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Article

Asymmetric Distribution of Fish Diversity in Inflows of the Black Irtysh River (Central Asia, Kazakhstan)

by
Nadir Shamilevich Mamilov
1,
Nazym Sapargaliyevna Sapargaliyeva
1,
Erlan Kegenov
1,
Gulnar Bolatovna Kegenova
1,*,
Igor Nikolaevich Magda
2,
Oleg Efimovich Lopatin
2,
Gulnaz Barinova
3,
Sayat Ermukhanbetovich Sharakhmetov
1,
Aliya Gabdullina
1,4,
Gleb Bolbotov
4,
Valentin Rudoi
5 and
Vladimir Vorobyov
4
1
Department of Zoology, Histology and Cytology, Faculty of Biology and Biotechnology, Al-Farabi Kazakh National University, Al-Farabi Avenue, 71, Almaty 050040, Kazakhstan
2
Institute of Zoology, Al-Farabi Avenue, 93, Almaty 050060, Kazakhstan
3
Institute of Animal Science and Veterinary, NCJSC «S. Seifullin Kazakh Agrotechnical Research University», Zhenis Avenue, 62, Astana 010000, Kazakhstan
4
Katon-Karagay State National Natural Park, Zhampeisov Street, 16A, Katon-Karagay 070908, Kazakhstan
5
Department of Ecology, Biochemistry, and Biotechnology, Faculty of Biology, Altai State University, Lenina Avenue, 61, Barnaul RF-656049, Russia
*
Author to whom correspondence should be addressed.
Diversity 2025, 17(9), 648; https://doi.org/10.3390/d17090648
Submission received: 25 June 2025 / Revised: 3 September 2025 / Accepted: 8 September 2025 / Published: 15 September 2025
(This article belongs to the Section Freshwater Biodiversity)
Browse Figures
Versions Notes

Abstract

The diversity and distribution of fish in the water bodies of the Black Irtysh basin was studied in connection with six abiotic environmental parameters (surface flow velocity, width and depth of rivers, connection with the main river, and bottom and maximum temperature). The current species diversity of fish consists of 29 native and 11 alien species. The diversity of right and left tributaries varies greatly as a result of differences in abiotic parameters. The presence of a hydrological connection of reservoirs by itself does not guarantee the distribution of species throughout the basin. Rare indigenous Siberian Taimen Hucho taimen and Markakol Lenok Brachymystax savinovi were regularly found in the right tributaries of the Irtysh River and Lake Markakol, respectively. New fish species to the Kazakh section of the Black Irtysh River have been discovered: Tibetan stone loach Triplophysa stolickai, Severtsov’s loach Triplophysa sewerzowi and alien fish species as asp Leuciscus aspius, Chinese false gudgeon Abbottina rivularis, sunbleak Leucaspius delinetaus, and Misgurnus sp.

