Composition and structure of macroinvertebrate communities of lakes in different altitudinal zones of Russian Altai
DOI: 10.5281/zenodo.7714667

Composition and structure of macroinvertebrate communities of lakes in different altitudinal zones of Russian Altai

Institute for Water and Environmental Problems, Siberian Branch of the Russian Academy of Sciences (IWEP SB RAS), 1 Molodezhnaya st. Barnaul, 656038, Russia
Institute for Water and Environmental Problems, Siberian Branch of the Russian Academy of Sciences (IWEP SB RAS), 1 Molodezhnaya st. Barnaul, 656038, Russia; Altai State University, 61 Lenina pr., Barnaul, 656049, Russia
Institute for Water and Environmental Problems, Siberian Branch of the Russian Academy of Sciences (IWEP SB RAS), 1 Molodezhnaya st. Barnaul, 656038, Russia
Altitudinal environmental gradient benthic invertebrates mountain lakes


In 2002-2020, the composition and structure of benthic invertebrate communities from 36 lakes of the Russian Altai located at various (low, mid and high) altitudes were studied. Low-mountain and lowland lakes have a similar zoobenthos structure. With height, the taxonomic structure becomes more complicated, and the dominant taxa of macroinvertebrates change. The peculiar feature of bottom zoocenoses in mid- and high-altitude lakes is high frequency of occurrence and large contribution to the total biomass of crustaceans of the family Gammaridae. We described the trophic structure of zoobenthos and identified five main trophic groups. In terms of species number, the collector-detritus feeder and predator groups dominated in the trophic structure of all lakes. By biomass, the growing proportion of filter feeders and shredders was observed with increasing height.

Corresponding author: Olga N. Vdovina (

Academic editor: A. Matsyura | Received 26 August 2022 | Accepted 18 September 2022 | Published 22 November 2022

Citation: Vdovina ON, Yanygina LV, Bezmaternykh DM (2022) Composition and structure of lake macroinvertebrate communities in different altitudinal zones of Russian Altai. Acta Biologica Sibirica 8: 531–555.


altitudinal environmental gradient, benthic invertebrates, mountain lakes.


Worldwide, about 20% of lakes are located at an altitude above 1000 m (Verpoorter et al. 2014). In terms of physical and biological properties, there is a big difference between high- and low-mountain lakes (Moser et al. 2019). Most mountain lakes are located in severe climate with low water temperatures, a prolonged period of snow and ice cover, and are characterized by poor food conditions. Compared to low-mountain lakes, high-mountain lake ecosystems are usually distinguished by low species richness (Fureder et al. 2006; Fjellheim et al. 2009) and simplified food webs, often consisting of less than three trophic levels (McNaught et al. 1999). To what extent an altitude affects the characteristics of a lake basin and related biogeochemical processes or how these environmental conditions influence biological properties and structures of communities are still poorly studied (Loria et al. 2019). The altitude gradient includes changes in various environmental factors, such as temperature, organic matter content, substrate type, macrophyte diversity (Hoffman et al. 1996; Nyman et al. 2005; Collado and De Mendoza 2009). When studying the altitude influence, the greatest attention is paid to temperature effects on distribution of macroinvertebrate groups, including Chironomidae (Larocque et al. 2001; Nyman et al. 2005; Bitusik et al. 2006), Oligochaeta (Dumnicka, Galas 2002; Collado and De Mendoza 2009; De Mendoza and Catalan 2010), and Trichoptera (Solem and Birks 2000) in different-altitude lakes. In mountain lakes, environmental factors are closely associated with altitude that generates complicated relationships and an inability to distinguish the impact of individual variables.

The studies of the macroinvertebrate faunas of mountain lakes throughout Europe (Fjellheim et al. 2009) demonstrate the presence of the same species, but different taxonomic composition. They also emphasize the importance of detailed knowledge of the regional faunas. Currently, the problem of identifying the formation patterns of benthic invertebrates in specific regions remains relevant.

Altai is the highest mountain region in southern Siberia; the ridges of its central and eastern parts rise above 3–4 km and are covered with eternal snow and glaciers. In the Russian Altai, there are more than 3,500 lakes, but the area of only 75 of them exceeds 1 km2 (Gvozdetsky and Mikhailov 1978). Most of the glaciers high-mountain lakes located in the subalpine zone at the very edge of the snow are almost completely devoid of life. With descending from Altai peaks, the fauna of lakes becomes similar to that of lowland ones (Zhadin and Gerd 1961). It became known about the entry of air pollutants into European mountain regions from industrially developed lowlands at the end of the last century (Brittain and Milner, 2001). Air pollution poses a threat to alpine ecosystems because many organisms in these areas exist at the limit of their tolerance to chemical and physical environmental factors (Fjellheim et al. 2000). Unlike mountain lakes of Europe, the anthropogenic load on the mid- and high Altai mountains is quite low or completely absent because of their remoteness and inaccessibility. It allows the latter as reference areas for indicating environmental changes.

Macroinvertebrates of the Altai region have been investigated since the beginning of the XX century (Gundrizer et al. 1982; Johansen 1981). In expeditions to the upper reaches of the Ob River, attention was traditionally paid to studying lakes, and primarily, Lake Teletskoye (Koveshnikov 2014). Since these lakes are situated in regions that are difficult to reach, the zoobenthos data are often fragmentary (with only evaluated fish forage reserves) or even missing altogether (Vesnina et al. 2012; Zalozny and Vorob’ev 2006; Krylova 2016; Popov et al. 2003).

In this study, we examine taxonomic richness, composition and trophic groups of macroinvertebrate communities in lakes located at different heights (321–2899 m asl). We assumed that with altitude climatic conditions would become more severe, species richness of benthic invertebrates decrease, feeding selectivity of organisms remains the same, shredders and filter feeders replace predators and collectors-detritus feeders.

Materials and methods

In 2002, 2003, 2007, 2008, 2018 and 2020, bottom macroinvertebrate communities from the Russian Altai lakes located at various (low, medium, high) altitudes were studied within the framework of implemented complex limnological expeditions. We investigated 36 lakes, where collected and analyzed 108 quantitative and 28 qualitative samples of zoobenthos. The analyzed material was selected and processed using the standard methods (Guidelines 1992; Wetzel, Likens 2000). Qualitative samples were taken with a water net or scraper, while quantitative ones were taken with a GR-91 bottom grab (70 cm2 mouth area). Bottom samples (boulders and pebbles) were collected using a hydrobiological net (with subsequent calculation of the area of stones by their projection in a plane), then washed through a nylon gauze with a mesh size of 350х350 μm. The animals were isolated and fixed in 70% ethanol. When taken two to three times, quantitative samples were combined into an integrated sample. For a more complete accounting of the zoobenthos composition, the samples were collected manually in various biotopes. The soil taken by a bar dredge was washed through a kapron bag with a 320-μm mesh. The samples were examined portion-wise, and the organisms found were placed in test tubes with 70% ethyl alcohol. After drying on a filter paper, we weighed the organisms on a torsion balance. Each species was determined by the "Key to Freshwater Invertebrates of Russia and Adjacent Lands" (1992–2004).

