population-level shifts in acid tolerance over the last century in a boreal shield lake recovering from acidification. Shifts in mean and variance of ecological tolerances of species that occupy ecosystems recovering from anthropogenic stress are important to consider because of their potential to influence eco-evolutionary dynamics at community and ecosystem levels. In a laboratory experiment, we compared juvenile survival of Leptodiaptomus minutus copepods hatched from resting eggs from three time periods (80- to 100-year- old: pre-industrial, 20- to 50-year- old: lake acidification, and 8- to 10-year- old: recovery of lake-water back to pH ≥ 6.0) under several pH treatments. Mitochondrial DNA was used to confirm species identity and to test for population bottlenecks as a possible mechanism to explain our results. We expected that nauplii hatched from eggs deposited prior to industrialization (lake-water pH ≈ 6.0) and from the period of pH recovery (lake-water pH ≥ 6.0) would have lower mean and more variable survival at acidic pH compared to nauplii hatched from the period of peak lake acidification (lake-water pH ≈ 4.7). Our results, which are likely a combination of both genetic and environmental effects, suggest support for this hypothesis. Nauplii hatched from eggs deposited during the period of acidification in George Lake had reduced variation in pH tolerance compared to the recovery period. This was likely driven by strong selection rather than genetic drift because we found no evidence of a population bottleneck. However, we could not detect differences in the variance of naupliar survival between pre-industrial and acidic time periods. Trends in mean naupliar survival from different time periods matched findings from other field-based studies that detected a relationship between lake acidification history and acid tolerance in L. minutus.     

Tolerance of Ceriodaphnia quadrangula and Diaphanosoma brachyurum(Crustacea: Cladocera) to experimental soft water acidification

We analysed in how far the decrease of pH, that is part of the ongoing restoration of the softwater Lake Windsborn (conductivity below 30 S cm^–1), may in future influence the occurrence of the two cladoceran species Ceriodaphnia quadrangula and Diaphanosoma brachyurum. In the field, the abundance of Ceriodaphnia was positively correlated with lake water pH, whereas there was no correlation between abundance and pH for D. brachyurum. Experiments on the tolerance against acidification included dynamic (24 h) and static tests (24, 48 h, 30 d), and were conducted with acidified lake water. C. quadrangula tolerated a slight acidification to pH 5.2, but not pH 4.8, whereas the NOEC value seems to be between pH 4.2 and 4.5 for D. brachyurum. Differences between the experimental NOEC values and field data may be explained by diurnal pH fluctuations and the low ion content of Lake Windsborn which puts an additional physiological challenge to its inhabitants.     

DISTRIBUTION OF THE GENUS CYCLOTELLA IN RELATION TO pH IN PRECAMBRIAN SHIELD LAKES: IMPLICATIONS FOR DIATOM-INFERRED pH1

Recent research on relationships between diatoms and pH suggests that the genus Cyclotella exhibits a strong relationship with lake acidity, being almost totally absent below pH 5.5. This decline has been used as an indicator of lake acidification in paleolimnological studies. In this study C. stelligera V.H. and C. kützingiana Thwaites were abundant in Precambrian Shield lakes with pH as low as 4.5. Cyclotella comta (Ehr.) Kütz. was found in lakes of pH < 5.5, but maximum abundance was observed in lakes of pH > 5.5. Cyclotella michiganiana Skv. was found in lakes of pH < 6.0. These results indicate that the use of C. stelligera, C. kützingiana, and possibly C. comta, in paleolimnological investigations of lake acidification, should be approached with caution. These taxa may exhibit a decline in abundance with decreasing lakewater pH, but this is partially a morphometric effect not necessarily related to anthropogenic acidification.     