1. Introduction

The study of the distribution of species is of great scientific and practical importance. From a scientific point of view, biogeography helps to understand the history of the formation of the relief of our planet, the evolution of biological diversity, and the adaptation of various organisms to the conditions of a particular habitat [1,2,3]. The practical importance of studying the distribution of animals and plants is the rational use of available natural resources, the preservation of the well-being of natural ecosystems, the prevention of outbreaks of vector-borne diseases, etc. [4,5,6]. Fresh waters are inhabited by an exceptionally diverse fish fauna [7,8,9]. Fish perform a variety of ecosystem services [10,11]. Therefore, the preservation of the natural diversity of fish is necessary to maintain the well-being of reservoirs, the environment, and humans [12]. The negative impact of humans is rapidly reducing the natural diversity of freshwater fish [8,12,13], which constantly increases the need for knowledge about the current distribution and condition of fish [14,15,16].
The Irtysh River is located in the center of Asia. This Irtysh River is a cross-border river that runs through the territory of the People’s Republic of China, Kazakhstan, and Russia. The length of the river from its source from People’s Republic of China to its confluence with the large Siberian river Ob exceeds spanning 4200 km, of which 1698 km is on the territory of the Republic of Kazakhstan [17,18,19]. The main territory of the basin serves as a living space for approximately 15 million people. The river is the main source of water in the region, which brings its highest value in social, economic, industrial, energy generation, and irrigation activities [20,21,22,23]. There are several large dams on the river section from Lake Zaisan to the Semipalatinsk city, which have significantly changed the natural hydrological regime of this river.
The Irtysh River is of great importance for the Russian Federation and the Republic of Kazakhstan; therefore, studies of fish fauna are constantly conducted here [24,25,26,27,28,29,30,31]. However, in the most published works, information is provided only on the largest reservoirs such as the Irtysh River itself, Lake Zaisan, Bukhtarma reservoir, and Lake Markakol. The effectiveness of water bodies in specific conditions depends on the preservation of local forms, and the integral ecosystem function across a large area depends on the preservation of all intraspecific diversity.
The upper section of the Irtysh River is called the Black Irtysh. There is a difference between geographical borders and fish distribution in this section of the river. Geographers and some biologists call the section of the Irtysh River from its sources to Lake Zaisan the Black Irtysh [19]. The diversity of the native fish fauna of this section of the Irtysh River consists of fish from Siberian rivers and the basin of Balkhash lake. This peculiarity serves as the basis for the allocation of the Irtysh River as a separate biogeographic region [31,32,33,34]. The features of the fauna of fish and aquatic invertebrates make it possible to expand the boundaries of this section, including Lake Zaisan itself and the section of the Irtysh River located below [33,34,35,36,37,38,39]. Based on a comparative study of the fish fauna of various sections of the Irtysh River, this sector of the river was called the Gorno-Irtysh region by Russian ichthyologists [39].
The intentional introduction of various fish species was initiated in the basins of the Irtysh and Ob rivers in the second half of the last century [40,41]. Biological invasions are now considered one of the main threats to the diversity of native fish species [42,43,44,45]. Despite the introduction of alien fish into the Irtysh basin, it continues [25,45,46,47,48,49].
The Black Irtysh is experiencing the least anthropogenic stress [50] and, accordingly, is of great importance for preserving the diversity of native fish. Therefore, the purpose of our study was to study the current diversity of fish in the Black Irtysh basin within the Republic of Kazakhstan from the border with the People’s Republic of China to the confluence of the Tikhaya River.

2. Materials and Methods

2.1. Study Area and Sampling

The field study was conducted annually during the warm season (from May to September) from 2010 to 2014 and 2020 to 2024. A detailed physical and geographical description of the Black Irtysh River was published recently [50], so we will not repeat it. The geographical coordinates of sampling sites are given in Table 1 and shown in Figure 1. All investigated rivers and lakes on the northern (right) side the river (Figure 2) have a permanent connection with the Irtysh River. The rivers on the southern (left) side are less wide and deeper (Figure 3). The tributaries Kendyrlik, Karatuma, Kargyba, and Shet-Karasu constantly bring their waters to the Irtysh River. The Uzunbulak, Shagan, Ashysu, Bogaz, Shet-Kandysu, and Karasu rivers are lost in the steppe, not reaching the main river. The main abiotic environmental characteristics used as factors that impact fish distribution were the maximum width and depth of water bodies, the speed of the current, the maximum water temperature at the surface, the ground at the bottom, and the connectivity [51]. The rivers’ velocity was measured using the float method (surface velocity, m/s). The surface water temperature was measured using TB-3-M1 (Steklopribor, Chernozavodskoe, Ukraina) and HI 98501 (Checktemp, Hanna Instruments, Cluj Napoka, Transilvania).
All types of electrofishing were banned in the Republic of Kazakhstan. Therefore, fishing was carried out using seine nets (20, 5, and 1 m in length, each with a 1.2 m height and a mesh size of 5 mm) in rivers, brooks, and shallow zones of lakes (up to about 1.2–1.3 m depth). The nets were dragged through the water by two people in such manner that the lower part of the net touched the bottom, where the upper part remained on the surface (Figure 4, left). About 100 m2 of the rivers were cached per each tentative. The rivers were sampled 10 times in such a way. This method is not the best, but it allows providing data on the total diversity, dominant species, and relative abundance of fish [52,53]. The method is traditionally used in Kazakhstan and so makes it possible to compare data obtained for different water bodies and by different researchers [54,55]. Gill nets (mesh sizes of 20, 30, 40, 50, and 60 mm, each with a length of 25 m and a depth of 2 m, working overnight) were used in the lakes, and the hook-and-line angling (spinning rods and fly rods) method was applied in all water bodies. Pictures of fish were performed using a GoPro Hero 4 with a case for underwater photography, and a Canon PowerShot G9 on land. In total, 2 to 5 km of each river was inspected in the mountainous, foothill, and plain parts. Each appropriate site in the river was photographed for about 40 min per visit. In addition, the fisherman’s catch of boat seine nets (Figure 4, right) and gill nets were investigated in the Zaisan Lake.
The species descriptions given in [56,57,58,59,60,61,62,63] were used to determining fish. Taxonomical names of the fish are given according to the Eschmeyer’s Catalog of Fishes [64].