The lakes studied are located at various heights, ie low mountain - 1–4, middle-mountains 5–26, high-mountains 27–36 (Fig. 1). For the Russian temperate zone of the northern hemisphere, the following altitudes have been accepted: low mountains – up to 1000 m, middle mountains – 1000–2000 m, highlands – more than 2000 m (Gvozdetsky 1977). The area of the lakes studied is within 0.01–4.52 km2, most of them do not exceed 1 km2 (Table 1). These lakes are characterized by low water salinity and low concentrations of macro and microcomponents. The hydrogen index corresponds to neutral or slightly acidic waters (pH = 5.8–8.6). Lake waters are pure and oxygen-saturated (Frolova et al. 2011; Zarubina and Fetter 2019, 2020). As compared to the data of the 30s – 40s of the XX century, the dissolved oxygen concentration, pH and salinity of water have changed insignificantly (Alekin 1935; Gundrizer 1950; Johansen 1950).

A detailed description of the lakes studied is given in our previous work (Yanygina and Krylova 2006, 2008; Vdovina and Bezmaternykh 2020). The affiliation of macroinvertebrates with a certain trophic group was defined according to the Moog O. and Hartmann A. classification (2017), which is a revised version of the Cummins classification (1973) for insects (Cummins et al. 2008). Dominant species were identified on frequency of their occurrence (Bakanov, 1987). Discriminant analysis was performed to reveal the composition of differences in the macroinvertebrate species observed for low, mid and high mountain lakes. Analysis of variance (one-way ANOVA) was employed in assessing the impact of various factors (lake size, substrate type, altitude, altitudinal zone) on relative abundance of macrozoobenthos trophic groups.

Figure 1.Map of the lakes studied. Low mountain lakes (<1000 meters above sea level): 1. Aya; 2. Manzherokskoe; 3. Kolyvanskoye; 4. Beloye middle mountain lakes (1000–2000 m): 5. Verkhny Itykul'; 6. Nizhny Itykul'; 7. Saygonysh; 8. Maly Saygonysh; 9. Kubyshka; 10. Arsoyok; 11. El'dengem; 12. Tugunrluachekkol'; 13. Maldu; 14. Tashtu; 15. Unnamed lake in the Kayry River basin; 16. Sundruk; 17. Ayukol‘; 18. Bezymyannoye; 19. Igistu-Kul'; 20. Maly Uzenkol'; 21. Pridorozhnoye; 22. Podkova; 23. Verkhnee Mul'tinskoye; 24. Srednee Mul'tinskoye; 25. Nizhnee Mul'tinskoye; 26. Poperechnoye High-mountain lakes (> 2000 m): 27. Yakhansoru; 28. Unnamed lake near Uzunkel'; 29. Kandash; 30. Sostukel'; 31. Argamdzhi; 32. Bol'shoye Tarkhatinskoye; 33. Zerlyukol'-Nur; 34. Krasnoye; 35. Maloye Tarkhatinskoye; 36. T’eply klyuch.

Lakes Coordinates Altitude, m asl Altitudinal belt Area, km2 Substrate
1 Aya 51°54'14"N 85°51'12"E 321 Premontane subtaiga 0.06 Detritus, gray silt
2 Manzherok 51°49'16"N 85°48'45"E 373 Premontane subtaiga 0.33 Muddy with detritus
3 Kolyvanskoye 51°21'56"N 82°11'38"E 332 Premontane steppe 4.52 Silt and sand
4 Beloye 51°17'40"N 82°38'47"E 537 Premontane forest-steppe 2.83 Silt and pebbles, stones
5 Verkhny Itykul' 51°22'32"N 88°46'57"E 1664 Mountain-taiga 0.99 Stones, sand
6 Nizhny Itykul' 51°20'57"N 88°48'37"E 1661 Mountain-taiga 2.77 Stones, sand
7 Saygonysh 51°13'37"N 88°27'39"E 1612 Mountain-taiga 0.81 Silted sand
8 Maly Saygonysh 51°13'34"N 88°27'17"E 1627 Mountain-taiga 0.02 Silted stones, detritus
9 Kubyshka 51°13'44"N 88°25'54"E 1728 Mountain-taiga 0.02 Macrophytes, detritus
10 Arsoyok 51°12'32"N 88°22'2"E 1473 Mountain-taiga 0.17 Stones, detritus, sand
11 El'dengem 51°15'58"N 88°21'46"E 1736 Mountain-taiga 0.06 Silt, detritus
12 Tugunrluachekkol' 51°17'25"N 88°17'08"E 1688 Mountain-taiga 0.05 Stones
13 Maldu 51°16'30"N 88°10'42"E 1649 Mountain-taiga 0.02 Stones
14 Tashtu 51°16'02"N 88°07'06"E 1765 Mountain-taiga 0.11 Stones
15 unnamed lake in river basin Kayry river 51°16'03"N 88°02'50"E 2082 Podgolets-subalpine 0.01 Silted sand, pebbles
16 Sundruk 51°12'33"N 87°06'51"E 1990 Podgolets-subalpine 0.37 Sand, pebbles, stones
17 Ayukol‘ 51°16'347"N 87°56'51"E 2015 Podgolets-subalpine 0.06 Stones
18 Bezymyannoye 50°28'00"N 87°42'35"E 2192 Podgolets-subalpine 0.01 Silt
19 Igistu-Kul' 50°30'53"N 87°40'22" 1823 Mountain-taiga 0.66 Boulders
20 Maly Uzenkol' 50°28'15"N 87°36'37"E 1991 Mountain-taiga 0.24 Boulders
21 Pridorozhnoye 50°24'32"N 87°35'60"E 1838 Mountain-taiga 0.01 White silt over boulders and pebbles
22 Podkova 50°27'38"N 87°37'00"E 1971 Mountain-taiga 0.26 Silt
23 Verhnee Mul'tinskoye 49°55'03"N 85°50'42"E 1797 Podgolets-subalpine 0.39 stones
24 Srednee Mul'tinskoye 49°59'00"N 85°49'47"E 1646 Mountain-taiga 0.92 Stones
25 Nizhnee Mul'tinskoye 50°00'11"N 85°49'50"E 1627 Mountain-taiga 1.73 Stones
26 Poperechnoye 49°55'27"N 85°53'25"E 1883 Podgolets-subalpine 0.43 Stones
27 Yakhansoru 51°06'49"N 88°53'33"E 1949 Podgolets-subalpine 0.52 Stones
28 unnamed lake near Uzunkel' 51°03'20"N 88°37'21"E 1957 Podgolets-subalpine 1.09 Stones
29 Kandash 51°22'35"N 88°08'40"E 1945 Podgolets-subalpine 0.07 Stones
30 Sostukel' 51°03'01"N 88°54'29"E 2023 Podgolets-subalpine 0.22 Stones
31 Argamdzhi 49°19'03"N 87°55'31"E 2376 Montane tundra-steppe 0.09 Black silt with detritus
32 Bol'shoye Tarkhatinskoye 49°34'14"N 88°23'02"E 2320 Montane tundra-steppe 0.48 Silt with detritus
33 Zerlyukol'-Nur 49°34'32"N 88°23'19"E 2321 Montane tundra-steppe 1.61 Silt
34 Krasnoye 49°24'24"N 88°02'18"E 2329 Montane tundra-steppe 0.24 Boulders
35 Maloye Tarkhatinskoye 49°17'49"N 88°01'38"E 2333 Montane tundra-steppe 0.05 Silt, sand, pebbles
36 Teply klyuch 49°24'24"N 88°02'18"E 2899 Golets-alpine 0.03 Silted fine gravel
Table 1.Main features of the studied lakes


