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Article

Ecological Approaches in the Process of Formation of the Bolshoe Toko National Park, Yakutia

by
Sophia Barinova
1,*,
Viktor A. Gabyshev
2,
Olga I. Gabysheva
2,
Anna P. Ivanova
2 and
Petro M. Tsarenko
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
M.G. Kholodny Institute of Botany, Tereshchenkovska st. 2, 01601 Kyiv, Ukraine
*
Author to whom correspondence should be addressed.
†
This author has passed away.
Diversity 2025, 17(9), 625; https://doi.org/10.3390/d17090625
Submission received: 4 August 2025 / Revised: 2 September 2025 / Accepted: 4 September 2025 / Published: 5 September 2025
(This article belongs to the Section Biodiversity Conservation)
Browse Figures
Versions Notes

Abstract

The creation of a new protected area, especially on permafrost territory, along with the adoption of legislative measures, requires a thorough assessment of its ecological diversity and condition. In the planned Bolshoe Toko National Park (Yakutia, Northeastern Russia), the main protected area will be a unique deep-water mountain lake of glacial origin, Bolshoe Toko Lake. Our aim was to study the species composition of algal communities of Bolshoe Toko Lake by combining our new and previously known data on the flora of algae and cyanobacteria of the lake. For the first time by analyzing environmental parameters, we identified factors and hotspots of diversity of the lake ecosystem. In the planktonic microflora of the lake, 479 species belonging to six taxonomic phyla were identified. This allows us to talk about a biodiversity hotspot at Bolshoe Toko Lake. The presence of rare, new endangered and critically endangered species in the flora of the lake confirms the need to create a national park. Bioindication analysis and contour maps of ecological factors made it possible to assess the current sustainability of the ecosystem when developing a plan for the creation of a new protected area and to identify potential problem areas and factors affecting the ecosystem. One such factor is the development of the coal basin, which is already having a noticeable impact on the lake environment.

1. Introduction

Preservation and development of the system of specially protected natural areas (SPNA), in connection with its increased role in maintaining the ecological balance and protecting the natural potential of the territory, is one of the declared priorities of the state environmental policy of the Russian Federation. Yakutia, with its still slightly or completely undeformed huge territory, occupies an important place in the global ecosystem. At the same time, the issue of preserving biodiversity is becoming increasingly relevant as the industrial development of the territory of South Yakutia. The development of transport highways, mineral deposits (such as gold, iron ore, and mineral coal) [1], and associated structures will undoubtedly affect the quantitative and qualitative characteristics of the flora and fauna of the region. Of particular concern, both previously and now, is the industrial development of Elga Coal Mine, located in close proximity to the state nature reserve of regional significance “Bolshoe Toko” in the Neryungri District. Biological components of ecosystems are exposed to risks of both direct (removal of soil and vegetation cover in development zones, mechanical destruction of soils by all-terrain vehicles, airborne pollution by gas and dust emissions and oil spills, unregulated shooting of game animals and birds, etc.) and indirect influence (increased fire safety in the area, activation of geomorphological processes, disturbance of animals and birds, alienation of their feeding grounds and disruption of their migration routes, etc.).
The currently existing Bolshoe Toko Nature Reserve was created earlier as a republican zoological reserve based on the Resolution of the Council of Ministers of the Yakut Autonomous Soviet Socialist Republic dated 4 April 1984 No. 129; then, in accordance with the Resolution of the Government of the Republic of Sakha (Yakutia) dated 27 November 1997 No. 515, without changing the borders, it was transferred to the category of “resource reserve” and, according to the Resolution of the Government of the Republic of Sakha (Yakutia) dated 5 December 2014 No. 437, also without changing the borders, it still functions in the category of “state nature reserve”. Thus, the natural ecosystems of the future park have been under a protective regime for almost 40 years.
Among a number of current environmental and compensation measures to minimize the negative impact of industrial development of the Elga Coal Mine on the biological components of ecosystems, the main one is recognized as strengthening the protection regime of this territory, which is a valuable natural standard and gene pool of rare species of animals and plants, by raising the status of the existing protected area in the region.
The experience of the functioning of protected areas in Yakutia has shown that the most accepted category of protected areas by the local population at the moment is the national park, which has a more lenient regime for regulating human activity in the area of traditional nature management compared to reserves.
The creation of the Bolshoe Toko National Park is envisaged by the federal project “Conservation of Biological Diversity and Development of Ecotourism” of the national project “Ecology”, the passport of which was approved by the minutes of the meeting of the project committee for the national project “Ecology” dated 21 December 2018 No. 3. The national project “Ecology” is being implemented in pursuance of the Decree of the President of the Russian Federation dated 7 May 2018 No. 204 “On the national goals and strategic objectives of the development of the Russian Federation for the period up to 2024”. The planned deadline for the creation of a protected area of federal significance according to the national project is 2022. Its area should be at least 263 thousand hectares.
Lake Bolshoe Toko is located within the boundaries of the planned national park and is included in the list of unique lakes of Yakutia. Elga Coal Mine is located 15 km northwest of the lake. There is evidence that Lake Bolshoe Toko is already experiencing negative impacts from coal mining at the present stage [2]. In the future, this may further increase due to the possible start of development of vast licensed mineral deposits in adjacent territories and may cause deterioration of the environmental situation on the lake (mineral deposits, including gold and coal, are located in the upper reaches of the main lateral tributaries of the lake).
Algae, in addition to having a high biodiversity of communities, are also known to be good indicators of surface water quality and are used in state monitoring systems for natural and polluted environmental locations [3,4].
The results of algological studies conducted on Lake Bolshoe Toko prior to this study are limited. The species composition of algae in Lake Bolshoe Toko was partially studied in the 1980s, but the list has not been published [5]. A study of samples from the sediment surface along spatial habitat gradients in Lake Bolshoe Toko revealed that the diatom assemblage is dominated by planktonic species (Pliocaenicus, Cyclotella, and Aulacoseira), as well as non-planktonic taxa, such as Achnanthidium [6]. Diatom species richness and diversity are higher in surface sediments in the northern part of the basin, associated with shallower waters and the availability of benthic and periphytic niches. Another study uses diatom-specific metabarcoding, applying a short rbcL marker combined with next-generation sequencing and morphological identification to analyze the diatom diversity in modern sediment samples of 17 intra-lake sites [7]. The researchers hypothesized that Lake Bolshoe Toko represents a unique biodiversity hotspot in Yakutia. Based on the study of the lake’s bottom sediments, including diatom valves in sediment cores, a reconstruction of the late Quaternary climate was performed. The authors conclude that a lag of deposited organic carbon concentrations behind changes in diatom alpha diversity reveals that species richness can augment the production and thus sequestration of organic matter in comparable lake systems [2].
The results of partial preliminary processing of this algological material were published by us on the flora of Heterokontophyta using the methods of electron scanning and transmission microscopy. A total of 162 species and varieties of pennate [8] and 10 centric diatoms [9] as well as 17 species and varieties of scale chrysophyte algae [10] were identified. The species lists of Heterokontophyta from these publications became part of this study. The Bolshoe Toko lake morphology was not changed by glaciation [11], and such ancient lakes often demonstrate an unusually high degree of biodiversity and endemism [12]. We have previously identified a significant number of rare species of diatoms and silica-scaled chrysophytes for the lake flora, including endemic species [10,13]. Extinction risks due to both global climate change and local anthropogenic impact are highly likely for such specific habitats [14], as well as for endemic [15] and ecologically specialized species [16]. The problem of the disappearance of endemic species [17], an increase in the proportion of cosmopolitan species, and the emergence of newly introduced species [18,19] has long been noted for a number of ancient lakes that were considered to be largely inviolate. In this regard, the relevance of organizing a stricter protection regime for Lake Bolshoe Toko and the issues of its justification are beyond doubt.
The study of the species composition of algae of other phyla of algae and cyanobacteria of Lake Bolshoe Toko was previously conducted in the 1980s, but the list was not isolated from the flora of the region and was not published [5]. The data on the abundance and biomass of algae and cyanobacteria of the plankton of Lake Bolshoe Toko obtained by us and presented in this study are also pioneering and have not been published before. To approve the IUCN conservation status, it is necessary to conduct a study and assessment of diversity on the primary screening and, guided by the developed normative documents, to give recommendations on the organization of monitoring. We followed these steps in our work.
We hypothesize that if biodiversity is high compared to nearby water bodies and the ecological approach shows that the ecosystem is intact, then Lake Bolshoe Toko can be presented as an IUCN protected area.
The aim of this study was to investigate the species composition of the algal and cyanobacterial communities of Lake Bolshoe Toko by combining our new and previously known data on the lake’s diversity, and to perform contour mapping and bioindication analysis for the first time to identify the factors and sources of potential impact on the lake’s ecosystem.

