Phytoplankton Diversity, Abundance and Toxin Synthesis Potential in the Lakes of Natural and Urban Landscapes in Permafrost Conditions
by
, , , and
Sophia Barinova
1,*
,
Viktor A. Gabyshev
2
,
Olga I. Gabysheva
2,
Yanzhima A. Naidanova
3 and
Ekaterina G. Sorokovikova
3
1
Institute of Evolution, University of Haifa, Mount Carmel, 199 Abba Khoushi Ave., Haifa 3498838, Israel
2
Institute for Biological Problems of Cryolithozone Siberian Branch of Russian Academy of Science (IBPC SB RAS), Lenin Av. 41, Yakutsk 677980, Russia
3
Limnological Institute of the Siberian Branch of the Russian Academy of Sciences, 3 Ulan-Batorskaya Str., Irkutsk 664033, Russia
*
Author to whom correspondence should be addressed.
Land 2025, 14(4), 721; https://doi.org/10.3390/land14040721
Submission received: 26 February 2025
/
Revised: 21 March 2025
/
Accepted: 26 March 2025
/
Published: 27 March 2025
(This article belongs to the Special Issue Vegetation Spatial Dynamics and Plant Diversity Trends of Natural and Urban Landscapes)
Abstract
:The region of Eastern Siberia that we have been studying is situated in Yakutia in the permafrost area. We studied five lakes of various geneses, located both in the urbanized territory of Yakutsk city and its suburbs and in natural landscapes at a distance from the impacted area. All lakes were found to have high levels of ammonium nitrogen, total phosphorus and total iron. The lakes’ plankton was found to contain 92 species of algae and cyanobacteria. Cyanobacteria in most lakes accounted for 53 to 98% of the biomass. In one of the natural lakes, 95% of the total biomass was Dinoflagellata. Bioindication, statistics and ecological mapping methods revealed correlations between cyanobacterial production intensity, landscape runoff and lake trophic state. Potentially toxic cyanobacteria containing microcystin and saxitoxin synthesis genes were found in four lakes. Our previous studies established that cyanobacterial harmful algal bloom (CyanoHABs) with microcystin production are characteristic only for lakes in urbanized areas that experience the input of nutrients and organic matter due to anthropogenic runoff. This study indicates that CyanoHABs are possible in lakes in natural areas that are permafrost-dune-type lakes according to their genesis. For the first time in the region, potentially toxic cyanobacteria with saxitoxin synthesis genes have been found. Dune-type lakes do not freeze to the bottom during winter due to taliks underneath them, which provides advantages for cyanobacteria vegetation. Dune-type lakes are very common in the permafrost area, so the extent of CyanoHAB’s distribution in this region may be underestimated.
1. Introduction
The anthropogenic impact on aquatic ecosystems with the expansion of the scale of urban economy, agricultural, and industrial production in the modern world is steadily increasing. Due to the increase in the flow of biogenic and organic substances of anthropogenic origin into water bodies, the rates of primary production and the process of eutrophication are accelerating, and researchers are increasingly noting the phenomenon of “algae bloom” of continental water bodies. By “algae bloom”, we mean the phenomenon of accumulation of phytoplankton biomass in the water column, leading to a visible change in its color and the appearance of accumulations (foam) on the surface of the reservoir.
Blooms may pose a direct threat to human and animal health, particularly when toxin-producing cyanobacteria are dominant. Microcystins (MCs) are the most prevalent and extensively studied cyanotoxins within temperate freshwaters. These toxic cyclic heptapeptides inhibit the activity of liver protein phosphatases 1 and 2A and have potent hepatotoxicity and tumor promotion activity [1]. MCs are produced by planktonic cyanobacteria of the genera Microcystis, Planktothrix, and Dolichospermum. Blooms of Dolichospermum and Aphanizomenon species can also be accompanied by the production of neurotoxins—anatoxin-a and saxitoxins [2]. Given the risk of human poisoning, the World Health Organization has recommended monitoring phytoplankton and cyanotoxins in water bodies and has developed standards for safe concentrations of various cyanotoxins in drinking and recreational waters [3].
Yakutia is a region with a low population density, which is 0.3 individuals per km2. Our research was conducted in the middle reaches of the Lena River basin, in the vicinity of the city of Yakutsk, which is the most densely populated part of Yakutia, where 42% of its population lives. Yakutsk is the largest city in the world located in the permafrost zone. For various socio-economic reasons, due to internal migration processes, local population growth is noted here. The number of residents of Yakutsk has increased by 43% over the past 20 years, currently amounting to 385 thousand people [4]. With population growth, the degree of pressure on water bodies associated with human activities is steadily increasing.
The features of phytoplankton vegetation in lakes located in the middle reaches of the Lena River basin in Central Yakutia have been studied by various authors since the second half of the last century [5,6,7,8,9,10]. According to long-term observations, it is known that in the summer, cyanobacteria dominate both in abundance and in the number of species in the plankton of those lakes in the region that are influenced to varying degrees by anthropogenic factors. The results of our previous studies have shown that despite the harsh climatic conditions (the short growing season and continuous distribution of permafrost), a number of lakes in urbanized areas of this region are characterized by the phenomenon of cyanobacterial harmful algal blooms (CyanoHAB) [11,12]. Thanks to molecular genetic studies, microcystin producers were identified in water bodies of this region of the cryolithozone. The relevance of CyanoHAB research is due to the seriousness of the detrimental consequences for the environment, economy, and human health, which the occurrence of this phenomenon in freshwater bodies leads to [1,2,3]. Therefore, one of the goals of this study was to determine the presence of potentially toxic cyanobacteria, which are producers of microcystins and saxitoxins, in the region’s water bodies.
The increased content of biogenic and organic substances, which is one of the drivers of CyanoHAB occurrence, can be caused not only by anthropogenic factors but also by natural causes, as well as global climatic phenomena. Thus, it is known that due to the insufficient drainage of permafrost soil and additional runoff from the catchment area due to intensive thawing processes, cryolithozone reservoirs are often over-enriched with carbon and organic matter [13]. In addition, permafrost degradation may lead to the mobilization of previously frozen organic carbon and nutrients and their entry into surface waters [14]. Therefore, another goal of this work was to test the hypothesis of the possibility of CyanoHAB occurrence in lakes of this cryolithozone region that are not subject to anthropogenic impact.
2. Materials and Methods
2.1. Site Description
The study area is located in Yakutia at the 62nd parallel north latitude in the middle reaches of the Lena River in a zone of continuous permafrost. The Siberian anticyclone that forms in the center of Asia in winter has a great influence on the climatic conditions of the region. Its powerful spur occupies the whole of Eastern Siberia. The climate features are significantly influenced by frequent intrusions of air masses from the Arctic Ocean with a very low water vapor content in summer. The climate is sharply continental with long, severe winters and short, hot summers. The region is home to the “pole of cold” (the village of Oymyakon), where the lowest temperatures in the Northern Hemisphere of the Earth are recorded. The frost-free period for the study area reaches 90 days [15]. The ice-free period on water bodies, and consequently the growing season, is limited to 120–125 days [16]. The average annual air temperature for the sampling area ranges from −2.4 to −8.7 °C, and the maximum temperature in summer is from 22.2 to 25.0 °C [17].
The work was carried out on five different types of lakes (Table 1, Figure 1 and Figure 2). Some of the lakes are located on the floodplain terrace of the Lena River and are river lakes (oxbow lake origin), representing channels separated from the river. Ierelyakh Lake is of thermokarst origin; its basin was formed as a result of the thawing of underground ice of permafrost. Bolshaya Chabyda Lake is a type of dune lake, the formation of which is caused by the advance of sand on the channel and runoff-trough network under the influence of winds at the turn of the Neopleistocene and Holocene. Dachnoe Lake is of artificial origin (Table 1). Some of the studied reservoirs are located within the city of Yakutsk and its suburbs. Observations were also carried out on lakes located outside the zone of anthropogenic impact.
2.2. Sampling
Field observations were carried out on 11 and 22 July 2024, during the summer low water period at the peak of the growing season. Samples were collected from the surface water layer (0–0.3 m). Phytoplankton for subsequent qualitative and quantitative analysis was collected using an Apstein net made of Sefar Nitex material with a mesh size of 15 μm (Sefar Group JSC, Thal, Switzerland). Net samples for molecular genetic analysis were fixed with 70% ethanol (final concentration). For hydrochemical analysis, water samples were collected by scooping and sent to the laboratory for immediate analysis.
2.3. Phytoplankton Analysis and Examining the Chemical Composition of Water
An Olympus BH-2 light microscope (Olympus Corporation, Tokyo, Japan) was used to examine phytoplankton samples. The quantitative counting of microalgae cells was performed in a 0.01 cm3 Nageotte counting chamber. Biomass was calculated by multiplying the number of cells by their volume, in accordance with [18]. The identification of microalgae species was performed in accordance with the international handbooks [19,20,21,22,23,24,25,26,27]. The relevance of data on the taxonomic affiliation of species was checked on the portal algabase.org [28].
Chemico-analytical work is executed by standard methods [29,30] using instruments listed below: Multitest IPL-101 device (LLC NPP “SEMIKO”, Novosibirsk, Russia), spectrophotometer PE-5300VI (GK “EKROS”, Saint Petersburg, Russia), Fluorat-02-2M device (LLC “Lumex-Marketing”, Saint Petersburg, Russia) and atomic-absorption spectrometer AAnalyst 400 (PerkinElmer Inc., Waltham, MA, USA).
The index diversity of Shannon–Wiener [31] of the phytoplankton community was calculated in the Biodiversity Pro 2.0 program [32]. The calculation of similarity and network graphs was performed as the network analyses in JASP on the botnet package in R [33]. Pearson correlation coefficients were calculated in the Wessa.net program [34].
The saprobity index of the algae community was calculated on the basis of the species-specific index saprobity S and the cell abundance of each indicator species [35,36].
Bioindicator analysis was performed according to [37] with species-specific ecological preferences of the revealed indicator taxa [35]. The BioDiversity Pro 2.0 program was used for similarity calculation [32].
The WESI index [35,36,37] was calculated to assess the influence of toxic pollution on the aquatic ecosystems by the equation
where WESI is an aquatic ecosystem state index, Rank Index S is the rank number from 1 to 9 of calculated for each community index S in the water quality class of [37], Rank N-NO3 is the rank number from 1 to 9 of the defined N-NO3 concentration for each sampling point in the water quality class of [37]. The index values vary from 0 to 5. If the index value is below one, then the ecosystem is exposed to toxic pollution, which inhibits photosynthesis.