Responses of insects,especially Chironomidae (Diptera), and mites to 130 years of acidification in a Scottish lake

About 130 years of anthropogenic acidification of Round Loch of Glenhead, SW Scotland, has resulted in successively decreased stability, diversity, productivity and survival rate of the non-biting midge (Diptera: Chironomidae) fauna. Similar trends have also been observed among mayflies, caddis-flies and water mites. The first effects of anthropogenic acidification on the insect and mite fauna, as evidenced by palaeolimnological analyses of²¹⁰Pb-dated sediment cores, occurred as early as around 1850, i.e. earlier than in any other lake hitherto studied. The drop in the lake's pH was first indicated by decreased stability and changes in species composition of chironomids and mayflies in the littoral zone. Major changes in the profundal chironomid fauna did not take place until about 1950, when mean lake pH dropped below 5.0. At the same time, the littoral insect fauna became even more unstable and the first significant elimination of species occurred. Comparison with insect fauna of other lakes suggests that a pH of less than 5 might be critical for the ecological conditions in many acidified lakes. None of the twelve most common chironomid species present prior to the acidification of the lake had disappeared after 120–130 years of considerable acidification, and they are all common in oligotrophic lakes with a pH of 6.5–7.0. This is in contrast to the effects of heavy acidification on other aquatic animal groups. Chironomids are probably more sensitive to lake trophic status than lake acidity.     

Plant Exclusion of a Herbivore; Crayfish Population Decline caused by an Invading Waterweed

The population of the crayfish Astacus astacus was recorded in Lake Steinsfjorden (southeast Norway) over the period 1979–1996 by studies of total catch effort (baited traps) and catch per unit effort over the season. Over the years 1979–1987, total population of exploitable crayfish (>9.5 cm) was 194,000 (±62,000 SD), and the catches were evenly distributed over the shallow areas. After 1987 there was a sudden decrease in population size, reaching a level of 92,000 (±22,000 SD) for the 10 years that followed. From 1977 onwards the Canadian pondweed Elodea canadensis invaded the lake and established dense covers over large parts of the shallow areas. The crayfish were gradually excluded from areas covered by Elodea, while the yield from remaining areas remained relatively constant. Direct observation by test-fishing and diving confirmed the virtual absence of crayfish within the Elodea stands. Dense stands may directly interfere with movement of adult crayfish, and also strong fluctuations in O₂ and pH were recorded within the stands, probably acting as a stress factor. Although anoxia was not observed in this survey, it may occur during years of massive dieback of Elodea. Thus, while crayfish are able to feed on Elodea, they are unable to control the rapid growth over large areas of this invader, and somewhat paradoxically the herbivore is spatially excluded by its potential resource.     

The influence of productivity and width of littoral zone on the trophic position of a large-bodied omnivore

Omnivory is common in many food webs. Omnivores in different habitats can potentially change their feeding behaviour and alter their trophic position and role according to habitat conditions. Here we examine the trophic level and diet of the omnivorous signal crayfish (Pacifastacus leniusculus) in gradients of trophic status and lake size, both of which have been previously suggested to affect trophic position of predators separately or combined as productive space. We found the trophic position of omnivorous crayfish to be positively correlated with lake trophic status, but found no evidence for any influence of lake size or productive space on crayfish trophic position. The higher trophic position of crayfish in eutrophic lakes was largely caused by a shift in crayfish diet and not by an increase in trophic links in basal parts of the food web. Hence, our results support the “productivity hypothesis,” suggesting that food chains can be longer in more productive systems. Furthermore, stable isotope data indicated that larger crayfish are more predatory than smaller crayfish in lakes with wider littoral zones. Wider littoral zones promoted the development of intrapopulation differences in trophic position whereas narrow littoral zones did not. Hence, differences in habitat quality between and within lakes seem to influence the trophic positions of omnivorous crayfish. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Changes in heart rate and circadian cardiac rhythm as physiological biomarkers for estimation of functional state of crayfish <Emphasis Type="Italic">Pontastacus leptodactylus</Emphasis> Esch. upon acidification of the environment

The changes in heart rate and circadian cardiac rhythm of crayfish Pontastacus leptodactylus Esch. kept in a lightning regime that is close to natural under optimal or low pH values were studied. The heart rate was registered in real time using an original noninvasive fiberoptic method. Upon acidification, disorders in circadian cardiac rhythm and organism reaction (by heart rate) in the suspension test were detected. The characteristics of cardiac activity are considered criteria for estimating the crayfish’s functional state at normal and stress conditions caused by the changes in the quality of the environment.     

