The Saline Lakes of Saskatchewan I. Background and Rationale for Saline Lakes Research

The historical background and utilization of southern Saskatchewan is described. The utilization of the saline lakes is discussed and the rationale for studying them is outlined.     

A Comparative Study of Primary Production and Related Factors in Four Saline Lakes in Victoria,Australia

The purpose of the study was to compare the primary plankton productivities of lakes of different salinities and to determine the causative factors involved in their production rates. Four lakes (specific conductivity —mS cm⁻¹ at 18°C) were initially chosen: Coragulac (9), Red Rock (25), Corangamite (38), Pink (250). Sampling and production measurements were made every two to three weeks. Three lakes were dominated by specific phytoplankton blooms: Red Rock (Anabaena spiroides), Corangamite (Nodularia spumigena). Pink (Dunaliella salina). Coragulac Lake had more diverse populations. Red Rock Tarn had some of the highest production values ever recorded. Extremely high soluble phosphate and inorganic carbon concentrations were the most important causative factors. Pink Lake had very low production rates. High salinity and low nutrient concentrations were limiting factors. The other lakes were intermediate in production and nutrient levels. Zooplankton populations were also determined.     

Bacterial Communities of Three Saline Meromictic Lakes in Central Asia

Meromictic lakes located in landlocked steppes of central Asia (~2500 km inland) have unique geophysiochemical characteristics compared to other meromictic lakes. To characterize their bacteria and elucidate relationships between those bacteria and surrounding environments, water samples were collected from three saline meromictic lakes (Lakes Shira, Shunet and Oigon) in the border between Siberia and the West Mongolia, near the center of Asia. Based on in-depth tag pyrosequencing, bacterial communities were highly variable and dissimilar among lakes and between oxic and anoxic layers within individual lakes. Proteobacteria, Bacteroidetes, Cyanobacteria, Actinobacteria and Firmicutes were the most abundant phyla, whereas three genera of purple sulfur bacteria (a novel genus, Thiocapsa and Halochromatium) were predominant bacterial components in the anoxic layer of Lake Shira (~20.6% of relative abundance), Lake Shunet (~27.1%) and Lake Oigon (~9.25%), respectively. However, few known green sulfur bacteria were detected. Notably, 3.94% of all sequencing reads were classified into 19 candidate divisions, which was especially high (23.12%) in the anoxic layer of Lake Shunet. Furthermore, several hydro-parameters (temperature, pH, dissolved oxygen, H₂S and salinity) were associated (P< 0.05) with variations in dominant bacterial groups. In conclusion, based on highly variable bacterial composition in water layers or lakes, we inferred that the meromictic ecosystem was characterized by high diversity and heterogenous niches.     

Seasonal changes in the chemistry and biology of a meromictic lake (Big Soda Lake,Nevada, U.S.A.)

Big Soda Lake is an alkaline, saline lake with a permanent chemocline at 34.5 m and a mixolimnion that undergoes seasonal changes in temperature structure. During the period of thermal stratification, from summer through fall, the epilimnion has low concentrations of dissolved inorganic nutrients (N, Si) and CH₄, and low biomass of phytoplankton (chlorophyll a ca. 1 mgm ^-3). Dissolved oxygen disappears near the compensation depth for algal photosynthesis (ca. 20 m). Surface water is transparent so that light is present in the anoxic hypolimnion, and a dense plate of purple sulfur photosynthetic bacteria (Ectothiorhodospira vacuolata) is present just below 20 m (Bchl a ca. 200 mgm^-3). Concentrations of N H₄ ⁺, Si, and CH₄ are higher in the hypolimnion than in the epilimnion. As the mixolimnion becomes isothermal in winter, oxygen is mixed down to 28 m. Nutrients (NH₄ ⁺, Si) and CH₄ are released from the hypolimnion and mix to the surface, and a diatom bloom develops in the upper 20 m (chlorophyll a > 40 mgm^-3). The deeper mixing of oxygen and enhanced light attenuation by phytoplankton uncouple the anoxic zone and photic zone, and the plate of photosynthetic bacteria disappears (Bchl a ca.10mgm^-3). Hence, seasonal changes in temperature distribution and mixing create conditions such that the primary producer community is alternately dominated by phytoplankton and photosynthetic bacteria: the phytoplankton may be nutrient-limited during periods of stratification and the photosynthetic bacteria are light-limited during periods of mixing.     