WESI = Rank Index S/Rank N-NO3
Statistical maps were created in the Statistica 12.0 program based on the GPS coordinates of sampling points for each biological and chemical variable.
2.4. DNA Extraction, Cloning, Sequencing and Phylogenetic Agnalysis
DNA was extracted using a DNK-Sorb kit (Amplisens, Moscow, Russia). Samples were screened for the presence of genes encoding microcystin and saxitoxin production using primers to regions of the aminotransferase (mcyE) and polyketide synthase (sxtA) genes [38,39]. DNA of the toxin-producing strains Microcystis aeruginosa BN23 and Dolichospermum lemmermannii 20/24BBG isolated from Lake Baikal were used as controls. PCR products of expected size (approximately 470 bp for mcyE, and 555 for sxtA) were subcloned into E. coli XL1BL cells using the CloneJET PCR Cloning Kit (Thermo Fisher Scientific Baltics UAB, Vilnius, Lithuania). Plasmid inserts were sequenced using vector-specific primers on the Nanofor-05 genetic analyzer (Syntol, Moscow, Russia) and BrilliantDye Terminator reagents (NimaGen, Nijmegen, The Netherlands). Sequences were analyzed with Chromas (https://chromas.software.informer.com/, accessed on 1 February 2025) and then aligned with MEGA 10.2 [40]. All unique sequences from this study have been deposited in GenBank under accession numbers PV232687-PV232698 for mcyE and PV232699-PV232704 for sxtA.
The data set was analyzed by neighbor-joining (NJ), and maximum likelihood (ML) methods using MEGA 10.2 software to infer trees and estimate branch support. Evolutionary distances for the NJ tree were calculated by the Kimura-2 parameter model and for ML tree—by the Tamura-3 model. The tree topology was evaluated by bootstrap support in 1000 replicons.
3. Results
3.1. Physico-Chemical Characteristics of the Studied Lakes’ Water
The upper water layer from which the sampling was carried out was warmed up quite well in all the studied lakes (Appendix A Table A1). All the studied lakes have a slightly alkaline reaction. The oxygen regime corresponds to normal values. The concentration of dissolved oxygen was higher in the lakes located outside the city of Yakutsk.
The content of oxidation-resistant organic matter (COD) is high in all the surveyed lakes. High color indices are typical for the Ytyk-Kyuyol and Dachnoe lakes. The waters of the other objects are colored weaker.
The surveyed lakes belong to the hydrocarbonate class, calcium–magnesium group, type II. The waters of the city lakes (Ytyk-Kyuyol and Saysary Lakes) are medium-hard and fresh and have increased mineralization. The other surveyed objects (Dachnoe, Ierelyakh and Bolshaya Chabyda Lakes) are fresh and medium-mineralized and have a hardness index of “soft–medium-hard”.
The content of nitrite nitrogen, nitrate nitrogen, and mineral phosphorus is characterized by relatively low values. The maximum concentration of nitrite nitrogen was found in Dachnoe Lake. The content of mineral phosphorus does not exceed 0.02 mg L−1, and silicon content does not exceed 3.8 mg L−1.
All the lakes surveyed showed high levels of ammonium nitrogen, total phosphorus and total iron. The highest concentrations of ammonium nitrogen were found in Ytyk-Kyuyol and Dachnoe Lakes; the highest levels of total phosphorus were found in Bolshaya Chabyda, Dachnoe, and Saysary Lakes.
3.2. Composition of Phytoplankton Community and Dominant Species
In the plankton of the lakes, 92 species and varieties of algae and cyanobacteria were identified, and one identification was made at the genus level. The highest species diversity of phytoplankton was found in Ytyk-Kyuyol Lake (Appendix A Table A2). The smallest number of species was found in Dachnoe Lake (Figure 3a). Five species were found in all the studied lakes: Botryococcus braunii, Ceratium hirundinella, Pandorina morum, Pseudopediastrum boryanum and Ulnaria acus. In three lakes, representatives of Chlorophyta predominated in terms of the number of species: Saysary, Ytyk-Kyuyol and Ierelyakh Lakes. In Dachnoe Lake, the phylum richest in terms of the number of species was Cyanobacteria; in Bolshaya Chabyda Lake, it was the Heterokontophyta phylum with diatoms (Figure 3b).
During sampling, water bloom was observed in the Bolshaya Chabyda, Dachnoe and Ytyk-Kyuyol Lakes. In the other lakes, we did not notice any signs of water blooming visually. The maximum level of phytoplankton biomass was observed in Bolshaya Chabyda and Dachnoe Lakes. Cyanobacteria in most lakes accounted for 53% to 98% of the phytoplankton biomass (Table 2). The only exception was Ierelyakh Lake, where 95% of the total biomass was Dinoflagellata.
In the plankton of Bolshaya Chabyda, Dachnoe and Ytyk-Kyuyol Lakes, two representatives of the genus Microcystis predominated in biomass, M. aeruginosa and M. flos-aquae, making up 60–70% of the total biomass of phytoplankton (Table 3, Figure 4). In Dachnoe Lake, together with Microcystis, Aphanizomenon flos-aquae was codominant; in Saysary and Bolshaya Chabyda Lakes, representatives of the genus Dolichospermum also dominated. Only in Ierelyakh Lake was Ceratium hirundinella dominant.
Table 4 presents the summary data on species richness, abundance and biomass of phytoplankton for the five lakes studied and shows the calculated Shannon indices based on the Appendix A Table A2 data. Table 4 presents the summary data on the species’ richness, abundance and biomass of phytoplankton for the five studied lakes and also shows the calculated Shannon indices based on the Appendix A Table A2 data. The Saprobity Index S and the Ecosystem State Index WESI were also calculated for the communities of each of the five lakes. The summary data show that the abundance and biomass of the studied lakes phytoplankton correlate well (R = 0.88, p = 0.02), as do the Shannon Index and Saprobity Index (R = 0.82, p = 0.04) and the number of species and Saprobity Index S (R = 0.85, p = 0.03). This indicates a sufficient equilibrium of the ecosystems of the studied lakes. The saprobity index S fluctuated between lakes within the limits of class 3 water quality. The WESI index was >1 and showed the absence of toxic effects on the process of photosynthesis. Consequently, the studied lakes can be considered to be at the initial stage of eutrophication, both under the influence of urban conglomerate and its waste and under the influence of natural factors in the permafrost zone.
3.3. Bioindicators
The response of phytoplankton communities to the main environmental indicators was analyzed using the bioindication method. Figure 5 shows that the percentage composition in all five lakes was dominated by species preferring a planktonic–benthic habitat (Figure 5a, Appendix A Table A3). There were few indicators of temperature conditions, but a significant gradient in their composition in the lakes is visible (Figure 5b). Thus, in Saysary, Ytyk-Kyuyol and Bolshaya Chabyda Lakes, temperate water indicators (eterm and temp) prevailed, while the communities of Ierelyakh and Dachnoe Lakes were significantly enriched in warm-water species. Both lakes were either in the zone of anthropogenic impact or differed in genesis by dune origin with taliks and heating above permafrost. Freshwater species predominated among salinity indicators, combining (up to 50% of indicator composition) with halophiles in Saysary and Ierelyakh Lakes, and in Bolshaya Chabyda Lake, they constituted up to 90% of indicators in phytoplankton (Figure 5c). Oxygen saturation was moderate in Saysary Lake, and in the remaining lakes, it decreased to a minimum in Dachnoe Lake (Figure 5d), corresponding to stagnant waters.
The pH indicators showed that the waters of Saysary Lake were slightly alkaline; Ierelyakh Lake was neutral; and Ytyk-Kyuyol, Dachnoe and Bolshaya Chabyda Lakes were acidified (Figure 6a). All the studied lakes had autotrophic phytoplankton communities according to the composition of nutrition type indicators (Figure 6b). The trophic status indicators were distributed unevenly across the lakes (Figure 6c). Mesotrophic species predominated in Saysary Lake; meso-eutrophic species predominated in Ytyk-Kyuyol, Ierelyakh and Bolshaya Chabyda Lakes; and up to 95% of eutrophic water indicators were in Dachnoe Lake. An assessment of the saturation with organic matter in the studied lakes based on the species-specific saprobity index S showed that the waters of all lakes belong to water quality class 3 (Figure 6d).
3.4. Comparative Analysis
The comparative analysis included the construction of trees with similar chemical and biological data in the lakes studied. Thus, the tree for chemical data showed a high similarity (up to 75%) of environmental indicators for the studied lakes, grouping the data into two clusters: (1) Saysary and Ytyk-Kyuyol and (2) the remaining lakes (Figure 7), which can be considered the result of regional commonality, despite the differences in anthropogenic impact and genesis.
The comparison of biological data (Figure 8) separated Ierelyakh Lake from the rest at the level of 20% similarity, and communities had similarity above 50% in Ytyk-Kyuyol, Dachnoe and Bolshaya Chabyda Lakes.
The most obvious differences in the state of the ecosystems of the studied lakes were observed in the quantitative indicators of phytoplankton. To compare the abundance of cells in communities, the lakes were designated by the degree of their potential load from “control” to “Antrop”, reflecting the position on the landscape relative to the sources of pollution. The comparative plot of the JASP shows the correlation of species abundance of phytoplankton in the five studied lakes (Figure 9). It can be seen that the phytoplankton of the lakes Ierelyakh and Bolshaya Chabyda, located on a landscape rise, is similar in quantitative data. Another pair of lakes close in number is those lakes that we consider to be under anthropogenic influence, Ytyk-Kyuyol and Dachnoye. A separate place with the greatest differences in phytoplankton is Saysary Lake, confirming its peculiarity in the urban landscape.
3.5. Species–Environmental Relationships
At the next stage, the RDA analysis of the relationship between the main environmental parameters and the biomass of taxonomic divisions in the studied lakes was carried out. Figure 10 shows that the environmental parameters are grouped into three main sets. Oxygen, nutrients and phosphorus stimulate the development of cyanobacteria in Dachnoe and Bolshaya Chabyda Lakes. An increase in saturation with dissolved salts (TDS) has a stimulating effect on the development of euglenophytes and green algae in Saysary and Ytyk-Kyuyol Lakes. Dinoflagellates, developing in bulk in Ierelyakh Lake, preferred waters with the absence of ammonium, an indicator of fresh organic pollution.