2. Materials and Methods

2.1. Site Description

Lake Bolshoe Toko is of tectonic origin, surrounded by the slopes of the Stanovoy Range up to 1500 m high in the south and by gentle spurs with heights of up to 1000–1100 m in the east and west (Figure 1 and Figure 2b,c). The northern part of the lake basin is a moraine processed by a glacier; many small lakes are located here. The height of the lake above sea level is 903.8 m, the greatest length is 15.4 km, the width is 7.5 km, and the length of the coastline is 51 km [11]. The surface area of the lake is 82.6 km2, and the volume of water is 2.51 km3. The greatest depths are concentrated in the southwestern part, where the maximum depth is 71 m, while the northern part of the lake is shallow (up to 10 m). The average depth is 30.5 m. The lake is a flowing lake: in the southern part, from the spurs of the Stanovoy Range, the Utuk River flows into it, originating at an altitude of 1880 m above sea level (a.s.l.), and at the northeastern end of the lake is the source of the Mulam River. The surface of the lake was divided by us into four quadrants in accordance with the inflowing and outflowing rivers and the heterogeneity of the landscape and the coastline (Figure 1).
There are two bays in the western part of the lake: Rybachy Bay and Nameless Bay. One sampling station for us is located on Okunevoe Lake, a small lake that, in some years, temporarily connects with Bolshoe Toko Lake during the seasonal rise in water level. The ice-free period lasts 146 days. The maximum ice thickness at the end of April reaches 136 cm. Winters are long; the average temperature in January, the coldest month, is −31.4 °C; the absolute lowest temperature is −65 °C; and the average annual temperature reaches −10.2 °C. Summers are short and hot, and the maximum temperature is 34 °C. The total annual precipitation fluctuates from 276 to 579 mm. According to the zoning principles of D.I. Shashko [20], the climate of the Tokinskaya Basin is characterized as sharply continental, cold–temperate, and humid (continentality coefficient 296, humidity coefficient 1.5; sum of active temperatures > 10 °C is 1032°C). The Tokinskaya Basin is characterized by continuous distribution of permafrost. Its thickness varies from 40–50 to 350 m in watershed areas.
Elga Coal Mine, one of the largest coking coal reserves in Asia and the world, is located 15 km northeast of Lake Bolshoe Toko. According to oral information from the ranger of the resource reserve I.N. Butyak, in recent years, as a result of the work of Elga Coal Mine, coal dust has been falling on part of the territory, which is sometimes clearly visible on the surface of the snow cover. The satellite images at our disposal help to obtain an idea of the distribution of coal dust in the study area (Figure 2). Figure 2a shows a satellite image of the territory taken in winter (22 March 2022, source: Google Earth computer program Version 10.88.0.3 Multi-threaded using “view a map over time” function). It is evident that the territory of the Elginskoye coal deposit is contaminated with coal dust, which is possibly dispersed in the direction of the prevailing winds (Figure 2d).

2.2. Sampling

Sampling was carried out from Bolshoe Toko Lake and the neighboring small Okunevoe Lake using an Apshteyn net (SEFAR NITEX filter fabric, 15 μm mesh size) (Sefar Group JSC, Thal, Switzerland) from 7 to 22 July 2015 (Figure 1) from the surface water horizon (0–0.3 m). A total of 110 algological plankton samples were collected (Appendix A, Table A1), half of which were for qualitative composition, while the other half were quantitative samples with a volume of 30 L. All algological samples were fixed by adding formalin. Samples for hydrochemical analysis in the amount of 6 were collected by scooping up to 2 L plastic containers of water. Water temperature during sampling was recorded using a Checktemp electronic thermometer (HANNA Instruments, Woonsocket, RI, USA). In the pelagic zone of Bolshoe Toko Lake, water transparency was measured using a Secchi disk (Appendix A, Table A1).

