Changes in diatom-inferred pH and add neutralizing capacity in a dilute, high elevation, Sierra Nevada lake since A.D. 1825

SUMMARY. 1. We developed equations which describe relationships between the relative abundance of diatom assemblages in the surface sediments of twenty-seven Sierra Nevada lakes and the present pH and acid neutralizing capacity in these lakes.
2. These equations were employed to reconstruct pH and acid neutralizing capacity at twenty-four depth intervals in a 20 cm long sediment core from Emerald Lake, a dilute, high elevation lake, located in Sequoia National Park, California, U.S.A.
3. The diatom-inferred pH and acid neutralizing capacity values indicated that neither pH nor acid neutralizing capacity exhibited significant trends since about A.D. 1825. We conclude from our diatom analysis that Emerald Lake has not been affected by acidic deposition.     

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.     

Responses of phytoplankton and epilithon during acidification and early recovery of a lake

1. Lake 302S in the Experimental Lakes Area of Canada was acidified from pH 6.7 (1981) to 5.1 (1986). The pH was further reduced to 4.5 in 1987 and held at that level until 1991. From 1992 to 1995, the pH was allowed to increase to a target value of 5.8.
2. The response of the phytoplankton community to decreasing pH from 6.0 to 5.1 was similar to that observed in another experimentally acidified lake (223) and in other atmospherically acidified lakes. Acidification affected species diversity of both the phytoplankton and epilithon. Phytoplankton diversity was positively correlated with pH. Epilithic algal diversity was more variable and did not correlate with pH.
3. Phytoplankton biomass was enhanced by acidification as the assemblage shifted from a dominance of chrysophytes to large dinoflagellates ( Gymnodinium sp. and Peridinium inconspicuum ). Epilithon biomass was unaffected, but dominance shifted from filamentous cyanophytes ( Lyngbya ) to acidophilic diatoms ( Tabellaria quadriseptata and Anomoeonis brachysira ).
4. The only taxon to be similarly affected in both the phytoplankton and epilithon was the cyanobacteria, being significantly reduced below pH 5.1. During early recovery (pH 5.5–5.8), cyanobacteria increased and species present prior to acidification recolonized both habitats.
5. In the early stages of recovery, planktonic and benthic assemblages remained more similar to acidified than natural assemblages, but more profound change began at pH > 5.5.     

Changes in the crustacean zooplankton community of Harp Lake, Canada, following invasion by Bythotrephes cederstrœmi

1. Detecting the impacts of invading Bythotrephes cederstrœmi (Crustacea, Onychopoda, Cercopagididae) on zooplankton in North American lakes has been hampered by the brevity of pre-invasion data, and by the difficulty of distinguishing the effects of the invader from other stresses. The data from Harp Lake in Ontario, Canada, circumvent these difficulties. Bythotrephes appeared in the lake in 1993. There is a 15-year pre-invasion data set, and no significant complicating concurrent stresses.
2. The species composition and the size structure of the crustacean zooplankton community of Harp Lake changed after the invasion. Several small species either declined dramatically in abundance (e.g. Bosmina longirostris , Tropocyclops extensus ) or disappeared ( Chydorus sphaericus , Diaphanosoma birgei , Bosmina ( Neobosmina ) tubicen ). In contrast the abundance of the larger cladocerans Holopedium gibberum and Daphnia galeata mendotae and the hypolimnetic copepod Leptodiaptomus sicilis increased. Several univariate and all multivariate summarizations of zooplankton abundance, biomass and size structure highlighted the uniqueness of the post-invasion community.
3. The alterations in the zooplankton community could not be attributed to changes in lake acidity, thermal regimes, penetration by ultraviolet light, nutrient status, fish stocking or the abundances of native invertebrate predators, but they were correlated with Bythotrephes abundance, both within and among years. Hence, we hypothesize that the invasion by Bythotrephes has significantly altered the crustacean zooplankton community of Harp Lake.     

