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Article

Invertebrate Assemblages in Some Saline and Soda Lakes of the Kulunda Steppe: First Regional Assessment and Ecological Implications

by
Larisa Golovatyuk
1,2,*,
Timur Kanapatskiy
3,
Olga Samylina
3,
Nikolay Pimenov
3,
Larisa Nazarova
4,* and
Anna Kallistova
3
1
Papanin Institute for Biology of Inland Waters, Russian Academy of Sciences, Nekouzsky District, 152742 Borok, Russia
2
Institute of Ecology of the Volga River Basin, Samara Federal Research Scientific Center, Russian Academy of Sciences, Komzina Str. 10, 445003 Tolyatti, Russia
3
Winogradsky Institute of Microbiology, Federal Research Center of Fundamentals of Biotechnology, Russian Academy of Sciences, 119071 Moscow, Russia
4
Laboratory of Biophysics of Ecosystems, Institute of Biophysics, Siberian Branch of Ruissian Academy of Sciences, Akademgorodok 50-50, 660036 Krasnoyarsk, Russia
*
Authors to whom correspondence should be addressed.
Water 2025, 17(15), 2330; https://doi.org/10.3390/w17152330
Submission received: 17 June 2025 / Revised: 19 July 2025 / Accepted: 1 August 2025 / Published: 5 August 2025
(This article belongs to the Section Biodiversity and Functionality of Aquatic Ecosystems)
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Abstract

The taxonomic composition and structure of invertebrate assemblages in five lakes from the Kulunda steppe, located in an arid region of southwestern Siberia (Russia), were studied. The lakes varied greatly in their total salinity (5 to 304 g L−1) and carbonate alkalinity (0.03 to 4.03 mol-eq L−1). The invertebrate fauna was characterized by low diversity. Only five taxa of macrozoobenthos and two taxa of planktonic invertebrates were identified. As water salinity increased, the taxonomic diversity of the studied lakes decreased, and at salinities > 276 g L−1, monodominant assemblages were formed. The high numbers and biomass of aquatic organism provide a rich food supply for native and migratory waterfowl. The low taxonomic diversity of the invertebrate assemblages of the lakes makes them vulnerable to any negative external impact. The climate in the Kulunda steppe demonstrates a long-term aridization trend. If this continues in the future, then over time, this may lead to the gradual salinization of lakes and a further decrease in the taxonomic diversity of hydrobiological assemblages. This emphasizes the ecological importance of the studied territory and the necessity for its inclusion in the list of sites protected by the Ramsar Convention.

1. Introduction

At present, various natural effects of climate change have an impact on hydrobiological communities in many areas across Eurasia [1,2,3]. These impacts include changes in temperature, precipitation patterns, and water availability. Climate-related stressors are among the most important environmental factors causing changes in the composition and structure of hydrobiological communities [4,5,6,7]. Climatic fluctuations reinforced by anthropogenic impacts intensify negative processes in natural ecosystems [8], such as, e.g., salinization of fresh waters that is observed in many regions across the world [9,10]. We hypothesize that increasing salinity leads to a decrease in taxonomic richness and promotes the monodominance of hydrobiological assemblages.
Examining the current state of the aquatic fauna in naturally saline water bodies is crucial for determining which species can be used as water salinity indicators, evaluating the composition of hydrobiological communities that are continuously exposed to varying salinities [11], and spotting “early warning signals” of ecosystem change. This knowledge fosters a better understanding of the possible ways water ecosystems can develop under the influence of salinity [12], as soda lakes worldwide (e.g., Lake Magadi, Mono Lake) show similar ecological constraints [13,14]. In most parts of the world, there is a lack of information about the unique diversity of saltwater organisms and their ability to tolerate this environment, as well as the patterns of community formation [10,11,12]. This is especially important, considering that saline lakes are present worldwide and that their estimated total volume (1.04 × 105 km3) is comparable to that of freshwater lakes (1.24 × 105 km3) [15].
Kulunda steppe is located in the southern part of the West Siberian Plain (Altai Krai of the Russian Federation). Many lakes of different origin, morphometry, hydrology and hydrochemistry are located in this region. Most of them are brackish, saline, or hypersaline. Soda lakes, which are double-extreme environments with high salinity (up to saturation) and high pH (up to 11) due to the high content of soluble carbonates in brines, are located in the south [16]. The Kulunda lakes belong to the wetlands of Russia [17]. The lakes in this region provide important habitats for both resident and migratory waterfowl (shelduck, crane, coot swan), whose food source is largely determined by the invertebrate organisms inhabiting these lakes [18]. Studies of the invertebrate fauna of the Kulunda steppe lakes began in the second half of the 20th century [19]; however, to date, the fauna of only nine lakes has been described [20]. In total, 37 taxa have been identified in these nine lakes including larvae of the Scathophagidae gen., Ceratopogon sp. and Setacera sp. [21], Diptera from the genus Ephydra (Isachenko, 1951), and larvae, pupae, and adults of E. glauca [22]. Most attention has been traditionally paid to the large brackish lakes [23,24], while information about benthic communities of small saline and hypersaline lakes is absent or only fragmentary data is available [20]. The macrozoobenthos communities of the unique soda lakes are also poorly studied, although back in the 1930s B.L. Isachenko, the founder of microbiological studies of soda lakes, noted a huge number of Artemia, larvae and pupae of Ephydra in the Tanatar lake brines, as well as a high biomass of insect remains, including winged ants [25].
Saline lake water bodies, including soda lakes, are inhabited by invertebrate communities morphologically and physiologically adapted to high salinity and pH [12]. However, the ecological preferences and tolerances of many salinity-resistant species are not fully understood [26]. Therefore, when studying such lakes, it is very important to identify the benthic species based on all life stages and to register the ecological conditions of their habitats.
This study aims to quantify the diversity, structure, and distribution patterns of invertebrate assemblages along a salinity gradient in five lakes of the Kulunda steppe.