The Saline Lakes of Saskatchewan III. Chemical Characterization

The salinity of Saskatchewan saline lakes varies up to 342%₀. Highly saline lakes are widely distributed. Some reach saturation at high summer temperatures but salt precipitation occurs on cooling. Saline lakes tend to increase in salinity with time but the greatest changes occur in the shallow more saline lakes. The lakes are dominated by sodium, magnesium and sulphate. The cation sequences are dominated by sodium or magnesium while calcium and potassium are proportionally much smaller. Chloride or bicarbonate-carbonate are secondary to sulphate among the anions. Ion sequences tend to be characteristic of certain physiographic regions. Lake types were magnesium (sodium) sulphate, sodium sulphate and sodium (magnesium) sulphate in order of abundance of lakes. Phosphorus and ammonia were high in concentration but nitrate and silica were scarce. The lakes were all alkaline between pH 7.8 and 9.8. The evolution of closed saline lakes is briefly discussed.     

Diversity of Carbon Monoxide-Oxidizing Bacteria in Five Lakes on the Qinghai-Tibet Plateau,China

Carbon monoxide-oxidizing (COX) bacteria play an important role in controlling the flux of carbon monoxide among natural reservoirs, and thus studying their diversity in natural environments is of great significance to understanding the carbon cycle. In this study, the COX bacterial diversity was investigated in five lakes (Erhai Lake, Gahai Lake1, Gahai Lake2, Xiaochaidan Lake, Lake Chaka) on the Qinghai-Tibet Plateau and its correlation with environmental variables of the lakes was explored. Phylogenetic analyses showed that the CO-oxidizers were dominated by Proteobacteria and Actinobacteria in the Qinghai-Tibet Plateau lakes, and their relative abundance varied with salinity: in the freshwater Erhai Lake, the COX bacteria in the water were dominated by the Betaproteobacteria, in contrast to the Actinobacteridae dominance in the sediment; in the saline and hypersaline lakes of Gahai Lake1, Gahai Lake2 and Xiaochaidan Lake, alphaproteobacterial COX bacteria were dominant in the water, whereas Actinobacteridae and alphaproteobacterial COX bacteria were dominant in the sediment. In the hypersaline Lake Chaka, an unknown COX bacterial clade and alphaproteobacterial COX bacteria were dominant in the water and sediment, respectively. Statistical analyses showed that salinity, pH, and major ions (e.g., K⁺, Na⁺, Ca²⁺, Mg²⁺, SO₄ ^2-, and Cl^-) were important factors affecting the COX bacterial community compositions in the investigated lakes. Overall, our results provided insights into the COX bacterial diversity in Qinghai-Tibetan lakes.     

Seasonal Changes in the Structure of the Anoxygenic Phototrophic Bacterial Community in Lake Mogilnoe,a Relict Lake on Kil'din Island in the Barents Sea

An anaerobic phototrophic bacterial community in Lake Mogilnoe, a relict lake on Kil'din Island in the Barents Sea, was studied in June 1999 and September 2001. Irrespective of the season, the upper layer of the anaerobic zone of this lake had a specific species composition of sulfur phototrophic bacteria, which were dominated by the brown-colored green sulfur bacterium Chlorobium phaeovibrioides. The maximum number of sulfur phototrophic bacteria was observed in June 1999 at a depth of 9 m, which corresponded to a concentration of bacteriochlorophyll (Bchl) e equal to 4.6 mg/l. In September 2001, the maximum concentration of this pigment (3.4 mg/l) was found at a depth of 10 m. In both seasons, the concentration of Bchl a did not exceed 3 μg/l. Purple sulfur bacteria were low in number, which can be explained by their poor adaptation to the hydrochemical and optical conditions of the Lake Mogilnoe water. In June 1999, the water contained a considerable number of Pelodictyon phaeum microcolonies and Prosthecochloris phaeoasteroides cell chains, which was not the case in September 2001. A 16S rDNA-based phylogenetic analysis of pure cultures of phototrophic bacteria isolated from the lake water confirmed that the bacterial community is dominated by Chl. phaeovibrioides and showed the presence of three minor species, Thiocystis gelatinosa, Thiocapsa sp., and Thiorhodococcus sp., the last of which is specific to Lake Mogilnoe.     