3.6. Statistical Mapping
The spatial landscape analysis of the distribution of the main environmental parameters and biological data in the lakes studied was carried out using statistical mapping (Figure 11 and Figure 12). Thus, Figure 11a shows the distribution of the heights of the studied lakes and potential pollutants: the city dump and the sand pit. It is evident that the dump is located above Ierelyakh, Dachnoe and Saysary Lakes, which may therefore be under the influence of various uncontrolled substances flowing down during the disposal of garbage and organic waste. The distribution of inorganic phosphates, as the main limiting nutrient for phytoplankton in lakes, is shown in Figure 11b, where only Bolshaya Chabyda Lake stands out with its minimum concentrations. It also differs in genesis, being a dune-type lake. The distribution of nitrates corresponds to the direction of runoff from the city dump, being at a maximum in Ytyk-Kyuyol Lake (Figure 11c). A similar distribution was found for ammonium in lake waters, which was highest in Dachnoe Lake, where the impact of the city landfill runoff and anthropogenic pollution from the population was combined (Figure 11d).
The mapping of biological parameters is shown in Figure 12. Phytoplankton species richness was highest in Ytyk-Kyuyol Lake, where the impact of a municipal landfill with nutrient-bearing runoff was previously identified (Figure 12a). Plankton community abundance was highest in Dachnoe Lake, where the impact of both the landfill and human population was combined (Figure 12b). Biomass was highest in Bolshaya Chabyda Lake (Figure 12c), with both phytoplankton production parameters coinciding with landscape depression (Figure 11a) and cyanobacterial biomass (Figure 12d). Eutrophication was highest in Dachnoe Lake (Figure 12e), and organic pollution as measured by the Saprobic Index S was higher in Ytyk-Kyuyol Lake, located in a landscape depression (Figure 12f).
3.7. Cyanotoxin Synthesis Genes
PCR using total DNA from phytoplankton samples and primers to the stxA gene gave a positive result in samples from Saysary and Ytyk-Kyuyol lakes. The 14 clones obtained from the lakes represented 6 different genotypes. The sequences had 98–99% similarity with the stxA genes of the Aphanizomenon, Dolichospermum and Anabaenopsis strains from water bodies in Germany, France, Spain, Lithuania and Australia and 99% with the sequence of the D. lemmermannii strain from Lake Baikal, Russia, located in southern Eastern Siberia. On the phylogenetic tree, sequences from the lakes of Yakutia formed a separate cluster with bootstrap support above 89% (Figure 13).
Fragments of microcystin synthesis genes mcyE were detected by PCR in the Ytyk-Kyuyol, Dachnoe, and Bolshaya Chabyda Lakes. Thus, cyanobacteria in Ytyk-Kyuyol Lake contained both microcystin and saxitoxin synthesis genes. Comparative analysis of sequences from the lakes of Yakutia with those available in the GenBank database revealed a high homology of 10 amplicons with mcyE genes of toxic representatives of the genus Microcystis and two with Dolichospermum. Two mcyE gene fragments obtained from Dachnoe and Ytyk-Kyuyol Lakes (Dach 24/07-1, Y-K 24/07-2) were 100% identical to the sequences of M. aeruginosa FCY-26 and M. aeruginosa NIES-843 strains isolated from lakes in Korea and Japan. Eight mcyE gene fragments from Yakutian lakes share up to 99% identity with the Microcystis aeruginosa (NIES-88, K-139, NIES-843) and M. viridis NIES-102 strains isolated during toxic blooms from Lakes Kasumigaura and Kawaguchi in Japan. Two mcyE sequences obtained from Ytyk-Kyuyol and Bolshaya Chabyda Lakes (BCh 24/07-1, Y-K 24/07-5) show 98–99% homology with numerous sequences of Dolishospermum-species from Scandinavian lakes, for example D. lemmermannii NIVA-CYA 270/1 (Arefjordvatnet Lake, Norway) and 99% with D. lemmermannii 2/23M from Lake Baikal. On the phylogenetic tree, the obtained clones were grouped together with related sequences into well-supported clades of the genera Microcystis and Dolichospermum (Appendix A Figure A1).
4. Discussion
The increased color and concentration of total iron, ammonium nitrogen, total phosphorus and difficult-to-oxidize organic matter (COD) noted for all the lakes studied are due to insufficient drainage of permafrost soils and additional runoff from the catchment area due to intensive thawing processes [13]. Thus, the trophic state of all the lakes studied is increased. However, the biological availability of biogenic substances may not be so high, despite their increased content. For example, it is known that in waters rich in oxygen and iron, the proportion of biologically available phosphorus is only 8% of the total phosphorus [45].
Among the studied lakes, two are beyond the influence of anthropogenic pollution sources: Bolshaya Chabyda and Ierelyakh Lakes. But only in one of them, Ierelyakh Lake, Dinoflagellata dominated. Dinoflagellate blooms are safe in fresh waters because neurotoxin production is only known in marine species belonging to the genera Alexandrium, Gymnodinium and Pyrodinium [46]. In the plankton of the other studied lakes, the dominant species were representatives of cyanobacteria. At the same time, in the non-impacted Bolshaya Chabyda Lake, the biomass of cyanobacteria was the highest of all the studied lakes, and the water bloom was visually noted. A number of authors who studied the lake phytoplankton of this region have repeatedly noted that the mass development of cyanobacteria is characteristic here only of water bodies experiencing the influx of organic matter and nutrients associated with household wastewater from villages or agricultural enterprises. Thus, the results of the work of L. Kopyrina et al., based on long-term observations of the species composition of nine thermokarst lakes located in the middle section of the Lena River, showed that cyanobacteria dominated only in lakes near villages in the most urbanized and densely populated region of Yakutia, and in lakes of adjacent areas, far from populated areas, Chlorophyta and Bacillariophyta species dominated [6]. Our previous study on the cyanobacteria plankton community of 17 lakes of various geneses, with varying degrees of anthropogenic pressure, located near the city of Yakutsk, near small villages, and at different distances from them, shows the impact of urbanized areas, characterized by increased cyanobacteria cell abundance [7]. A study conducted on the scale of the Eurasian continent found that the species diversity and proportion of cyanobacteria in phytoplankton increase under the influence of anthropogenic eutrophication [47]. The same study notes that the species richness of cyanobacteria clearly increases with an increase in the duration of the ice-free period in water bodies in the direction from north to south but does not show a response to a decrease in the average water temperature during the growing season, which, according to the author, indicates a high resistance of cyanobacteria to climatic stress. Therefore, the biodiversity of cyanobacteria in the area of our studies is significantly lower than in neighboring areas with a milder climate and a long growing season [48].
Bolshaya Chabyda Lake, unlike most previously studied lakes in the cryolithozone, is not of thermokarst origin but of dune-type genesis. Thermokarst lakes have a shallow depth, no more than 1.3 m, and most of these lakes freeze to the bottom in winter. Dune-type lakes also have a shallow depth, up to 1.5–2 m. But deep taliks are characteristic under them, due to which these lakes do not freeze to the bottom [49]. It is known that lake sediments are the main wintering ground for the Microcystis population, maintaining their viability [50,51,52,53]. Microcystis colonies, controlling their own buoyancy, are able to sink to the bottom, where the temperature is usually higher than on the surface, and settle in the sediments and safely survive the winter period. In the spring, when the water column warms up and penetrates solar radiation increases, the wintering benthic population of Microcystis rises into the water and begins to grow. The presence of a deep talk under the dune-type Bolshaya Chabyda Lake and the fact that this lake does not freeze to the bottom in winter create an advantage for a number of cyanobacteria species, which leads to their mass vegetation and bloom in the summer.
In our previous study, we obtained the first data on the distribution of cyanobacterial toxins and carried out the first molecular genetic detection of cyanotoxin producers in the plankton of some lakes in this region [8]. The main producers of microcystins were identified as two species: Microcystis aeruginosa and M. flos-aquae. And during year-round monitoring carried out on Ytyk-Kyuyol Lake, the presence of intracellular and extracellular MCs in the lake ice was recorded [9]. However, the mcyE genes belonging to representatives of the genus Dolichospermum detected in this study in Ytyk-Kyuyol and Bolshaya Chabyda Lakes allow us to conclude that the spectrum of microcystin-producing species in the lakes of Yakutia is wider and, in addition to Microcystis aeruginosa and M. flos-aquae, probably includes D. lemmermannii. A similar ratio of toxic Microcystis and Dolichospermum genotypes (9:1) was previously observed during CyanoHAB in Lake Kotokel, Russia, Eastern Siberia [54]. Isolation of strains showed that the microcystin-producing species in Lake Baikal were M. aeruginosa and D. lemmermannii [55]. That is, despite the differences in the hydrochemical and climatic characteristics of the reservoirs of Yakutia and the south of Eastern Siberia, they had a similar composition of cyanobacteria-producing microcystins.
Among the most potent cyanotoxins are saxitoxin and its variants, neurotoxic alkaloids, also known as paralytic shellfish toxins (PSTs) [12]. In this work, cyanobacteria containing the sxtA gene encoding polyketide synthase responsible for the initiation of saxitoxin synthesis were identified for the first time in the plankton of Saysary and Ytyk-Kyuyol Lakes [39]. The GenBank database contains a small number of sxtA gene sequences of planktonic cyanobacteria—three species of Aphanizomenon and Dolichospermum and one species of Anabaenopsis (Figure 13). Among the probable saxitoxin producers in the species composition of Lake Saysary are Aphanizomenon flos-aquae and Dolichospermum lemmermannii; D. planctonicum was also recorded in Ytyk-Kyuyol Lake (Appendix A Table A2). The phylogenetic relationships between these genera based on the sxtA gene, as well as the 16S rRNA gene, are not resolved, and the identification of saxitoxin-producing species in the lakes of Yakutia requires further research [56]. In some lakes in the temperate zone where CyanoHABs producing PSTs are found (Scandinavian lakes, Lake Baikal), D. lemmermannii was identified as the putative PSTs-producing species [57,58,59], while toxic genotypes of Aphanizomenon spp. and D. planctonicum are common in lakes in Lithuania and Germany [39,60].