2. Materials and Methods

2.1. Study Area

Kulunda steppe (70–250 m a.s.l.) is located between the Ob and Irtysh rivers, mainly on the territory of the Altai Krai in Russia and partly in northern Kazakhstan (Figure 1a). It is a part of the Eurasian steppe belt with a prevalence of grass communities interspersed with pine forests [27,28].
The climate of this region is continental, Dfb (snow, fully humid, warm summers) according to Köppen–Geiger classification [29]. It is characterized by long and cold winters (mean January T = −17–19 °C) and short warm summers (mean July T = 19–21 °C). According to meteorological observation (Station Klyuchi, 52.27 N, 79.20 E, alt 143 m a.s.l.), mean annual temperature (Tann) since the beginning of observations in 1951 is 2.9 °C (Figure 2). Annual precipitation varies between 150 and 900 mm per year. The average annual precipitation since the beginning of observations in 1966 is 354 mm per year (Figure 2). Most precipitation occurs in the summer.
An analysis of climate change in the Kulunda steppe has shown that there is a noticeable increase in the average annual temperature that became especially prominent after 1982. On the contrary, a negative trend is observed in the annual precipitation, which particularly decreased after 1978 (Figure 2). Taken together, these processes lead to a noticeable increase in the degree of aridity of the forest-steppe and steppe landscapes of the Altai Krai [30].
Our investigation included five lakes (Figure 1b): Petukhovskoe Sodovoe (52°6′20″ N, 79°9′22″ E), Lomovoe (51°42′38″ N, 79°43′12″ E), Tanatar VI (51°37′08″ N, 79°48′53″ E), Tanatar IV (51°37′59″ N, 79°51′11″ E), and Gorchina 1 (51°40′19″ N, 79°54′20″ E). These lakes were chosen to represent a salinity gradient and low anthropogenic impact. They experience less anthropogenic impact than nearby lakes: soda extraction is not conducted here (unlike in the Tanatar I–III lakes), Artemia harvesting does not occur (except for possible poaching), recreational activities (swimming, mud therapy) are not practiced, and other forms of active human exploitation are absent. Nevertheless, given the proximity of settlements (the village of Severka for Petukhovskoe Lake, and the village of Malinovoe Ozero for the others), the studied lakes cannot be considered unaffected by human activity. All lakes included in this study are located in the dry steppe region of ancient deltas of paleorivers with chestnut soils and solonetzes [16]. The lakes are endorheic, fed mainly by groundwater [23], and experience only a minor anthropogenic load [23].

2.2. Sampling and Laboratory Methods

The lakes Tanatar VI, Petukhovskoe Sodovoe, and Gorchina 1 were sampled twice, in July 2021 and July 2022. The lakes Tanatar IV and Lomovoe were sampled once, in July 2021 and July 2022, respectively.
Lake water pH was measured using a field pH meter 320/Set-1 (WTW GmbH, Weilheim, Germany). Total salinity values were measured in the field using a portable refractometer (ATAGO ATCS/MillE, ATAGO CO., Ltd., Tokyo, Japan) and are presented in promille (‰, parts per thousand). For samples collected in 2022, the total dissolved solids (TDSs) in g L−1 were additionally measured by the gravimetric method. Data on total salinity and TDS obtained by the two methods gave comparable results, so we used the values of total salinity expressed in promille (‰). The salinity classification of lakes is given according to the Venice System (1958) with modifications for inland water bodies [31]. The total carbonate alkalinity and concentrations of the anions CO32− and HCO3− were assessed by titration with a 1 M or 0.1 M HCl solution, depending on the hydrochemical type and total salinity of the lake. To prepare acid solutions, 1 N standard titers were used. Measurements were performed three times. The measurement errors did not exceed 5%. Other main ions (Na+, SO42−, and Cl−) were measured only for samples collected in 2022 by capillary electrophoresis using the “Kapel-103PT” system.
The hydrobiological study included an investigation of macrozoobenthos as well as planktonic crustaceans from the genus Artemia. The qualitative macrozoobenthos samples were taken from the lakes Tanatar IV (2 samples), Tanatar VI (5 samples), Petukhovskoe Sodovoe (5 samples), and Gorchina 1 (5 samples) in 2021. Quantitative macrozoobenthos samples were taken from the lakes Petukhovskoe Sodovoe, Tanatar VI, Gorchina 1, and Lomovoe in 2022 in two repetitions from four stations (in the northern, southern, eastern, and western parts of the lakes) in each of the studied lakes. Quantitative plankton samples were taken from the lakes Petukhovskoe Sodovoe, Tanatar VI, Gorchina 1, and Lomovoe in one repetition from four stations. The water depth at the sampling stations was 0.4–0.5 m.
The benthic invertebrates were collected using a handle blade trawl. The handle blade trawl had a 20 cm mouth width with a blade at the lower side (pulling way 50 cm). The collected sediments were washed through a net with a mesh size of 300–333 µm and preserved with a 4% formaldehyde solution [32]. Ten liters of water were filtered through a plankton net to account for the abundance and biomass of planktonic Artemia crustaceans. Insect imago was caught using a special net and preserved in 96° alcohol. In total, 49 qualitative and quantitative samples of benthos and 16 samples of plankton were processed, and 47 adult insects were caught and identified. The Diptera larvae were reared in the laboratory in order to identify them at the species level. The reared pupae and adults were preserved in 96° alcohol. Artemia were identified morphologically using adult specimens. All identifications followed special taxonomic keys [22,33,34,35,36,37].