Seasonal changes in the structure of the anoxygenic photosynthetic bacterial community in Lake Shunet,Khakassia

Seasonal studies of the anoxygenic phototrophic bacterial community of the water column of the saline eutrophic meromictic Lake Shunet (Khakassia) were performed in 2002 (June) and 2003 (February–March and August). From the redox zone down, the lake water was of dark green color. Green sulfur bacteria predominated in every season. The maximum number of green sulfur bacteria was 10⁷ cells/ml in summer and 10⁶ cells/ml in winter. A multi-syringe stratification sampler was applied for the study of the fine vertical distribution of phototrophs in August 2003; the sampling was performed every 5 cm. A 5-cm-thick pink-colored water layer inhabited by purple sulfur bacteria was shown to be located above the layer of green bacteria. The species composition and ratio of purple bacterial species depended on the sampling depth and on the season. In summer, the number of purple sulfur bacteria in the layer of pink water was 1.6 × 10⁸ cells/ml. Their number in winter was 3 × 10⁵ cells/ml. In the upper oxygen-containing layer of the chemocline the cells of purple nonsulfur bacteria were detected in summer. The maximum number of nonsulfur purple bacteria, 5 × 10² cells/ml, was recorded in August 2003. According to the results of the phylogenetic analysis of pure cultures of the isolated phototrophic bacteria, which were based on 16S rDNA sequencing, green sulfur bacteria were close to Prosthecochloris vibrioformis, purple sulfur bacteria, to Thiocapsa and Halochromatium species, and purple nonsulfur bacteria, to Rhodovulum euryhalinum and Pinkicyclus mahoneyensis.     

Anoxygenic phototrophic bacterial community of Lake Shira (Khakassia)

The anoxygenic phototrophic bacterial community of the brackish meromictic Lake Shira (Khakassia) was investigated in August 2001, July 2002, and February–March 2003. In all the periods of investigation, the prevailing microorganisms were purple sulfur bacteria similar to Lamprocystis purpurea in morphology and pigment composition. Their highest number (3 × 10⁵ cells/ml) was recorded in July 2002 at the depth of 15 m. According to 16S rRNA gene analysis, the strain of purple sulfur bacteria isolated in 2001 and designated ShAm01 exhibited 98.6% similarity to the type strain of Thiocapsa roseopersicina and 97.1–94.4% similarity to the type strains of Tca. pendens, Tca. litoralis, and Tca. rosea. The minor microorganisms of the anoxygenic phototrophic bacterial community within the period of investigation were nonsulfur purple bacteria phylogenetically close to Rhodovulum strictum (98.3% similarity, strain ShRb01), Ahrensia kielensis (of 93.9% similarity, strain ShRb02), Rhodomicrobium vannieli (of 99.7% similarity, strain ShRmc01), and green sulfur bacteria, phylogenetically close to Chlorobium limicola (of 98.7% similarity, strain ShCl03).     

The Saline Lakes of Saskatchewan IV. Primary Production by Phytoplankton in Selected Saline Ecosystems

Primary production by phytoplankton, efficiency of photosynthesis, and chlorophyll-a concentrations were determined for seven saline lakes that varied widely in ionic concentration and composition. The investigations were done during the summer months of 1972 and 1973. Productivity ranged from 0.001 to 11.135 g C m⁻³ day⁻¹ and 0.053 to 7.968 g C m⁻² day⁻¹. Highest productivities were measured in two lakes that supported blooms of Aphanizomenon flos-aquae and Nodularia spumigena, respectively. Species of Cyanophyceae, Bacillariophyceae and Chlorophyceae dominated the phytoplankton of the study lakes. Active chlorophyll-a ranged from 0.01 to 116 mg m⁻³. Integral photosynthetic efficiency estimates were <1% except during phytoplankton blooms when they were considerably higher. The overall range of 0.03 to 3.8% is concordant with estimates for other lacustrine ecosystems. The extinction of light caused by photo-synthetic processes, or in situ efficiency, was <1% in the trophogenic zone for most lakes but, it was considerably higher during blooms. In situ efficiencies invariably increased with depth in ail lakes.     