2.3. Phytoplankton Analysis and Examining the Chemical Composition of Water

An Olympus BH-2 light microscope (Olympus, Tokyo, Japan) was used to study the algological samples. Species identification was performed using identification guides [21,22,23,24,25,26,27]. The taxonomic affiliation of species was clarified using data from the Algabase.org portal [28]. The number of phytoplankton cells was counted using a 0.01 mL Nageotte chamber in triplicate. Phytoplankton biomass was calculated using the volumetric counting method [29].
Chemical analysis of water samples was performed using generally accepted methods [30]. The water pH was measured using a potentiometric method with a Multitest IPL-101 pH meter/ion meter (LLC NPP “SEMIKO”, Novosibirsk, Russia). A titration method with iodometric determination was used to measure oxygen concentration and biological oxygen demand (BOD5). Water salinity (TDS) was calculated as the sum of ions using the following methods: turbidimetry for sulfate anions; flame spectrophotometry for potassium and sodium cations using an AAnalyst 400 atomic-absorption spectrometer (PerkinElmer Inc., Waltham, MA, USA); mercurimetric titration for chloride ions; and titration for calcium, magnesium, and bicarbonate ions. The hardness of water was determined by complexometric titrations using eriochrome black T as an indicator. A photometric method with a PE-5300VI spectrophotometer (GK “EKROS”, Saint Petersburg, Russia) was applied to determine water color, silicon content, chemical oxygen demand (COD), and nutrient concentrations. Nessler’s reagent, Griess reagent, salicylic acid, ammonium molybdate, and sulfosalicylic acid were used for the measurement of ammonium ion, nitrite ion, nitrate ion, phosphate ions, and total iron, respectively. A combined reagent composed of ammonium molybdate and ascorbic acid was used to determine total phosphorus content. The fluorimetric method with a Fluorat-02-2M device (LLC “Lu-mex-Marketing”, Saint Petersburg, Russia) was used to establish phenols concentration.

2.4. Statistical Analysys

The index diversity of Shannon–Wiener [31] of phytoplankton community was calculated using the Biodiversity Pro 2.0 program [32]. Calculation of similarity of the network graphs was performed as the network analyses in JASP (Jeffreys’s Amazing Statistics Program 0.16.4.0) on the bootnet package in ‘R’ [33]. Pearson correlation coefficients were calculated in the Wessa.net program v1.0.13 [34]. Saprobity index of algae community was calculated on the basis of species-specific saprobity index s and cell abundance of each indicator species [35,36]. Bioindicator analysis was performed according to [35] with species-specific ecological preferences of revealed indicator taxa [35].
The aquatic ecosystem state index WESI [36] was calculated to assess the toxic pollution influence on the aquatic ecosystems by the following equation:
WESI = Rank Index S/Rank N-NO3
where WESI is an aquatic ecosystem state index, Rank Index S is the rank number from 1 to 9 calculated for each community index S in the water quality class of [36], and Rank N-NO3 is the rank number from 1 to 9 of defined N-NO3 concentrations for each sampling point in the water quality class of [36]. 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.
Contour maps were created in the Statistica 12.0 program based on the GPS coordinates of sampling points for each biological and chemical variable. Redundancy discriminant analysis (RDA) was performed with the CANOCO 4.5 program for calculation of biological dominated variables as dependent and environment variables as independent relationships [37]. Rare and endangered species are marked based on data on the distribution of well-studied diatoms in Central Europe [38] in comparison with IUCN (International Union for Conservation of Nature) criteria [39].

3. Results

3.1. Water Physical and Chemical Data

The waters of the studied lakes have no taste or smell and have a high color (30°–40°) (Supplement Table S1). The water temperature varies at the observation stations from 15.1 to 24.5 °C, with the northern, shallow part of the lake warmed up more than the southern part, where the maximum depths are noted (Figure 3b). The hydrogen index indicates a shift in the water reaction to the slightly acidic side (6.27–6.43). The BOD index does not exceed 1.64 mgO L−1 and is characterized by low values. The oxygen regime is favorable—the concentration is 8.90–9.79 mgO L−1 at 98–104% saturation. Water transparency according to the Secchi disk at individual stations varied from 3 to 5 m, and water temperature during sampling at observation points varied from 13.5 to 30.4 °C (Appendix A, Table A1). Oxygen concentration was 8.3–9.16 mg L−1 (Supplement Table S1).
The level of total dissolved solids (TDS) is very low (22–41 mg L−1), which characterizes the water as “low mineralized”. The highest value of water mineralization is noted in Rybachy Bay of Bolshoe Toko Lake (Figure 3a), which can be caused by the influence of groundwater or the release of underground springs there. According to the value of total hardness, the water is “very soft”, and according to the ratio of its main ions, it belongs to the hydrocarbonate class, calcium group, and corresponds to type III.
The contents of phenols, anionic surfactants, and oil products are characterized by insignificant values. Most biogenic elements are characterized by low contents. The concentration of silicon did not exceed 2.34 mg L−1, ammonium ion–0.52 mg L−1, total phosphorus–0.016 mg L−1, phosphate ion–0.008 mg L−1, and nitrite ion–0.009 mg L−1. In the northwestern part of Bolshoe Toko Lake, the maximum values of nitrite ion content were noted (Figure 3c), which may be caused by the influence of Elga Coal Mine. The highest content of phosphate ions was found in the southern and southeastern parts of Bolshoe Toko Lake (Figure 3d). Relatively high concentrations were noted for three components—oxidation-resistant organic matter (by COD value), total iron, and color indices. The concentration of COD is 24–36 mg L−1, total iron–0.51–0.70 mg L−1, and color–30°–40°.
Figure 3 shows the distribution of the main environmental parameters across the lake. It is evident that the increased concentration of mineral salts is confined to the southeastern part, quadrant SE (Figure 3a). The water temperature is higher in the shallow northern part of the lake (Figure 3b). The nitrate influx was predominantly from the northwestern shore (Figure 3c), and phosphates had increased concentrations in the southern and southeastern parts of the lake (Figure 3d).

3.2. Phytoplankton

3.2.1. Species Richness in Community

A total of 479 species and intraspecies of algae and cyanobacteria were identified in the planktonic surface samples collected during the study period (Table 1, Tables S2, and S3). Heterokontophyta strongly predominates among the six phyla. This phylum currently unites several classes that previously constituted a taxonomic phylum. The Class column in Table 1 shows that diatoms also predominate, but the role of chrysophytes in the plankton is also great, as are green, zygnematalean algae and cyanobacteria and indicating a natural community characteristic of cold waters. At the same time, a large number of diatom species opens opportunities both for broad bioindication analysis and for identifying threatened and rare species when assessing the current state of the flora before the end of the conservation process.
Contour mapping of the distribution of the main diversity indices is presented in Figure 4. Overall species richness was highest in the northeastern part of the lake, NE (Figure 4a), and this coincides with the distribution of diatoms (Figure 4b). Cyanobacteria were richest in the SE and NE parts, where the shore was enriched by the lakes and bays (Figure 4c). Dinoflagellate species were rich in the NW communities, where a few small, permanent streams come into the lake.
The diversity indices calculated by us for the surface area of Lake Bolshoe Toko (Sp./Area) were 5.8 species and intraspecies per square kilometer, and the index of intraspecific variability of the identified diversity (Ssp./Sp.) was 1.17.