2.3. Data Analyses

The distribution of species in the lakes was analyzed using the calculated frequency of occurrence (F, %) of species across all samples [38]. The Shannon’s diversity index [39] and Simpson index [40] were calculated using the Paleontological Statistics (PAST) version 3.0 [41]. Relationships between the significant environmental variables and the individual RDA axes were examined through correlation coefficients and t-values. Both DCA and RDA were performed using CANOCO 4.5 [42].
The graph is based on open-access meteorological data for the Klyuchi weather station (52.27 N, 79.20 E, alt 143 m a.s.l.) [43]. Thin gray lines represent the data (mean annual measured values); black lines represent Loess 0.3 smooth of the data; dashed gray lines represent the mean data values for the periods of observations: 2.9 °C for the T (1951–2022), and 354 mm year−1 for P (1966–2022).

3. Results

3.1. Hydrochemical Characteristics of the Investigated Lakes

The hydrochemical parameters and sediment characteristics of the studied lakes are given in Table 1. During the study period, Lake Tanatar IV was characterized as myxosaline, intermediate between oligo- and mesosaline (5‰). Lake Tanatar VI was also characterized as myxosaline. However, the salinity in this lake increased from 22‰ in 2021 (polysaline) to 34‰ in 2022 (eusaline). Lakes Petukhovskoe Sodovoe, Gorchina 1, and Lomovoe were characterized as hypersaline during both years (43 to 304‰). Our observation has shown that there was a noticeable increase in the total salinity in the three studied lakes in 2022 compared to 2021 (Table 1). The lakes Tanatar IV, Tanatar VI, Petukhovskoe Sodovoe, and Gorchina 1 belong to the soda (carbonate) type, and only Lake Lomovoe belongs to the saline (chloride) type. In all lakes except for Lake Lomovoe, we observed alkaline water with pH values of 9.6–10.2. The total alkalinity was the highest in Lake Gorchina 1 in July 2022 (4.03 mol-eq L−1).

3.2. Hydrobiological Characteristics

During the study period, five species of macrozoobenthic organisms from two orders were found in both qualitative and quantitative samples. These included five species from the order Diptera (families Ceratopogonidae, Ephydridae, Chironomidae, and Muscidae), and one species from the order Heteroptera (family Corixidae). Additionally, two species from the order Anostraca (family Artemiidae) were identified in plankton samples (Table 2).
Qualitative macrozoobenthos samples included water bugs Paracorixa concinna concinna (Lake Tanatar IV), larvae of the biting midge Palpomyia schmidti (Lake Petukhovskoe Sodovoe), and chironomids Microchironomus deribae (lakes Tanatar IV and Tanatar VI).
Four invertebrate taxa were identified in the quantitative samples of macrozoobenthos from Lake Tanatar VI. Four other taxa were found in Petukhovskoe Sodovoe Lake. One zooplankton taxon was found in each of Lakes Tanatar VI, Petukhovskoe Sodovoe, Lomovoe, and Gorchyna 1. Benthic invertebrates were not found in any of these last two lakes.
Insects P. schmidti, C. riethi, and E. glauca had a frequency of occurrence of over 50% in benthic samples (Table 2). Larvae of P. schmidti and E. glauca were permanent residents only in Lakes Tanatar VI and Petukhovskoye Sodovoye. Biting midge Culicoides riethi was found only in Lake Tanatar VI with an occurrence rate of 88%.
With the exception of Lake Tanatar IV, Artemia sp. was found in all the lakes. However, in lakes Tanatar VI and Petukhovskoe Sodovoe, only one specimen of Artemia sp. was found, and it was only in one sample from each lake. The frequency of occurrence of Artemia sp. reached 100% in Lake Gorchina 1, and population of Artemia parthenogenetica was also found in Lomovoe Lake with a frequency of 100%. The identification of crustaceans in the soda lakes Tanatar VI, Petukhovskoe Sodovoe, and Gorchina 1 was limited to the genus level (Artemia sp.). Artemia sp. identification was limited by lack of genetic markers.
In Tanatar VI and Petukhovskoe Sodovoe lakes, the average macrozoobenthos abundance was 7 thous. ind. m−2 and 8.03 thous. ind. m−2, respectively, while the corresponding biomass was 11.7 g m−2 and 9.03 g m−2, as shown in Table 3. Insect larvae belonging to the families Ceratopogonidae and Ephydridae dominated the assemblages. The proportion of biting midges in the overall abundance of macrozoobenthos ranged from 84% or 5.88 ind. m−2 (Lake Tanatar VI) to 98% or 7.87 ind. m−2 (Lake Petukhovskoe Sodovoe). The greater part of the biomass was produced by biting midges (Lake Petukhovskoe Sodovoe, 76% or 6.86 g m−2) and shore flies (Lake Tanatar VI, 77% or 9 g m−2) (Figure 3).
The insects P. schmidti (Lake Petukhovskoe Sodovoe), C. riethi (Lake Tanatar VI), and E. glauca (Lake Tanatar VI) constitute the greatest part of the abundance and biomass of macrozoobenthos in the studied lakes (Figure 3). The larvae of P. schmidti inhabited mainly sandy bottom sediments (fine, medium-grained sand). C. riethi and E. glauca had high abundances in gray silt.
Very dense populations of planktonic crustaceans from the genus Artemia were found in Gorchina 1 and Lomovoe lakes; the average abundance of these crustaceans ranged from 8.0 thous. ind. m−3 to 39.0 thous. ind. m−3. The highest abundance of 70.4 thous. ind. m−2 was found in the Lake Lomovoe. The dominance indices of Artemia in the zooplankton of these lakes reached 100%.
The distribution of benthic and planktonic taxa along the axes reflects the influence of the hydrochemical parameters of the water on their distribution (Figure 4). The most tolerant species, Artemia parthenogenetica and Artemia sp., which can survive in higher salinity, are located in the right center of the biplot and correspond to the (S) axis. The two-winged Ephydra glauca is located on the left in the centrum of the biplot and is associated with high pH values. Palpomyia schmidti and Lispe sp., located in the lower left part of the biplot, prefer lower values of salinity, alkalinity (A) and pH. The most distant from all axes Microchironomus deribae and Culicoides riethi show the greatest sensitivity to all the factors under consideration.
The calculated Shannon and Simpson diversity indices, which are two of the most commonly used indices in hydrobiological and ecological research, showed low values in all the studied lakes. The indices ranged from 0 to 0.72 for the Shannon index and from 0 to 0.48 for the Simpson index. The indices were the lowest in the Gorchina 1 and Lomovoe hypersaline lakes (Table 3).