In the already classic studies of the CyanoHAB response to the temperature regime of water bodies, researchers associated the threats of the consequences of this phenomenon with an increase in temperature due to the danger of global climate change [61]. Modern studies of cyanobacterial blooms have identified a wider scale of this phenomenon and demonstrated evidence that the mass development of cyanobacteria occurs both with weak water heating and under ice [62]. However, for northern regions, such as the cryolithozone, where the vegetation season is limited to 120–125 days of the ice-free period in water bodies and where lakes are covered with ice for up to 7 months during the harsh winter, the danger of CyanoHAB remains underestimated.
There is increasing evidence that microcystin plays a role in mitigating oxidative stress damage [63] by protecting key proteins involved in photosynthesis and carbon fixation [64,65]. Recent studies in which M. aeruginosa strains were subjected to cold shock during cultivation under continuous light revealed a protective function of microcystin under oxidative stress (the concentration of the toxin in cells doubled) [66]. Although the water in the lakes we studied was well warmed up at the time of sampling, it should be considered that the growing season in the region is very short, and the period of good water warming is very limited and occurs only at its peak [9]. It is obvious that phytoplankton vegetates for a significant part of the season at weak water heating. In addition, for the studied region, given its high-latitude geographical location, in the summer period, a long daylight period is typical. We can assume that in the lakes of Yakutia, suboptimal water temperature can stimulate microcystin synthesis in toxigenic genotypes, which are likely to have a competitive advantage over non-toxic ones. In addition, CyanoHABs may also be characteristic of other water bodies in high-latitude regions with a harsh climate, where similar studies have not been conducted so far.
The detection of potentially toxic cyanobacteria in urban and suburban lakes of Yakutsk used for recreational purposes may pose a threat to the population and require the monitoring of biological indicators and phytoplankton with subsequent statistical mapping of quantitative results. In areas of population and industrial concentration in the cryolithozone, the remote sensing of lake blooms during the summer period of maximum temperatures can be recommended. And for blooming lakes, where cyanobacterial toxins have not been previously determined, the summer screening of cyanotoxins by molecular methods is recommended.
5. Conclusions
All the examined lakes were found to have high contents of ammonium nitrogen, total phosphorus and total iron. The lakes’ plankton had 92 species of algae and cyanobacteria. Cyanobacteria in most lakes accounted for 53% to 98% of the biomass. The only exception was the non-impacted Ierelyakh Lake, where 95% of the total biomass was Dinoflagellata. The studied lakes can be considered to be at the initial stage of eutrophication, both under the influence of urban conglomerate and its waste and under the influence of natural factors in the permafrost zone. CyanoHAB was first noted in the natural lakes of the cryolithozone of this region, which is probably due to the peculiarities of the lake of the dune-type genesis, which does not freeze to the bottom in winter. In the plankton of Bolshaya Chabyda, Dachnoe and Ytyk-Kyuyol Lakes, two representatives of the genus Microcystis, M. aeruginosa and M. flos-aquae, dominated by biomass; mcyE genes related to Microcystis spp. were detected in these lakes, indicating that the blooms were toxic. For the first time in the region, potentially toxic cyanobacteria with saxitoxin synthesis genes have been found in Saysary and Ytyk-Kyuyol Lakes. The discovery of potentially toxic cyanobacteria in the urban and suburban lakes of Yakutsk requires monitoring.
Author Contributions
Conceptualization, S.B. and V.A.G.; methodology, E.G.S. and V.A.G.; software, S.B. and E.G.S.; validation, S.B., V.A.G.; formal analysis, E.G.S., O.I.G. and Y.A.N.; investigation, Y.A.N. and O.I.G.; resources, V.A.G.; data curation, S.B.; writing—original draft preparation, V.A.G., S.B. and E.G.S.; writing—review and editing, E.G.S., S.B. and V.A.G.; visualization, S.B., E.G.S. and O.I.G.; supervision, V.A.G.; project administration, V.A.G.; funding acquisition, V.A.G. and E.G.S. All authors have read and agreed to the published version of the manuscript.
Funding
This research was carried out within the state assignment of the Ministry of Science and Higher Education of the Russian Federation (theme No. FWRS-2021-0023, reg. No. AAAA-A21-121012190038-0; theme No. FWRS-2021-0026, reg. No. AAAA-A21-121012190036-6). Molecular biological works were financially supported by state assignment No. 0279-2021-0015.
Data Availability Statement
The original contributions presented in the study are included in the article, further inquiries can be directed to the corresponding author.
Acknowledgments
We are grateful to the Israeli Ministry of Aliyah and Integration for partial support of this work. Sequencing was carried out in The Shared Research Facilities for Physical and Chemical Ultramicroanalysis LIN SB RAS. The authors express their gratitude to Viktoria Filippova, research fellow at the Institute for Biological Problems of the Siberian Branch of Russian Academy of Sciences, for providing photographs of the sampling sites.
Conflicts of Interest
The authors declare no conflicts of interest.
Abbreviations
The following abbreviations are used in this manuscript:
| JASP | Jeffreys’s Amazing Statistics Program |
| WESI | Water Ecosystem State Index |
| CCA | Canonical correspondence analysis |
| TDS | Total dissolved solids |
| TSS | Total suspended solids |
| COD | Chemical oxygen demand |
Appendix A
Table A1.
Physical and chemical variables of water in the studied lakes in the permafrost zone of Yakutia based on samples collected in 2024. Abbreviated station code names as in Table 1.
Table A1.
Physical and chemical variables of water in the studied lakes in the permafrost zone of Yakutia based on samples collected in 2024. Abbreviated station code names as in Table 1.
| Variables | Sa | Yt | Ie | Da | BC |
|---|---|---|---|---|---|
| Water temperature, °C | 26.90 | 25.50 | 23.80 | 23.40 | 22.10 |
| pH | 8.44 | 8.28 | 8.47 | 7.98 | 8.41 |
| O2, mg L−1 | 7.31 | 11.37 | 14.63 | 20.15 | 20.96 |
| COD, mg L−1 | 104.90 | 118.30 | 118.00 | 104.70 | 142.90 |
| Pt-Co units | 44 | 75 | 47 | 85 | 36.00 |
| TDS, mg L−1 | 609.96 | 512.86 | 337.28 | 235.09 | 346.14 |
| Hardness, mmol. L−1 | 5.84 | 4.80 | 3.12 | 2.48 | 2.77 |
| Ca2+, mg L−1 | 49.70 | 49.70 | 19.24 | 30.46 | 42.00 |
| Mg2+, mg L−1 | 40.82 | 28.19 | 26.24 | 11.66 | 8.19 |
| Na+, mg L−1 | 60.00 | 50.60 | 30.40 | 14.30 | 38.00 |
| K+, mg L−1 | 8.24 | 7.72 | 12.10 | 4.97 | 8.04 |
| HCO3−, mg L−1 | 332.00 | 274.00 | 189.00 | 130.00 | 150.00 |
| Cl−, mg L−1 | 100.00 | 77.35 | 54.20 | 25.70 | 12.21 |
| SO42−, mg L−1 | 19.20 | 25.30 | <10.0 | 18.00 | 87.70 |
| N-NH4+, mg L−1 | 0.71 | 0.85 | 0.60 | 0.92 | 0.66 |
| N-NO2−, mg L−1 | 0.006 | 0.009 | 0.006 | 0.021 | 0.010 |
| N-NO3−, mg L−1 | 0.10 | 0.16 | 0.12 | 0.15 | 0.12 |
| P-PO43−, mg L−1 | 0.02 | <0.02 | <0.02 | 0.02 | <0.02 |
| Ptot, mg L−1 | 0.80 | 0.60 | 0.40 | 0.90 | 1.84 |
| Si, mg L−1 | 1.88 | 3.55 | 1.51 | 3.88 | 1.93 |
| Fetot, mg L−1 | 0.40 | 0.57 | 0.53 | 0.57 | 0.35 |
Table A2.
Average abundance (thou. cells per L) of algae and cyanobacteria species from the studied lakes in the permafrost zone of Yakutia, based on samples collected in 2024. Abbreviated station code names as in Table 1.
Table A2.
Average abundance (thou. cells per L) of algae and cyanobacteria species from the studied lakes in the permafrost zone of Yakutia, based on samples collected in 2024. Abbreviated station code names as in Table 1.
