Littoral benthic macroinvertebrates of mountain lakes in the Tatra Mountains (Slovakia,Poland)

Littoral benthic macroinvertebrates of 45 mountain lakes in the Tatra Mountains were sampled using a semi-quantitative method in September 2000. A total of 32,852 specimens were identified to 93 taxa belonging to 14 higher taxonomic groups. Multivariate statistics (CCA, RDA) and nine biotic metrics (AQEM/STAR) were used to explain relationships between macroinvertebrate assemblages and environmental variables. Up to 57% of the ecological position of littoral macroinvertebrate assemblages were explained by variance of environmental variables divided into chemical, trophic, physical, catchment and location. Five types of Tatra lakes were recognized using CCA: A — strongly acidified lakes (small catchment, low pH, high concentration of TP, DOC, highest amount of POM in littoral); B — alpine acidified lakes (low amount of POM, low values of biotic metrics); C — alpine non-acidified lakes (high value of diversity index, predominance of Diptera); D — subalpine acidified lakes (high values of biotic metrics: number of families, proportion of crenal and rhithral taxa/total taxa); E — subalpine non-acidified lakes (high values of biotic metrics: number of families, number of genera, BMWP score, number of taxa and abundance of EPT taxa). RDA was used to design five levels of macroinvertebrate taxa acidification tolerance. The Tatra Acidification Index (TAI) was established to assess the acidification status of the lakes in the Tatra Mts.     

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.     

Subfossil chironomids (Diptera,Chironomidae) in three Tatra Mountain lakes (Slovakia) on an acidification gradient

Three lakes were studied in the High Tatra Mountains at altitudes from 2000 to 2157 m a.s.l., which represent three categories of acidity status recognised in the Tatra lakes in the 1980s: non-acidified, acidified, and strongly acidified. Subfossil chironomid remains from dated sediment cores covering ca. throughout 200 years were analysed. The chironomid thanatocoenoses of all cores layers reflected ultra-oligotrophic non-acidified conditions in L’adové pleso. Nevertheless, the finding of the acid-tolerant species Zalutschia tatrica in littoral samples in 1980s indicates that the littoral zone was more influenced by acidification than deeper areas of the lake. Three stages of lake developmental history can be distinguished in the acidified lake Vy?né Wahlenbergovo pleso based on the chironomid subfossil record: the pre-acidification stage before the 1920s, the anthropogenic acidification stage between the 1920–1980s, and the recovery of the lake from the end of the 1980s. The period of acidification was accompanied by a decline in chironomid numbers, while chironomid fauna composition was unchanged during the whole investigated history of the lake. None of the chironomid taxa present prior to the acidification period disappeared during the peak of acidification. The most apparent change in the sediment record taken from the strongly acidified lake Starolesnianske pleso was recognised in the layers corresponding to the period 1960–1980. It is characterised by the disappearance of the dominant acid-sensitive Tanytarsus lugens group and the dominance of acidtolerant Tanytarsus gregarius group. The most recent chironomid assemblage probably benefits from the amount of food resources as a result of increased lake productivity induced by acidification.     

Restructuring of Fish Assemblages in Swedish Lakes Following Amelioration of Acid Stress through Liming

A large-scale liming program was initiated in Sweden in the mid-1970s to ameliorate the effects of acidification and to restore acidified lakes and rivers to conditions present before acidification. I compared the fish fauna composition and biomass among limed, acidified, and consistently circumneutral lakes. The study focused on (1) recolonization and fish assemblage development and (2) fish biomass and assemblage composition. Time-series data from 14 limed, 5 acid, and 10 circumneutral lakes, and a data set of 323 limed and 346 nonlimed circumneutral lakes, are analyzed. Long-term monitoring revealed that fish species richness increased after liming and was of the same magnitude in limed lakes as in nonlimed circumneutral lakes after 10–20 years. Species proportions differed between limed and nonlimed lakes, and the relative biomass of Perca fluviatilis (Eurasian perch) was in general higher, whereas cyprinid and pelagic species were less important in the limed lakes than in the nonlimed circumneutral lakes. Recolonization of fish species was found to be a crucial factor for the development of the fish assemblages after liming, and in several limed lakes the reintroduction of fish species has accelerated the recovery of pre-acidification fish assemblages.     