2.2. Descriptive Statistics

The Bray–Curtis index was used to compare similarity of fish assemblages from different sites [65,66]. The UPGMA cluster analysis method was used to graphically represent the obtained similarity matrix.
The correlation between the abiotic parameters of water and fish species, as well as the pairwise presence/absence of fish species, was calculated from the correlation matrix using the Pearson’s linear correlation coefficient and Kendall’s tau coefficient. Both correlation coefficients were considered significant at the 0.5 level and 95% confidence level (p < 0.05) [67]. A multifactor estimation of the relationship of fish with the 6 environmental variables as maximal observed summer temperature (C), stream velocity (velocity), water bodies observed width (width) and depth (depth), dominant type of ground on bottom of the rivers (Rbed), and connection with the Irtysh River or Zaisan Lake (Con) was performed using canonical correspondence analysis (CCA) [68]. The significance of differences was estimated based on the Kruskal–Wallis (multiple comparisons) test. To reduce the effects of rare species, only species caught in two or more sites were included in the CCA. The significance of the canonical axes was assessed by the Monte Carlo permutation test (p < 0.05). All calculations were performed using the PAST 4.07b software version [69].

3. Results

The identified species list is presented in Table 2. One species of jawless vertebrates as lamprey, 29 native, and 11 alien fish species were found. Fishermen caught a lamprey on 14 June 2022 in the Bukhtarma River. Unfortunately, they damaged this specimen so that we could not identify this lamprey to the species. It could be the Siberian brook lamprey Lethenteron kesslerii or Arctic lamprey Lethenteron camtschaticum. All other species were represented in large numbers. All the species were represented by adult and young fishes that indicated satisfactory conditions for they existence and reproduction.
Indigenous E. luceus, L. baicalensis, R. lacustris, P. ujmonensis, Siberian gudgeon, Spotted thicklip loach, Spined loach, European perch and alien Prussian carp, common carp, and freshwater bream are distributed throughout the Black Irtysh basin. Among the studied species, only the distribution of Siberian gudgeon and European perch does not correlate significantly with any of the six studied abiotic factors (Supplementary Figures S1 and S2). A comparison of the diversity of fish in the right (southern) and left (northern) tributaries of the Irtysh River showed large differences in species composition related to the abiotic conditions of the reservoirs. The southern tributaries are the distribution boundary of such species as P. poljakowi, G. dybowskii, T. dorsalis, T. sewerzowi, T. stolickai. Sculpins C. sibiricus and C. dzungaricus, H. taimen, B. lenok, and T. arcticus did not enter these reservoirs. Barbatula toni is widespread in the northern tributaries and only occasionally in the left the Zhineshkesu River and Ujdene Reservior. The T. sewerzowi and the stolickai were discovered in the Black Irtysh basin for the first time. Also, alien species such as L. aspis, A. rivularis, L. delinetaus, and Misgurnus sp. were recorded for the Kazakh part of the Black Irtysh for the first time.
The L. aspius, A. rivularis, A. alburus, and A. delinetaus are found exclusively in right tributaries. Misgurnus sp. was found not only in the right-bank tributaries of the Kendirlik and Karatuma rivers but also in the high-altitude Shanghin Lake on the left bank of the Irtysh.
Indigenous Siberian Taimen H. taimen is listed as vulnerable in the International Red List [70], and Markakol Lenok B. savinovi is listed as endangered [71]. Both species were regularly found in the right tributaries of the Irtysh River and Lake Markakol, respectively.
Cluster analysis of the similarity of fish assemblages showed a clear division of the three groups (Figure 5).
Some of their characteristic species inhabiting the right-bank tributaries are the H. taimen, B. lenok, and B. toni (Supplementary Figure S3). The G. dybowskii, P. percnurus, and P. poljakowii minnow represent species that inhabit exclusively the left tributaries of the Irtysh River (Supplementary Figure S4).
The CCA showed a division of the groups depending on the studied environmental characteristics (Figure 6). H. taimen, B. lenok, B. savinovi, T. arcticus, and B. toni tend to distribute in water bodies with the highest current velocity and low temperature. The axes 1–3 of CCA explained 81.5% of the total variability, and the analysis was statistically significant at the Monte Carlo test (Trace = 0.8916; p = 0.001).