4. Discussion

The invertebrate assemblages of the five investigated Kulunda saline lakes have been studied for the first time. All these lakes are representative of the common regional type of water bodies, with total salinity ranging from myxosaline (5‰) to hypersaline (304‰). Prior to our study, hydrobiological investigations of macrozoobenthos took place only at nine other lakes of the Kulunda steppe: Baklanye, Dolgoye, Gorkoye-Lebedyanskoye, Krivoe, Kulundinskoye, Mostovoye, Batovoye, Chernokovo, and Lena [21,45,46]. In total, 37 taxa have been identified in these nine lakes and included larvae of the Scathophagidae gen., Ceratopogon sp. and Setacera sp. [21], Diptera from the genus Ephydra [21], and larvae, pupae, and adults of E. glauca [22]. On the other hand, research has already been performed on populations of the planktonic crustacean Artemia in the variety of saline lakes of Altai Krai [35,47,48,49,50,51]. To the best of our knowledge, however, we have demonstrated the emergence of Artemia in lakes Tanatar VI and Gorchina 1 for the first time.
The data on the taxonomic composition of the macrozoobenthos in the saline lakes of Altai Krai was expanded by our study, which included insects from the families Ceratopogonidae, Chironomidae, and Ephydridae that were identified by all life stages. This helped to accurately identify the collected specimens to the species level and, in turn, more precisely describe their ecological features based on literature and our own data.
The biting midges P. schmidti and chironomids M. deribae have been recorded for the first time in the lakes of the Kulunda steppe and throughout the Altai Krai, according to a comparison of our findings with previously published [19,21,45,46].
P. schmidti was found in Tanatar VI and Petukhovskoe Sodovoe lakes in the salinity range of 34–60‰ (TDS = 31.6–60.5 g L−1). This species is halophilic and is distributed worldwide in the steppe and semi-desert regions [34,52]. It represents a Sahara-Arabian element in the Palearctic region, and our discovery of this species in the steppe Altai lakes expands its known distribution, which previously included Mongolia, Iraq, Iran, Hungary, Tajikistan, Kazakhstan, Azerbaijan, south of European Russia, Ukraine, Slovakia, and Spain [34]. Earlier studies revealed that P. schmidti larvae could survive in a salinity range of 8.3 to 78 g L−1, with the highest abundance under a salinity range of 21 to 32 g L−1 [53]. P. schmidti dominated the benthic assemblages and had the highest abundance (48,000 ind./m2) in our study of the saline rivers in the Lake Elton basin under salinity of 27 g L−1 [54]. In the studied Kulunda lakes, this species was the most abundant (15,496 ind./m2) under a total salinity of 60‰ (TDS = 60.5 g L−1). It was the first time that such a high abundance of P. schmidti under such high salinity was discovered. Earlier it was shown by FA-biomarker analysis that the larvae of P. schmidti selectively consumed algae, primarily diatoms, and ignored bacteria and detritus [55].
Another Ceratopogonidae species, Culicoides riethi, found only in the Lake Tanatar VI (total salinity 34‰, TDS = 31.6 g L−1) belongs to the euryhaline ecological group; however, it frequently reaches high densities in saline waters [56]. This species is known from Russia, the middle and southern regions of Western Europe, North Africa, Iran, Mongolia, North China, and Japan [57,58]. It was found in benthic assemblages of the hypersaline lake Baskunchak [59]. It is also widely distributed in saline rivers from the hypersaline Lake Elton basin, where its larvae and pupae inhabit environments with salinities ranging from 4 to 31.7 g L−1 [53]. The upper salinity limit of the Elton population is similar to the salinity of Tanatar VI Lake. The Culicoides riethi was not detected in the investigated Kulunda lakes at higher salinities during this study. The ordination demonstrated that it showed greater sensitivity to salinity than a number of other species living in these lakes.
Shore flies E. glauca (Ephydridae) were found in Tanatar VI and Petukhovskoe Sodovoe lakes at a total salinity range of 34–60‰ (TDS = 31.6–60.5 g L−1). The species was more abundant in Tanatar VI Lake, where it was detected in all samples examined, at a salinity of 34‰ (TDS = 31.6 g L−1). This halophilic insect species usually inhabits coastal zones of salt lakes and seas and is known from Europe, Kazakhstan, Uzbekistan, Ukraine, and Turkey [57]. Previously, it was already recorded in the lakes of the Altai Krai [22]. Ephydridae species are often associated with riparian habitats and extremely high salinity [60,61,62,63]. Larvae of the genus Ephydra feed on cyanobacteria, unclaimed by other invertebrates [64,65]. Larvae have functional anal organs on the last segment of the abdomen (also called anal papillae), which can change the “relief” and surface area of their surface in order to regulate the flow of ions from the environment into the insect’s hemolymph [66]. Both adult flies and their larvae are an invaluable source of food for birds, reptiles, and predatory arthropods that inhabit the shores of saline lakes [63].
Chironomids M. deribae, which was found only in two samples from the lake Tanatar VI, inhabit inland saline rivers [54], alkaline-saline lakes [67], estuaries, and coastal zones of seas [68]. This species was found earlier in waters with salinity ranging from 4.6 g L−1 to 42 g L−1 in the southeast of the European part of Russia [53] and the Netherlands [69]. We found the larvae of this chironomid species in the oligosaline and mesosaline sections of the rivers from the Lake Elton basin, where the optimum salinity for the species was 10.6 g L−1 [70]. The species has been found in various biotopes like ditches, small and large ponds, cattle pools, lakes, on sandy and clay sediments [69].
The Heteroptera P. concinna concinna was found only in a qualitative sample taken from the myxosaline lake Tanatar IV (5‰). This is a euryhaline subspecies of insect, found in various fresh and saline water bodies of Northern and Central Eurasia, where they often reach large abundances. There is evidence of P. concinna concinna dominance in highly saline steppe lakes in Western Siberia [71]. In saline rivers of the Lake Elton basin, they live under a salinity of 26 g L−1 [53]. In Kulunda lakes with higher salinity, P. concinna concinna was not found during this study.
The fly Lispe sp. was found only in the Petukhovskoe Sodovoe lake under a total salinity of 60‰ (TDS = 60.5 g L−1). The genus belongs to the family Muscidae, which is widely distributed worldwide except for Antarctica and New Zealand, and includes ca. 200 species [63,64]. Both larvae and imago of Lispe sp. inhabit semi-aquatic environments, ranging from flowing to standing and from freshwater to saltwater. Lispe sp. is known to be exceptionally tolerant to variable environmental conditions [72,73].
Thus, the invertebrate assemblages of the studied water bodies consist mainly of halophilic macrozoobenthos species, enriched by euryhaline taxa (P. concinna concinna, C. riethi) and planktonic halobionts (Artemia parthenogenetica, Artemia sp.). These salt-resistant species are widely distributed, not only in various aquatic ecosystems (lakes, rivers, and shallow sea bays) throughout Eurasia but also in saline-soda lakes on other continents [67]. Earlier genetic studies showed that the bisexual populations of Altai lakes may belong to A. urmiana [74,75]. According to recent data, some bisexual populations in the region may belong to the species A. sinica or even A. sorgeloosi [51]. Thus, species identification of bisexual Artemia was not determined in our study since genetic examination is highly desirable.