The impact of water-level regulation on littoral macroinvertebrate assemblages in boreal lakes

Regulation of lake water level for power production and flood control is among the major anthropogenic disturbances in boreal aquatic ecosystems. In Finland, over 300 lakes, representing one third of the total inland water area of the country, are artificially regulated. To study the effects of regulation on lake littoral macroinvertebrate communities, samples were taken from upper stony littoral and from lower soft bottom littoral habitats of 11 lakes with different regulation amplitudes (wintertime fall in water level 1.19–6.75 m). Twelve unregulated (wintertime fall in water level 0.11–0.55 m) lakes with otherwise similar characteristics were used as a reference. Non-metric Multidimensional Scaling ordinations showed that the composition of macroinvertebrate assemblages was strongly associated with the amplitude of water level regulation. Taxon richness also decreased with increasing intensity of regulation. Freezing and flushing of sediments in late winter are probably the most important factors leading to the impoverished littoral macroinvertebrate fauna. Invertebrates with long life cycle seem to be particularly vulnerable to unnatural water level fluctuation. Our results show that regulation of water level has a major impact on functionally significant lake littoral macroinvertebrates.     

Acidification and the structure of crustacean zooplankton in mountain lakes: The Tatra Mountains (Slovakia,Poland)

Species composition of planktonic Crustacea in 102 lakes in the West and High Tatra Mountains, studied during the peak of anthropogenic acidification (1978–1996), is presented in this work. Zooplankton of the Tatra lakes have been studied since the middle of the 19^th century, which later enabled the recognition of lake acidification and the assessment of its effect on the plankton community of lake ecosystems. In the pre-acidification period, the distribution of zooplankton was determined namely by the lake altitude and orientation (north vs. south) and by the catchment character. Crustacean zooplankton in larger lakes consisted of a limited number of species, with Acanthodiaptomus denticornis and Daphnia longispina dominating lakes in the forest zone, and Arctodiaptomus alpinus, Cyclops abyssorum, Daphnia longispina, Daphnia pulicaria, and Holopedium gibberum dominating lakes in the alpine zone. Ceriodaphnia quadrangula, Daphnia obtusa, Daphnia pulex, and Mixodiaptomus tatricus occurred in lakes with high concentrations of dissolved organic matter and in strongly acidified waters. Anthropogenic acidification has caused drastic changes in both the chemistry and biology of the Tatra lakes. Based on their status during the acidification peak, lakes were divided into three categories: non-acidified (with no change in the species composition of crustacean zooplankton due to the acidification), acidified (planktonic Crustacea disappeared in lakes with meadow-rocky catchments), and strongly acidified lakes where original Crustacea in meadow-rocky catchment lakes disappeared and were replaced by populations of the acid-tolerant littoral species Acanthocyclops vernalis, Chydorus sphaericus, and Eucyclops serrulatus. The acidification-induced processes of oligotrophication and toxicity of aluminium played a key role in the extinction of species. Despite the first signs of biological recovery observed in the early 2000s, acidification remains the most important factor governing the structure of plankton in the Tatra lakes.     

Classification of the Tatra Mountain lakes (Slovakia) using chironomids (Diptera,Chironomidae)

Chironomid assemblages in thirty-three mountain lakes situated above tree line in the Slovakian part of the Tatra Mountains were studied during 2000–2002. Chironomid species/taxa, collected as pupal exuviae, were correlated with physical, chemical, and lake morphometry variables of 22 lakes. Two-way indicator species analysis (TWINSPAN) was used to classify the lakes into four distinct groups: higher situated alpine lakes, lower situated alpine lakes, subalpine lakes and acidified lakes. Presence/absence of eight taxa was identified as indicative for this classification. In discriminant function analysis, pH, dissolved organic carbon, altitude and lake area were the most significant variables reflecting differences among groups of lakes. This model of four variables allowed 77% success in the prediction of group membership. A multiple regression model with lake area, concentration of magnesium and total phosphorus accounted for 37% of the variance in taxa richness. Lakes with greater area contained more chironomid taxa than smaller ones. Lakes with higher alkalinity and higher trophic status tend to support more taxa. Canonical correspondence analysis (CCA) indicated that most variation in the composition of chironomid assemblages was related to pH and to altitude. The results can be used as reference data for long-term monitoring of the Tatra lakes, especially in connection with a recovery from acidification and global climatic change.     

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.     

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.     