4. Discussion

The results of the observations allowed us to establish differences in the diversity of fish in the right and left tributaries of the Black Irtysh River. The methods of morphological recognition of species of minnows, loaches, and gudgeons lag behind their molecular genetic identification [72,73,74,75,76]. Therefore, the list we have provided does not pretend to be comprehensive and taxonomically accurate. So, at the moment, the habitat of G. acutipinnatus in Lake Markakol is beyond doubt. However, the taxonomic status of gudgeons inhabiting other bodies of water in the Irtysh basin is debatable [74,75]. According to the shape of the lips, the length of the barbels, and the location of the scales on the lower side of the body, all the studied gudgeons were most similar to G. sibiricus. Loach B. toni observed in our samples were well distinguished from B. altayensis by widely separated nostrils. All the individuals we captured had their nostrils widely dilated. However, we do not exclude the possibility of the existence of another species of B. altayensis in the surveyed area [72].
Morphological peculiarities of fish from Lake Zaisan and the Irtysh River have served as the basis for the identification of endemic subspecies of grayling T. arcticus brevicephalus Mitrofanov 1971, spotted thicklip loach as T. strauchii zaisanikus Menschikov 1937, and T. strauchii ruzsky Kessler 1874. A wide variety of loaches and minnows inhabiting the reservoirs of the Kazakh part of the Irtysh basin also require careful study [59,60,61,62,77,78,79,80]. Existence in the basin of the Kazakh part of the Irtysh River T. dorsalis and P. poljakowii were identified about 20 years ago [77,78,79], but for some reason these species were not included in the recent lists of basin species [30,31]. Barbatula nuda and Cobitis elongatoides are also indicated for the Chinese part of the Black Irtysh [81]. Thus, for an informed decision on the taxonomic status of many discovered species, comparative morphological and genetical analyses of a large group of species are necessary, which is beyond the scope of this study.
In this paper, we were more interested in the functional diversity of fish. Such requirements of fish lifestyle and attitude to the flow velocity, diet, spawning substrate, and the possibility of re-spawning throughout the year are well known for most of the fish we have discovered [60,61,62,63]. Based on these data, we compiled a table of the basic requirements of the fish species we discovered (Supplementary Table S1) and analyzed it using the principal component method (Figure 7).
Based on the results of the PCA, we identified six functional groups of fish. The first group (I) consists of fish that prefer moderate or fast water flows and feed mainly on benthos. In the second group (II), there are only Cottus that live on a fast current and feed on benthos. The third group (III) includes fish species that prefer the flow of water and animal food. The tench T. tinca belongs to a separate group (IV), because it prefers stagnant water and is not a predator. The fifth (V) and sixth (VI) groups combine native and alien species that live in slow-flowing rivers with well-developed aquatic vegetation and consume a variety of foods.
The applied methods of fish sampling do not allow us to establish the presence of all rare species in the investigated water bodies. Such species known for the Black Irtysh basin are the native Siberian sturgeon Acipenser baeri, sterlet Acipenser ruthenus, and nelma Stenodus leucichthys nelma [30,31], as well as the alien red eye Scardinius erythrophthalmus, grass carp Ctenopharyngodon idella, silver carp Hypophthalmichthys molitrix, bighead carp Hypophthalmichthys nobilis, Coregonus peled, Coregonus albula, and Perccottus glenii [25,31,47,48,49]. The presence of these species in the lower part of the Irtysh River or in the territory of the People’s Republic of China was confirmed by direct observation or DNA barcoding [25,26,27,28,29,30,31,46,47,81,82,83]. Also for this part of the river, Tian et al. [81] identified Hypomesus olidus (Pallas, 1814), Micropercops swinhonis (Günther, 1873), and Hedinichthys minuta (Li, 1966), which have not yet been identified in Kazakhstan. Another dangerous alien species, the rotan Perccottus glenii, is spread in the Russian part of the Irtysh River basin [84,85,86], but unlike sunbleak Leucaspius delineatus and bleak Alburnus alburnus, it has not yet been found in the Black Irtysh basin. Since there are a number of dams impassable for fish on the Irtysh River, it can be assumed that humans directly participated in the settlement of sunbleak and bleak. Both of these species appeared in the Black Irtysh basin after the deliberate introduction of Europena whitefish Coregonus lavaretus, whitefish Coregonus lavaretus, vendace Coregonus albula, and peled Coregonus peled. Juveniles of all four species are quite similar in appearance and ecology, which greatly facilitates the penetration of sunbleak and bleak during transplants from pond farms with the whitefishes. Rotan differs from them both in appearance and ecology. In addition, the rotan is a predator that is being fought on pond farms. Nevertheless, the appearance of this species in the reservoirs of the Black Irtysh seems very likely to us.
Tian et al. [81] indicate two alien species of loach, Misgurnus dabryanus (Dabry de Thiersant, 1872) and Misgurnus anguillicaudatus (Cantor, 1842), for a section of the Irtysh River basin within the People’s Republic of China. The discovery of the alien Nikolsky loach M. nikolsky in the Ob River basin was first reported more than 15 years ago [87]. According to the external morphological features, the fish we caught are similar to M. anguillicaudatus; however, an analysis using molecular genetic data is necessary to accurately determine the taxonomic status of these fish. Asp L. aspius, Chinese false gudgeon A. rivularis, bleak A. alburnus and belica L. delinetaus do not cause difficulties in species identification.
Naturalization of new alien species is an indicator of unfavorable changes in freshwater ecosystems [44]. Based on hydrochemical and hydrobiological data, the ecological condition of the reservoirs of the Black Irtysh was assessed as satisfactory [50]. It is likely that a direct change in the hydrological regime due to the withdrawal of more water for agriculture in the last few dry years has created the opportunity for the successful naturalization of new alien species. In the future, construction of dams on the tributaries of the Black Irtysh River could significantly worsen the habitat conditions for valuable indigenous species as H. taimen, T. arcticus, and B. lenok.
Our results are generally an example of the fact that the presence of a hydrological connection of reservoirs does not guarantee the distribution of fish species throughout the basin. The diversity of freshwater fish reflects the diversity of habitats in each water body [7,88]. Therefore, the presented results serve as a basis for monitoring the conservation of the natural diversity of fish and the water ecosystems of the Upper Irtysh.