Changes in the crustacean zooplankton community of Harp Lake, Canada, following invasion by Bythotrephes cederstrœmi

1. Detecting the impacts of invading Bythotrephes cederstrœmi (Crustacea, Onychopoda, Cercopagididae) on zooplankton in North American lakes has been hampered by the brevity of pre-invasion data, and by the difficulty of distinguishing the effects of the invader from other stresses. The data from Harp Lake in Ontario, Canada, circumvent these difficulties. Bythotrephes appeared in the lake in 1993. There is a 15-year pre-invasion data set, and no significant complicating concurrent stresses.
2. The species composition and the size structure of the crustacean zooplankton community of Harp Lake changed after the invasion. Several small species either declined dramatically in abundance (e.g. Bosmina longirostris , Tropocyclops extensus ) or disappeared ( Chydorus sphaericus , Diaphanosoma birgei , Bosmina ( Neobosmina ) tubicen ). In contrast the abundance of the larger cladocerans Holopedium gibberum and Daphnia galeata mendotae and the hypolimnetic copepod Leptodiaptomus sicilis increased. Several univariate and all multivariate summarizations of zooplankton abundance, biomass and size structure highlighted the uniqueness of the post-invasion community.
3. The alterations in the zooplankton community could not be attributed to changes in lake acidity, thermal regimes, penetration by ultraviolet light, nutrient status, fish stocking or the abundances of native invertebrate predators, but they were correlated with Bythotrephes abundance, both within and among years. Hence, we hypothesize that the invasion by Bythotrephes has significantly altered the crustacean zooplankton community of Harp Lake.     

Assessing the health of a zooplankton community in a small Precambrian Shield lake during recovery from experimental acidification

Acidified lakes recover chemically relatively quickly following the reduction or cessation of acidic inputs. Although fish, invertebrate, and phytoplankton communities are reported to begin to return to preacidification states in chemically-improving lakes, the process and extent of biological recovery are not well-documented. The experimental acidification of Precambrian Shield Lake 223 (27.4 ha surface area; 14.4 m maximum depth) in the Experimental Lakes Area in northwestern Ontario, provides an opportunity to compare the zooplankton community prior to acidification with that during progressive acidification and during chemical recovery. Acidified with sulfuric acid from pH 6.47 (ice-free season mean) in 1976 to pH 5.0 (1981 to 1983), Lake 223 has been allowed to recover in steps of pH 5.5 (1984 to 1987), pH 5.8 (1988 to 1990), and pH 6.11 (1991). Total zooplankton biomass showed no trend to increase or decrease during the acidification and recovery, but species composition changed. Compared with species composition at pH 6.13 early in acidification in 1977, the recovering community at pH 6.11 in 1991 had the previously-dominant cladoceran species present in very low numbers and had two newly-appearing cladoceran species. The community had lost one species of calanoid and gained none and lost two species of cyclopoids and gained two. It appeared to lose four species of rotifiers and gain seven. In nearby unmanipulated reference Lake 239 (56.1 ha; 30.4 m), species shifts were recorded but they involved rarer species, not dominants as in Lake 223. Although the zooplankton community in 1991 is in a new state with respect to species composition, static measures of total community biomass, contribution to biomass by the four main taxonomic groups, per cent smilarity to the preacidification community (for crustaceans), and biomass of herbivores do not indicate impairment of community health. Lowered species diversity for both crustaceans and rotifers partially returned to preacidification levels. Nevertheless, the rotifer community in 1991 was more dissimilar to the preacidification community than was the crustacean community, and carnivore biomass appeared to be depressed in Lake 223. The Lake 223 zooplankton community at pH 6.11 in 1991 appears to be in a state of flux.     