The taxonomic richness of invertebrates in the Kulunda steppe lakes is poor, which is primarily due to the high salinity of the water. The low taxonomic diversity of the macrozoobenthos of saline lakes has been repeatedly reported earlier [76,77]. Thus, just 11 species of invertebrates were identified during a previous long-term study of macrozoobenthos from other salt lakes in the Kulunda steppe [46]. Similarly, a two-year investigation of five salt lakes in Crimea revealed only eight macrozoobenthos species [78]. The high alkalinity of the studied lakes may have an impact on species richness, as was earlier shown for the soda lakes of Ethiopia, where only seven benthic species were discovered [67].
It has been shown that in the range from 0.18 to 30 g L−1, certain taxonomic groups of organisms sensitive to this environmental factor disappear from the macrozoobenthos community, such as mayflies, caddis flies, molluscs, and leeches [79]. As a result, the total number of species decreases from 156 in freshwater to only 10 at a salinity level of 30 g L−1 [79].
Though our investigation included a limited number of lakes and samples, a decrease in the species richness of benthic assemblages with the increase in salinity could be noted. The least mineralized myxosaline lake Tanatar VI had the greatest taxonomic richness (five species). In the lakes with salinity above 276‰, benthic invertebrates were not recorded at all, and only planktonic crustaceans from the genus Artemia were found. It was shown that the species and functional diversity of zooplankton decreased with increasing salinity, whereas their abundance, biomass, and daily secondary production did not [80]. Our investigation is based only on mid-summer sampling. In the future, it is necessary to plan multi-season studies of these lakes.
Only one group of consumers, namely crustaceans from the genus Artemia, is usually present in biological communities of hypersaline lakes with salinities of 184–340 g L−1 [46,81]. Numerous studies of lakes with varying degrees of salinity have demonstrated a decrease in taxonomic richness of macrozoobenthos and zooplankton with an increase in water salinity [46,76,77,78,79,80,81,82]. This finding is supported by our current investigation of the saline lakes in the Kulunda steppe.
Regarding taxonomic richness, Diptera predominate in the Kulunda steppe lakes, which is consistent with other previously studied water bodies of the region [46] and with saline or soda lakes in other parts of the world [12,67,83,84,85,86,87,88]. This can be explained by the resistance of a number of Diptera species to extreme environments [64,89]. For instance, the presence of long respiratory tubes that enable insects to breathe atmospheric air determines the morphological adaptations to low-oxygen hypersaline conditions of the larvae and pupae of some dipterans, including species of the genus Ephydra. Some Diptera species that receive atmospheric air through open tracheae retain a relatively impenetrable cuticle, like their terrestrial ancestors [90]. Apparently, the respiratory strategy strongly affects osmotic permeability and, probably, the response of species to salinization [90,91]. Short life cycle, high fertility of adults, capacity for active dispersal, increased mobility, and use of the same substrate as food by both larvae and adults are among ecological adaptations that allow them to survive in harsh environments [64]. Furthermore, several species of the family Ephydridae inhabit highly salinized environments and consume cyanobacteria that are not claimed by other aquatic insect species. This allows them to avoid food competition and increases their chances for survival in extreme conditions [64,65].
The fauna of the lakes with a total salinity of 34 to 60 g L−1 were not identical. The benthic community of the lake Tanatar VI (34 g L−1) is dominated by the halophilic E. glauca and is supplemented by two euryhaline species, whereas Petukhovskoe Sodovoe (60 g L−1) is predominantly inhabited by the halophilic species P. schmidti and E. glauca and one euryhaline species. This is due to differences in salinity tolerance and the association of dominant species with different types of sediments, which include the biting midges P. schmidti and C. riethi described in detail above.
There are not only differences in the composition of dominant species in the lakes with different levels of salinity; we also observed a high (up to 100%) degree of dominance reflected by low diversity indices. In the hypersaline lakes (Gorchina 1 and Lomovoe), monodominant complexes of aquatic organisms were formed. The development of monodominant complexes in the Kulunda steppe lakes was previously indicated for zooplankton assemblages [80].
The highest values of the diversity indices among the studied lakes were characteristic of the lake Tanatar VI with the lowest total salinity, and they decreased with the increase in total salinity. This reflects a simplification of the structure of benthic assemblages, which leads to a decrease in ecosystem stability. A similar pattern was found in the lakes from different regions of the world with a wide salinity gradient [81].
The abundance and biomass of macrozoobenthos in the lakes Tanatar VI and Petukhovskoe Sodovoe, with a total salinity of 34 to 60 g L−1, were high due to the mass development of Diptera larvae, which are valuable food sources for native and migratory waterfowl [92]. The observed high abundance of Diptera larvae in these lakes was comparable with, or even exceeded, numbers known from other bird-feeding sites for migratory birds in brackish-water habitats of Europe [93,94,95,96]. This makes the investigated lakes a potential feeding hotspot for native and migratory waterfowl comparable, e.g., to the central and eastern parts of Siwash Bay [96].
Low abundance and biomass of macrozoobenthos were observed previously during the 2007–2011 study of the meso- and hypersaline lakes of the Ob-Irtysh interfluve, which also included water bodies from the Kulunda steppe. The abundance varied from 0.1 to 1.5 thous. ind. m−2, and the biomass varied from 0.07 to 4.28 g m−2 [46]. These parameters were five and two times lower, respectively, than the macrozoobenthos abundance and biomass values that we discovered in the lakes with comparable total salinity. It demonstrates the wide variations in hydrobiont abundance and biomass which can be found in the saline lakes of the Kulunda steppe. This may be caused by the Diptera life cycles, whose mass hatching can markedly deplete macrozoobenthos communities [54].
The abundance and biomass of crustaceans from the genus Artemia in the studied lakes Lomovoe and Gorchina 1 were similar to the average seasonal values previously discovered in several saline lakes in the Kulunda steppe [35].
The climate in the Kulunda steppe demonstrate a long-term aridization trend (Figure 2) [30]. Given continued aridization, we project further salinization, though regional variability may affect exact rates. This can lead to the gradual salinization of lakes and a further decrease in the taxonomic diversity of hydrobionts. This is one of the major climate-related threats for many arid regions of the world that leads to a reduction in the stability of these ecologically vulnerable natural ecosystems [54,97].