Taxonomic composition. In low mountain lakes, 103 species of macrozoobenthos of seven classes were recorded: Oligochaeta (15 species), Hirudinea (4), Phylactolaemata (1), Gastropoda (5), Acari (11), Crustacea (1) and Insecta (66) (Table 2). Among insects, the maximum species richness fell on Diptera (37 species, of which 30 are chironomids). The rest of the species represented dragonflies, mayflies, true bugs, caddisflies, and beetles. Tubificidae (Limnodrilus hoffmeisteri – 66%) and Chironomidae (60%) showed the highest frequency of occurrence. The genera Cricotopus (33%), Procladius (33%), Chironomus (26%) and Ablabesmyia (26%) were the most often detected chironomids. In low-mountain lakes, the species richness of the zoobenthos was relatively high (8.5±2.3 species per sample), the Shannon species diversity index reached on average 1.7±0.3 bits/ind.

Macrozoobenthos of middle mountain lakes included 94 species of 10 classes: Demospongiae (1 species), Turbellaria (1), Nematoda (1), Oligochaeta (9), Hirudinea (4), Bivalvia (2), Gastropoda (5), Acari (2), Crustacea (4), and Insecta (65). Amphibiotic insects accounted for 69% of the identified taxa, most of them (39 species) belong to the Diptera. Mayflies, beetles, caddisflies, stoneflies, and net-winged insects were also present. In Diptera, chironomid larvae (36 species) prevailed, mainly from the subfamily Orthocladiinae. Larvae of the genera Cricotopus (28%), Synorthocladius (25%) and Eukiefferiella (23%) were the most common. In other taxa, the Gammaridae family made up 40% and Naididae – 30%. The species richness of benthic invertebrates was low, i.e., 0-11 species in a sample (on average 5.1±0.3), the Shannon diversity index varied within 0–2.85 (on average 1.4±0.1 bits/ind.).

In high mountain lakes, a total of 36 species of macrozoobenthos were recorded of 7 classes (Table 2): Demospongiae (1 species), Nematoda (1), Oligochaeta (2), Hirudinea (2) Bivalvia (2), Crustacea (2), and Insecta (26). Among insects, dipterans demonstrated the highest species richness (17 species, chironomids). Beetles, mayflies, caddisflies, and true bugs were also found. Chironomids were observed in 86% of the samples, among them Chironomus (44%) and Stictochironomus (27%), as well as the Cladotanytarsus A (27%) predominated. For other taxa, crustaceans of genus Gammarus prevailed (50%). Taxonomically, the benthic communities in the studied lakes were not rich (on average 4.4±0.6 species per sample); the Shannon diversity index varied from 0 to 3.04 (on average 1.11±0.2 bits/ind.).