Sulfate reduction and methanogenesis in the Shira and Shunet meromictic lakes (Khakasia, Russia)

The biogeochemical and molecular biological study of the chemocline and sediments of saline meromictic lakes Shira and Shunet (Khakasia, Russia) was performed. A marked increase in the rates of sulfate reduction and methanogenesis was revealed at the medium depths of the chemocline. The rates of these processes in the bottom sediments decreased with depth. The numbers of the members of domains Bacteria, Archaea, and of sulfate-reducing bacteria (SRB) were determined by fluorescence in situ hybridization with rRNA specific oligonucleotide probes labeled with horseradish peroxidase and subsequent tyramide signal amplification. In the chemocline, both the total microbial numbers and those of Bacteria were shown to increase with depth. The archaea and SRB were present in almost equal numbers. In the lake sediments, a drastic decrease in microbial numbers with depth was revealed. SRB were found to prevail in the upper sediment layer and archaea in the lower one. This finding correlated with the measured rates of sulfate reduction and methanogenesis.     

Microbiological and Isotopic-Geochemical Investigations of Meromictic Lakes in Khakasia in Winter

Microbiological and isotopic-geochemical investigations of the brackish meromictic lakes Shira and Shunet were performed in the steppe region of Khakasia in winter. Measurements made with a submersed sensor demonstrated that one-meter ice transmits light in a quantity sufficient for oxygenic and anoxygenic photosynthesis. As in the summer season, in the community of phototrophic bacteria found in Lake Shira, the purple sulfur bacteria Amoebobacter purpureus dominated, whereas, in Lake Shunet, the green sulfur bacteria Pelodictyon luteolum were predominant. Photosynthetic production, measured using the radioisotopic method, was several times lower than that in summer. The rates of sulfate reduction and production and oxidation of methane in the water column and bottom sediments were also lower than those recorded in summer. The process of anaerobic methane oxidation in the sediments was an exception, being more intense in winter than in summer. The data from radioisotopic measurements of the rates of microbial processes correlate well with the results of determination of the isotopic composition of organic and mineral carbon (δ¹³C) and hydrogen sulfide and sulfate (δ³⁴S) and suggest considerable seasonal variations in the activity of the microbial community in the water bodies investigated.__________Translated from Mikrobiologiya, Vol. 74, No. 4, 2005, pp. 552–561.Original Russian Text Copyright © 2005 by Savvichev, Rusanov, Rogozin, Zakharova, Lunina, Bryantseva, Yusupov, Pimenov, Degermendzhi, Ivanov.     

Biological fractionation of stable carbon isotopes at the aerobic/anaerobic water interface of meromictic water bodies

Mass-spectrometric investigation of carbon isotope composition (δ¹³C) was carried out for suspended organic matter and dissolved mineral compounds for the water column of some meromictic water bodies differing in salinity and trophic state. As a rule, a more pronounced carbon isotope fractionation (resulting from the metabolism of phytoplankton and anoxygenic phototrophic bacteria) was revealed in the zones of enhanced oxygenic and anoxygenic photosynthesis. Carbon isotope fractionation at the border between oxidized and reduced waters depends both on the activity of microbial communities and on the dominant species of phototrophic microorganisms. Analysis of the distribution profiles of the isotopic composition of suspended organic matter and dissolved mineral carbon revealed active mineralization of the organic matter newly formed via anoxygenic photosynthesis in the monimolimnion by microbial communities, resulting in the release of isotopically light carbon dioxide. Mineral carbon in the anaerobic zones of highly productive meromictic water bodies is therefore enriched with the light ¹²C isotope.     

The Phototrophic Community Found in Lake Khilganta (an Alkaline Saline Lake Located in the Southeastern Transbaikal Region)