| Species | Sa | Yt | Ie | Da | BC |
|---|---|---|---|---|---|
| Charophyta | |||||
| Closterium acutum var. linea (Perty) West & G.S.West | 5.6 | ||||
| Closterium leibleinii Kützing ex Ralfs | 0.0002 | ||||
| Cosmarium formosulum Hoff | 0.1 | 0.0008 | 0.002 | ||
| Staurastrum boreale West & G.S.West | 1.5 | 0.6 | |||
| Staurastrum gracile Ralfs ex Ralfs | 0.004 | ||||
| Staurastrum manfeldtii Delponte | 0.0007 | ||||
| Staurastrum tetracerum Ralfs ex Ralfs | 1.6 | 0.02 | |||
| Chlorophyta | |||||
| Actinastrum hantzschii Lagerheim | 363.9 | 162.3 | |||
| Ankistrodesmus arcuatus Korshikov | 180.7 | ||||
| Ankistrodesmus fusiformis Corda | 158.4 | ||||
| Botryococcus braunii Kützing | 366.6 | 161.1 | 53.7 | 620.7 | 1521.9 |
| Coelastrum astroideum De Notaris | 60.2 | 22.4 | 0.3 | ||
| Desmodesmus armatus (Chodat) E.H.Hegewald | 599.6 | 59.6 | 0.3 | 3.1 | |
| Desmodesmus spinosus (Chodat) E.Hegewald | 522.2 | 2.2 | 154.6 | ||
| Dicellula geminata (Printz) Korshikov | 30 | ||||
| Kirchneriella lunaris (Kirchner) Möbius | 160.4 | ||||
| Lagerheimia genevensis (Chodat) Chodat | 200.4 | ||||
| Lagerheimia subsalsa Lemmermann | 211.5 | ||||
| Lemmermannia tetrapedia (Kirchner) Lemmermann | 306.7 | 94 | |||
| Micractinium pusillum Fresenius | 164 | 52.9 | |||
| Monoraphidium contortum (Thuret) Komárková-Legnerová | 283.5 | 92.1 | |||
| Monoraphidium griffithii (Berkeley) Komárková-Legnerová | 142.7 | 53.7 | |||
| Mucidosphaerium pulchellum (H.C.Wood) C.Bock, Proschold & Krienitz | 164 | 54 | 2.4 | 17.4 | |
| Oocystis borgei J.W.Snow | 90.1 | ||||
| Oocystis lacustris Chodat | 0.4 | ||||
| Pandorina morum (O.F.Müller) Bory | 135.2 | 22.5 | 133.2 | 14.6 | 38.3 |
| Parapediastrum biradiatum (Meyen) E.Hegewald | 7.9 | ||||
| Pediastrum duplex Meyen | 14.7 | 5.3 | 0.2 | ||
| Pseudopediastrum boryanum (Turpin) E.Hegewald | 23.8 | 42.4 | 0.9 | 4.1 | 15.9 |
| Scenedesmus obtusus f. disciformis (Chodat) Compère | 70.4 | 0.6 | |||
| Scenedesmus obtusus Meyen | 177.7 | 14.8 | |||
| Scenedesmus quadricauda (Turpin) Brébisson | 481 | ||||
| Schroederia setigera (Schröder) Lemmermann | 67.4 | 19.7 | |||
| Selenastrum bibraianum Reinsch | 25.6 | ||||
| Tetradesmus lagerheimii M.J.Wynne & Guiry | 497.3 | 282.8 | 376.5 | ||
| Tetradesmus obliquus (Turpin) M.J.Wynne | 52.2 | ||||
| Tetraëdron caudatum (Corda) Hansgirg | 13.2 | ||||
| Tetraëdron minimum (A.Braun) Hansgirg | 29 | 11.6 | |||
| Treubaria planctonica (G.M.Smith) Korshikov | 363.5 | ||||
| Cyanobacteria | |||||
| Anathece clathrata (West & G.S.West) Komárek, Kaštovský & Jezberová | 6250.4 | 35,419.2 | |||
| Aphanizomenon flos-aquae Ralfs ex Bornet & Flahault | 3322.8 | 530.5 | 22,091.8 | ||
| Aphanocapsa conferta (West & G.S.West) Komárková-Legnerová & Cronberg | 8.7 | ||||
| Aphanocapsa delicatissima West & G.S.West | 980.8 | ||||
| Aphanocapsa incerta (Lemmermann) G.Cronberg & Komárek | 8575.2 | 255.7 | 5540.5 | ||
| Aphanocapsa planctonica (G.M.Smith) Komárek & Anagnostidis | 3434.5 | ||||
| Chroococcus turgidus (Kützing) Nägeli | 2 | ||||
| Dolichospermum sp. | 373.7 | ||||
| Dolichospermum lemmermannii (Richter) P.Wacklin, L.Hoffmann & J.Komárek | 158.1 | 8 | 10.3 | 2015.2 | |
| Dolichospermum perturbatum (H.Hill) Wacklin, L.Hoffmann & Komárek | 173.6 | ||||
| Dolichospermum planctonicum (Brunnthaler) Wacklin, L.Hoffmann & Komárek | 32.4 | 87.7 | |||
| Dolichospermum sigmoideum (Nygaard) Wacklin, L.Hoffmann & Komárek | 382.4 | ||||
| Dolichospermum spiroides (Klebahn) Wacklin, L.Hoffmann & Komárek | 241.7 | ||||
| Lyngbya cincinnata (Itzigsohn) Compère | 0.6 | ||||
| Merismopedia tranquilla (Ehrenberg) Trevisan | 635.6 | ||||
| Microcystis aeruginosa (Kützing) Kützing | 4847.8 | 1736.1 | 13,621.8 | ||
| Microcystis flos-aquae (Wittrock) Kirchner | 23,607.9 | 76,325.1 | 49,904.9 | ||
| Microcystis wesenbergii (Komárek) Komárek ex Komárek | 320.4 | 553.8 | 300 | 2353.8 | |
| Oscillatoria anguina Bory ex Gomont | 0.7 | ||||
| Oscillatoria limosa C.Agardh ex Gomont | 0.3 | ||||
| Oscillatoria princeps Vaucher ex Gomont | 17.4 | ||||
| Phormidium ambiguum Gomont | 76.7 | ||||
| Phormidium chalybeum (Mertens ex Gomont) Anagnostidis & Komárek | 97 | ||||
| Snowella lacustris (Chodat) Komárek & Hindák | 954.9 | 9.5 | |||
| Woronichinia naegeliana (Unger) Elenkin | 1923.1 | 7777.8 | |||
| Dinoflagellata | |||||
| Ceratium hirundinella (O.F.Müller) Dujardin | 0.6 | 1.4 | 22.7 | 0.04 | 0.04 |
| Peridinium cinctum (O.F.Müller) Ehrenberg | 0.0004 | ||||
| Euglenophyta | |||||
| Lepocinclis acus (O.F.Müller) B.Marin & Melkonian | 5.4 | 1.6 | 0.01 | ||
| Lepocinclis oxyuris (Schmarda) B.Marin & Melkonian | 0.2 | 0.003 | |||
| Monomorphina pyrum (Ehrenberg) Mereschkowsky | 4.2 | 2.3 | |||
| Phacus longicauda (Ehrenberg) Dujardin | 0.2 | ||||
| Phacus orbicularis Hübner | 0.8 | ||||
| Strombomonas acuminata (Schmarda) Deflandre | 0.4 | ||||
| Trachelomonas dybowskii Dreżepolski | 0.4 | ||||
| Trachelomonas granulosa Playfair | 0.01 | ||||
| Trachelomonas hispida (Perty) F.Stein | 0.005 | ||||
| Trachelomonas planctonica Svirenko | 2 | ||||
| Trachelomonas woycickii Koczwara | 3.6 | ||||
| Heterokontophyta | |||||
| Acanthoceras zachariasii (Brun) Simonsen | 4.5 | ||||
| Amphora ovalis (Kützing) Kützing | 0.1 | ||||
| Aulacoseira granulata (Ehrenberg) Simonsen | 19.1 | 26.6 | |||
| Craticula cuspidata (Kützing) D.G.Mann | 0.4 | ||||
| Dinobryon divergens O.E.Imhof | 10.6 | 0.3 | 4.4 | ||
| Dinobryon sociale (Ehrenberg) Ehrenberg | 50.6 | ||||
| Frustulia saxonica Rabenhorst | 0.1 | ||||
| Navicula radiosa Kützing | 1.1 | ||||
| Nitzschia acicularis (Kützing) W.Smith | 57.2 | ||||
| Pseudostaurastrum limneticum (Borge) Guiry | 2.8 | ||||
| Rhoicosphenia abbreviata (C.Agardh) Lange-Bertalot | 0.1 | ||||
| Stauroneis phoenicenteron (Nitzsch) Ehrenberg | 0.01 | 0.02 | |||
| Surirella librile (Ehrenberg) Ehrenberg | 0.3 | ||||
| Tetraplektron laevis (Bourrelly) Ettl | 0.7 | ||||
| Ulnaria acus (Kützing) Aboal | 18.1 | 7 | 2.1 | 18.3 | 33.9 |
| Ulnaria ulna (Nitzsch) Compère | 28.1 |
Table A3.
Ecological preferences of algae and cyanobacteria species from the studied lakes in permafrost zone of Yakutia based on samples collected in 2024.
Table A3.
Ecological preferences of algae and cyanobacteria species from the studied lakes in permafrost zone of Yakutia based on samples collected in 2024.
| Taxa | Hab | T | OXY | HAL | pH | D | AUT-HET | TRO | Index S |
|---|---|---|---|---|---|---|---|---|---|
| Charophyta | |||||||||
| Closterium acutum var. linea (Perty) West & G.S.West | P-B | - | - | - | ind | - | - | m | 2.2 |
| Closterium leibleinii Kützing ex Ralfs | P-B | - | st-str | - | ind | - | - | e | 2.6 |
| Cosmarium formosulum Hoff | P-B | - | - | - | ind | - | - | me | 1.8 |
| Staurastrum boreale West & G.S.West | B | - | - | - | ind | - | - | m | - |
| Staurastrum gracile Ralfs ex Ralfs | P-B | - | st | i | acf | - | - | m | - |
| Staurastrum manfeldtii Delponte | B | - | - | - | ind | - | - | e | - |
| Staurastrum tetracerum Ralfs ex Ralfs | P-B | - | st-str | i | ind | - | - | om | 1.3 |
| Chlorophyta | |||||||||
| Actinastrum hantzschii Lagerheim | P-B | - | st-str | i | - | - | - | - | 2.3 |
| Ankistrodesmus arcuatus Korshikov | P-B | - | st-str | i | - | - | - | - | 2.1 |
| Ankistrodesmus fusiformis Corda | P-B | - | st-str | i | - | - | - | e | 2.0 |
| Botryococcus braunii Kützing | P-B | - | st | i | ind | - | - | - | 1.5 |
| Coelastrum astroideum De Notaris | P | - | st-str | - | - | - | - | e | 2.2 |
| Desmodesmus armatus (Chodat) E.H.Hegewald | P-B | - | st-str | - | - | - | - | e | 1.9 |
| Desmodesmus spinosus (Chodat) E.Hegewald | P-B | - | st-str | - | alf | - | - | - | 2.0 |
| Dicellula geminata (Printz) Korshikov | P-B | - | st | - | - | - | - | me | 2.2 |
| Kirchneriella lunaris (Kirchner) Möbius | P-B | - | st-str | i | - | - | - | e | - |
| Lagerheimia genevensis (Chodat) Chodat | P | - | - | i | - | - | - | - | 2.1 |
| Lagerheimia subsalsa Lemmermann | P-B | - | st-str | - | - | - | - | - | 1.2 |
| Lemmermannia tetrapedia (Kirchner) Lemmermann | P-B | - | st-str | i | ind | - | - | e | 2.0 |
| Micractinium pusillum Fresenius | P-B | - | st-str | - | - | - | - | m | 0.3 |