Taxon Low-mountains Middle-mountains High-mountains
Phylum Porifera
Classis Demospongiae
Familia Spongillidae
Spongillidae indet. 19, 21, 22, 25 25
Phylum Plathelminthes
Classis Turbellaria
Turbellaria indet. 24, 26
Phylum Nemathelminthes
Classis Nematoda
Mermithidae indet. 18, 22 31, 33
Phylum Annelida
Classis Oligochaeta
Familia Lumbriculidae
Lumbriculus variegatus (O.F. Müller) 10, 12, 23, 24
Familia Naididae
Chaetogaster sp. 20
Chaetogaster diaphanus (Gruithuisen) 3 5
Dero sp. 3
Nais barbata O.F. Müller 3, 4
Nais bretscheri Michaelsen 4
Nais communis Piguet 3, 4
Nais pardalis Piguet 3, 4
Nais pseudobtusa Piguet 4
Nais sp. 23, 25
Nais variabilis Piguet 5
Ophidonais serpentina (O.F. Müller) 3, 4
Ripistes parasita (Schmidt) 3
Stylaria lacustris (Linnaeus) 3, 4 20
Uncinais uncinata (Oersted) 3, 4
Familia Tubificidae
Limnodrilus claparedeanus Ratzel 3, 4
Limnodrilus hoffmeisteri Claparède 2, 3, 4 25 34
Tubifex tubifex (O.F. Müller) 3, 4 7 34, 36
Spirosperma ferox (Eisen) 3 10, 15
Classis Hirudinea
Erpobdella octoculata (L.) 1, 3 5, 10, 20, 21, 22 28
Erpobdella sp. 8, 16
Glossiphonia complanata (L.) 3, 4 5, 6, 22 30, 32
Helobdella stagnalis (L.) 1, 3 12
Hemiclepsis marginata (O.F. Müller) 3
Phylum Bryozoa
Classis Phylactolaemata
Plumatella repens (L.) 1
Phylum Mollusca
Classis Bivalvia
Euglesa sp. 7, 8, 9, 10, 11, 15
Sphaerium corneum (L.) 18 35
Sphaerium sp. 32, 33, 34, 35
Classis Gastropoda
Familia Planorbidae
Anisus acronicus (Férussac) 24
Anisus sp. 3, 4
Armiger sp. 4
Hippeutis euphaea (Bourguignat) 3
Familia Lymnaeidae
Lymnaea fontinalis (Studer) 3 24
Lymnaea ovata (Draparnaud) 3
Lymnaea stagnalis (L.) 19, 22
Lymnaea sp. 23
Familia Physidae
Physa sp. 25
Phylum Arthropoda
Classis Euchelicerata
Familia Arrenuridae
Arrenurus sinuator (Müller) 4
Arrenurus sp. 3
Familia Hydrachnidae
Hydrachna sp. 3, 4
Familia Lebertiidae
Lebertia sp. 14
Familia Mediopsidae
Mediopsis orbicularis (Müller) 4
Familia Pionidae
Forelia sp. 3
Piona coccinea Koch 3
Piona pusila (Neuman) 3
Piona sp. 4 10, 23, 24
Tiphys sp. 4
Familia Sperchontidae
Sperchon sp. 3
Familia Unionicolidae
Neumania sp. 4
Classis Crustacea
Familia Gammaridae
Gammarus c.f. barnaulensis Schellenberg 6, 8, 9, 11, 12, 14
Gammarus lacustris G.O. Sars 3 10, 13, 19, 21, 22, 24, 26 27, 32, 34, 35
Gammarus korbuensis Martynov 16
Gammarus sp. 5, 7 29
Classis Insecta
Ordo Neuroptera
Familia Sisyridae
Sisyra fuscata (F.) 9
Ordo Odonata
Familia Aeshnidae
Aeschna culumberculus Harris 3
Familia Coenagrionidae
Erythromma najas (Hansemann) 1
Erythroma sp. 3
Coenagrionidae indet. 4
Familia Corduliidae
Somatochlora sp. 2
Familia Gomphidae
Gomphidae indet. 3
Familia Lestidae
Sympecma fusca (Vanderlinden) 4
Sympecma paedisca (Brauer) 3
Ordo Plecoptera
Familia Nemouridae
Nemoura sp. 6
Familia Chloroperlidae
Chloroperlidae indet. 26
Ordo Ephemeroptera
Familia Ameletidae
Ameletus sp. 29
Familia Baetidae
Baetis sp. 5
Baitis gr. vernus 13
Baitis gr. rhodani 26
Cloeon dipterum L. 4 20
Familia Caenidae
Caenis lactea (Burmeister) 22
Caenis miliaria (Tshernova) 22
Caenis horaria L. 3, 4
Caenis sp. 1
Familia Ephemerellidae
Ephemerella (T.) lenoki Tshernova 24
Familia Haptageniidae
Ecdyonurus (A.) vicinus (Demoulin) 5
Ecdyonurus sp. 3 25
Heptagenia sp. 24, 25
Familia Leptophlebiidae
Leptophlebia (P.) strandii Eaton 11, 12, 24
Leptophlebia (N.) chocolate (Imanishi) 26
Ordo Heteroptera
Familia Corixidae
Corixa sp. 3
Familia Gerridae
Gerris lacustris (L.) 27
Familia Nepidae
Nepa cinerea L. 1
Familia Naucoridae
Ilyocoris cimicoides (L.) 3
Familia Pleidae
Plea minutissima Leach 3, 4
Ordo Trichoptera
Familia Apataniidae
Apatania zonella Zett 5
Apatania sp. 26
Apataniidae indet. 26
Familia Hydroptilidae
Agraylea multipunctata Curtis 4
Agrypnia pagetana Curt. 3
Agraylea sexmaculata Curtis 4
Oxyithira costalis Curt. 4
Familia Leptoceridae
Oecetis sp. 10, 20
Familia Limnephilidae
Limnephilus borealis Zett 27, 28
Limnephilus nigriceps Zett 27
Limnephilus rhombicus (L.) 24, 25
Limnephilus stigma Curtis 27
Familia Molannidae
Molanna albicans (Zetterstedt) 16
Familia Phryganeidae
Agrypnia obsoleta (Hagen) 14 28
Phryganea bipunctata Retzius 3
Familia Rhyacophilidae
Rhyacophila sp. 17
Ordo Coleoptera
Familia Chrysomelidae
Donacia sp. 9
Calerucella sp. 3
Chrysomelidae indet. 3
Plateumaris sp. 3
Prasocuris sp. 4
Familia Dytiscidae
Acilius canaliculatus (Nicolai) 27
Agabus sp. 27, 29
Dytiscus circumflexus Fabricius 29
Graphoderus sp. 2
Hydrotus sp. 4
Hygrotus sp. 8
Oreodytes sp. 24
Familia Hydraenidae
Hydraena sp. 3
Familia Hydrophilidae
Hydrophilidae indet. 1
Ordo Diptera
Diptera indet. 9
Familia Dixidae
Dixella aestivalis (Meigen) 9
Familia Chaoboridae
Chaoborus (C.) crystallinus (De Geer) 3
Chaoborus (C.) flavicans (Meigen) 2
Familia Simuliidae
Simulium sp. 23
Familia Ceratopogonidae
Bezzia (H.) bicolor (Meigen) 3, 4
Bezzia sp. 1
Mallochoholea setigera (Loew) 4
Palpomyia lineata (Meigen) 2, 3
Sphaeromias pictus (Meigen) 2
Familia Chironomidae
Ablabesmyia gr. monilis 3, 4 6, 7, 13, 17, 24, 25 27, 28
Ablabesmyia sp. 24 34
Chironomus plumosus (L.) 2, 3, 4
Chironomus sp. 3 15, 18, 20, 21, 22, 23, 25 32, 35
Cladotanytarsus mancus (Walker) 3, 4 20 32, 36
Cladotanytarsus gr. А 22 33, 34
Cladotanytarsus sp. 25
Corynoneura gr. edwarsi 3
Corynoneura scutellata 5, 14, 25
Corynoneura sp. 20
Cricotopus gr. laricomaris 30
Cricotopus sylvestris (Fabricius) 3, 4 24
Cricotopus tibialis (Meigen) 2
Cricotopus gr. tremulus 12, 24, 25
Cricotopus sp. 4 5, 6, 13, 14, 17, 20, 25 34
Cryptochironomus defectus (Kieffer) 3, 4 25 35
Diamesa bertrami Edwards 23
Diplocladius cultriger Kieffer 23
Dicrotendipes nervosus (Staeger) 4 25 34
Dicrotendipes setemmaculatus (Becker) 4
Endochironomus albipennis (Meigen) 1, 3
Endochironomus donatoris Shilova 3
Endochironomus stackelbergi Goetghebuer 4 5 34
Endochironomus tendens (Fabricius) 3
Eukiefferiella gr. devonica 23, 24, 25
Eukiefferiella sp. 23
Glyptotendipes glaucus (Meigen) 1, 2, 3, 4 9, 20
Glyptotendipes paripes (Edwards) 16
Hydrobaenus lugubris (Fries) 35
Microchironomus tener (Kieffer) 3
Micropsectra sp. 25
Orthocladius sp. 22, 24
Parachironomus varus Goethgebuer 3, 4
Paratanytarsus confusus Palmen 3, 4 23
Paratanytarsus sp. 12, 13, 14, 17
Polypedilum bicrenatum Kieffer 3
Polypedilum convictum (Walker) 3
Polypedilum cf. litofiles Akhrorov 1
Polypedilum nubeculosum (Meigen) 22
Polypedilum scalaenum (Schrank) 31
Polypedilum tetracrenatum Hirvenoja 3, 4
Procladius (H.) ferrugineus (Kieffer) 1, 2, 3, 4
Procladius (H.) choreus Meigen 3, 4 8, 11, 15
Procladius sp. 18, 19, 22 34, 36
Prodiamesa olivacea (Meigen) 23
Psectrocladius delatoris Zelentzov 9 28
Psectrocladius obvius (Walker) 4
Psectrocladius sp. 23, 25
Psecrocladius (P.) zetterstedti Brundin 22
Sergentia gr. coracina 16
Synorthocladius semivirens (Kieffer) 6, 10, 23, 24, 25, 26
Stictochironomus crassiforceps (Kieffer) 4 28, 36
Stictochironomus gr. histrio 35
Tanypus punctipennis Meigen 3, 4
Tanypus sp. 4 23, 24 36
Tanytarsus medius Reiss et Fittkau 23
Tanytarsus sp. 23, 24
Chironomidae pupae 4
Total species 103 94 36
Table 2.Taxonomic composition of macrozoobenthos. Note: 1–36, habitat numbers corresponding to Fig. 1; –, no record.

