Epilithic diatoms of mountain lakes of the Tatra Mountains (Slovakia)

In the framework of the international project EMERGE, species composition of epilithic diatoms of 34 selected high mountain lakes of Slovak part of the Tatra Mountains were investigated. In all, 127 taxa of diatoms belonging to 26 genera were recorded. Comparison of the epilithic assemblages of the investigated lakes showed differences both in relative abundance and taxa present in the individual lakes. On the basis of species composition and relative abundance of epilithic diatoms using hierarchical cluster analysis, two main groups of lakes were created. The first group includes 4 lakes which can be classified as strongly acidified and 9 lakes which are acidified or threatened by acidification. In this group, acidophilous species such as Achnanthes helvetica, A. marginulata, Aulacoseira distans and Tabellaria flocculosa prevailed. The second group includes 19 non-acidified lakes. The most common diatoms in these lakes were Achnanthes minutissima, A. helvetica, Cymbella minuta and Denticula tenuis which occur practically in all these lakes but with different abundances. The division of lakes due to their diatom species composition mostly confirmed the status of these lakes as judged from their water chemistry.     

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.     

Acidification of small mountain lakes in the High Tatra Mountains,Poland

A hydrochemical investigation was carried out on eight small High Tatra mountain lakes (Poland). When comparing recent data with those from the period 1935–1965, a constant process of acidification of the lakes is found. The average pH of precipitation is 4.8 in the study area, but the lakes are in two stages of acidification: weak (pH 6.0–6.5) and intermediate (pH 5.5–5.8). The differences are due to differences in water sources.     

Algal flora of lakes in the High Tatra Mountains (Slovakia)

Eighty seven from a total of 120 lakes in the Slovak part of the High Tatra Mts. have been visited since 1982. Their summer phytoplankton and algae growing on stones were collected, identified and documented. Some species that are interesting, rare, or not previously known in Slovakia have been found: Clastidium setigerum, Colacium calvum, Chroococcus subnudus, Chr. quaternarius, Coelastrum printzii, Coenocystis quadriguloides, Oocystis naegelii, Scopulonema polonicum, Thelesphaera alpina, Trochiscia prescottii. The number of algal species, found in the open water of the lakes, decrease with altitude and increase with pH.     

Chlorophyll-phosphorus relationship in acidified lakes of the High Tatra Mountains (Slovakia)

Concentrations of total phosphorus (TP) and chlorophyll a (CHLA) were measured in 28 lakes in the High Tatra Mountains (Slovakia) from 1983 to 1990. The relationship between log CHLA and log TP in the Tatra lakes is similar to relationships developed for lakes in other regions, but variation is higher. A part of this variation is caused by acidification of the lakes. In the lakes with pH between 4.9 and 6.3 the CHLA concentrations are often extremely low while TP concentrations decreased, but not as drastically.     

Chemical characteristics of lakes in the High Tatra Mountains,Slovakia

The chemistry of 53 lakes at various stages of acidification and inhabited (at the presence and/or in the past) by pelagic Crustacea was studied in September 1984. Ten of these lakes were investigated in detail biannually (July and October 1987–1990). The July results reflect the influence of snowmelt and were compared with the October ones. The most important anion was sulphate with the average values of 98 and 104 μeq 1^-1 in 1984 and 1987–1990, respectively. High concentrations of nitrate (21–56 μeq 1^-1) were observed in lakes above the treeline. Mean relative composition of cations does not differ between July and October; small changes are in the mean relative composition of anions. Acidification of lakes, expressed as a decrease in alkalinity, is 100 μeq l^-1, and is equal to the increase in the sum of sulphate and nitrate. The values of total phosphorus and COD are the lowest in the range of pH 5–6.5. Alkalinity, sulphate, nitrate and pH do not show any trend with time over the last ten years.     

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.     