3.2.2. Abundance and Biomass

The abundance and biomass data of the plankton surface samples are presented in Supplement Tables S4 and S5. Statistical comparison of abundance indices revealed high heterogeneity of the data (Figure 5), making it difficult to estimate the abundance distribution over the lake surface. The same statistical JASP was constructed for phytoplankton biomass.
However, contour mapping allows us to identify areas of the water area that are most favorable for the development of plankton as a whole, as well as each of the phyla (Figure 6). The distribution map of the total abundance (Figure 6a) coincides with the distribution of cyanobacteria (Figure 6b), and the maximum development is confined to the northeastern part of the lake with a low indented shore and Lake Okunevoe, which is connected to Lake Bolshoe Toko. Dinoflagellates are more abundant at stations closer to the northwestern shore (Figure 6c). The distribution of biomass over the surface of the lake is very similar to the abundance maps (Figure 6d–f).

3.2.3. Bioindicators

The bioindicator analysis was based on the species-specific ecology of the detected phytoplankton species (Supplementary Table S2). It is evident that 440 out of 479 taxa were indicator taxa, which is more than 80% of the species composition, and most of them belong to diatoms, where indicator systems are most developed. The abundance of indicator species of each ecological group was estimated in terms of their abundance (Supplement Table S6) in communities at each station of the lake surface. The most striking indicators were mapped. Figure 7a shows the distribution of eutrophic species over the lake surface, where it is evident that eutrophication is expressed in the northwestern and northeastern parts, where the water is shallow and the shore is indented. Pollution indicators related to the fifth water quality class were present in the northwestern and western parts of the lake (Figure 7b). The saprobity indices S calculated based on the abundance and species-specific properties were within the range of water quality classes 2–3; the distribution of their values is shown in Figure 7c. The map shows that their highest value, showing the places of organic pollution influx, are located in the southern water area, where the Utuk River flows in and the coastline is heavily indented, causing water stagnation. The calculated WESI is based on the nitrate nitrogen concentration data and the calculated saprobity indices S at each station where these data are available, and a contour map of the distribution of its values was constructed (Figure 7d). It is evident that the highest values are at stations near the shallow Olenek Lake, which most of the time merges with the main water body of Lake Toko. That is, self-purification processes of water are active here. In other parts of the lake, self-purification activity decreases but has never been less than 1, which indicates natural processes and the absence of a toxic impact on Lake Toko.
Statistical calculation of the similarity of the composition of bioindicators at the stations of the surface of Lake Toko showed that with all the diversity, the most similar indicator properties are the northeastern and southeastern parts, united in cluster 1 in Figure 8. The parts of the lake in the northwest and southwest have different species composition of indicator species of phytoplankton from cluster 1 (cluster 2). This allows dividing the water area of the lake into two quite different parts, western and eastern, in accordance with the direction of water flow in the water body of the lake, since the Utuk River flows in from the south and the Mulam River flows out in the north of the lake basin.

3.3. Species–Environment Relationships

The relationships between the number of species in phyla in the phytoplankton communities of Lake Bolshoe Toko and environmental parameters were analyzed using RDA analysis. The calculations were based on the data in Supplement Tables S1, S3 and S7 and are shown in Table S8. Figure 9 shows that elevated salt concentrations and water temperatures stimulated the development of green algae, dinoflagellates, and cyanobacteria diversity while leading to nitrate depletion and increased Heterokontophyta development. The presence of phosphates and phenols is associated with the saprobic index S, but these parameters had elevated values where there was a lower diversity of diatoms. Interestingly, the WESI was higher where non-diatom diversity was more pronounced, indicating that self-purification processes were more active where high-diversity communities developed.

3.4. Threat Categories of IUCN for Diatom Algae

Among the 479 identified species of algae and cyanobacteria in the plankton of Lake Bolshoe Toko, a new representative of the relict diatom flora was found and described for the first time for science—Pliocaenicus bolshetokoensis Genkal, Gabyshev & Kulikovskiy; in addition, a rare species was found—Discostella guslyakovyi Genkal, Bondarenko & Popovskaya. Six species of diatoms and two species of scaly chrysophytes from Lake Bolshoe Toko were noted for the flora of Russia for the first time: Symbolura cf. lura Miho et Krammer, Fragilaria perminuta (Grunow) Lange-Bertalot, Gomphonema parallelistriatum Lange-Bertalot & E.Reichardt, Pinnularia cf. neomaior var. inflata Krammer, Pinnularia cf. stidolphii Krammer, Planothidium cf. distinctum (Messikommer) Lange-Bertalot, Mallomonas inornata K.H. Nicholls, and Synura synuroidea (Prowse) Pusztai, Certnerová, Skaloudová & Skaloud. These species belong to diatoms and golden algae of the phylum Heterokontophyta, for which there is information on threatened and Red Book species. Table 2 shows the distribution of the identified diatom species (297) out of 479 species of algae and cyanobacteria in the phytoplankton of Lake Bolshoe Toko (Supplement Table S2). As can be seen from the table, the known information on threatened diatom species is limited to 146 species. Most of them belong to the LEAST CONCERN category (66). But it is important that the numbers of ENDANGERED (32) and CRITICALLY ENDANGERED (10) species are high, which together with VULNERABLE (2) and NEAR THREATENED (14) species make up a total of 58 diatom species, which is 39.7% of the identified threatened species. However, it should be considered that the 151 species in the Bolshoe Toko list that remained unassessed by IUCN categories should also be included in Table 2 as NOT EVALUATED. These 151 “not assessed” species together with 15 known species that are “not evaluated” category, altogether form a more representative number of not evaluated species in the studied lake diversity (Figure 10). Thus, the high percentage of threatened species in the planned reserve may be lower, but this category at the same time requires attention and should be monitored during monitoring of the protected area.