Discriminant analysis (Fig. 2) suggested significant differences in the macroinvertebrate structure of lakes located in different altitudinal zones (Wilks’ λ = 0.01, F=10.01, p<0.0001). In the study groups, the families Tubificidae (Wilks’ λ = 0.03; F=15.80; p=0.0001), Euglesidae (Wilks’ λ =0.03; F=16.25; p=0.0001), Caenidae (Wilks’ λ = 0.024; F=11.17; p=0.0006) and the subfamily Chironominae (Wilks’ λ =0.03; F=16.25; p=0.0001) made the greatest contribution to lakes’ difference (Wilks’ λ =0.025; F=11.66; p=0.0005).

Figure 2.The discriminant analysis of the macroinvertebrate composition of the Russian Altai lakes located in different altitudinal zones. 1 – low-mountains (<1000 meters above sea level), 2 – medium mountain ranges (1000–2000 m), 3 – high mountain ranges (> 2000 m).

Trophic groups. Based on the type of feeding, five major trophic groups of macroinvertebrates from the study lakes were identified as follows: (1) Collectors-detritus feeders, facultative filter feeders (hereinafter – Collectors-detritus feeders); (2) Collectors-obligate filter feeders; (3) Scrapers; (4) Shredders; (5) Predators.

Here, the greatest species richness falls on detritus collectors eating detritus from the ground surface (Table 3). This group is represented by larvae of diptera of the family Chironomidae, mayflies, and oligochaetes. A large number of species belong to predators with predominance of leeches, flatworms, dragonfly larvae, bedbugs of the families Pleidae, Gerridae, Nepidae and Notonectidae, beetles of the family Dytiscidae, diptera larvae from the families Chaoboridae and Ceratopogonidae. In these communities, predatory larvae from caddisflies of the families Leptoceridae and Rhyacophilidae, chironomids of the subfamily Tanypodinae, and megalopterans of the family Sialidae were also identified. The collector-obligate filter feeder group was formed mainly by bivalve mollusks. Among shredders, the Gammaridae and beetles of the Chrysomelidae prevailed. It is hard to attribute the feeding spectrum of many species to a specific trophic group unambiguously; therefore, scrapers and some shredders include organisms with a mixed type of feeding in appropriate proportions (Moog and Hartmann 2017).

Trophic groups Lakes, N/M±m
Low-mountains Middle-mountains High-mountains
Collectors-detritus feeders 38.9/1.71±0.55 46/0.31±0.07 15.2/0.79±0.29
Shredders 8.4/0.19±0.18 10.9/1.23±0.37 2.7/0.37±0.17
Scrapers 7.4/0.13±0.07 11.4/0.22±0.1 1.9/0.04±0.02
Predators 39.3/0.54±0.31 17.6/0.29±0.09 12.3/0.16±0.06
Collectors-obligate filter feeders 3/0.09±0.05 6.1/0.16±0.07 3.9/0.55±0.25
Others 6/0.06±0.03 2/0.06±0.05
Table 3.Trophic groups of macrozoobenthos in lakes from different altitudinal zones. Note: N – number of species; M±m – mean and error of mean biomass of trophic groups, g/m2 – no records.

By the biomass of zoobenthos, collectors-detritus feeders andpredators dominated in low mountain lakes. In midmountain lakes, shredders (represented by the Gammaridae) dominated in biomass, collectors-detritus feeders and predators subdominated, the remaining groups were represented equally. In high mountain lakes, the role of shredders decreased, collector-detritus feeders were again dominated by biomass, and the role of collector-obligate filter feeders increased as well.

ANOVA revealed that the distribution of the lakes trophic groups in the studied depends on height, altitudinal zone, and substrate type (Table 4). The size of the lake is less important; this indicator significantly affects only the proportion of scrapers in benthic biomass. The proportion of collectors-detritus feeders and shredders depends on height, altitudinal zone, and soil type, whereas scrapers – on lake size (p=0.011), height (p <0.001) and substrate (p=0.001). The proportion of collector-obligate filter feeders is influenced solely by substrate type (p=0.024), while predators are influenced by the altitudinal zone (p <0.001).

Variable Effect R F p
Collectors-detritus feeders altitudinal zones 0.51 6.67 ≤0.001
lake area 0.02 0.04 0.848
altitude 0.47 10.98 ≤0.001
substrate 0.47 11.06 ≤0.001
Collectors-obligate filter feeders altitudinal zones 0.22 0.94 0.444
lake area 0.12 1.14 0.288
altitude 0.25 2.61 0.080
substrate 0.31 3.90 0.024
Scrapers altitudinal zones 0.26 1.52 0.206
lake area 0.36 11.19 0.001
altitude 0.42 8.36 ≤0.001
substrate 0.33 4.71 0.011
Shredders altitudinal zones 0.52 6.77 ≤0.001
lake area 0.04 0.11 0.746
altitude 0.44 9.24 ≤0.001
substrate 0.35 5.26 0.007
Predators altitudinal zones 0.42 3.93 ≤0.001
lake area 0.01 0.02 0.900
altitude 0.17 1.19 0.309
substrate 0.08 0.23 0.795
Table 4.One-way analysis of the variance of functional feeding group of lakes in different altitudinal zones of the Russian Altai (figures in italic indicate significance at p<0.05)