Preimaginal stages of Diptera (excluding Chironomidae and Simuliidae) of mountain lakes,their inlets and outlets in the Tatra Mountains (Slovakia)

The occurrence of preimaginal stages of Diptera (excluding Chironomidae and Simuliidae) in the Tatra Mountain lakes, their inlets and outlets, and the influence of acidification on the presence of these taxa were assessed. At 21 sites, 18 Diptera taxa were recorded in 2000 and 2001. The most frequent were Dicranota sp. (73%), Pedicia (C.)rivosa (50%) and Wiedemannia sp. (14%). Diptera were ordinated by RDA; relationships between taxa and environmental variables were significantly determined by temperature and habitat characteristics (altitude, habitat, alkalinity, Ca²⁺, pH, temperature, conductivity). The RDA diagram confirmed the existence of four Diptera assemblage groups corresponding with four habitat types. Group 1 — alpine and subalpine oligotrophic lakes at higher altitudes threatened by acidification, with the acid tolerant taxa Tipula rufina, T. alpium, Dicranota sp. and Rhypholophus sp. Group 2 — subalpine oligotrophic lakes not threatened by acidification with the acid sensitive taxa Wiedemannia sp., Hemerodromia sp., Tipula (S.) benesignata, Orimarga sp., Chionea sp., Pedicia (C.) rivosa and Bazarella subneglecta which prefer higher temperature. Group 3 — subalpine oligotrophic lakes, with the acid sensitive species Berdeniella illiesi, which prefers cold water. Group 4 — outlets of submontane oligotrophic lakes with characteristic taxa Tricyphona sp. and Tipula goriziensis. Acidified alpine and subalpine Tatra lakes are situated mostly at higher altitudes (1705–2145 m. a.s.l.) and have specific taxonomic composition.     

Factors governing nutrient status of mountain lakes in the Tatra Mountains

1. Nutrient and chlorophyll a levels, and bacterial numbers of 84 glacial lakes in the Tatra Mountains (Slovakia and Poland, Central Europe) were determined to assess the impact of catchment vegetation and water acidity on lake trophic status. 2. Catchment vegetation was the crucial factor governing nutrient content of lakes. 3. Concentrations of organic carbon, organic nitrogen, and chlorophyll a, and bacterial numbers were tightly correlated with total phosphorus (TP) content. Their levels were the highest in forest lakes, then decreased in alpine lakes with decreasing amount of catchment vegetation and soil cover, and were the lowest in lakes situated in bare rocks. 4. The above pattern was further modified by lake water acidity. Concentrations of TP, organic carbon, and chlorophyll a were lower in alpine lakes with pH between 5 and 6 than in more or less acid alpine lakes. Zooplankton was absent in all alpine lakes with pH between 5 and 6. 5. Nitrate concentrations followed an inverse trend to TP; lowest values were in forest lakes, then increased with decreasing amount of catchment soils and vegetation. Within the lakes of the same type of catchment vegetation, nitrate concentrations were negatively correlated to TP. N‐saturation of catchment areas and lake primary production were dominant processes controlling nitrate levels in lakes and nitrate contribution to lake acidification.     

Radiometric dating of sediment records from mountain lakes in the Tatra Mountains

Sediment cores from nine different lakes in the Tatra Mountains, collected as part of the EU funded AL:PE, MOLAR and EMERGE projects investigating natural environmental records stored in remote mountain lake sediment sequences, were dated radiometrically by ²¹⁰Pb and ¹³⁷Cs. At five sites, D?ugi Staw G?sienicowy and Zielony Staw G?sienicowy on the Polish side of the Tatra Mountains and Starolesnianske pleso, Ni?né Terianske pleso, and L’adové pleso on the Slovak side of the Tatra Mountains, the cores were sectioned at close intervals and analysed in detail to produce a high resolution chronology. For the remaining four sites, Zmarzly Staw G?sienicowy (Poland), and Vel’ké Hincovo pleso, Vy?né Temnosmre?inské pleso, Vy?né Wahlenbergovo pleso (Slovakia), it was sufficient to establish a low resolution sketch chronology and only a few samples were analysed from each core. At L’adové pleso, multiple cores were collected in order to establish spatial distribution of sediments over the bed of the lake. Cores from all sites had good records of the fallout radionuclides from which it was possible to construct reliable chronologies of the recent sediments.     