Studies on zooplankton in two acidified high mountain lakes in the Alps

Being located in remote areas, alpine lakes are good indicators of regional and global pollution, and are particularly sensitive to atmospheric depositions. When situated in areas where acidic rocks dominate, they are sensitive to acidification. In the framework of an international project partially funded by the EU, a two-year study on zooplankton was carried out on two lakes, Lake Paione Inferiore (LPI) and Lake Paione Superiore (LPS), selected because of their susceptibility to acidification. In particular, LPS is permanently acidified, with pH ranging between 5.3 and 6.2, and LPI is acidified during the ice melt, when pH drops to 6.1 units. In addition, LPI is subjected to further anthropogenic interference, since fish (Onchorynchus mykiss) have been repeatedly introduced into the lake during the last thirty years. Literature information is available on the species composition and the seasonal dynamics of the zooplankton communities of the two lakes before these anthropogenic disturbances took place. Previously, the two lakes had a similar species composition and abundant zooplankton. An analysis of the present-day situation reveals significant changes compared to the past. Both acidification and the introduction of fish are responsible for the marked alterations observed. By reconstructing of the past cladocera assemblages and by analysing the literature on lakes with different impacts of the two factors (pH and fish), we can estimate the relative importance of the two different anthropogenic disturbances on species composition and abundance of the zooplankton communities of the two lakes. This revised version was published online in August 2006 with corrections to the Cover Date.     

Variation in calanoid copepod resting egg abundance among lakes with different acidification histories

The maintenance of species and genetic diversity within zooplankton egg banks may be crucial to the re-establishment of zooplankton communities following historical disturbance, such as anthropogenic acidification which globally caused widespread damage to ecological communities. Despite this, no other study has described basic characteristics of zooplankton egg banks among lakes with different acidification histories, such as variation in resting egg concentration. Theoretically, habitats with frequent periods of harsh environmental conditions are expected to select for resting egg production or prolonged dormancy in zooplankton, which would increase the size of the resting egg bank in lake sediments. In this study, we compared abundances of viable and inviable calanoid copepod resting eggs among three freshwater lakes with different acidification histories. While Swan Lake underwent major chemical and biological changes from acid and metal deposition, Teardrop and Bat lakes were relatively unaffected by historical acidification and had comparatively constant, but different pH over time. We also tested the effect of age on the viability of resting eggs. As predicted, higher numbers of viable resting eggs were found in recent sediments from acid-recovering Swan Lake compared to study lakes with relatively temporally constant environments (Teardrop and Bat lakes) when the total number of eggs was held as a covariate. We detected this result in spite of similar pelagic abundances of Leptodiaptomus minutus, the dominant species in zooplankton communities of these lakes. This pattern did not necessarily hold for inviable egg concentrations since these eggs were more abundant in both Swan and Bat lakes compared to Teardrop Lake in older sediments (1939–1951, 1800s). Within study lakes, the abundance of viable resting eggs declined with increased egg age. Further study is required to test mechanisms underlying these patterns. Handling editor: S. I. Dodson     

Contrasting responses of two cladocerans to changes in the nutritional value of nannoplankton

SUMMARY. 1. The potential tor competition in the zooplankton of small Araucana Lake (Andes, Argentina) was tested experimentally.
2. Laboratory enrichment of lake water with nutrients resulted in an increased abundance of nannoplankton with the cyanobacteria Gloeocapsa minor predominating.
3. The dominant species of zooplankton showed contrasting responses to enrichment: the abundance of Bosmina longirostris increased, whereas that of Ceriodaphnia dubia decreased relative to the corresponding controls.
4. The clearance rates for various nannoplankton by these cladocerans were similar, resulting in high niche overlap as estimated by the Pianka index.
5. Life table experiments showed that when fed on a pure suspension of lake flagellates (10³ cells ml⁻¹) Ceriodaphnia had a significantly higher average intrinsic rate of natural increase ( r ) than on a suspension of Gloeocapsa (0.28 v . 0.146); in the case of Bosmina no significant differences for the two foods were found (0.182 and 0.141, respectively).
6. The contrasting responses of the animals to enrichment are explained by the differing suitability of particular nannoplankton species as food for different cladoceran species. The seasonal succession in the species composition and food quality of nannoplankton may be important for the coexistence of competing zooplankters in lakes.     