Physiological responses of Daphnia pulex to acid stress

Background  

Acidity exerts a determining influence on the composition and diversity of freshwater faunas. While the physiological implications of freshwater acidification have been intensively studied in teleost fish and crayfish, much less is known about the acid-stress physiology of ecologically important groups such as cladoceran zooplankton. This study analyzed the extracellular acid-base state and CO₂ partial pressure (P _CO2), circulation and ventilation, as well as the respiration rate of Daphnia pulex acclimated to acidic (pH 5.5 and 6.0) and circumneutral (pH 7.8) conditions.     

Response of predatory zooplankton populations to the experimental acidification of Little Rock Lake, Wisconsin

To assess the effects of lake acidification on large predatoryzooplankton, we monitored population levels of four limnetictaxa for 6 years in a lake with two basins, one of which wasexperimentally acidified (2 years at each of three levels: pH5.6, 5.2 and 4.7). Concentrations of phantom midge (Chaoborusspp.), the most abundant large predator, remained similar inthe treatment and reference basins until the fourth year (pH5.2) when they increased in the treatment basin. In contrast,Epischuru lacustris and Leptodora kindtii disappeared from limneticsamples, and water mites declined to near zero upon acidification.Treatment basin populations of E.Iacustris declined sharplyduring the second year of acidification. The nature of the declinesuggested sensitivity of an early life stage during the firstyear at pH 5.6. Leptodora kindtii showed no population responseat pH 5.6, but declined to essentially zero at pH 5.2. Treatmentbasin populations of water mites fluctuated until decliningin the fifth and sixth years (pH 4.7). These changes indicatea variety of direct and indirect responses to lake acidification.     

Reciprocal Relationships Between Exotic Rusty Crayfish,Macrophytes, and <Emphasis Type="Italic">Lepomis</Emphasis> Species in Northern Wisconsin Lakes

Abstract The non-native rusty crayfish (Orconectes rusticus) has invaded many lakes of northern Wisconsin, profoundly changing littoral zones in the process. There are other lakes that have been invaded, but do not exhibit these changes. We hypothesized that endogenous feedbacks could form involving rusty crayfish, the macrophytes they destroy, and Lepomis species whose abundance is positively related to macrophyte abundance and also consume juvenile crayfish. We assessed this proposal with long-term data from one lake, a regional comparative study, and a case study of Lepomis predation on crayfish. Through time and across lakes, abundances of rusty crayfish, littoral macrophytes and species of the genus Lepomis were related in a fashion that indicated a set of feedbacks that regulate the abundance of all three. Intense predation on juvenile crayfish by abundant Lepomis is capable of maintaining some crayfish populations at low abundance. Thus, some lakes display profound ecological changes where crayfish achieve high abundance, and others sustain crayfish at low abundance. Consequently, lakes invaded by rusty crayfish may take on the appearance of alternative ecological regimes. Direct experimentation is necessary to determine if, and under what conditions, a lake can exist in either regime. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of Experimental Acidification on Phyto-, Bacterio-and Zooplankton in Enclosures of a Highly Humic Lake

In situ enclosure experiments were performed in a highly humic lake to examine the effects of acidification on phyto-, bacterio- and zooplankton. The acidity of three enclosures was adjusted with H₂SO₄ to pH 3.5, 4 and 5 and one enclosure and the lake served as controls. The diversity of plankton as well as the mean concentration of chlorophyll α, primary production and respiration of plankton decreased with increasing acidity. Bacterial density was slightly lower in the pH 3.5 enclosure than in the other enclosures and in the lake, but there were no differences in the morphological type or mean volume of the cells between different treatments. In the acidified enclosures Cryptomonas ovata and Chlamydomonas spp. were the dominant phytoplankters, while Dinobryon and Mallomonas species seemed to be most sensitive to acidity. Keratella cochlearis, Kellicottia longispina and Bosmina longispina were the most tolerant zooplankton to acidity.     