Littoral benthic macroinvertebrates of alpine lakes (Tatra Mts) along an altitudinal gradient: a basis for climate change assessment

Littoral benthic macroinvertebrates were studied in three alpine lakes in the High Tatra Mountains (Slovakia) located at different elevations: 2157, 1940 (alpine zone) and 1725 m (sub-alpine zone). The study sites were selected in order to obtain a gradient in thermal regimes and particular organic matter (POM). Differences in the faunal composition of lakes were tested for the ability of these differences to indicate climatic changes, and species/taxa were identified that could be used for the purposes of monitoring and climate change assessment. Macroinvertebrates were sampled quantitatively during the ice-free seasons of 2000 and 2001, and lake surface water temperature (LSWT) and POM were measured. LSWT and POM were negatively correlated with elevation, whereas ice cover was positively correlated with elevation. A total of 60 oligostenothermic macroinvertebrate species/taxa were collected belonging to ten higher taxonomic groups. Statistical analysis showed trends in several biotic metrics with altitude. More specifically, there was a clear increase in the number of species/taxa, genera, and higher taxonomic groups, as well as an increase in the Shannon–Wiener diversity with decreasing altitude. On the contrary, evenness and density of benthic macroinvertebrates did not show any clear relationship with altitude. Gatherers of detrital particles dominated the assemblages’ trophic structures, but no distinct changes in the proportion of functional feeding groups along the altitudinal gradient were found. While the non-insect fauna of the lakes was rather uniform across the elevational gradient, the insect fauna composition was highly correlated with altitude, as confirmed by Detrended Correspondence Analysis. Aquatic insects, in particular chironomids and caddisflies, can therefore be used as good indicators of temperature changes. Our results suggest that under warmer conditions, non-insect benthic macroinvertebrates will remain more or less stable, while aquatic insects will undergo an increase in the number of thermophilic species typical for lower altitudes. These colonizers will increase the diversity of alpine lakes, while the extinction of cold stenothermal species will lead to impoverishment of the native fauna. An indirect impact on benthic macroinvertebrates through changes in food sources is likely, and changes in trophic structure of the littoral assemblages can be expected.     

Macroinvertebrate indicators of lake acidification: analysis of monitoring data from UK, Norway and Sweden

Although the acid sensitivity of many invertebrate species in lakes is well known, methods for assessment of lake acidification based on macroinvertebrate samples are less developed than for rivers. This article analyses a number of existing metrics developed for assessment of river acidification, and evaluates their performance for assessment of lake acidification. Moreover, new species-based indicators of lake acidification were developed and tested. The selected dataset contains 668 samples on littoral macroinvertebrates from 427 lakes with almost 60% of the samples from Sweden and the rest from UK and Norway. Flexible, non-parametric regression models were used for explorative analyses of the pressure–response relationships. The metrics have been assessed according to their response to pH, the degree of non-linearity of the response and the influence of humic compounds. Acid-sensitive metrics often showed a threshold in response to pH between 5.8 and 6.5. Highly acid-tolerant metrics were typically dominant across the whole pH range. Humic level had a positive effect for most acid-sensitive metrics. Generally, most metrics showed a more non-linear response pattern for the humic lakes than for clear lakes. The significant relationship between these macroinvertebrate metrics and acidification shows that there is a potential for developing further the assessment systems for ecological quality of lakes based on these metrics, although the metrics explained a low % of the variation (<30%). In order to improve the predictive power of the biotic metrics across the acidified part of Europe, further harmonization and standardisation of sampling effort and taxa identification are needed.     

Biological recovery of the Bohemian Forest lakes from acidification

A limnological survey of eight small, atmospherically acidified, forested glacial lakes in the Bohemian Forest (?umava, Böhmerwald) was performed in September 2003. Water chemistry of the tributaries and surface layer of each lake was determined, as well as species composition and biomass of the plankton along the water column, and littoral macrozoobenthos to assess the present status of the lakes. The progress in chemical reversal and biological recovery from acid stress was evaluated by comparing the current status of the lakes with results of a survey four years ago (1999) and former acidification data since the early 1990s. Both the current chemical lake status and the pelagic food web structure reflected the acidity of the tributaries and their aluminium (Al) and phosphorus (P) concentrations. One mesotrophic (Ple?né jezero) and three oligotrophic lakes (?erné jezero, ?ertovo jezero, and Rachelsee) are still chronically acidified, while four other oligotrophic lakes (Kleiner Arbersee, Prá?ilské jezero, Grosser Arbersee, and Laka) have recovered their carbonate buffering system. Total plankton biomass was very low and largely dominated by filamentous bacteria in the acidified oligotrophic lakes, while the mesotrophic lake had a higher biomass and was dominated by phytoplankton, which apparently profited from the higher P input. In contrast, both phytoplankton and crustacean zooplankton accounted for the majority of plankton biomass in the recovering lakes. This study has shown further progress in the reversal of lake water chemistry as well as further evidence of biological recovery compared to the 1999 survey. While no changes occurred in species composition of phytoplankton, a new ciliate species was found in one lake. In several lakes, this survey documented a return of zooplankton (e.g., Cladocera: Ceriodaphnia quadrangula and Rotifera: three Keratella species) and macrozoobenthos species (e.g., Ephemeroptera and Plecoptera). The beginning of biological recovery has been delayed for ～20 years after chemical reversal of the lakes.     