The effect of anthropogenic acidification on the hydrofauna of the lakes of the West Tatra Mountains (Slovakia)

Fourteen West Tatra lakes were studied, of which one could be considered to be recently anthropogenically acidified and eight others classified as acidification-endangered. In the anthropogenically acidified lake, the zooplankton assemblage has been substantially altered (three mountain-lake crustacean species have been eliminated). Several littoral macrobenthic species sensitive to acidification have either been eliminated from the acidified and acidification-endangered lakes or occur only sporadically. The effect of acidification has so far not been observed on the benthic fauna of the lakes medial which is probably due to the higher pH below the surface of sediment. In comparison with the High Tatra, acid depositions have had a less pronounced effect on the lakes of the West Tatra.     

The Contribution of Ciliated Protozoa to Zooplankton Biomass in an Acidic, Subtropical Lake

Ciliated protozoa accounted for up to 50% of the mean daily zooplankton biomass in McCloud Lake, a small (5 ha), oligotrophic, acidic (pH 4.7) lake in north-central Florida. Food resources (algae and bacterioplankton) were limiting for crustacean and rotifer zooplankton during much of the year. Myxotrophic ciliates were a dominant component of the planktonic food web. Stentor niger , an uncommon species in the plankton of lakes, dominated the ciliate assemblage and usually comprised >90% of total ciliate biomass. Stentor niger always contained high densities of photosynthetic zoochlorellae and contributed an estimated 30% to the total autotrophic biomass.     

Late fall phytoplankton dynamics in three lakes, Rocky Mountain National Park

We studied phytoplankton population dynamics during the month preceding formation of ice cover in three small subalpine lakes in Rocky Mountain National Park, Colorado, U.S.A. The outflow from Emerald Lake, which is surrounded by talus, flows into Dream Lake, which is surrounded by sub-alpine forest. Nymph Lake is a lower seepage lake with abundant macrophytes in summer. The major ion concentrations in the three lakes were similar during the study, although Emerald and Dream Lakes had higher concentrations of nitrate and silica than Nymph Lake. A principal component analysis (PCA) showed that the phytoplankton in Emerald and Dream Lakes were distinct from the phytoplankton in Nymph Lake. The species composition changed in each lake during the late fall. The patterns of change in Emerald and Dream Lakes were similar on the PCA diagram despite the greater abundance of diatoms in Dream Lake and the decreasing flow from Emerald Lake into Dream Lake during the fall. In Nymph Lake, a progressive shift in species distribution occurred with a decrease in the most abundant chlorophyte, Chlamydomonas sp., and increases in several species, including two chrysophytes and the diatom Eunotia sp. The marked change in species composition in all three lakes suggests that phytoplankton populations are influenced by changes in water temperatures and incident solar radiation that occur during the late fall. We also compared these data with phytoplankton data for two fall periods from two other hydrologically connected Rocky Mountain lakes. PCA analysis showed that the difference between years was greater than the change during the fall and that the fall species composition in these two lakes was distinct from that in Emerald and Dream Lakes or in Nymph Lake. Studying phytoplankton dynamics in alpine and sub-alpine lakes may offer clues as to how these ecosystems may respond to projected climate changes in the Rocky Mountain region, such as warmer temperatures and later formation of ice-cover.     