The taxonomic structure of the studied lakes is characterized by the dominance of chironomids and oligochaetes that is typical for the mountain lakes (Manca et al. 1998; Krno et al. 2006; Boggero and Lencioni 2006; Loskutova 2011). The predominance of various subfamilies of chironomids and families of oligochaetes was observed at different altitudes. In low mountain lakes, among oligochaetes – Tubificidae (66%), while among chironomids - Chironominae (60%) and Tanypodinae (50%) were detected most frequently, which is typical for lowland lakes of western Siberia (Bezmaternykh and Vdovina 2020). In mid-mountain lakes, the number of macroinvertebrate taxa increased with height. Here, Orthocladiinae were more common among chironomids (58%) and Naididae – among oligochaetes (30%). The predominance of species of the subfamily Orthocladiinae in the chironomid fauna is typical for the mountain lakes (Fureder et al. 2006). Above 2000 m, the number of species and taxa diversityof benthic communities declined. In zoobenthos, chironomids of the subfamily Chironominae (73%) predominated. Oligochaetes of the Naididae family fell out of the dominant complex. The same phenomenon was marked in alpine lakes of the Pyrenees (Collado and de Mendoza 2009; de Mendoza and Catalan 2010). A peculiar feature of bottom zoocenoses from the mid- and high-mountain lakes is the high frequency of gammarid occurrence (40 and 50%, respectively) as well as their large contribution to the total biomass that is also characteristic of the mountain and alpine lakes of the Altai-Sayan mountain country (Vershinin 1979; Lepneva 1933). Among other peculiarities of the taxonomic composition of the bottom communities is the low diversity of mollusks that reside in these and other lakes adjacent to Lake Teletskoye. This indicator is associated with an unfavorable slightly acid hydrochemical composition of waters (Johansen 1950).

As for the species composition of macroinvertebrate communities, the group of collectors-detritus feeders is most abundant, which is confirmed by other researchers (Baturina et al. 2014; Kurashov 1994; Yakovlev 2000, 2005; Timm and Mols 2005); predators are in the second position by species number in all lakes. The ratio of trophic groups by biomass varies significantly. If in low-mountain lakes the dominant collectors-detritus feeders account for more than 50% of macroinvertebrate biomass, above 1000 to 2000 m asl they (together with predators) take the second place,thus giving the pas to shredders. At high altitudes, collector-detritus feeders dominate, collectors-obligate filter feeders join them as a subdominant group, and shredders move to the third position. The cluster analysis suggests the best similarity of the trophic structure of macrozoobenthos in mid- and high-mountain lakes, and the least one for low-mountain lakes (Figure 3). With the transition from low- to mid-mountain lakes, an increase in average biomass of shredders and scrapers is observed. It is also true for the southwestern Scandinavian watercourses located at different heights (Brönmark et al. 1984). This is most likely due to the fact that allochthonous organic matter predominates in mountain reservoirs and trophic groups focus on the consumption of certain food resources. Shredders are actively involved in disposal of large plant residues being the detritus suppliers to filter feeders, scrapers and collectors-detritus feeders (Cummins 1973; Wallace et al. 1996). With height, the average biomass of the filter feeders also increases, although this fact has not been confirmed by any reliable dependencies. This distribution of filter feeders is explained by the type of substrate in lakes. In lowland reservoirs, the soils are mainly represented by silts and water contains a lot of suspended inorganic substances. With height, a rocky substrate prevails, and the amount of suspended matter decreases that facilitates the feeding of filter feeders.

Our main hypothesis that with an altitude increase climatic conditions become more severe and species richness of macroinvertebrates declines has been confirmed. Similar trends for some lakes and rivers that cover wide altitudinal gradients also reveal an almost linear decrease in local macroinvertebrate taxa with increasing altitude. This pattern is true for alpine lakes (Fjellheim et al. 2000), lakes of the Perineas(de Mendoza and Catalan 2010), alpine rivers (Jacobsen et al. 2020), rivers of Nepal (Suren 1994), Ecuador (Jacobsen 2004) and Colorado (Harrington et al. 2015). The decrease in species diversity with increasing altitude depends on many factors and is primarily associated with changing environmental conditions. The species distribution response to environmental changes is mainly determined by their ability to settle and survive under suitable conditions (Lavergne et al. 2010). In mountain lakes, only extreme temperature fluctuations are responsible for the reduction in species richness (Fureder et al. 2006; Havens et al. 2015). As a result, species with short life cycles appear (Belle and Goedkoop 2020). It is apparent that organic matter content and aquatic vegetation are essential drivers of changes in species diversity along the altitude gradient (Hoffman et al. 1996; Nyman et al. 2005; Bigler et al. 2006; Kernan et al. 2009; Collado and de Mendoza 2009). As is well known, the ability to use available food resources is of great importance to organisms. Therefore, eurybiont species with their small body size and broader food preferences are more likely to find suitable habitats in extreme conditions (Angert et al. 2011). It should be noted that, a decrease in species diversity may be induced by various environmental factors like snow melting rates (Obertegger et al. 2007), short summers, intense ultraviolet radiation (Hansson et al. 2007; Miller and McKnight 2015), water level fluctuations, substrate type (Bretschko 1995; de Mendoza and Catalan 2010), etc. The altitude gradient is a set of interrelated environmental factors with different impacts on the composition and structure of hydrobionts communities; distinguishing the influence of individual variables is hardly possible.

Figure 3.The cluster analysis of the trophic spectrum and species composition in study lakes.

Therefore, the taxonomic and trophic structures of the lakes zoobenthos in the studied lakes are differed, while the structure of zoobenthos in low-mountain and lowland lakes is similar. With height, the taxonomic structure becomes more complicated, and the dominant taxa change. The fauna of mid- and high-mountain lakesis specific due to the presence of homotopic species of the family Gammaridae and low diversity of molluscs. In the lakes under study, we have identified five major trophic groups of benthic invertebrates. By species number, collectors-detritus feeders and filter feedersdominated in trophic structure. By biomass, the trophic groups of various altitude lakes were represented differently; the proportion of filter feeders and shredders increased. Recent studies have paid great attention to the effect of the altitude gradient on benthic invertebrate communities. The study of this factor in remote areas of the Russian Altai characterized by low anthropogenic loads makes it possible to determine major trends in natural dynamics of communities, to use these lakes as the reference areas indicating the environmental changes and to develop the programs for monitoring of the transformed lake ecosystems.


The authors thank the researchers from IWEP SB RAS and Dr. N.I. Ermolaeva in person for her assistance with field trip arrangements and sampling.

This work has been supported by the Russian Science Foundation, grant 22-27-20134 (


Al'ekin OA (1935) Lakes of the Katun' Alps. Study of lakes in the USSR. Leningrad, 153-235. (in Russian)

Angert AL, Crozier LG, Rissler LJ, Gilman SE, Tewksbury JJ, Chunco AJ (2011) Do species' traits predict recent shifts at expanding range edges? Ecol Lett 14(7): 677-689.