4. Discussion

The species composition of the lake’s planktonic algal communities is highly diverse and includes 479 species from six taxonomic phyla. The data we obtained on the planktonic flora of the surface horizon of Lake Bolshoe Toko indicate the uniqueness of the lake’s aquatic biota. Hotspot is a term used broadly to define a high burst of diversity in a particular area. Clear criteria have been defined for identifying the spatial boundaries of hotspots [40]. Defining hotspot areas is necessary to prioritize resources put towards global conservation. However, when defining hotspot areas to address local conservation issues, downscaling approaches may help to focus conservation efforts [41,42,43]. Therefore, a burst of diversity of Bolshoe Toko—a lake in a permafrost zone—can also be described as a hotspot of phytoplankton diversity. For example, for 70 mountain lakes of this region, including the Baikal region, Transbaikalia and southern Yakutia, located at an altitude of 330 to 1900 m a.s.l. in the Khamar–Daban, Eastern Sayan, Kodar, Udokan, Kalarsky, and Baikalsky ranges, the largest of which is Lake Oron, only 196 species and intraspecific taxa were identified in the planktonic algal flora [44]. Findings of a number of diatom species indicate that Lake Bolshoe Toko, having suffered little from the glacier, served as a refugium for them. These are, for example, representatives of the genus Pliocaenicus, which were widespread in Pliocene water bodies (about 5 million years ago) and found refuge in the Northern Hemisphere in some mountainous areas [45,46], including Lake Bolshoe Toko. All these facts indicate the validity of the decision to create a national park in this territory.
Coal dust contains a number of chemical components, which may include heavy metals that form toxic compounds: Pb, Cd, Zn, Cu, and Hg [47]. Heavy metal compounds can accumulate in ecosystems over time and have a long-term negative effect on terrestrial and aquatic biota. To some extent, the presence of this trend is confirmed by data on the assessment of anthropogenic impact obtained from the analysis of cores from the bottom sediments of Lake Bolshoe Toko. The analysis of these materials shows, in particular, an increase in the concentration of mercury in the bottom sediments of the lake by 1.6 times since the mid-19th century, which researchers associate with the influence of atmospheric precipitation [2]. In general, the values of the studied variables of the chemical composition of Lake Bolshoe Toko vary within the range of values characteristic of background stage of taiga–permafrost landscapes of the humid zone [48]. At the present stage, the lake is predominantly under the influence of natural factors and is not transformed.
Lake Khamra, the closest lake to Lake Bolshoe Toko studied for diatoms, is located ca. 30 km northwest of the Lena River in southwest Yakutia (Republic of Sakha, Lensky District), at an elevation of 340 m a.s.l. Its mirror area is 4.6 km2 and its maximum water depth is 22.3 m, which is comparable with the parameters of Lake Bolshoe Toko [49]. The diatom flora of Lake Khamra is represented by 280 species [50], which is also comparable with the diatom flora of the potential reserve studied by us. However, despite the similarity of location, landscape, and climate, the joint flora had only a small number of species in common, only 48, with a total list of 334. This demonstrates the high individuality of the studied flora of both Lake Bolshoe Toko and Lake Khamra, which are similar in parameters, and as a result is also the basis for preserving Lake Bolshoe Toko. There are no other closely located lakes with similar parameters for which the flora of algae and cyanobacteria is known in the region, which especially increases the value of the studied species composition and the intention to preserve it in the protected area.
The highest number of species in the surface-layer communities of Lake Toko was observed in its northwestern part, mainly due to the high diversity of diatoms. This can be associated with the high overgrowth of this shallow area with aquatic plants, which serve as an excellent substrate for the development of not only planktonic but also attached forms. Since the eutrophication of lakes, especially cold-water ones, begins with the overgrowth of shallow areas with aquatic vegetation [51], in the future, it is the NE part of the lake that should be monitored as the highest priority.
In the southern and southeastern parts of Lake Toko (SE), where the Utuk River flows into the lake and where the coastline is heavily indented, filamentous cyanobacteria developed in large numbers, primarily from the genera Dolichospermum (Dolichospermum flos-aquae (Bornet & Flahault) P. Wacklin, L. Hoffmann & Komárek; Dolichospermum lemmermannii (Richter) P. Wacklin, L. Hoffmann & J. Komárek); and Limnothrix planctonica (Wołoszyńska) Meffert, indicating an increased influx of nutrients that are quickly absorbed by phytoplankton. The central part of the lake is inhabited by a rather poor complex of species, dominated by golden flagellates Mallomonas papillosa K. Harris & D.E. Bradley and green algae Botryococcus braunii Kützing. Particular attention is drawn to the significant species richness of dinoflagellates in the communities of the NW shore of Lake Toko. Peridinium inconspicuum var. contactum Er. Lindemann was most represented here. Such a distribution, revealed in the example of contour maps, turned out to be typical for fairly large lakes, such as many northern [51,52] and southern lakes Kinneret and Sevan [53,54].
Dinophytes are usually associated with some adverse impact, which can be assumed for Lake Toko. It is in those places where they predominated in phytoplankton communities that there is a runoff of small and drying streams directed from the mine, the impact of which is clearly visible in Figure 1. Dinophytes are flagellates that obtain nutrition by photosynthesis, and they are also facultative heterotrophs, capable of surviving under conditions of toxic suppression of photosynthesis activity. Thus, their development in coastal NW complexes can be associated with the coal plant and the spread of its dust in the southeast direction. This is also confirmed by the reduced values of the WESI near the northwest shore. In general, the importance of wind-induced atmospheric transfer has already been considered by us using the example of the impact of salty sea winds in the Kostyanoy Nos Nature Reserve [55], as well as the low-temperature impact in the Arctic on water bodies of the Ust-Lensky Nature Reserve and Kotelny Island [56,57]. Using environmental mapping, visually unobservable impacts of oil and gas production facilities, as well as diamond mining in permafrost conditions, were identified [58,59].
The number of taxa per unit surface area of Lake Bolshoe Toko (Sp./Area) was 5.8 species but the number of intraspecific taxa per square kilometer was greater than previously calculated for protected areas in the permafrost zone of Yakutia (0.05–2.14 taxa km−2) [56]. those calculated earlier Such a high index for the created reserve, even at the preliminary stage of studying the diversity of algae and cyanobacteria, confirms the hypothesis that its flora is a biodiversity hotspot.
The index of intraspecific variability of the identified diversity in the Bolshoe Toko Lake (Ssp./Sp.) was 1.17, which is also higher than for the reserves of Yakutia [56] and in the latitudinal aspect is located between the index for the algal flora of the British Isles (1.15 Ssp./Sp.) and Georgia (1.19 Ssp./Sp.) [60]. This confirms the dependence of intraspecific variability on the latitude of the water body and provides important direction for monitoring the diversity of the created reserve.
According to our data, the geographical factors, as well as the influence of such climatic factors as northeast winds bringing coal dust from coal mining areas [56], were significant for the formation of algal communities in permafrost lakes. Comparative floristry of diatoms in Arctic lakes from Spitsbergen to Chukotka statistically reliably established the high individuality of the species composition in lakes of the permafrost zone [56]. The methods of bioindication and comparative analysis that we used also made it possible to emphasize the high sensitivity of ecosystems in these lakes.
High diversity appears to be a property of permafrost water body ecosystems inherent in ecotones, as was previously found for scaly chrysophytes [61]. Since a large number of species were recorded in Lake Bolshoe Toko, the future reserve can be considered not only an ecotone but also a diversity hotspot. Ecotones are boundary areas of different landscapes where a noticeable increase in organism diversity is observed [62], a phenomenon best studied for plant communities. Until now, the term “ecotone” was used for algal communities very rarely, such as for chrysophyte algae from riverine habitats [63] or for diatom sanctuaries [56]. Based on this, a hypothesis was formed about the introduction of species from temperate latitudes into Arctic waters under the conditions of global climate change [64]. That is, an atypically high level of species diversity of these hydrobionts may reflect early signs of climate warming. Thus, we have identified an important role of climatic, morphometric, and other environmental variables associated with the geographical location of the reservoir in the formation of the composition of algae and cyanobacteria communities in permafrost zone reservoirs. The results of the study are important for developing the basis for monitoring the biodiversity of non-impact, ecologically vulnerable areas in the context of global climate change.
While research in the Arctic region is extremely difficult, there is some information about the diversity of diatoms in lakes of the permafrost zone. A comparative analysis of the diatom species composition in Lake Bolshoe Toko (277) and other Arctic lakes showed that the known flora in the lakes of the Bolshezemelskaya tundra [65] is closest to our data. Thus, the number of species varied in these lakes from 146 in Lake Vanutkiny to 470 in Lake Korotaikha. These lakes are located in the territory included in the IUCN list. Other lakes in the territory of IUCN sites in Arctic Chukotka were also rich in species, from 166 in Lake Ekitiki to 288 in Lake Ervynaygytgyn [66,67]. In eastern Yakutia, lake floras had significantly lower diversity, from 33 to 137 species [56], and in nine studied lakes in western Yakutia they did not exceed 67 species, as in Lake Aalaah [68]. Diatoms in the northernmost known lakes of Svalbard Island [69] comprised from 42 to 79 species. These results place the algal flora of Lake Bolshoe Toko in the ranks of objects with extraordinary diversity, and meaning it can be classified as a biodiversity hotspot in the permafrost zone.
Among the rare or endangered species identified for Lake Bolshoe Toko, the most prevalent are those that prefer moderate temperatures and slightly acidic or neutral environments free of organic pollutants. Therefore, aquatic communities in clean, freshwater lakes with a neutral reaction of the environment, located in the permafrost zone and characterized by the highest level of biodiversity, are the most vulnerable, which must be monitored when monitoring the newly created IUCN protected natural area. The proportions of diatom diversity and rare species diversity were found to be correlated [70].