Cladoceran assemblages, seasonal succession and the importance of a hypolimnetic refuge

SUMMARY. 1. We investigate the importance of a refuge from fish predation to the abundance, species composition and seasonal succession of zooplankton. Thirty lakes representing a range of depths were sampled twice in summer for physical/chemical parameters and zooplankton community structure.
2. We define the refuge from centrarchid predators to be that space between the thermocline and the zone of anoxia. As lakes vary in rate of oxygen depletion from the hypolimnion. the refuge size and lake depth are independent: refuge size decreases during the summer period.
3. Lake depth and refuge size independently explain variation among lakes in zooplankton species composition, but seasonal community change within lakes is best predicted by loss of refuge size.
4. Refuge size also explains the substantial variation in the relative dominance of the two major daphnid species. Lakes possessing a large refuge are dominated by D. pulicaria ; those with a small refuge are dominated by the smaller, D. galeata mendotae . We suggest that lakes of intermediate refuge size, which are characterized by high species diversity, represent a more equitable balance of predation and competition.     

ZOOPLANKTON OF LAKES MUTANDA, BUNYONYI AND MULEHE

Quantitative samples of zooplankton from three lakes in the Kigezi District of Uganda have been studied. The systematics of the zooplankton are considered, and some of the identifications given by Worthington & Ricardo (1936) in a previous study of one of these lakes are revised.
Lake Mulehe is the shallowest of the three lakes and contains the largost standing crop of zooplankton. This is in agreement with chemical data which indicato that the supply of nutrient salts in Lake Mulehe is higher than in the other two lakes.
In October 1962 the zooplankton of Lake Mutanda was characterized by the relative abundance of three species of Daphnia which were not found in the samples from the other lakes, although two of these species were present in Lake Bunyonyi in 1931. Rotifers were sparse in Lako Mutands, but here dominant by Tetramastix opoliensis. Lake Bunyonyi was richer in rotifers, but here the dominant species was Keratella tropica , whilo in Lake Mulehe the dominant rotifer was Synchaeta pectinata.
The zooplankton of Lake Mutanda in October 1962 was similar in composition to that of Lake Bunyonyi in 1931, but in 1962 the zooplankton of Lake Bunyonyi was more like that of Lake Mulehe. The possible causos of this change are discussed.     

Environmental factors influencing zooplankton species composition of lakes in north-central Ontario,Canada

To assess the relative importance of lake chemistry, morphometry and zoogeography on limnetic zooplankton, we collected zooplankton, water, and morphometric data from 132 headwater Canadian Shield lakes in 6 regions across north-central Ontario. A subset of these lakes (n = 52) were fished with gill nets. We clustered lakes based on their zooplankton species composition (presence/absence). Discriminant analysis was employed to determine how well lake characteristics could predict zooplankton community types. Correct classification of zooplankton communities for three models ranged from 72 to 91%. Lake size, lake location, and buffering capacity were ranked as the most important factors separating lake groups. Fish abundance (CPUE) was not significant in distinguishing between zooplankton communities. Though the range of lake sizes was limited (1–110 ha), larger lakes tended to support more species. Lake location (zoogeography) also influenced species composition patterns. Although Algoma lakes tended to be larger (\-x = 18.0 ha, other lakes \-x = 2.5 ha), they supported relatively depauperate zooplankton communities. Buffering capacity was ranked third in the discriminant analysis models, but pH and alkalinity were not significantly different between lake groups.     

Long-term impact of an invertebrate predator, Heterocope septentrionalis, on an arctic pond zooplankton community

1. A long-term field experiment to determine what forces structure and maintain the large-bodied zooplankton assemblages in ponds which lack visual predators was undertaken at the Toolik Lake Research Station in arctic Alaska. Predatory copepods, Heterocope septentrionalis, were introduced to a pond that lacked both fish and invertebrate predators, but contained small- to medium-sized zooplankton, Bosmina longirostris , Daphnia pulex , and Diaptomus pribilofensis. A second pond, which also lacked any predators but had a population of B. longirostris , but no other abundant species, served as a reference population for B. longirostris.
2. The introduction of H. septentrionalis caused the extinction of D. pulex within 1 year and drove B. longirostris to extremely low densities within 4 years. The population of B. longirostris in the reference pond fluctuated throughout the study period, but remained above five individuals per litre. This experiment demonstrates that H. septentrionalis , under natural conditions, made a numerical response to abundant prey and had the capacity to eliminate or reduce vulnerable prey populations of small- to medium-sized zooplankton.     