Bakanov AI (1987) Quantitative assessment of dominance in ecological communities. Moscow, 64 pp. (in Russian)

Baturina MA, Loskutova OA, Shchanov VM (2014) Structure and distribution of zoobenthos of the Kharbey Lake System. J Sib Fed Univ Biol 7: 332-356.

Belle S, Goedkoop W (2020) Functional diversity of chironomid communities in subarctic lakes across gradients in temperature and catchment characteristics. Limnology 22(7).

Bezmaternykh DM, Vdovina ON (2020) Composition and structure of macrozoobenthos of lakes in different natural zones and subzones of Western Siberia. Limnology 21: 3-13.

Bigler C, Heiri O, Krskova R, Lotter AF, Sturm M (2006) Distribution of diatoms, chironomids, and cladocera in surface sediments of thirty mountain lakes in south-eastern Switzerland. Aquatic Sciences 68: 154-171.

Bitusik P, Svitok M, Kolosta P, Hubkova M (2006) Classification of the Tatra Mountain lakes (Slovakia) using chironomids (Diptera, Chironomidae). Biologia, Bratislava 61: 191-201.

Boggero A, Lencioni V (2006) Macroinvertebrates assemblages of high altitude lakes, inlets and outlets in the southern Alps. Archiv fur Hydrobiologie 165: 37-61.

Bretschko G (1995) Opportunities for high alpine research, the lake Vorderer Finstertaler See as an example (Kühtai, Tirol, 2237 m a.s.l.). Limnologica 25: 105-108.

Brittain JE, Milner AM (2001) Special issue: glacier-fed rivers - unique lotic ecosystems. Freshwater Biology 46: 1571-1874.

Brönmark Ch, Herrmann J, Maimgvist B, Otto Ch, Sjöstrom P (1984) Animal community structure as a function of stream size. Hydrobiologia 112: 73-79.

Collado R, de Mendoza G (2009) Environmental factors and distribution of littoral oligochaetes in Pyrenean lakes. Advances in Limnology 62: 215-244.

Cummins KW (1973) Trophic relation in aquatic insects. Ann. Rev. Entomol 8:183-206.

Cummins KW, Merritt RW, Berg MB (2008) Ecology and distribution of aquatic insects: An introduction to the aquatic insects of north America. Kendall Hunt Publication, Dubuque, 105-122.

de Mendoza G, Catalan J (2010) Lake macroinvertebrates and the altitudinal environmental gradient in the Pyrenees. Hydrobiologia 648: 51-72.

Dumnicka E, Galas J (2002) Factors affecting the distribution of Oligochaeta in small high mountain ponds (Tatra Mts., Poland). Archiv fur Hydrobiologie 156: 121-133.

Fjellheim A, Boggero A, Halvorsen GA, Nocentini AM, Rieradevall M, Raddum GG, Schnell ØA (2000) Distribution of benthic invertebrates in relation to environ mental factors. A study of European remote alpine lake ecosystems. Verhandlugen der Internationale Vereinigung fur Theoretische und Angewandte Limnologie 27: 484-488.

Fjellheim A, Raddum GG, Vandvik V, Lniceanu C, Boggero A, Brancelj A, Stuchlik E (2009) Diversity and distribution patterns of benthic invertebrates along alpine gradients. A study of remote European freshwater lakes. Advances in Limnology 62: 167-190.

Frolova NL, Povalishnikova EU, Efimova LE (2011) Complex studies of water bodies in the Altai Mountains (the Multy river basin as a case study) - 75 years later. Bulletin of RAS. Geographical Series 2. 113-126. (in Russian)

Fureder L, Ettinger R, Boggero A, Thaler B, Thies H (2006) Macroinvertebrate diversity in alpine lakes: effects of altitude and catchment properties. Hydrobiologia 562: 123-144.

Guidelines for hydrobiological monitoring of freshwater ecosystems (1992) Hydrometeoizdat. St. Petersburg, 318 pp. (in Russian)

Gundrizer AN, Ioganzen BG, Kafanova VV, Petlina AP (1982) Ichthyology and hydrobiology in Western Siberia. Tomsk, TGU, 318 pp. (in Russian)

Gvozdetsky NA (1977) Some ideas on possible systematic research development in physical geography. Geogr. Iss. 104: 7-13. (in Russian)

Harrington RA, Poff NL, Kondratieff BC (2015) Aquatic insect β-diversity is not dependent on elevation in Southern Rocky Mountain streams. Freshw. Biol. 61: 195-205.

Havens KE, Pinto-Coelho RM, Beklioğlu M, Christofersen KS, Jeppesen E, Lauridsen TL, Erdoğan Ş (2015) Temperature efects on body size of freshwater crustacean zooplankton from Greenland to the tropics. Hydrobiologia 743(1): 27-35.

Hoffman RL, Liss WJ, Larson GL, Deimling EK, Lomnicky GA (1996) Distribution of nearshore macroinvertebrates in lakes of the Northern Cascade Mountains, Washington, USA. Archiv fur Hydrobiologie 136: 363-389.

Ioganzen B. G. Hydrobiological studies of water bodies of Western Siberia. Essays on the history of hydrobiological research in the USSR. Proc. of the Russian Hydrobiological Society 24: 171-184. (in Russian)

Ioganzen BG (1950a) On typology of reservoirs in the Chulcha river basin and fish breeding possibilities. Studies of Siberian Reservoirs 111: 143-150. (in Russian)

Ioganzen BG (1950б) Freshwater mollusks of the Chulcha river basin. Studies of Siberian Reservoirs 111: 137-142. (in Russian)

Jacobsen D (2004) Contrasting patterns in local and zonal family richness of stream invertebrates along an Andean altitudinal gradient. Freshw. Biol. 49: 1293-1305.

Jacobsen D, Wiberg-Larsen P, Brodersen K, Hansen S, Lindegaard C, Friberg N, Dall P, Kirkegaard J, Skriverf J, Toman M (2020) Macroinvertebrate communities along the main stem and tributaries of a pre-Alpine River: composition responds to altitude, richness does not. Limnology 84: 1-8.

Kernan M, Ventura M, Bitusik P, Brancelj A, Clarke G, Velle G, Raddum GG, Stuchlık E, Catalan J (2009) Regionalisation of European mountain lake ecosystems according to their biota: environmental versus geographical patterns. Freshwater Biology 54: 2470-249.

Key to freshwater invertebrates of Russia and adjacent lands (1992-2004): Vol 1-6. Zoological institute of RAS, St. Petersburg.

Koveshnikov MI (2014) Zoobenthos of water bodies of the Biya river basin. Saaabrücken: LAP LAMBERT Academic Publishing, 278 pp. (in Russian)

Krno I, Sporka F, Galas J, Hamerlık L, Zatovicova S, Bitusik P (2006) Littoral benthic macroinvertebrates of mountain lakes in the Tatra Mountains (Slovakia, Poland). Biologia, Bratislava 61 (Suppl. 18): 147-166.