Contrasts between dystrophic and clearwater lakes in the long‐term effects of acidification on cladoceran assemblages

1. Most studies on zooplankton responses to acidification have focused on clearwater lakes with a dramatic acidification history. The role of dissolved organic carbon (DOC) in moderating zooplankton responses to acidification in naturally acidic, dystrophic lakes is less well understood and is partially impeded by a lack of baseline data. 2. Cladocera leave identifiable remains preserved in lake sediments that can be used to provide information on pre‐industrial species assemblages and their responses to environmental stressors such as acidification. Therefore, we used palaeolimnological approaches to track cladoceran assemblage responses to acidification since c.1850 (inferred from sedimentary diatom assemblages) in three acidified lakes in Kejimkujik National Park (Nova Scotia, Canada) that differ markedly in DOC content. These include two highly dystrophic lakes (Kejimkujik and Pebbleogittch lakes), and one clearwater lake (Beaverskin Lake). 3. In dystrophic Pebbleogittch Lake, an increase in the acid‐tolerant, jelly‐clad, pelagic taxon Holopedium glacialis occurred coincident with diatom‐inferred pH (DI‐pH) declines, but no other notable cladoceran assemblage shifts occurred. Similarly, Cladocera assemblages did not appear to respond to lakewater acidification in dystrophic Kejimkujik Lake. 4. In contrast, in the clearwater Beaverskin Lake, several observed shifts in cladoceran assemblage corresponded to DI‐pH declines, including an increase in the proportion of littoral taxa and an increase in Hill’s N2 species diversity. This may indicate increased water clarity as a result of acidification‐related decreases in DOC, which may have enhanced growth of emergent aquatic macrophytes and improved visibility for planktivorous fish, leading to increased predation on pelagic taxa. Species shifts within the littoral assemblage of Beaverskin Lake may reflect the differing tolerances of littoral taxa to low pH and aluminium toxicity. 5. Overall, our results suggest that cladoceran assemblages in naturally acidic, dystrophic lakes may be resilient against additional pH declines related to industrial emissions of acidifying agents, as dystrophic lakes are less vulnerable to increased aluminium toxicity and acidification‐induced increases in water clarity and often have a pre‐industrial cladoceran assemblage already adapted to acidic conditions.     

Acidification and alkalinization of lakes by experimental addition of nitrogen compounds

Fertilization of a small lake with ammonium chloride for four years as part of a eutrophication experiment caused it to acidify to pH values as low as 4.6. Implications for acidification of lakes via precipitation polluted with ammonium compounds are discussed.When phosphate was supplied with the ammonium, biological nitrogen uptake, apparently by phytoplankton, was the main mechanism causing acidification. When ammonium was applied without phosphate, it accumulated to high concentrations in solution, after which nitrification caused rapid acidification. In both cases, the whole-lake efficiency of acidification was low, averaging about 13% of the potential acidification of supplied ammonium chloride (Table 2).Subsequent application of phosphate plus sodium nitrate for two years caused the pH of the lake to increase. The efficiency of alkalinization was higher than for acidification, averaging 69% of the potential alkalinization of the supplied sodium nitrate.     

DIURNAL ACID METABOLISM IN ISOETES HOWELLII FROM A TEMPORARY POOL AND A PERMANENT LAKE

Water chemistry and titratable acidity and malic acid levels in Isoetes howellii leaves were sampled every 6 hr from plants in a seasonal pool and an oligotrophic lake. Plants in the seasonal pool showed a diurnal fluctuation of ~ 300 μequivalents titratable acidity g⁻¹ fresh wt; daytime deacidification was 75% complete by noon and nighttime acidification was 45% complete by midnight. Late in the season after the pool had dried, emergent leaves showed only a very weak tendency to accumulate acid at night. Plants from the oligotrophic lake had a diurnal change of ~100 μeq g⁻¹ fresh wt, daytime deacidification was only 45% complete by noon but nighttime acidification was 80% complete by midnight. Water characteristics were distinctly different between these two systems. In the seasonal pool there were marked diurnal changes in temperature, pH, oxygen and carbon dioxide. Free-CO₂ levels were an order of magnitude greater in the early morning than in the late afternoon. In contrast, the conditions in the oligotrophic lake showed no marked diurnal fluctuation, though total inorganic carbon levels were extremely low relative to other aquatic systems.     