5. Conclusions

The study of the modern fauna in the lakes located in the Kulunda steppe revealed that the variety of species of aquatic organisms in soda and hypersaline lakes was limited to a small number of specialized taxa, and was not higher than five species of macrobenthos and two taxa of planktonic invertebrates. The species of the euryhaline, halophilic, and halobiont ecological groups inhabited the lakes. With an increase in water salinity, there was a decrease in the taxonomic diversity of biological communities, leading to development of monodominant hydrobiont complexes at extremely high salinity. The biting midges Palpomyia schmidti Goetghebuer, 1934, and chironomids Microchironomus deribae (Freeman, 1957) were discovered for the first time in the study region. Findings of Diptera species previously unknown in this region significantly expanded our knowledge about their distribution and ecology. The halophilic species P. schmidti was found for the first time at a total salinity of 60‰, which is twice as high as the previously recognized optimum for this species.
The high numbers and biomass of aquatic organisms provide a rich food supply for native and migratory waterfowl. The low taxonomic diversity of the invertebrate communities of the lakes makes them vulnerable to any negative external impact. This highlights the ecological significance of the area under study. Therefore, we recommend including Lakes Petukhovskoe Sodovoe and Tanatar IV in the Ramsar candidate list.

Author Contributions

Conceptualization, L.G., L.N. and O.S.; methodology, L.G., O.S. and A.K.; software, L.G.; validation, O.S., L.G., L.N., N.P. and T.K.; formal analysis, O.S., L.G., L.N. and A.K.; investigation, T.K., O.S. and L.G.; resources, O.S., L.G., L.N. and N.P.; data curation, L.N.; writing (original draft preparation, review and editing) L.G., L.N. and O.S.; visualization, L.G., L.N., O.S. and T.K.; supervision, N.P.; project administration, L.G., O.S., L.N., T.K. and N.P.; funding acquisition, N.P. All authors have read and agreed to the published version of the manuscript.

Funding

Field work was supported by the Russian Science Foundation grant No 22-14-00038-C, https//rscf.ru/project/22-14-00038, accessed on 15 June 2025, data analysis was supported by state assignment of Research Center of Biotechnology. Sample processing was carried out within the framework of the state research assignments No 124032500016-4, 1024032600218-3-1.6.20.

Data Availability Statement

The data presented in this study will be openly available in PANGEA upon article publication.

Acknowledgments

The authors thank Yu.V. Litty (Winogradsky Institute of Microbiology RAS) for obtaining the ion composition data.

Conflicts of Interest

The authors declare no conflicts of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.