Krylova EN (2016) Oligochaetes of mountain lakes of the Altai Republic. In: Biodiversity, environmental problems of the Altai Mountains and adjacent regions: present, past, future: Proceed.IV Int. Sc. Conf. (Russia), September 2016. Gorno-Altaisk, 105-106. (in Russian)

Kurashov EA (1994) Meiobenthos as a component of lake system. St. Petersburg, Alga-fond, 223 pp.

Larocque I, Hall RI, Grahn E (2001) Chironomids as indicators of climate change: a 100-lake training set from a subarctic region of northern Sweden (Lapland). Journal of Paleolimnology 26: 307-322.

Lavergne S, Mouquet N, Thuiller W, Ronce O (2010) Biodiversity and climate change: integrating evolutionary and ecological responses of species and communities. Annu Rev Ecology Evol S 41: 321-350.

Lepneva SG (1933) The bottom fauna of mountain lakes in the basin of Lake Teletskoye. Studies of Lakes of the USSR. Leningrad, 135-168. (in Russian)

Loria KA, McKnight D, Ragar DM, Johnson PT (2020) The life aquatic in high relief: shifts in the physical and biological characteristics of alpine lakes along an elevation gradient in the Rocky Mountains, USA. Aquatic Sciences 82: 11.

Loskutova OA (2011) Zoobenthos in different-type lakes at the western slope of the Circumpolar Urals (the Maly Patok river basin). Izvestiya of Samara Scientific Center, RAS. 1(5): 1124-1126. (in Russian)

Manca M, Ruggiu D, Panzani P, Asioli A, Mura G, Nocentini A (1998). Report on a collection of aquatic organisms from high mountain lakes in the Khumbu Valley (Nepalese Himalayas). Memorie dell'Istituto Italiano di Idrobiologia 57: 77-98.

McNaught AS, Schindler DW, Parker BR, Paul AJ, Anderson RS, Donald DB, Agbeti M (1999) Restoration of the food web of an alpine lake following fsh stocking. Limnol Oceanogr 44(1): 127-136.

Miller MP, McKnight DM (2015) Limnology of the Green Lakes Valley: phytoplankton ecology and dissolved organic matter biogeochemistry at a long-term ecological research site. Plant Ecol Divers 8(5-6): 689-702.

Moog O, Hartmann A (2017) Fauna Aquatica Austriaca, 3. Edition 2017. BMLFUW, Wien

Moser KA, Baron JS, Brahney J, Oleksy IA, Saros JE, Hundey EJ, Strecker AL et al (2019) Mountain lakes: eyes on global environmental change. Glob Planet Chang 178: 77-95.

Nyman M, Korhola A, Brooks S, (2005) The distribution and diversity of Chironomidae (Insecta: Diptera) in western Finnish lapland, with special emphasis on shallow lakes. Global Ecology and Biogeography 14: 137-153.

Obertegger U, Flaim G, Braioni MG, Sommaruga R, Corradini F, Borsato A (2007) Water residence time as a driving force of zooplankton structure and succession. Aquat Sci 69(4): 575-583.

Popov PA, Ermolaeva NI, Kipriyanova LM, Mitrofanova EYu (2003) The state of hydrobiocenoses in Altai high mountains. Sib. Ecol. J. 2: 181-192. (in Russian)

Solem JO, Birks HH (2000) Late-glacial and early Holocene Trichoptera (Insecta) from Krakenes Lake, western Norway. Journal of Paleolimnology 23: 49-56.

Suren AM (1994) Macroinvertebrate communities of streams in western Nepal: effects of altitude and land use. Freshwat. Biol. 32: 323-336.

Timm H, Mols T (2005) Macrozoobenthos of Lake Verevi. Hydrobiologia 547: 185-195.

Vdovina ON, Bezmaternykh DM (2020) Benthic macroinvertebrate communities of high- and mid- mountain lakes of the Russian Altai. Fish Farming and Fisheries 2: 4-13. (in Russian)

Verpoorter C, Kutser T, Seekell DA, Tranvik LJ (2014) A global inventory of lakes based on high-resolution satellite imagery. Geophys Res Lett 41(18): 6396-6402. L060641

Vershinin VK, Konovalova OS, Fomenko LA (1979) Zoobenthos of some Altai reservoirs and its role in the introduced peled' nutrition. Biological resources of the Altai territory and ways of their rational use. Barnaul, 123-124. (in Russian)

Vesnina LV, Zelentsov NV, Ryzhakova OG (2012) Fisheries characteristics of alpine lake Zerlyukol'-Nur in Kosh-Agachsky region, the Altai Republic. Bull. of Novosib. St. Agr. Univ. 4: 45-48. (in Russian)

Wallace JB, Grubauch JW, Whiles MR (1996) Biotic indices and stream ecosystem processes: results from an experimental study. Ecolog. Applic. 6: 140-151.

Wetzel RG, Likens GE (2000) Limnological Analyses. Springer, 429 pp.

Yakovlev VA (2000) Trophic structure of zoobenthos as an ecological indicator for aquatic ecosystems and a water quality index. Water Resour 27: 211-218. (in Russian)

Yakovlev VA (2005) Freshwater zoobenthos of northern Fennoscandia (diversity, structure and anthropogenic dynamic) Part 2. Apatity, Kola Science Centre RAS, 145 pp. (in Russian)

Yanygina LV, Krylova E N (2008) Zoobentos of high-altitude water bodies in the basin of Lake Teletskoye. World of Science, Culture, Education. 4: 18-20. (in Russian)

Yanygina LV, Krylova EN (2006) Zoobenthos-based bioindication of the ecological state of Altai foothill reservoirs Polzunovsky Vestnik. 2-1: 365-368. (in Russian)

Zaloznyy NA, Vorob'ev DS (2006) Oligochaetes and leeches of Western Siberia reservoirs (field/laboratory collection and processing of material). Tomsk, 216 pp. (in Russian)

Zarubina EYu, Fetter GV (2019) Production and destruction of organic matter in mountain lakes of the Russian Altai. In: Proc. of XII Congress of Hydrobiological Society of RAS (Russia), September, 2019. Petrozavodsk, 165. (in Russian)

Zarubina EYu, Fetter GV (2020) On hydrological and geochemical characteristics of alpine lakes in the Multa river basin (Gorny Altai). Bulletin of the Altai Branch of RGS. 4(59): 74-82. (in Russian)

Zhadin VI, Gerd SV (1961) Rivers, lakes and reservoirs of the USSR, their fauna and flora. Moscow, 600 pp. (in Russian)