Ecological impact of introduced fish in high altitude lakes: a case of study from the European Alps

Fish introduction is a major threat to alpine lake biota leading to the loss of native species and to the degeneration of natural food-webs. This study provides an extensive investigation on the impact of the introduced fish Salvelinus fontinalis on the native communities of alpine lakes in the Gran Paradiso National Park. We compared the macroinvertebrate and zooplankton communities of six stocked and nine fishless lakes with a repeated sampling approach during the summers 2006–2009. The impact of fish presence on alpine lake fauna is often mediated by the strong seasonality governing these ecosystems, and it dramatically affects the faunal assemblage of littoral macroinvertebrates and the size, structure, and composition of the pelagic zooplankton community with a strong selective predation of the more visible taxa. Direct ecological impacts include a decrease or extinction of non-burrower macroinvertebrates and of large zooplankton species, while small zooplankton species and burrower macroinvertebrates were indirectly advantaged by fish presence. Due to the existence of a compensation between rotifers and crustaceans, fish presence does not affect total zooplankton biomass and diversity even if fish are a factor of ecological exclusion for large crustaceans. These compensatory mechanisms are a key process surrounding the impact of introduced fish in alpine lakes.     

Rapid and unexpected effects of piscivore introduction on trophic position and diet of perch (Perca flavescens) in lakes recovering from acidification and metal contamination

1. Yellow perch (Perca flavescens) are often the only surviving fish species in acidified lakes. We studied four lakes along a gradient of recovery from acidification and that had different food web complexities. All had abundant yellow perch, two had low piscivore abundance, one had a well‐established piscivore population and one was manipulated by introducing piscivorous smallmouth bass (Micropterus dolomieu). We hypothesised that there would be strong effects on perch abundance, behaviour and diet induced by the presence of piscivores. 2. In the manipulated lake, the bass reduced yellow perch abundance by 75% over a 2‐year period. Concomitantly, perch use of the pelagic habitat fell from 48 to 40%. 3. In contrast to findings from less disturbed systems, yellow perch in the littoral zone of the manipulated lake did not strongly shift from zooplankton to benthic food sources after the arrival of piscivores. Diet analysis using stable carbon isotopes revealed a strong continued reliance on zooplankton in all lakes, independent of the degree of piscivory. The failure to switch to benthos in the refuge area of the littoral zone is most likely related to the depauperate benthos communities in these formerly acidified lakes. 4. Yellow perch in lakes recovering from acidification face a considerable ecological challenge as the necessary switch to benthic diet is hindered by a low abundance of benthos. The arrival of piscivores in these recovering lakes imposes further restrictions on perch access to food items. We infer that future recovery of perch populations (and higher trophic levels) will have to be preceded by the re‐establishment of diverse benthic macroinvertebrate communities in these lakes.     

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.     

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.     

Littoral macroinvertebrates of acidified lakes in the Bohemian Forest

Mountain lakes in the Bohemian Forest, on both the Czech and German sides, were atmospherically acidified mainly in the 1960s–1980s and have since been recovering from acidification. In 2007, we performed the first complete study on littoral macroinvertebrates in all eight lakes. The goals of the study were to 1) compare macroinvertebrates in the lakes during the process of recovery and 2) investigate relations between the occurrence of taxa and water chemistry. Lake water pH varied from 4.6 to 5.7, concentrations of dissolved reactive Al and labile Al ranged from 118–601 and 11–470 μg L^?1, respectively, and DOC concentrations were < 6 mg L^?1. Altogether 73 taxa were identified from all lakes; a positive relationship was found between pH and the number of macroinvertebrate taxa. The highest number of taxa was found in the least acidic lakes Laka and Grosser Arbersee, including the mollusk Pisidium casertanum. In contrast, the lowest diversity was found in the most acidified ?ertovo jezero. Cluster analyses of macroinvertebrates and water chemistry suggested pH as the key factor influencing the occurrence of macroinvertebrate taxa. An interesting finding was the occurrence of the boreo-montane water beetle Nebrioporus assimilis in Prá?ilské record of this species in the Czech Republic since 1960.     