5. Conclusions

  • The fish diversity of the right-bank and left-bank tributaries of the Black Irtysh River significantly differ in species composition. Seven fish species were found only in the right tributaries and only eight species in the left tributaries. The size of reservoirs and the speed of the current are the main abiotic factors affecting the distribution of fish in the Black Irtysh basin. Thus, the presence of a hydrological connection of the water bodies by itself does not guarantee the distribution of species throughout the basin.
  • A rare species like the taimen H. taimen and a local endemic as B. savinovi are still preserved in right-bank tributaries and Markakol Lake.
  • Fish species new to the Kazakh section of the Black Irtysh River have been discovered: Tibetan stone loach T. stolickai and Severtsov’s loach T. sewerzowi and alien fish species including asp L. aspius, Chinese false gudgeon A. rivularis, sunbleak L. delinetaus, and Misgurnus sp., whereas Misgurnus sp. Are related to the first functional group (I) and others to the fifth (V) and sixth (VI) groups. The appearance of new alien species indicates unfavorable changes in the ecosystems of the Black Irtysh.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/d17090648/s1, Figure S1. Kendall’s tau. Figure S2. Person’s correlation. Figure S3. Fishes typical for the right tributaries. Figure S4. Fish species specific for the left tributaries. Table S1. Fish requirements.