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Figure 1. Geographical position of the studied lakes: (a) location of the Kulunda steppe in the south of the West Siberian Plane, (b) location of the studied lakes near villages Severka and Malinovoe ozero (Altai Krai).
Figure 1. Geographical position of the studied lakes: (a) location of the Kulunda steppe in the south of the West Siberian Plane, (b) location of the studied lakes near villages Severka and Malinovoe ozero (Altai Krai).
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Figure 2. Mean annual air temperature (T, °C) and annual precipitation (P, mm year −1) according to the meteorological data (Station Klyuchi, 52.27 N, 79.20 E, alt 143 m a.s.l.). The grey lines show the observed data; the black lines are fitted using LOESS smoothing with a span of 0.5. The dotted lines show the mean values for the entire observation period for T (°C) and P (mm per year).
Figure 2. Mean annual air temperature (T, °C) and annual precipitation (P, mm year −1) according to the meteorological data (Station Klyuchi, 52.27 N, 79.20 E, alt 143 m a.s.l.). The grey lines show the observed data; the black lines are fitted using LOESS smoothing with a span of 0.5. The dotted lines show the mean values for the entire observation period for T (°C) and P (mm per year).
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Figure 3. Structure of macrozoobenthos assemblages in the studied lakes of the Kulunda steppe (July, 2022, quantitative samples).
Figure 3. Structure of macrozoobenthos assemblages in the studied lakes of the Kulunda steppe (July, 2022, quantitative samples).
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Figure 4. RDA biplot illustrating the relationship between five environmental variables and taxonomic composition of macrozoobenthos and plancton. A—total alkalinity, S—salinity, pH—hydrogen index.
Figure 4. RDA biplot illustrating the relationship between five environmental variables and taxonomic composition of macrozoobenthos and plancton. A—total alkalinity, S—salinity, pH—hydrogen index.
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Table 1. Hydrochemical parameters and sediment characteristics of the studied lakes.
Table 1. Hydrochemical parameters and sediment characteristics of the studied lakes.
LakeTanatar IVTanatar VIPetukhovskoe SodovoeGorchina 1Lomovoe
Year20212021202220212022202120222022
Total salinity, ‰52234436082276304
TDS, g L−1n/dn/d31.6n/d60.5n/d243.4337.2
pH9.99.99.810.210.210.310.27.57
Total alkalinity, mol-eq L−10.060.210.350.500.741.24.030.03
Na+, g L−1n/dn/d15.93n/d3.51n/d122.45157.25
CO32−, g L−11.204.809.0013.8020.4034.80106.200
HCO3−, g L−11.223.053.052.443.662.4429.891.83
Cl−, g L−1n/dn/d7.06n/d1.38n/d22.1163.9
SO4−, g L−1n/dn/d1.62n/d8.49n/d1.3771.99
Chemical type **Soda
Na-HCO3-CO3		Soda
Na-HCO3-CO3		Soda
Na-HCO3-CO3		Soda
Na-HCO3-CO3		Saline
Na-Cl
SedimentsGray silt with whit inclusions of soda concretions and medium-grained sand. Black concretions and plant fibers. Possible presence of carbonate detritusFine, medium-grained sand with fragments of microalgaeFine organic dark brown-greenish silt with frequent inclusions of translucent salt crystals. Small concretions of soda (up to 1 mm). Presence of carbonate detritus
Note: ** the chemical type is indicated in accordance with the classification developed in [44].
Table 2. Taxonomic composition and frequency of occurrence: % of the total number of macrozoobenthos samples for the lakes Tanatar IV, Tanatar VI, Petukhovskoe Sodovoe and % of the total number of plancton samples for Tanatar VI, Petukhovskoe Sodovoe, Gorchina 1 and Lomovoe of hydrobionts in the studied lakes from Kulunda steppe in 2021–2022. Life stages: L for larvae; P for pupae. Ecological groups (“Ecology” in the table) in relation to salinity: E for euribiontic, H for halophilic, and HB for halobiontic species. For the lake Tanatar IV, only qualitative macrozoobenthos samples were taken.