Recovery from acidification in boreal lakes inferred from macroinvertebrates and subfossil chironomids

Acidification of waters and soils caused by emissions and the long-range transport of air pollutants has been a serious worldwide problem during the last decades. The extent of the acidification problem in Finnish acid-sensitive forest lakes was examined in the Acidification Research Project (HAPRO) in the mid-1980s. The recent decline in the emissions of air pollutants has resulted in the chemical recovery of watersheds in many regions, and the present work on the recovery processes in acidified Finnish headwater lakes (REPRO) was launched to examine whether the chemical recovery has already been accompanied by biological recovery. The patterns of recovery were studied by re-sampling littoral macrozoobenthos in a subset of the previously sampled HAPRO lakes. Paleolimnological samples were taken in order to assess the possible dependence of lacustrine chironomid communities on the changing degree of acidification. Acid sensitive and moderately acid sensitive benthic species revealed slight recovery in the formerly most acidic (pH 5.5) but recently recovered lakes. The most significant factors affecting the response of benthic communities were increased mean lake pH and decreased labile aluminium concentration. Paleolimnological chironomid analysis revealed a slight response along the pH gradient, but also significant structural similarity between the present and pristine chironomid assemblages. This implies that no major changes in chironomid communities of these acidic lakes have occurred during the past centuries. The alternative future trends and threats to biological recovery in small headwater lakes are discussed.     

Macroinvertebrate Diversity in Alpine Lakes: Effects of Altitude and Catchment Properties

Lakes are common features of alpine landscapes, and the attention given to alpine lakes has increased recently in response to increased recognition of the important role that these freshwaters play as sensible indicators of climate change. Despite this general research interest, there is nevertheless a general lack of information about zoobenthos especially of lakes in the Alps, and only few published data are available, which has made it nearly impossible to draw general conclusions in respect to benthic community structure, profundal and/or littoral food webs. This paper aims to explore the relationships between main environmental/catchment properties of 55 lakes and their littoral benthic fauna across three regions of the Alps. We provide updated information on relative abundance, species richness, distribution and ecology of macroinvertebrates which occur and are typical in the littoral of high-mountain lakes of the Alps. These lakes were located in the Lago-Maggiore Watershed (Italy and Switzerland), in South Tyrol (Italy) and in North/East Tyrol (Austria), between 1840 and 2796 m a.s.l., in catchments undisturbed by human activities. As the studied lakes are situated above the tree line, they were characterised by low nutrient levels indicating an oligotrophic status. Lake water chemistry corresponded closely to the geo-lithology of the catchment and some parameters (especially nutrient concentrations) differed between the regions. The macroinvertebrates were dominated by insects which to a high degree were chironomid larvae and pupae. Other insect orders were typical cold stenotherm species of Ephemeroptera, Plecoptera and Trichoptera. Non-insect macroinvertebrates contributed to the 144 taxa found. Other than lake size and catchment area, the faunal parameters exhibited a clearer pattern along altitude. Macroinvertebrates per sample increased with higher elevation, reached their maximum in lakes between 2400 and 2600 m a.s.l., but decreased strongly above 2600 m. The altitudinal pattern of species richness and Shannon diversity resembled each other being highest between 2001 and 2200 m a.s.l., but decreased when going lower and higher, respectively. Various patterns and trends along altitude were also evident when individual groups were analysed within the individual sampling regions. The somewhat conflicting trends of various biocoenotic indices let assume that factors other than altitude are also responsible for the structure of faunal assemblages in the littoral of alpine lakes. Six variables (“bare rocks” and “nitrate”, “alkalinity”, “ammonia” and “peat bog”) were selected by the CCA analysis where these three groups of lakes were identified: (1) lakes with a higher alkalinity (higher pH, conductivity, ion concentration), a higher relative vegetation cover (compared to the “bare rocks” on the opposite side) and lower nitrate levels; (2) lakes with a higher portion of “bare rocks” in their catchments and higher nitrate levels; and (3) a smaller group of lakes with higher ammonia levels and a boggy environment. Geographical patterns seemed to have weak effects on the presence of taxa while catchment properties had evident impacts on macroinvertebrate communities in these lakes. In this way, the present study contributes to the overall understanding of environmental settings and effects on high mountain lake ecosystems and assists in refining research and conservation strategies for an important landscape aspect in the Alps.