Author Contributions

Conceptualization, methodology, investigations, data analysis, original draft preparation, supervision, N.S.M., N.S.S., and G.B. (Gulnaz Barinova); field investigation and data curation, E.K., I.N.M., O.E.L., S.E.S., G.B. (Gleb Bolbotov), A.G., V.R., V.V.; formal analysis and resources, N.S.M., N.S.S., A.G.; visualization, N.S.M., E.K., A.G., G.B. (Gleb Bolbotov); project administration, E.K., G.B.K., A.G. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Institutional Review Board Statement

Ethical review and approval were waived for this study due to the fish catching and research being performed in accordance with the laws of the Republic of Kazakhstan “On the protection, reproduction, and use of fauna” from 9 July 2004 No. 593-II (as amended by the Law of the Republic of Kazakhstan, dated 3 July 2013 No. 124-V, dated 16 May 2014 No. 203-V, dated 29 September 2014 No. 239-V, dated 29 December 2014 No. 269-V ZRK, dated 29 October 2015 No. 376-V ZRK, dated 29 March 2016 No. 479-V, dated 15 June 2017 No. 73, dated 2 April 2019 No. 241, dated 28 October 2019 No. 268, dated 23 February 2021 No. 11, dated 30 April 2021 No. 34, dated 24 November 2021 No. 75, dated 14 July 2022 No. 141), from 30 December 2021 No. 97-VII ZRK “On the responsible treatment of animals”, and the order of the Minister of Agriculture of the Republic of Kazakhstan dated 10 November 2004 No. 652. “On approval of the Rules for the use of animals, with the exception of rare and endangered animals, in scientific, cultural, educational, and aesthetic purposes, including for the creation of zoological collections”, and Rules for the organization and conduct of scientific activities and scientific research in environmental institutions, approved by the Order of the Minister of Agriculture of the Republic of Kazakhstan dated 13 March 2012 No. 25-02-01/94.

Data Availability Statement

The original contributions presented in the study are included in the article; further inquiries can be directed to the corresponding author.

Acknowledgments

The authors are deeply grateful to Mitrofanov I.V., Baymukanov M.T., and Timirkhanov S.R for scientific advice. We also height appreciate Korneliyuk A.I., Amanbayev Zh., Dorzhanov N., Budkeev D., Tikenov S., and Chelyshev A. for your help in organizing and collecting the source data. We highly appreciate the time spent and the valuable recommendations of our anonymous reviewers, who were able to significantly improve this manuscript.