Table 2. Taxonomic composition and frequency of occurrence: % of the total number of macrozoobenthos samples for the lakes Tanatar IV, Tanatar VI, Petukhovskoe Sodovoe and % of the total number of plancton samples for Tanatar VI, Petukhovskoe Sodovoe, Gorchina 1 and Lomovoe of hydrobionts in the studied lakes from Kulunda steppe in 2021–2022. Life stages: L for larvae; P for pupae. Ecological groups (“Ecology” in the table) in relation to salinity: E for euribiontic, H for halophilic, and HB for halobiontic species. For the lake Tanatar IV, only qualitative macrozoobenthos samples were taken.
TaxaLife Stages/EcologyTanatar IVTanatar VIPetukhovskoe SodovoeGorchina 1Lomovoe
Order Anostraca
Family Artemiidae
Artemia parthenogeneticaHB 100
Artemia sp.HB 2525100
Klass Insecta
Order Heteroptera
Paracorixa concinna concinna (Fieber, 1848)(♂)/E100
Order Diptera
Family Ceratopogonidae
Palpomyia schmidti Goetghebuer, 1934(L, P♂)/H 87100
Culicoides (Monoculicoides) riethi Kieffer, 1914(L, P♂)/E 87
Family Chironomidae
Microchironomus deribae (Freeman, 1957)(L, P♂)/H1008
Family Ephydridae
Ephydra glauca Meigen, 1830(L, P♂)/E 9261
Family Muscidae
Lispe sp.L, P 46
Number of species 25411
Table 3. Structural indicators of invertebrate assemblages in the investigated Kulunda lakes (July, 2022).
Table 3. Structural indicators of invertebrate assemblages in the investigated Kulunda lakes (July, 2022).
ParameterLake
Tanatar VIPetukhovskoe SodovoeGorchina 1Lomovoe
Abundance of macrozoobenthos, thous. ind. m−2Mean ± Δ7.0 ± 2.528.03 ± 3.98
Min–max0.13–18.12.29–15.65
Abundance of plancton, thous. ind. m−3Mean ± Δ0.01 ± 00.01 ± 08.0 ± 1.2639 ± 12.6
Min–max0.010.014.16–12.227.6–70.4
Biomass of macrozoobenthos, g m−2Mean ± Δ11.7 ± 8.119.03 ± 3.95
Min–max1.61–18.421.74–15.9
Biomass of plancton, g m−3Mean ± Δ0.001 ± 00.001 ± 04.62 ± 4.42201 ± 92
Min–max0.0010.0012.15–6.2839.1–363
Dominant species of macrozoobenthos (d, %), calculated by abundance/biomass C. riethi
(d = 74/18),
E. glauca
(d = 11/78)		P. schmidti
(d = 92/76)
Dominant species of plancton (d, %), calculated by abundance/biomass Artemia sp.
(d = 100/100)		A. parthenogenetica (d = 100/100)
Shannon index (H) 0.07–0.720.06–0.1400
Simpson index (I) 0.02–0.480.02–0.0500
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MDPI and ACS Style

Golovatyuk, L.; Kanapatskiy, T.; Samylina, O.; Pimenov, N.; Nazarova, L.; Kallistova, A. Invertebrate Assemblages in Some Saline and Soda Lakes of the Kulunda Steppe: First Regional Assessment and Ecological Implications. Water 2025, 17, 2330. https://doi.org/10.3390/w17152330

AMA Style

Golovatyuk L, Kanapatskiy T, Samylina O, Pimenov N, Nazarova L, Kallistova A. Invertebrate Assemblages in Some Saline and Soda Lakes of the Kulunda Steppe: First Regional Assessment and Ecological Implications. Water. 2025; 17(15):2330. https://doi.org/10.3390/w17152330

Chicago/Turabian Style

Golovatyuk, Larisa, Timur Kanapatskiy, Olga Samylina, Nikolay Pimenov, Larisa Nazarova, and Anna Kallistova. 2025. "Invertebrate Assemblages in Some Saline and Soda Lakes of the Kulunda Steppe: First Regional Assessment and Ecological Implications" Water 17, no. 15: 2330. https://doi.org/10.3390/w17152330

APA Style

Golovatyuk, L., Kanapatskiy, T., Samylina, O., Pimenov, N., Nazarova, L., & Kallistova, A. (2025). Invertebrate Assemblages in Some Saline and Soda Lakes of the Kulunda Steppe: First Regional Assessment and Ecological Implications. Water, 17(15), 2330. https://doi.org/10.3390/w17152330

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