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.     

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.     

River morphology of mountain streams influenced by an extreme windstorm in the High Tatra Mountains (northern Slovakia)

In this study we describe how the river morphology of seven High Tatras streams has been influenced in different ways by a severe windstorm that occurred in November 2004. A control site situated in an undisturbed area is compared with six sites in windstorm damaged areas. In 2009–2010 River Habitat Survey (RHS) was used to assess the character of 500 m lengths of stream and to derive associated morphological indices (HQA, HMS sensu Raven 1998, RHS indices sensu Vaughan 2010). Large amounts of large woody debris in the channel influenced the streams despite most of it having been removed after the windstorm. Streams situated in deforested valley slopes were probably more affected by erosion; they had higher amounts of transported inorganic matter and also depositional bar features. Some changes in land-use had occurred with streams situated on the south-east side of the High Tatras having a more uniform banktop vegetation structure than the other streams. Distinct secondary succession of bank top vegetation was observed along windstorm influenced streams. The highest percentage of shrubs occurred where impacted streams remained untouched. Morphologically, the streams most affected by windstorm are those situated in deforested steep valley slopes that are affected by erosion and siltation.     

Ammonium uptake in alpine streams in the High Tatra Mountains (Slovakia)

Uptake of NH _inf4 ^sup+ -N by streambed biota of mountain brooks was studied in the alpine zone of the High Tatra Mountains. Experiments were performed involving in situ dosing of ammonium directly to the acidified stream and incubations of ammonium and streambed bryophytes in enclosures within a range of pH from 4.45 to 8.10.NH _inf4 ^sup+ -N uptake length decreased with decreasing stream discharge, while comparable values of discharge-normalized uptake lengths were found during two in situ experiments.Maximum uptake rates of NH _inf4 ^sup+ -N obtained during the incubation of bryophytes (6 to 11 mg m^–2 h^–1) were comparable with results of two in situ experiments (8 and 12 mg m^–2 h^–1). The average NH _inf4 ^sup+ -N uptake rates observed during incubations lasting 3 to 5 hours (4.3 mg m^–2 h^–1) were not related to the pH of stream water. Nitrification of about 50% of the NH _inf4 ^sup+ -N added was observed in non-acidified streams, but was negligible in acidified streams. Significant photoinhibition of nitrification was observed in non-acidified streams during enclosure experiments.     

Macroinvertebrates of inlets and outlets of the Tatra Mountain lakes (Slovakia)

The composition of invertebrates of inlets and outlets of selected West Tatra and High Tatra lakes (Slovakia, 49°10′ N, 20°00′ E) was analyzed. Twenty-seven sampling sites representing 11 inlets and 16 outlets of 19 mountain lakes were investigated. In total, 10,692 aquatic invertebrates from 27 samples were sorted, 7,314 from inlets and 3,378 from outlets. Altogether, 110 zoobenthic taxa (81 in inlets and 85 in outlets) belonging to 12 higher taxonomic groups were identified. The most constant benthic groups found both in lets and outlets, at more than half of all sites, were Oligochaeta (93% of all sites), Chironomidae (78%), Trichoptera (63%), Plecoptera (59%) and Turbellaria (56%). The most abundant groups were Chironomidae (4,697 ind.), Oligochaeta (2,473 ind.) and Turbellaria (2,236 ind.). No statistically significant differences were found between the benthic communities of inlets and outlets. On the contrary, significant differences (P < 0.05) were found in some parameters of benthic communities of inlets situated in different altitudes (< 1800 m a.s.l., 1800–2000 m a.s.l., > 2000 m a.s.l), outlets of different altitudes, and finally, outlets of lakes with different areas (< 5 ha and ≥ 5 ha). Cluster analysis relatively clearly separated outlets of subalpine lakes from the alpine lakes. Also, outlets of lakes with the lowest pH were separated from the other sites. In the case of inlets, West Tatra sites were separated from the High Tatra sites.     