Loss of diversity and degradation of wetlands as a result of introducing exotic crayfish

The introduction of the alocthonous Louisiana red swamp crayfish (Procambarus clarkii) in Chozas (a small shallow lake situated in León (North-West Spain)) in 1996 switched the clear water conditions that harboured an abundant and a quite high richness of plants, invertebrates, amphibians and birds to a turbid one followed by strong losses in abundance and richness in the aforementioned groups. Crayfish exclusion experiments done in Chozas previous to this work confirmed the role of crayfish herbivorism on macrophyte destruction that had a trophic cascade effect on the wetland ecosystem. Direct and indirect effects of crayfish introduction on Chozas lake communities have been evaluated and compared with previous conditions before 1996 or with other related lakes in which crayfish were no present. Crayfish had a main role in submerged plant destruction and a potential effect on amphibia and macroinvertebrate population decrease. Plant destruction (99 % plant coverage reduction) was directly related to invertebrates (71 % losses in macroinvertebrate genera), amphibia (83 % reductions in species), and waterfowls (52 % reduction). Plant-eating birds were negatively affected (75 % losses in ducks species); nevertheless, fish and crayfish eating birds increased their presence since the introduction. Introduction of crayfish in shallow plant-dominated lakes in Spain is a main risk for richness maintenance in these endangered ecosystems. This revised version was published online in July 2006 with corrections to the Cover Date.     

PHOTOSYNTHESIS OF ALGAL CULTURES AND PHYTOPLANKTON FOLLOWING AN ACID pH SHOCK1

In acidifying lakes, pH decreases abruptly in response to acid precipitation events. We tested the hypothesis that, in comparison to a circumneutral lake, phytoplankton photosynthesis in an acidifying lake is less sensitive to a rapid decrease in pH (acid pH shock). Phytoplankton in Plastic Lake, which is undergoing acidification, was characterized by a predominance of Pyrrophyta, and phytoplankton photosynthesis decreased to a lesser extent in response to an acid pH shock than the photosynthesis of populations from St. Nora Lake, a circumneutral lake located nearby, in which Pyrrophyta were not abundant. Rates of phytoplankton photosynthesis in acid pH shock experiments were significantly correlated with hydrogen ion but not with dissolved inorganic carbon (DIC) concentrations. Depression of photosynthesis following an acid pH shock occurred in axenic cultures of Chlorella pyrenoidosa Chick but was not observed in axenic cultures of the acidophilic alga Chlorella saccharophila (Krug.) Nadson or in three species isolated from Plastic Lake. However, the three isolates were not acidophilic during growth. We conclude that phytoplankton in acidifying lakes consists predominantly of species which are tolerant to acid pH for short periods (hours) but cannot grow at these pHs.     

Concentrations and distribution of Fe,Zn and Cu in tissues of the white sucker (Catostomus commersoni) in relation to elevated levels of metals and low pH

This study examines whether the process of lake acidification influences the accumulation of Fe, Zn and Cu in the tissues of the white sucker (Catostomus commersoni). Concentrations of Fe, Zn and Cu were measured in the liver, kidney and muscle of white sucker sampled from 4 acidic (pH range 4.8–5.3), 1 slightly acidic (pH = 5.8) and 3 circumneutral (pH = 6.3, 6.4) lakes located in south-central Ontario, Canada. Pearson product-moment correlation coefficients were used to determine relationships between average elemental concentrations in the 3 tissues and both sediment and water metal concentrations plus lake pH, DOC and alkalinity. Despite the 1000-fold difference in H+ concentration among the 8 study lakes, tissue concentrations of Fe, Zn and Cu did not correlate with lake pH. Average Fe, Zn and Cu tissue concentrations did not correlate with metal concentrations in lake water. Only Zn concentrations in the liver and muscle were correlated with Zn concentrations in the sediment (r = 0.83 and r = 0.88, P < 0.05). Iron and Cu were regulated by the white sucker over a wide range of lake pH and metal concentrations in both the water and sediment. In contrast, Zn tissue concentrations were correlated with sediment Zn concentrations, the latter are thought to result from Zn inputs of anthropogenic origin.     

Between‐lake variation in the trophic ecology of an invasive crayfish

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