| Monoraphidium contortum (Thuret) Komárková-Legnerová | P-B | - | st-str | i | - | - | - | - | - |
| Monoraphidium griffithii (Berkeley) Komárková-Legnerová | P-B | - | st-str | i | - | - | - | e | 2.5 |
| Mucidosphaerium pulchellum (H.C.Wood) C.Bock, Proschold & Krienitz | P-B | - | st-str | i | ind | - | - | - | 1.8 |
| Oocystis borgei J.W.Snow | P-B | - | st-str | i | ind | - | - | e | 1.7 |
| Oocystis lacustris Chodat | P-B | - | st-str | hl | - | - | - | - | - |
| Pandorina morum (O.F.Müller) Bory | P | - | st | i | - | - | - | m | - |
| Parapediastrum biradiatum (Meyen) E.Hegewald | P-B | - | - | i | alb | - | - | - | 2.9 |
| Pediastrum duplex Meyen | P | - | st-str | i | ind | - | - | e | - |
| Pseudopediastrum boryanum (Turpin) E.Hegewald | P-B | - | st-str | i | ind | - | - | e | 2.1 |
| Scenedesmus obtusus f. disciformis (Chodat) Compère | P | - | - | - | - | - | - | e | 1.9 |
| Scenedesmus obtusus Meyen | P-B | - | st-str | - | - | - | - | e | 1.8 |
| Scenedesmus quadricauda (Turpin) Brébisson | P | - | - | i | - | - | - | - | 2.1 |
| Schroederia setigera (Schröder) Lemmermann | P | - | st-str | i | alf | - | - | e | 1.7 |
| Selenastrum bibraianum Reinsch | P-B | - | st-str | - | - | - | - | e | 1.7 |
| Tetradesmus lagerheimii M.J.Wynne & Guiry | P-B | - | st-str | i | ind | - | - | e | 2.15 |
| Tetradesmus obliquus (Turpin) M.J.Wynne | P-B | - | st-str | i | ind | - | - | ot | 2.4 |
| Tetraëdron caudatum (Corda) Hansgirg | P-B | - | st-str | i | ind | - | - | e | 2.0 |
| Tetraëdron minimum (A.Braun) Hansgirg | P-B | - | st-str | i | alf | - | - | e | 2.1 |
| Treubaria planctonica (G.M.Smith) Korshikov | P | - | st | - | - | - | - | - | 1.9 |
| Cyanobacteria | |||||||||
| Anathece clathrata (West & G.S.West) Komárek, Kaštovský & Jezberová | P-B | - | - | hl | - | - | - | me | 1.8 |
| Aphanizomenon flos-aquae Ralfs ex Bornet & Flahault | P-B | - | - | hl | alb | - | - | m | 1.95 |
| Aphanocapsa conferta (West & G.S.West) Komárková-Legnerová & Cronberg | P | - | - | i | - | - | - | me | - |
| Aphanocapsa delicatissima West & G.S.West | P-B | - | - | i | - | - | - | m | - |
| Aphanocapsa incerta (Lemmermann) G.Cronberg & Komárek | P-B | - | - | i | - | - | - | me | 2.2 |
| Aphanocapsa planctonica (G.M.Smith) Komárek & Anagnostidis | P-B | - | - | i | - | - | - | o-e | - |
| Chroococcus turgidus (Kützing) Nägeli | P-B | - | aer | hl | alf | - | - | e | 0.8 |
| Dolichospermum sp. | - | - | - | - | - | - | - | - | - |
| Dolichospermum lemmermannii (Richter) P.Wacklin, L.Hoffmann & J.Komárek | P | - | - | i | - | - | - | e | - |
| Dolichospermum perturbatum (H.Hill) Wacklin, L.Hoffmann & Komárek | P | - | - | - | - | - | - | m | - |
| Dolichospermum planctonicum (Brunnthaler) Wacklin, L.Hoffmann & Komárek | P-B | - | st-str | - | - | - | - | e | 2.0 |
| Dolichospermum sigmoideum (Nygaard) Wacklin, L.Hoffmann & Komárek | P | - | - | i | - | - | - | e | 1.7 |
| Dolichospermum spiroides (Klebahn) Wacklin, L.Hoffmann & Komárek | P-B | - | st-str | i | - | - | - | e | 1.3 |
| Lyngbya cincinnata (Itzigsohn) Compère | - | - | - | - | - | - | - | - | - |
| Merismopedia tranquilla (Ehrenberg) Trevisan | P-B | - | - | i | ind | - | - | - | 2.3 |
| Microcystis aeruginosa (Kützing) Kützing | P-B | - | - | hl | acf | - | - | me | 2.2 |
| Microcystis flos-aquae (Wittrock) Kirchner | P-B | - | - | i | - | - | - | e | 1.6 |
| Microcystis wesenbergii (Komárek) Komárek ex Komárek | P-B | - | - | - | - | - | - | - | 2.3 |
| Oscillatoria anguina Bory ex Gomont | B | - | - | - | - | - | - | - | - |
| Oscillatoria limosa C.Agardh ex Gomont | P-B | - | st-str | hl | alf | - | - | - | - |
| Oscillatoria princeps Vaucher ex Gomont | P-B | - | st-str | - | ind | - | - | - | - |
| Phormidium ambiguum Gomont | B | eterm | st-str | i | ind | - | - | - | - |
| Phormidium chalybeum (Mertens ex Gomont) Anagnostidis & Komárek | P-B | - | st-str | - | - | - | - | e | 3.3 |
| Snowella lacustris (Chodat) Komárek & Hindák | P | - | - | i | alb | - | - | me | 1.6 |
| Woronichinia naegeliana (Unger) Elenkin | P | - | st | - | - | - | - | e | 1.8 |
| Dinoflagellata | |||||||||
| Ceratium hirundinella (O.F.Müller) Dujardin | P | - | st-str | i | - | - | - | e | 1.3 |
| Peridinium cinctum (O.F.Müller) Ehrenberg | P-B | - | st-str | i | - | - | - | - | 1.4 |
| Euglenophyta | |||||||||
| Lepocinclis acus (O.F.Müller) B.Marin & Melkonian | P | eterm | st | i | ind | - | - | - | 2.4 |
| Lepocinclis oxyuris (Schmarda) B.Marin & Melkonian | P-B | - | st-str | mh | ind | - | - | - | 2.3 |
| Monomorphina pyrum (Ehrenberg) Mereschkowsky | P-B | eterm | st-str | mh | ind | - | - | - | - |
| Phacus longicauda (Ehrenberg) Dujardin | P-B | - | st | i | ind | - | - | - | 2.8 |
| Phacus orbicularis Hübner | P-B | - | st-str | i | ind | - | - | - | - |
| Strombomonas acuminata (Schmarda) Deflandre | P | - | st-str | i | ind | - | - | - | - |
| Trachelomonas dybowskii Dreżepolski | P | - | - | - | - | - | - | - | 2.3 |
| Trachelomonas granulosa Playfair | P | eterm | - | - | - | - | - | - | 2.2 |
| Trachelomonas hispida (Perty) F.Stein | P-B | eterm | st-str | i | acf | - | - | - | 2.2 |
| Trachelomonas planctonica Svirenko | P | eterm | st-str | I | ind | - | - | - | 2.1 |
| Trachelomonas woycickii Koczwara | P | - | st | - | - | - | - | - | 2.3 |
| Heterokontophyta | |||||||||
| Acanthoceras zachariasii (Brun) Simonsen | P | - | st-str | i | ind | - | - | - | 1.4 |
| Amphora ovalis (Kützing) Kützing | B | temp | st-str | i | alf | sx | ate | e | 1.5 |
| Aulacoseira granulata (Ehrenberg) Simonsen | P-B | temp | st-str | i | alf | es | ate | e | 2.0 |
| Craticula cuspidata (Kützing) D.G.Mann | B | temp | st-str | i | alf | es | - | me | 2.45 |
| Dinobryon divergens O.E.Imhof | P-B | - | st-str | i | ind | - | - | - | 1.2 |
| Dinobryon sociale (Ehrenberg) Ehrenberg | P | - | - | i | - | - | - | - | 1.3 |
| Frustulia saxonica Rabenhorst | B | temp | st-str | hb | acf | - | ate | - | - |
| Navicula radiosa Kützing | B | temp | st-str | i | ind | sx | - | - | - |
| Nitzschia acicularis (Kützing) W.Smith | P-B | temp | st | i | alf | es | ats | om | 1.4 |
| Pseudostaurastrum limneticum (Borge) Guiry | P | - | st-str | - | - | - | - | e | - |
| Rhoicosphenia abbreviata (C.Agardh) Lange-Bertalot | B | temp | st-str | i | alf | es | ate | me | 1.9 |
| Stauroneis phoenicenteron (Nitzsch) Ehrenberg | P-B | temp | st-str | i | ind | - | - | - | - |
| Surirella librile (Ehrenberg) Ehrenberg | P-B | temp | st-str | i | alf | - | hne | - | - |
| Tetraplektron laevis (Bourrelly) Ettl | - | - | - | - | - | - | - | - | - |
| Ulnaria acus (Kützing) Aboal | P-B | warm | st-str | i | alf | es | ate | me | 1.85 |
| Ulnaria ulna (Nitzsch) Compère | P-B | temp | st-str | i | alf | es | ate | e | 2.4 |
Abbreviations: Habitat (Hab) (P—planktonic, P-B—plankto-benthic, B—benthic; aer—aerophytes); temperature (T) preferences (temp—temperate, eterm—eurythermic, warm—warm-water); oxygenation and water moving (Oxy) (aer—aerophiles, st-str—low streaming water, st—standing); salinity ecological groups (Hal) according to [41] (hb—oligohalobes-halophobes, i—oligohalobes-indifferent, hl—halophiles; mh—mesohalobes); pH preferences groups (pH) according to [42] (alb—alkalibiontes; alf—alkaliphiles, ind—indifferent; acf—acidophiles); organic pollution indicators according to [67] (D): sx—saproxenes; es—eurysaprobes; nitrogen uptake metabolism (Aut-Het) [43]: ats—nitrogen-autotrophic taxa, tolerating very small concentrations of organically bound nitrogen; ate—nitrogen-autotrophic taxa, tolerating elevated concentrations of organically bound nitrogen; hne—facultative nitrogen-heterotrophic taxa, needing periodically elevated concentrations of organically bound nitrogen; Trophic state indicators (Tro) [43]: (ot—oligotraphentic; om—oligomesotraphentic; m—mesotraphentic; me—mesoeutraphentic; e—eutraphentic; o-e—oligo to hypereutraphentic); Index S, species-specific index saprobity according to [44].
Table A4.
Average abundance (thou. cells per L) in ecological groups of indicators of phytoplankton in the studied lakes of the permafrost zone of Yakutia based on samples collected in 2024. Abbreviated station code names as in Table 1.
Table A4.