Hydrological processes in small catchments of mountain headwater lakes: The Tatra Mountains

This study evaluates runoff and different methods for the estimation of water balance and runoff genesis in four small alpine catchments, which lie outside the standard network of hydrological and climate networks. These test catchments, whose size ranges between 2.3 and 110 ha, are located above the timberline at elevations between 1,784 and 2,380 m. Their land surfaces consist of lakes, rock formations, debris deposits, and alpine meadows. Hydrological data were collected for the water year 2001. The catchments were instrumented by three automatic weather stations recording global and net solar radiation, air temperature, humidity, precipitation, and soil temperature. Lake water levels were registered with staff gauges and runoff evaluated from water storage calculations. Runoff genesis was investigated by means of chemical tracers (Rhodamine WT and Lithium chloride). Hydrological process estimations were made using standard methods including: the input of precipitation and snowmelt, both potential and actual evaporation, which was estimated by the approaches of Hamon, Priestley-Taylor, Penman and Grindley, and runoff calculated from the lake storage, were compared with results of the conceptual hydrological Brook90 model. The empirical results show that hydrological processes are governed by the temperature-dependent regime of high mountain snowmelt. However, the major differences in water yield and runoff genesis between watersheds were due to differences in the morphologies of the lakes and their basins, the soilvegetation complex, and runoff routing. Evaluating approaches to estimation, Penman’s combination of both aerodynamic and energy balance method provides the best fit to observed data with observed evapotranspiration being 78 to 99% of the potential calculated. The deterministic Brook90 model is effective for precipitation-runoff genesis studies in small headwater catchments. In the L’adové pleso basin predicted and observed water yield show close correlation. The annual sum of actual evapotranspiration calculated by BROOK90 (352 mm) corresponds closely to that estimated by the approach of Penman (386 mm).     

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.     

Concentration of nutrients in selected lakes in the High Tatra Mountains, Slovakia: effect of season and watershed

Concentrations of total phosphorus (TP), inorganic and organic nitrogen, organic matter, and chlorophyll-a were studied in ten mountain lakes at various stages of acidification, trophy, and type of watershed during each July and October from 1987 to 1990. Concentrations of TP and total organic matter were higher in July than in October. Concentrations of NH4₄ ⁺-N decreased and NO₃ ^–-N increased from July to October. The relative composition of total nitrogen (TN) and its concentration were strongly dependent on the type of watershed: the lowest TN concentrations were observed in lakes with forested watersheds, increasing above the timberline and reaching maximum values in acidified lakes with rocky watersheds. In the pool of TN, nitrate was most important in lakes above the timberline (70–86% of TN), and organic nitrogen in forest lakes (> 90% of TN). Lakes with rocky watersheds were characterized by high ratios of TN:TP (> 250 by mass). The concentration of chlorophyll-a varied widely, from 0.01 to 22.6 µg l^–1, without any consistent change between July and October, and were P limited.     

Biological indices applied to benthic macroinvertebrates at reference conditions of mountain streams in two ecoregions (Poland, the Slovak Republic)

The study was carried out from 2007 to 2010 in two ecoregions: the Carpathians and the Central Highlands. The objectives of our survey were to test the existing biological index metric based on benthic macroinvertebrates at reference conditions in the high- and mid-altitude mountain streams of two ecoregions according to the requirements of the EU WFD and to determine which environmental factors influence the distribution of benthic macroinvertebrates. Our results revealed statistically significant differences in the values of the physical and chemical parameters of water as well as the mean values of metrics between the types of streams at the sampling sites. RDA analysis showed that the temperature of the water, pH, conductivity, the stream gradient, values of the HQA index, and altitude were the parameters most associated with the distribution of benthic macroinvertebrate taxa and the values of the metrics. The values of biological indices should be considered according to the stream typology including altitude and geology. At the reference conditions, the suggested border values of biological indices are very harsh. The values of the biological indices of most sampling sites did not correspond to the requirements of the high status in rivers. The streams at altitudes above 1,200 m a.s.l. should be treated as another river type and new reference values should be established.     

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.