5. Conclusions

The process of creating a new protected area, especially in the permafrost area, includes a screening assessment of the diversity and ecology of the potential protected area as its background state. As a result of the assessment of the background diversity for Lake Bolshoe Toko, 479 phytoplankton species belonging to six taxonomic phyla were identified. This is a very high diversity in comparison with the available data on lakes in Yakutia, and it can be considered as a diversity hotspot. Among them, rare, new ENDANGERED and CRITICALLY ENDANGERED species were found, which confirms the need to create a reserve. The use of an integrated approach to assessing the diversity of plankton communities and their response to the impact of the environment of Lake Bolshoe Toko using bioindication and environmental mapping methods allowed not only to assess the current state of its ecosystem for planning a new protected area, but also to suggest areas of diversity hotspot and identify influencing factors, one of which is the already observed impact of coal mining.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/d17090625/s1, Supplement Table S1: Physical and chemical variables Lake Bolshoe Toko water, based on samples collected in 2015. Station numbers as in Appendix A, Table A1; Supplement Table S2: Distribution of phytoplankton species in Lake Bolshoe Toko with ecological preferences and Red List [29] and IUCN [30] categories. Abbreviations of threat categories as in Table 2. “1”, present; “0”, absent. Station numbers as in Appendix A, Table A1; Supplement Table S3: Number of species, average abundance (cells L−1), and average biomass (cells L−1) per phylum of phytoplankton in Lake Bolshoe Toko, based on samples collected in 2015. Station numbers as in Appendix A, Table A1; Supplement Table S4: Average abundance (cells L−1) of phytoplankton species from Lake Bolshoe Toko, based on samples collected in 2015. Station numbers as in Appendix A, Table A1; Supplement Table S5: Average biomass (mg L−1) of phytoplankton species from Lake Bolshoe Toko, based on samples collected in 2015. Station numbers as in Appendix A, Table A1; Supplement Table S6: Average abundance (cells L−1) in ecological groups of indicators of phytoplankton in Lake Bolshoe Toko, based on samples collected in 2015. Station numbers as in Appendix A, Table A1; Supplement Table S7: Number of species in ecological groups of indicators of phytoplankton in Lake Bolshoe Toko, based on samples collected in 2015. Station numbers as in Appendix A, Table A1; Supplement Table S8: Correlation matrix for RDA of biological (dependent) and environmental (independent) variables for Bolshoe Toko Lake.

Author Contributions

Conceptualization, S.B. and V.A.G.; methodology, A.P.I. and P.M.T.; software, S.B. and A.P.I.; validation, S.B. and V.A.G.; formal analysis, A.P.I., O.I.G. and P.M.T.; investigation, P.M.T. and O.I.G.; resources, V.A.G.; data curation, S.B.; writing—original draft preparation, V.A.G., S.B. and A.P.I.; writing—review and editing, A.P.I., S.B. and V.A.G.; visualization, S.B., A.P.I. and O.I.G.; supervision, V.A.G.; project administration, V.A.G.; funding acquisition, V.A.G. and A.P.I. 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).

Institutional Review Board Statement

Not applicable.

Data Availability Statement

Data are available upon request from the authors.

Acknowledgments

We are grateful to the Israeli Ministry of Aliyah and Integration for partial support of this work. We are grateful to Ivan Nikolaevich Butyak the ranger of the Bolshoe Toko Resource Reserve for help and full cooperation in carrying out field work on the Bolshoe Toko Lake.

Conflicts of Interest

The authors declare no conflicts of interest.

Abbreviations

The following abbreviations are used in this manuscript:
JASPJeffreys’s Amazing Statistics Program
WESIWater Ecosystem State Index
RDARedundancy discriminant analysis
TDSTotal dissolved solids
CODChemical oxygen demand
BODBiological oxygen demand