The structure of the phototrophic community found in Lake Khilganta (the Agin-Buryat Autonomous Area), a shallow saline soda lake (depth, 35–45 cm; water mineralization, 45 g/l; alkalinity, 30 mg-equiv/l; pH 9.5) has been studied. The bottom of the lake is covered with a 10- to 15-mm microbial mat, whose basis is formed by the filamentous cyanobacterium Microcoleus chthonoplastes. The mat exhibits pronounced layering and contains a significant amount of minerals. Six zones, which have characteristic colors and consistencies and are composed of intermittent layers, have been identified along the vertical profile. Live phototrophic bacteria have been found in the three upper zones. The bulk of the cyanobacteria is concentrated in the upper zone. In the lower zones, the development of purple bacteria has been observed. The diurnal dynamics of the vertical distribution of phototrophic microorganisms, which results from variations in the physicochemical environmental parameters, is described. Ectothiorhodospira sp. are dominant among the anoxyphotobacteria present. Their number, determined according to the inoculation method, is 10⁶–10⁷ cells/ml. The purple bacteria of the genera Allochromatium, Thiocapsa, and Rhodovulum are also present. Experiments with isolated pure cultures have shown that the anoxygenic photosynthetic bacteria of Lake Khilganta are halotolerant and alkalitolerant or alkaliphilic. In liquid enrichment cultures, at pH 9.5, the ratio of anoxyphotobacteria species is close to that observed in the lake. When the pH is increased to 10.4, it is Ectothiorhodospira, which is the most adapted to life under increased mineralization and alkalinity, that predominantly develops. Photosynthetic activity has been observed in the three upper mat zones and constitutes, on average, 1.5 g C/(m²h); the share of anoxygenic photosynthesis accounts for 75–95% of the total productivity. The main role in sulfide oxidation belongs to the phototrophic anoxyphotobacteria and cyanobacteria. In terms of the physicochemical conditions and structure of the phototrophic community, Lake Khilganta is similar to shallow saline water bodies of marine origin. The main differences consist in the increased alkalinity and in the consequent prevalence of alkaliphilic and alkalitolerant microorganisms and in the absence of representatives of the neutrophilic group of green sulfur bacteria.__________Translated from Mikrobiologiya, Vol. 74, No. 3, 2005, pp. 410–419.Original Russian Text Copyright © 2005 by Kompantseva, Sorokin, Gorlenko, Namsaraev.     

Saline lakes of the Paroo,inland New South Wales,Australia

Twenty-five lakes from fresh to crystallizing brine in the semi-desert of northwestern New South Wales, Australia, were studied regularly for 27 months. The lakes are small, shallow and ephemeral. Chemically waters are mainly of the NaCl type. Seventy-four species of invertebrate occur in saline waters (>3 g l^–1) with crustaceans such as Parartemia minuta, Apocyclops dengizicus, Daphniopsis queenslandensis, Diacypris spp. and Reticypris spp. dominant, particularly at higher salinities. The insects Tanytarsus barbitarsis and Berosus munitipennis are also important in meso- and hypersaline lakes. They are joined in hypo- and mesosaline waters by many others, including more beetles, odonatans, trichopterans, pyralids, notonectids, and corixids. Species richness declines with increasing salinity. There is a prominent inland faunal component mainly of crustaceans, including P. minuta, D. queenslandensis, R. walbu, Trigonocypris globulosa and Moina baylyi.     

Distribution, diversity, and activity of sulfate-reducing bacteria in the water column in Gek-Gel lake, Azerbaijan

The distribution and activity of sulfate-reducing bacteria (SRB) in the water column of the alpine meromictic Gek-Gel lake were studied. Apart from traditional microbiological methods based on cultivation and on measuring the process rates with radioactive labels, in situ fluorescent hybridization (FISH) was used, which enables identification and quantification without cultivating organisms. The peak rate of sulfate reduction, 0.486 μg S 1⁻¹ day⁻¹, was found in the chemocline at 33 m. The peak SRB number of 2.5×10⁶ cells/ml, as determined by the most probable number method on selective media, was found at the same depth. The phylogenetic affiliation of the SRB, as determined by FISH, revealed the predominance of the Desulfovibrio spp., Desulfobulbus spp., and Desulfoarculus spp./Desulfomonile spp. groups. The numbers of spore-forming Desulfotomaculum spp. increased with depth. The low measured rates of sulfate reduction accompanied by high SRB numbers and the predominance of the groups capable of reducing a wide range of substrates permit us to assume utilization of electron acceptors other than sulfate as the main activity of the SRB in the water column. Original Russian Text ? O.V. Karnachuk, N.V. Pimenov, S.K. Yusupov, Yu.A. Frank, Ya.A. Puhakka, M.V. Ivanov, 2006, published in Mikrobiologiya, 2006, Vol. 75, No. 1, pp. 101–109.     