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.     

Composition and Structure of Zooplankton Communities in Eighteen Arctic and Subarctic Lakes

The factors influencing the composition and structure of zooplankton communities in 18 lakes in the Canadian arctic and subarctic were determined during 1975 and 1976. Phytoplankton were consumed in very low numbers by all species (Diaptomus sicilis, Heterocope septentrionalis, Cyclops scutifer, Daphnia longiremis, Bosmina longirostris, Holopedium gibberum, Keratella cochlearis, Kellicottia longispina) and therefore differences in algal productivity among the lakes had little effect on the zooplankton. Variations in surface area, maximum water depth, pH and the ionic composition of the water were also unimportant in controlling the communities. However, low temperature exerted strong influence over the diversity, abundance, fecundity and vertical distribution of most species.     

Diversity of communities and quantitative parameters of hydrobionts in lakes of the Onon-Toreisk Plain

The diversity of communities and the quantitative parameters of hydrobionts of 17 lakes of the Onon-Toreisk Plain in Northeastern Mongolia during the low-water period are analyzed. The organisms revealed in the lakes include 35 species of phytoplankton, 35 species of zoobenthos, 31 species of zooplankton, and 14 species of hydrophytes. Chara virgata, a representative of charophytes, was revealed in Mongolia for the first time. The greatest species diversity of the communities of hydrobionts is seen in oligohaline lakes. The highest quantitative parameters of communities of phytoplankton and zoobenthos are typical for Gurmiin Lake.     

Acidification of lakes in Šumava (Bohemia) and in the High Tatra Mountains (Slovakia)

Acidification of lakes takes place when pH of rainwater is less than 4.5 and the catchments lie on sensitive geology. Both conditions are met for most lakes in Bohemia and Slovakia. Since 1978 we have studied mountain lakes in the Sumava and in the High Tatra Mountains. In Šumava the three lakes under study are of glacial origin. The catchments are small, with steep sides covered by spruce. The bedrocks are biotite-rich paragneiss, together with gneiss, quartzite and granite. In summer 1936 surface pH was 5.7–6.9 in the Lake Čertovo and 6.9–7.0 in the Lake Černé. Now the pH values are 4.3–4.8 in the two lakes and in the Lake Prášilské as well. Old reports on zooplankton are from the years 1871, 1892–96, 1935–37, 1947 and 1960. Since 1979 we have not found any planktonic Crustacea in the lakes Černé and Čertovo. Lake Prášilské is inhabited by Daphnia longispina and Cyclops abyssorum. In July 1989 the pH values were 4.4, 4.7 and 4.7, concentrations of labile monomeric Al were 0.83, 0.68 and 0.24 mg l^-1 in the lakes Čertovo, Černé and Prašilské, respectively. High levels of toxic Al compounds might be responsible for the extinction of planktonic Crustacea in the lakes Čertovo and Černé. All the three lakes are void of fish at present. In the High Tatra Mts. we examined more than 40 lakes above timberline in altitudes 1612–2145 m. They are all clearwater, naturally fishless lakes. The bedrock is granite. Owing to different levels of calcium the lakes are now in different stages of acidification. According to recent changes in the zooplankton they can be divided into three groups: (1) Species composition of planktonic Crustacea has not changed. (2) Planktonic Crustacea were present until 1973 but are absent now. (3) From the original species of Crustacea only Chydorus sphaericus is present. The three groups are well separated along the gradients of calcium, ANC and pH. They can be identified with the Henriksen's bicarbonate (our group 1), intermediate (our group 2) and acid (our group 3) lakes. We suppose that in the process of acidification the lakes of the group (2) have been shifted from oligotrophy to ultraoligotrophy.