Appendix A

Table A1. Information on the sample type and sampling locations at Lake Bolshoe Toko in 2015.
Table A1. Information on the sample type and sampling locations at Lake Bolshoe Toko in 2015.
Station No.WaterbodyDescription of
Sample Station		Sample DateWater Temperature, °CSecchi Depth, mLatitudeLongitude
1Bolshoe Toko LakePelagic zone20 July 201517.504.3056.04556130.8271
6Bolshoe Toko LakePelagic zone19 July 201518.104.5056.01276130.8489
8Bolshoe Toko LakePelagic zone19 July 201519.304.5056.03873130.9061
9Bolshoe Toko LakePelagic zone19 July 201519.304.5056.0441130.8808
11Bolshoe Toko LakePelagic zone19 July 201520.304.5056.06375130.9247
12Bolshoe Toko LakePelagic zone19 July 201522.305.0056.06867130.8973
13Bolshoe Toko LakePelagic zone19 July 201521.304.5056.1011130.9673
14Bolshoe Toko LakePelagic zone19 July 201517.604.5056.0809130.8832
15Bolshoe Toko LakeLittoral7 July 201519.90 56.07509130.8342
16Bolshoe Toko LakeLittoral7 July 201518.00 56.07445130.8346
17Bolshoe Toko LakeLittoral8 July 201518.10 56.08563130.8615
19Bolshoe Toko LakeLittoral8 July 201520.20 56.10141130.9106
20Bolshoe Toko LakeLittoral8 July 201520.60 56.10169130.9082
21Bolshoe Toko LakeLittoral8 July 201518.10 56.07989130.8461
22Bolshoe Toko LakeLittoral8 July 201516.70 56.07896130.847
23Bolshoe Toko LakeLittoral8 July 201513.50 56.0547130.8205
24Bolshoe Toko LakeLittoral8 July 201514.80 56.05364130.8196
25Bolshoe Toko LakeLittoral8 July 201517.50 56.0371130.8049
26Bolshoe Toko LakeLittoral8 July 201516.00 56.03712130.8067
27Bolshoe Toko LakeLittoral11 July 201517.70 56.01585130.8018
28Bolshoe Toko LakeLittoral11 July 201515.70 56.01406130.8176
29Bolshoe Toko LakeLittoral11 July 201516.70 56.00842130.8325
31Bolshoe Toko LakeLittoral11 July 201515.90 55.99805130.8456
32Bolshoe Toko LakeLittoral12 July 201515.10 56.01826130.8144
34Bolshoe Toko LakePelagic zone13 July 201518.105.0055.99742130.8742
35Bolshoe Toko LakeLittoral13 July 201519.20 56.00509130.9023
36Bolshoe Toko LakeLittoral13 July 201520.30 56.00506130.9011
37Bolshoe Toko LakeLittoral13 July 201520.70 55.99395130.883
38Bolshoe Toko LakeLittoral13 July 201521.20 55.9927130.8827
39Bolshoe Toko LakeLittoral13 July 201520.40 55.98794130.8562
40Bolshoe Toko LakeLittoral13 July 201521.30 55.98702130.8552
41Bolshoe Toko LakeLittoral13 July 201521.80 55.99039130.8411
43Bolshoe Toko LakeLittoral14 July 201517.10 56.02217130.8995
44Bolshoe Toko LakeLittoral14 July 201518.40 56.0245130.9022
46Rybachy Bay of Bolshoe Toko LakeLittoral14 July 201520.30 56.032130.9191
47Rybachy Bay of Bolshoe Toko LakePelagic zone14 July 201521.303.0056.02686130.9218
48Rybachy Bay of Bolshoe Toko LakeLittoral14 July 201524.20 56.02507130.9246
49Rybachy Bay of Bolshoe Toko LakeLittoral14 July 201520.50 56.02741130.9135
50Rybachy Bay of Bolshoe Toko LakePelagic zone14 July 201522.203.0056.0208130.9128
51Rybachy Bay of Bolshoe Toko LakeLittoral14 July 201524.60 56.01935130.9162
52Rybachy Bay of Bolshoe Toko LakeLittoral14 July 201522.10 56.02212130.9093
53Rybachy Bay of Bolshoe Toko LakeLittoral14 July 201523.50 56.01673130.9071
54Nameless Bay of Bolshoe Toko LakeLittoral15 July 201523.10 56.05244130.9485
55Nameless Bay of Bolshoe Toko LakeLittoral15 July 201523.40 56.0512130.9454
56Bolshoe Toko LakeLittoral15 July 201519.60 56.05351130.9395
57Bolshoe Toko LakeLittoral15 July 201523.20 56.06905130.9623
58Bolshoe Toko LakeLittoral15 July 201524.90 56.06803130.964
59Bolshoe Toko LakeLittoral15 July 201522.70 56.081130.9733
60Bolshoe Toko LakeLittoral15 July 201524.00 56.081130.9751
61Bolshoe Toko LakeLittoral15 July 201522.20 56.09388130.9761
62Bolshoe Toko LakeLittoral15 July 201522.10 56.09436130.9791
63Okunevoe LakeLittoral16 July 201524.50 56.05982130.9475
64Bolshoe Toko LakeLittoral18 July 201523.60 56.10671130.9372
65Bolshoe Toko LakeLittoral18 July 201530.40 56.10869130.9381
67Bolshoe Toko LakeLittoral22 July 201519.70 56.10508130.9759