The Influence of Salt Concentration on the Copy Number of Plasmid pSH1 Replicating in <Emphasis Type="Italic">Micrococcus</Emphasis> sp. 9

The study of heterotrophic bacteria isolated from the brackish waters of Lake Shira has shown that some of them contain plasmid pSH1 of approximately 2.7 kb in size. The number of plasmid copies in plasmid-containing strains cultivated at a minimal concentration of sodium chloride is found to be low, whereas the sub-culturing of these strains at high salt concentrations increases the plasmid number. The role of natural pSH1 plasmid in the osmotolerance of host bacteria is discussed.__________Translated from Mikrobiologiya, Vol. 74, No. 3, 2005, pp. 349–356.Original Russian Text Copyright © 2005 by Lobova, Zagrebel’nyi, Popova.     

Photosynthetic bacteria in meromictic lakes and stratified fjords of the Vestfold Hills,Antarctica

Thirteen meromictic lakes and two permanently stratified fjords in the Vestfold Hills, Antarctica, were surveyed in 1983 for photosynthetic bacteria. Burton Lake and Ellis Fjord were sampled throughout the year to determine seasonal variations. Physical and chemical parameters were recorded and related to the species present. The dominant species in waters with salinities of 100.7 g kg^–1 were Chlorobium vibrioforme and Chlorobium limicola with populations at the O₂–H₂S interface in the range 0.3 to 6.7 × 10⁶ ml^–1. Neither of these species was found at higher salinities. Thiocapsa roseopersicina and a Chromatium sp. were found in low numbers (< 10⁵ ml^–1) in most of the same waters as the Chlorobium spp. These bacterial phototrophs developed in a narrow band below the O₂–H₂S interface where both light and H₂S were available. Very low numbers (< 10² ml^–1) of Rhodopseudomonas palustris were found in both oxic and anoxic waters having salinity 148 g kg^–1. The dominance of the Chlorobium spp. is ascribed to their more efficient maintenance metabolism during the darkness, their faster growth at low light intensities (< 1 µE m^–2 s^–1) and the lack of selective filtering of incident light. The Chlorobium spp. grew well at –2 °C, but not –5°C in hypersaline waters. The concentration of H₂S had no apparent effect on the development of the bacterial flora. Viable cells were found to depths of 100 m in Ellis Fjord indicating that viability in total darkness could have been maintained for periods of the order of 1700 days.     

Abundance and Diversity of Sulfur-Oxidizing Bacteria along a Salinity Gradient in Four Qinghai-Tibetan Lakes,China

Sulfur-oxidizing bacteria (SOB) play important roles in the sulfur cycle and are widespread in a number of environments, but their occurrence and relationship to geochemical conditions in (hyper)saline lakes are still poorly understood. In this study, the abundance and diversity of SOB populations were investigated in four Qinghai-Tibetan lakes (Erhai Lake, Gahai Lake 1, Gahai Lake 2 and Xiaochaidan Lake) by using quantitative polymerase chain reaction (qPCR) and soxB gene- (encoding sulfate thiohydrolase) based phylogenectic analyses. qPCR analyses showed that in the studied lakes, the total bacterial 16S rRNA and soxB gene abundances in the sediments were distinctly higher than in the overlying waters. The 16S rRNA gene abundance in the waters ranged 5.27 × 10⁶–6.09 × 10⁸ copies per mL and 7.39 × 10¹⁰–2.9 × 10¹¹ copies per gram sediment. The soxB gene abundance in the waters ranged from 1.88 × 10⁴ to 5.21 × 10⁵ per mL and 4.73 × 10⁶–2.65 × 10⁷ copies per gram sediment. The soxB gene in the waters of the two hypersaline lakes (Gahai Lake 2 and Xiaochaidan Lake) was more abundant (2.97 × 10⁵ and 5.21 × 10⁵ copies per mL) than that in the two low-salinity lakes (1.88 × 10⁴ and 3.36 × 10⁴ copies per mL). Phylogenetic analysis showed that Alpha- and Betaproteobacteria were dominant SOB in the investigated lakes, and the composition of proteobacterial subgroups varied with salinity: in freshwater Erhai Lake and low-salinity Gahai Lake 1, the SOB populations were dominated by the Betaproteobacteria, whereas in hypersaline Lake Gahai 2 and Xiaochaidan Lake, the SOB populations were dominated by Alphaproteobacteria. Overall, salinity played a key role in controlling the diversity and distribution of SOB populations in the investigated Qinghai-Tibetan lakes.