Average abundance (thou. cells per L) in ecological groups of indicators of phytoplankton in the studied lakes of the permafrost zone of Yakutia based on samples collected in 2024. Abbreviated station code names as in Table 1.
| Variable | Sa | Yt | Ie | Da | BC |
|---|---|---|---|---|---|
| Habitat | |||||
| B | 76.676 | 1.502 | 0.780 | 0 | 2.318 |
| P-B | 7671.032 | 44,310.455 | 6568.461 | 106,745.137 | 104,459.438 |
| P | 1446.599 | 3034.493 | 175.086 | 7976.926 | 2534.584 |
| Temperature | |||||
| temp | 0 | 76.293 | 0.068 | 0.013 | 56.677 |
| eterm | 86.255 | 5.881 | 0.028 | 0 | 0 |
| warm | 18.145 | 7.031 | 2.109 | 18.281 | 33.867 |
| Oxygen | |||||
| aer | 0 | 0 | 0 | 0 | 1.962 |
| st-str | 3885.325 | 1917.010 | 31.691 | 131.160 | 656.791 |
| st | 874.278 | 2195.718 | 186.919 | 8413.130 | 1560.230 |
| Salinity | |||||
| hb | 0 | 0 | 0 | 0 | 0.065 |
| i | 3541.633 | 38,973.991 | 489.403 | 82,551.148 | 55,429.458 |
| hl | 3322.785 | 5378.215 | 6250.760 | 23,828.256 | 49,042.966 |
| mh | 4.188 | 2.496 | 0.003 | 0 | 0 |
| Water pH | |||||
| acf | 0 | 4847.756 | 0.009 | 1736.111 | 13,621.860 |
| ind | 1572.928 | 1365.943 | 57.407 | 642.309 | 1920.507 |
| alf | 636.709 | 114.592 | 4.671 | 18.281 | 245.941 |
| alb | 3322.785 | 1493.276 | 9.549 | 22,091.772 | 0 |
| Watanabe | |||||
| sx | 0 | 0 | 0 | 0 | 1.235 |
| es | 18.145 | 83.324 | 2.177 | 18.281 | 88.937 |
| Autotrophy-Heterotrophy | |||||
| ats | 0 | 57.165 | 0 | 0 | 0 |
| ate | 18.145 | 26.159 | 2.177 | 18.281 | 88.769 |
| hne | 0 | 0 | 0 | 0 | 0.291 |
| Trophic state | |||||
| ot | 0 | 0.052 | 0 | 0 | 0 |
| om | 0.002 | 0.057 | 0 | 0 | 0 |
| m | 3.622 | 0.136 | 0.134 | 2.480 | 0.138 |
| me | 0.018 | 14.389 | 0.902 | 2.309 | 49.075 |
| e | 2.401 | 5.252 | 0.025 | 84.201 | 5.674 |
| o-e | 0 | 3.434 | 0 | 0 | 0 |
| Class of Water Quality | |||||
| Class 1 | 164.49 | 52.932 | 0 | 0 | 0 |
| Class 2 | 378.917 | 674.433 | 77.0084 | 63.15 | 1528.872 |
| Class 3 | 6929.046 | 20,904.405 | 901.52 | 88,820.915 | 57,521.776 |
| Class 4 | 97.43 | 8.083 | 0.0002 | 0 | 0 |
Abbreviations: Habitat (P—planktonic, P-B—plankto-benthic, B—benthic; aer—aerophytes); Temperature preferences (temp—temperate, eterm—eurythermic, warm—warm-water); oxygenation and water moving (Oxy) (aer—aerophiles, st-str—low streaming water, st—standing); Salinity ecological groups according to [41] (hb—oligohalobes-halophobes, i—oligohalobes-indifferent, hl—halophiles; mh—mesohalobes); pH preferences groups (pH) according to [42] (alb—alkalibiontes; alf—alkaliphiles, ind—indifferent; acf—acidophiles); organic pollution indicators according to Watanabe et al. [67] (sx—saproxenes; es—eurysaprobes); nitrogen uptake metabolism (Autotrophy-Heterotrophy) [43]: ats—nitrogen-autotrophic taxa, tolerating very small concentrations of organically bound nitrogen; ate—nitrogen-autotrophic taxa, tolerating elevated concentrations of organically bound nitrogen; hne—facultative nitrogen-heterotrophic taxa, needing periodically elevated concentrations of organically bound nitrogen; Trophic state indicators [43]: (ot—oligotraphentic; om—oligomesotraphentic; m—mesotraphentic; me—mesoeutraphentic; e—eutraphentic; o-e—oligo to hypereutraphentic); Class of Water Quality according Index S [44].
Figure A1.
Maximum likelihood tree based on partial mcyE and ndaF sequences of planktonic cyanobacteria. Sequences from this study are bold. Numbers above the branches are bootstrap support as a percentage of 1000 replicates for ML and NJ methods. The bar indicates 5% sequence divergence.
Figure A1.
Maximum likelihood tree based on partial mcyE and ndaF sequences of planktonic cyanobacteria. Sequences from this study are bold. Numbers above the branches are bootstrap support as a percentage of 1000 replicates for ML and NJ methods. The bar indicates 5% sequence divergence.
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Figure 1.
Research area located on the left side of the Lena River near the city of Yakutsk. The map highlights the city’s territory in red and suburban areas in blue. The white arrows on the map denote the lakes that have been explored: (Sa) Saysary, (Yt) Ytyk-Kyuyol, (Ie) Ierelyakh, (Da) Dachnoe, (BC) Bolshaya Chabyda. Yellow arrows and numbers indicate sources of human-made impact outside the city’s and suburban territory: (1) municipal controlled landfill dump, (2) sand quarry. In the lower right corner, there is a world map, where the red dot indicates the geographical location of the research area.
Figure 1.
Research area located on the left side of the Lena River near the city of Yakutsk. The map highlights the city’s territory in red and suburban areas in blue. The white arrows on the map denote the lakes that have been explored: (Sa) Saysary, (Yt) Ytyk-Kyuyol, (Ie) Ierelyakh, (Da) Dachnoe, (BC) Bolshaya Chabyda. Yellow arrows and numbers indicate sources of human-made impact outside the city’s and suburban territory: (1) municipal controlled landfill dump, (2) sand quarry. In the lower right corner, there is a world map, where the red dot indicates the geographical location of the research area.
Figure 2.
View of the explored lakes: (a,b), Bolshaya Chabyda Lake and flakes of Microcystis sp. on the water surface; (c), Saysary Lake; (d), Dachnoe Lake; (e), Ierelyakh Lake; (f), Ytyk-Kyuyol Lake. (Photos by V. Filippova).
Figure 2.
View of the explored lakes: (a,b), Bolshaya Chabyda Lake and flakes of Microcystis sp. on the water surface; (c), Saysary Lake; (d), Dachnoe Lake; (e), Ierelyakh Lake; (f), Ytyk-Kyuyol Lake. (Photos by V. Filippova).
Figure 3.
Distribution of algae and cyanobacteria species in phyla in absolute terms (a) and as a percentage in the lake community (b) in the studied lakes of the permafrost zone of Yakutia.
Figure 3.
Distribution of algae and cyanobacteria species in phyla in absolute terms (a) and as a percentage in the lake community (b) in the studied lakes of the permafrost zone of Yakutia.
Figure 4.
Micrographs of the dominant phytoplankton species: (a) Aphanizomenon flos-aquae Ralfs ex Bornet & Flahault; (b) Ceratium hirundinella (O.F.Müller) Dujardin; (c) Dolichospermum lemmermannii (Richter) P.Wacklin, L.Hoffmann & J.Komárek; (d) Microcystis aeruginosa (Kützing) Kützing; (e) Microcystis flos-aquae (Wittrock) Kirchner. Scale bars: (a–c) 50 μm; (d) 20 μm; (e) 100 μm.
Figure 4.
Micrographs of the dominant phytoplankton species: (a) Aphanizomenon flos-aquae Ralfs ex Bornet & Flahault; (b) Ceratium hirundinella (O.F.Müller) Dujardin; (c) Dolichospermum lemmermannii (Richter) P.Wacklin, L.Hoffmann & J.Komárek; (d) Microcystis aeruginosa (Kützing) Kützing; (e) Microcystis flos-aquae (Wittrock) Kirchner. Scale bars: (a–c) 50 μm; (d) 20 μm; (e) 100 μm.
Figure 5.
Indicators’ abundance in studied lakes in the permafrost zone of Yakutia (on the basis of Appendix A, Table A4). Ecological group abbreviations: ((a), Habitat) (P—planktonic, P-B—plankto-benthic, B—benthic); temperature ((b), Temperature) preferences (temp—temperate, eterm—eurythermic, warm—warm-water); salinity ecological groups ((c), Salinity) according to [41] (hb—oligohalobes-halophobes, i—oligohalobes-indifferent, hl—halophiles; mh—mesohalobes); oxygenation and streaming ((d), Oxygen) (st—standing water, st-str—low streaming water, aer—aerophiles).
Figure 5.
Indicators’ abundance in studied lakes in the permafrost zone of Yakutia (on the basis of Appendix A, Table A4). Ecological group abbreviations: ((a), Habitat) (P—planktonic, P-B—plankto-benthic, B—benthic); temperature ((b), Temperature) preferences (temp—temperate, eterm—eurythermic, warm—warm-water); salinity ecological groups ((c), Salinity) according to [41] (hb—oligohalobes-halophobes, i—oligohalobes-indifferent, hl—halophiles; mh—mesohalobes); oxygenation and streaming ((d), Oxygen) (st—standing water, st-str—low streaming water, aer—aerophiles).
Figure 6.
Indicators’ abundance in the studied lakes in the permafrost zone of Yakutia (on the basis of Appendix A Table A4). Abbreviations: pH preference groups ((a), pH) according to [42] (alb—alkalibiontes; alf—alkaliphiles, ind—indifferent; acf—acidophiles); nitrogen uptake metabolism ((b), Autotrophy-Heterotrophy) [43]: ats—nitrogen-autotrophic taxa, tolerating very small concentrations of organically bound nitrogen; ate—nitrogen-autotrophic taxa, tolerating elevated concentrations of organically bound nitrogen; hne—facultative nitrogen-heterotrophic taxa, needing periodically elevated concentrations of organically bound nitrogen; trophic state indicators ((c), Trophic state) [43]: (ot—oligotraphentic; om—oligomesotraphentic; m—mesotraphentic; me—mesoeutraphentic; e—eutraphentic; o-e—oligo to hypereutraphentic); Index S ((d), Class of water quality), species-specific index saprobity according to [44].
Figure 6.