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Figure 1. Explored area and Bolshoe Toko Lake with sample stations divided into four parts: southwest (SW), northwest (NW), northeast (NE). and southeast (SE) (a). Map of sample stations (purple symbols, numbers of sample stations according to Appendix A, Table A1) with geographical grid (b). World map with red star denoting explored area location (c).
Figure 1. Explored area and Bolshoe Toko Lake with sample stations divided into four parts: southwest (SW), northwest (NW), northeast (NE). and southeast (SE) (a). Map of sample stations (purple symbols, numbers of sample stations according to Appendix A, Table A1) with geographical grid (b). World map with red star denoting explored area location (c).
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Figure 2. View of the Bolshoe Toko Lake: (a) Satellite image illustrating the spread of coal dust from the Elga Coal Mine (circled in red) in winter; (b) the southern part of Lake Bolshoe Toko against the backdrop of the slopes of the Stanovoy Range; (c) view from the shore of the shallow northeastern part of Lake Bolshoe Toko; (d) map of the direction and strength of prevailing winds in the study area.
Figure 2. View of the Bolshoe Toko Lake: (a) Satellite image illustrating the spread of coal dust from the Elga Coal Mine (circled in red) in winter; (b) the southern part of Lake Bolshoe Toko against the backdrop of the slopes of the Stanovoy Range; (c) view from the shore of the shallow northeastern part of Lake Bolshoe Toko; (d) map of the direction and strength of prevailing winds in the study area.
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Figure 3. Contour maps of distribution of the main environmental variables ((a) in TDS, mg L−1; (b) in water temperature, grad.; (c) in nitrate nitrogen, mg L−1; (d) in phosphate, mg L−1) at the Bolshoe Toko Lake according to Supplement Table S1. Black circles are the sampling stations.
Figure 3. Contour maps of distribution of the main environmental variables ((a) in TDS, mg L−1; (b) in water temperature, grad.; (c) in nitrate nitrogen, mg L−1; (d) in phosphate, mg L−1) at the Bolshoe Toko Lake according to Supplement Table S1. Black circles are the sampling stations.
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Figure 4. Contour maps of distribution of the main species richness data (a) in total number of species; (b) in number of Heterokontophyta species; (c) in number of Cyanobacteria species; and (d) in number of Dinoflagellata species at the Bolshoe Toko stations according to Supplement Table S3. Black circles are the sampling stations.
Figure 4. Contour maps of distribution of the main species richness data (a) in total number of species; (b) in number of Heterokontophyta species; (c) in number of Cyanobacteria species; and (d) in number of Dinoflagellata species at the Bolshoe Toko stations according to Supplement Table S3. Black circles are the sampling stations.
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Figure 5. JASP plot for the phytoplankton abundance similarity on the Bolshoe Toko stations of the lake surface according to Figure 1 and Supplement Table S4. The thickness of the lines is proportional to the strength of the connection. Blue lines—positive correlation; red lines—negative correlation. Sample station numbers placed in circles according to Appendix A, Table A1.
Figure 5. JASP plot for the phytoplankton abundance similarity on the Bolshoe Toko stations of the lake surface according to Figure 1 and Supplement Table S4. The thickness of the lines is proportional to the strength of the connection. Blue lines—positive correlation; red lines—negative correlation. Sample station numbers placed in circles according to Appendix A, Table A1.
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Figure 6. Contour maps of distribution of vegetation-level data: (a) in total phytoplankton abundance, cells L−1; (b) in cyanobacteria abundance, cells L−1; (c) in Dinoflagellata abundance, cells L−1; (d) in total phytoplankton biomass, mg L−1; (e), in cyanobacteria biomass, mg L−1; and (f) in Dinoflagellata biomass, mg L−1 at the Bolshoe Toko stations according to Supplement Table S3. Black circles are the sampling stations.
Figure 6. Contour maps of distribution of vegetation-level data: (a) in total phytoplankton abundance, cells L−1; (b) in cyanobacteria abundance, cells L−1; (c) in Dinoflagellata abundance, cells L−1; (d) in total phytoplankton biomass, mg L−1; (e), in cyanobacteria biomass, mg L−1; and (f) in Dinoflagellata biomass, mg L−1 at the Bolshoe Toko stations according to Supplement Table S3. Black circles are the sampling stations.
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Figure 7. Distribution of the eutrophic indicator species number (a); water quality class 5 indicators species number (b); saprobity index S (c); and WESI (d) at the Bolshoe Toko stations according to Supplement Table S7. Black circles are the sampling stations.
Figure 7. Distribution of the eutrophic indicator species number (a); water quality class 5 indicators species number (b); saprobity index S (c); and WESI (d) at the Bolshoe Toko stations according to Supplement Table S7. Black circles are the sampling stations.
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Figure 8. JASP plot for the phytoplankton species–indicators correlation on the Bolshoe Toko stations over the quadrants of the lake surface according to Figure 1 and Supplement Table S7. The thickness of the lines is proportional to the strength of the connection. Blue lines—positive correlation; red lines—negative correlation. Clusters marked as 1 and 2. Sample station numbers placed in circles according to Appendix A, Table A1.
Figure 8. JASP plot for the phytoplankton species–indicators correlation on the Bolshoe Toko stations over the quadrants of the lake surface according to Figure 1 and Supplement Table S7. The thickness of the lines is proportional to the strength of the connection. Blue lines—positive correlation; red lines—negative correlation. Clusters marked as 1 and 2. Sample station numbers placed in circles according to Appendix A, Table A1.
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Figure 9. RDA triplot for the phytoplankton species in taxonomic phyla, saprobity index S, WESI, and main environmental variables in the Bolshoe Toko lake stations according to Supplement Tables S1, S3, S7 and S8. Arrows with the red end—environmental variables; blue arrows—taxonomic phyla; empty circles—sampling stations.
Figure 9. RDA triplot for the phytoplankton species in taxonomic phyla, saprobity index S, WESI, and main environmental variables in the Bolshoe Toko lake stations according to Supplement Tables S1, S3, S7 and S8. Arrows with the red end—environmental variables; blue arrows—taxonomic phyla; empty circles—sampling stations.
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Figure 10. Distribution of the number of diatom species in the flora of the Bolshoe Toko Lake by IUCN threat categories. The order of the IUCN categories on the x-axis corresponds to threat reduction. The code of each IUCN category according to Table 2.
Figure 10. Distribution of the number of diatom species in the flora of the Bolshoe Toko Lake by IUCN threat categories. The order of the IUCN categories on the x-axis corresponds to threat reduction. The code of each IUCN category according to Table 2.
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Table 1. Species number in major phyla and classes of phytoplankton in the Bolshoe Toko Lake, Yakutia, 2015.
Table 1. Species number in major phyla and classes of phytoplankton in the Bolshoe Toko Lake, Yakutia, 2015.
PhylumNo. of SpeciesClassNo. of Species
Heterokontophyta309Bacillariophyceae264
Chlorophyta70Chlorophyceae51
Charophyta45Cyanophyceae44
Cyanobacteria44Zygnematophyceae42
Dinoflagellata8Chrysophyceae24
Euglenophyta3Trebouxiophyceae15
Total479Dinophyceae8
Mediophyceae8
Xanthophyceae6
Coscinodiscophyceae5
Ulvophyceae4
Euglenophyceae3
Klebsormidiophyceae3
Eustigmatophyceae1
Phaeophyceae1
Total479
Table 2. Threat categories for diatom species in the Bolshoe Toko Lake.
Table 2. Threat categories for diatom species in the Bolshoe Toko Lake.
IUCN CategoryIUCN CodeNo. of Red List CategoryRed List CategoryNumber of Species
EXTINCTEX1Extinct or Lost0
CRITICALLY ENDANGEREDCR2, 3Threatened with Extinction, Highly Threatened10
ENDANGEREDEN4, 5Threatened, Threat of Unknown Extent32
VULNERABLEVU6Extremely Rare2
NEAR THREATENEDNT7Near Threatened14
LEAST CONCERNLC9Not Threatened66
DATA DEFICIENTDD8Data Deficient7
NOT EVALUATEDNE0, 10Not established15
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Barinova, S.; Gabyshev, V.A.; Gabysheva, O.I.; Ivanova, A.P.; Tsarenko, P.M. Ecological Approaches in the Process of Formation of the Bolshoe Toko National Park, Yakutia. Diversity 2025, 17, 625. https://doi.org/10.3390/d17090625

AMA Style

Barinova S, Gabyshev VA, Gabysheva OI, Ivanova AP, Tsarenko PM. Ecological Approaches in the Process of Formation of the Bolshoe Toko National Park, Yakutia. Diversity. 2025; 17(9):625. https://doi.org/10.3390/d17090625

Chicago/Turabian Style

Barinova, Sophia, Viktor A. Gabyshev, Olga I. Gabysheva, Anna P. Ivanova, and Petro M. Tsarenko. 2025. "Ecological Approaches in the Process of Formation of the Bolshoe Toko National Park, Yakutia" Diversity 17, no. 9: 625. https://doi.org/10.3390/d17090625

APA Style

Barinova, S., Gabyshev, V. A., Gabysheva, O. I., Ivanova, A. P., & Tsarenko, P. M. (2025). Ecological Approaches in the Process of Formation of the Bolshoe Toko National Park, Yakutia. Diversity, 17(9), 625. https://doi.org/10.3390/d17090625

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