Indicators’ abundance in the studied lakes in the permafrost zone of Yakutia (on the basis of Appendix A Table A4). Abbreviations: pH preference groups ((a), pH) according to [42] (alb—alkalibiontes; alf—alkaliphiles, ind—indifferent; acf—acidophiles); nitrogen uptake metabolism ((b), Autotrophy-Heterotrophy) [43]: ats—nitrogen-autotrophic taxa, tolerating very small concentrations of organically bound nitrogen; ate—nitrogen-autotrophic taxa, tolerating elevated concentrations of organically bound nitrogen; hne—facultative nitrogen-heterotrophic taxa, needing periodically elevated concentrations of organically bound nitrogen; trophic state indicators ((c), Trophic state) [43]: (ot—oligotraphentic; om—oligomesotraphentic; m—mesotraphentic; me—mesoeutraphentic; e—eutraphentic; o-e—oligo to hypereutraphentic); Index S ((d), Class of water quality), species-specific index saprobity according to [44].
Figure 7.
Tree of similarity of chemical variables of the studied lakes in the permafrost zone of Yakutia (on the basis of Appendix A Table A1).
Figure 7.
Tree of similarity of chemical variables of the studied lakes in the permafrost zone of Yakutia (on the basis of Appendix A Table A1).
Figure 8.
Tree of similarity biological variables of studied lakes in permafrost zone of Yakutia (on the basis of Appendix A Table A4).
Figure 8.
Tree of similarity biological variables of studied lakes in permafrost zone of Yakutia (on the basis of Appendix A Table A4).
Figure 9.
JASP network plot of the correlation of species abundance of phytoplankton in five studied lakes in the permafrost zone of Yakutia. Control, unpolluted; Antrop, lakes under anthropogenic impact; City, lake within the city territory of Yakutsk. The thickness of the lines is proportional to the strength of the connection, indicated by the numbers.
Figure 9.
JASP network plot of the correlation of species abundance of phytoplankton in five studied lakes in the permafrost zone of Yakutia. Control, unpolluted; Antrop, lakes under anthropogenic impact; City, lake within the city territory of Yakutsk. The thickness of the lines is proportional to the strength of the connection, indicated by the numbers.
Figure 10.
RDA plot for phytoplankton biomass in phyla and major environmental variables of studied lakes in the permafrost zone of Yakutia.
Figure 10.
RDA plot for phytoplankton biomass in phyla and major environmental variables of studied lakes in the permafrost zone of Yakutia.
Figure 11.
Spatial distribution maps of environmental variables ((a), in altitude; (b), in phosphates; (c), in nitrates; (d), in ammonium) of the studied lakes in the permafrost zone of Yakutia (on the basis of Appendix A Table A1).
Figure 11.
Spatial distribution maps of environmental variables ((a), in altitude; (b), in phosphates; (c), in nitrates; (d), in ammonium) of the studied lakes in the permafrost zone of Yakutia (on the basis of Appendix A Table A1).
Figure 12.
Spatial distribution maps of major biological variables ((a), number of species; (b), phytoplankton abundance; (c), phytoplankton biomass; (d), cyanobacteria biomass; (e), eutraphentic species abundance; (f), index S value) of the studied lakes in permafrost zone of Yakutia (on the basis of Table 2 and Appendix A Table A3).
Figure 12.
Spatial distribution maps of major biological variables ((a), number of species; (b), phytoplankton abundance; (c), phytoplankton biomass; (d), cyanobacteria biomass; (e), eutraphentic species abundance; (f), index S value) of the studied lakes in permafrost zone of Yakutia (on the basis of Table 2 and Appendix A Table A3).
Figure 13.
Maximum likelihood tree based on partial sxtA sequences of 25 cyanobacteria. Sequences from this study are bold. Numbers above the branches are the bootstrap support as a percentage of 1000 replicates for ML and NJ methods. The bar indicates 1% sequence divergence.
Figure 13.
Maximum likelihood tree based on partial sxtA sequences of 25 cyanobacteria. Sequences from this study are bold. Numbers above the branches are the bootstrap support as a percentage of 1000 replicates for ML and NJ methods. The bar indicates 1% sequence divergence.
Table 1.
A concise description of the studied lakes.
| Code | Lake Name | Area of Lake, km2 | Depth, m | Latitude, N | Longitude, E | Altitude, m a.s.l. | Type of Origin |
|---|---|---|---|---|---|---|---|
| Sa | Saysary | 0.40 | 6.5 | 62°01′17.3″ | 129°41′39.6″ | 96 | R |
| Yt | Ytyk-Kyuyol | 0.79 | 3.0 | 62°01′22.02″ | 129°36′59.0″ | 108 | R |
| Ie | Ierelyakh | 0.10 | 1.2 | 62°06′46.20″ | 129°17′37.0″ | 258 | K |
| Da | Dachnoe | 0.23 | 3.5 | 62°07′29.30″ | 129°37′22.9″ | 122 | A |
| BC | Bolshaya Chabyda | 1.68 | 1.6 | 61°58′06.20″ | 129°22′42.5″ | 207 | D |
Note: Classification of lakes by their type of origin: (R) oxbow lakes, (K) thermokarst lakes, (D) dune lake and (A) artificial damming lake.
Table 2.
Average abundance and biomass of phytoplankton phyla in studied lakes in the permafrost zone of Yakutia.
Table 2.
Average abundance and biomass of phytoplankton phyla in studied lakes in the permafrost zone of Yakutia.
| Variable | Sa | Yt | Ie | Da | BC |
|---|---|---|---|---|---|
| Abundance, thou. cells L−1 | |||||
| Charophyta | 1.599 | 7.245 | 0.027 | 0.002 | 0.640 |
| Chlorophyta | 4730.839 | 1888.172 | 193.682 | 660.458 | 2603.003 |
| Cyanobacteria | 4748.504 | 45,345.389 | 6525.961 | 114,042.861 | 104,297.677 |
| Dinoflagellata | 0.585 | 1.410 | 22.708 | 0.041 | 0.041 |
| Euglenophyta | 13.143 | 7.500 | 0.030 | 0.407 | 0 |
| Heterokontophyta | 73.304 | 97.391 | 2.490 | 18.294 | 94.980 |
| Biomass, mg L−1 | |||||
| Charophyta | 0.0136 | 0.0179 | 0.0004 | 0.0001 | 0.0026 |
| Chlorophyta | 0.4276 | 0.1981 | 0.0396 | 0.0417 | 0.1953 |
| Cyanobacteria | 0.6309 | 1.4183 | 0.0020 | 3.8314 | 4.7496 |
| Dinoflagellata | 0.0220 | 0.0531 | 0.8550 | 0.0016 | 0.0015 |
| Euglenophyta | 0.0457 | 0.0423 | 0.0002 | 0.0012 | |
| Heterokontophyta | 0.0601 | 0.0703 | 0.0029 | 0.0242 | 0.1510 |
Note: Abbreviated station code names as in Table 1.
Table 3.
Dominant phytoplankton species of the studied lakes and their share in percentage of the total biomass (emphasized in bold and italicized).
Table 3.
Dominant phytoplankton species of the studied lakes and their share in percentage of the total biomass (emphasized in bold and italicized).
| Species | Sa | Yt | Ie | Da | BC |
|---|---|---|---|---|---|
| Aphanizomenon flos-aquae Ralfs ex Bornet & Flahault | 14 | 2 | - | 29 | - |
| Ceratium hirundinella (O.F.Müller) Dujardin | 2 | 3 | 95 | 0.04 | - |
| Dolichospermum sp. | 20 | - | - | - | - |
| Dolichospermum lemmermannii (Richter) P.Wacklin, L.Hoffmann & J.Komárek | 10 | 0.3 | - | 0.2 | 30 |
| Microcystis aeruginosa (Kützing) Kützing | - | 30 | - | 5 | 30 |
| Microcystis flos-aquae (Wittrock) Kirchner | - | 40 | - | 60 | 30 |
Note: “-”, species not found. Abbreviated station code names as in Table 1.
Table 4.
Sum of averaged variables of phytoplankton in studied lakes in the permafrost zone of Yakutia.
Table 4.
Sum of averaged variables of phytoplankton in studied lakes in the permafrost zone of Yakutia.
| Variable | Sa | Yt | Ie | Da | BC |
|---|---|---|---|---|---|
| Abundance, thou. cells L−1 | 9568 | 47,347 | 6745 | 114,722 | 106,996 |
| Biomass, mg L−1 | 1.2 | 1.8 | 0.9 | 3.9 | 5.1 |
| No of Species | 37 | 54 | 29 | 21 | 26 |
| Shannon H’ | 1.072 | 0.91 | 0.565 | 0.251 | 0.528 |
| Index S | 1.93 | 2.04 | 1.88 | 1.64 | 1.89 |
| Index WESI | 2.00 | 2.50 | 2.00 | 2.00 | 2.00 |
Note: Abbreviated station code names as in Table 1.
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Barinova, S.; Gabyshev, V.A.; Gabysheva, O.I.; Naidanova, Y.A.; Sorokovikova, E.G. Phytoplankton Diversity, Abundance and Toxin Synthesis Potential in the Lakes of Natural and Urban Landscapes in Permafrost Conditions. Land 2025, 14, 721. https://doi.org/10.3390/land14040721
AMA Style
Barinova S, Gabyshev VA, Gabysheva OI, Naidanova YA, Sorokovikova EG. Phytoplankton Diversity, Abundance and Toxin Synthesis Potential in the Lakes of Natural and Urban Landscapes in Permafrost Conditions. Land. 2025; 14(4):721. https://doi.org/10.3390/land14040721
Chicago/Turabian StyleBarinova, Sophia, Viktor A. Gabyshev, Olga I. Gabysheva, Yanzhima A. Naidanova, and Ekaterina G. Sorokovikova. 2025. "Phytoplankton Diversity, Abundance and Toxin Synthesis Potential in the Lakes of Natural and Urban Landscapes in Permafrost Conditions" Land 14, no. 4: 721. https://doi.org/10.3390/land14040721
APA StyleBarinova, S., Gabyshev, V. A., Gabysheva, O. I., Naidanova, Y. A., & Sorokovikova, E. G. (2025). Phytoplankton Diversity, Abundance and Toxin Synthesis Potential in the Lakes of Natural and Urban Landscapes in Permafrost Conditions. Land, 14(4), 721. https://doi.org/10.3390/land14040721
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