Conflicts of Interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

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Figure 1. Schematic map of the Black Irtysh basin. Study sites are indicated by numbers: 1, Uzunbulak brook; 2, Shagan; 3, Ashysu; 4, Shar; 5, Bogaz; 6, Kargyba; 7, Karasu Shet-Karasu, Shet-Kandysu; 8, Zaisan Lake; 9, Ujdene Reservior; 10, Karatuma, Kogeady, Beiyttibulak, Zhineshkesu, Saryeshki, and Kandyrlik; 11, Irtysh; 12, Kalzhyr; 13, Markakol Lake; 14, Topolevka; 15, Kurchum; 16 Shanghin Lake and Kara Kaba; 17, Bukhtarma; 18, Belaya; 19, Tikhaya.
Figure 1. Schematic map of the Black Irtysh basin. Study sites are indicated by numbers: 1, Uzunbulak brook; 2, Shagan; 3, Ashysu; 4, Shar; 5, Bogaz; 6, Kargyba; 7, Karasu Shet-Karasu, Shet-Kandysu; 8, Zaisan Lake; 9, Ujdene Reservior; 10, Karatuma, Kogeady, Beiyttibulak, Zhineshkesu, Saryeshki, and Kandyrlik; 11, Irtysh; 12, Kalzhyr; 13, Markakol Lake; 14, Topolevka; 15, Kurchum; 16 Shanghin Lake and Kara Kaba; 17, Bukhtarma; 18, Belaya; 19, Tikhaya.
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Figure 2. A typical water body of the right-bank of the Irtysh River.
Figure 2. A typical water body of the right-bank of the Irtysh River.
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Figure 3. A typical water body of the left-bank of the Irtysh River.
Figure 3. A typical water body of the left-bank of the Irtysh River.
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Figure 4. Fish sampling using hand seine net (left) and boat seine net (right).
Figure 4. Fish sampling using hand seine net (left) and boat seine net (right).
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Figure 5. Similarity of fish assemblages (Bray–Curtis coefficient, UPGMA). Left tributaries are shown with black; the Ujdene water reservoir is green; right tributaries shown with aqua and lakes with blue. The Black Irtysh River and Zaisan Lake are shown with red. Abbreviations of the water bodies are given by Table 1.
Figure 5. Similarity of fish assemblages (Bray–Curtis coefficient, UPGMA). Left tributaries are shown with black; the Ujdene water reservoir is green; right tributaries shown with aqua and lakes with blue. The Black Irtysh River and Zaisan Lake are shown with red. Abbreviations of the water bodies are given by Table 1.
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Figure 6. CCA plot of species (objects), sites, and environmental variables of water bodies in the Irtysh River basin: first axis horizontally, second axis vertically. Abbreviations of the fish species are given in Table 2. Blue dots with abbreviations present fish species, aqua dots present right side inflows, black dots present left side inflows, the red dotes present the Irtysh River and Zaisan Lake, and the green one present Ujdene Reservior. Abbreviations of the environmental arrows are Width—width of the water bodies in the sampling sites (m), Depth—maximal observed depth (m), Con—connection with the main river, C—maximal observed water temperature, Velocity—stream velocity (m/s), Rbed—type of ground on bottom of the rivers.
Figure 6. CCA plot of species (objects), sites, and environmental variables of water bodies in the Irtysh River basin: first axis horizontally, second axis vertically. Abbreviations of the fish species are given in Table 2. Blue dots with abbreviations present fish species, aqua dots present right side inflows, black dots present left side inflows, the red dotes present the Irtysh River and Zaisan Lake, and the green one present Ujdene Reservior. Abbreviations of the environmental arrows are Width—width of the water bodies in the sampling sites (m), Depth—maximal observed depth (m), Con—connection with the main river, C—maximal observed water temperature, Velocity—stream velocity (m/s), Rbed—type of ground on bottom of the rivers.
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Figure 7. Fish functional groups in space of the 1st and 2nd principal components. Fish species abbreviations were given in Table 1. Colored dotes indicate different species. The abbreviations of fish species are given in the Table 2. Roman numerals show the functional groups of fish. The full explanation is given in the text.
Figure 7. Fish functional groups in space of the 1st and 2nd principal components. Fish species abbreviations were given in Table 1. Colored dotes indicate different species. The abbreviations of fish species are given in the Table 2. Roman numerals show the functional groups of fish. The full explanation is given in the text.
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Table 1. List of the investigated water bodies and the main sampling sites.
Table 1. List of the investigated water bodies and the main sampling sites.
Water BodiesAbbreviationNEAbove Sea Level, m
Irtysh RiverIRT47°55′29″84°59′50″408
47°52′35″84°46′49″394
49°07′56″84°06′43″390
Zaisan LakeZAI47°40′22″84°27′05″437
47°52′14″84°46′49″437
Right (North) bank:
Shanghin Lake (=Bukhtarminskoe)LBU49°17′44″83°53′07″2068
Markakol LakeLMA48°47′32″86°01′25″1451
49°39′39″85°42′51″1450
Kalzhyr RiverKal48°00′39”85°11′01”411
48°29′52”84°13′49”862
Sarymsakty RiverSar49°10′01″85°36′35″1058
Bukhtarma RiverBuk49°15′15″85°22′13″739
49°27′31″85°03′59″591
49°47′44″84°08′24″407
Belaya RiverBel49°18′06″85°18′43″685
49°16′59”85°17′56”677
Kara Kaba RiverKab49°01′14″85°51′23″2140
Kurchum RiverKur48°37′20″83°54′48″476
Topolevka RiverTop48°54′45″85°50′18”2019
Tikhaya RiverTik50°24′36″83°30′36″749
Left (South) bank:
Kendyrlik RiverKen47°37′27″84°57′13″450
47°79′58″84°79′60″390
Saryeshki RiverSae47°37′01″85°00′30″453
Karatuma RiverKar47°30′50″84°56′59″489
Kogeady RiverKgd47°34′41″85°05′36″507
Shar RiverSHR49°14′35″81°49′20″602
Shagan RiverSha49°19′23”78°24′17”681
49°47′34”78°45′52”367
Ashysu RiverAsh49°43′31”78°52′58”403
Uzunbulak brookUzb49°45′51”78°5′54”794
Karasu RiverKsu47°33′38″83°38′58″647
Shet-Karasu RiverKKS47°33′49″83°39′05″645
Shet-Kandysu RiverSka47°12′24″84°27′59″730
Bogaz RiverBog47°87′11″82°00′21″764
Kargyba RiverKgb47°57′40″82°57′30″533
Beiyttibulak RiverBBk47°29′31″85°10′17″722
Ujdene ReserviorUjd47°22′34″84°47′27″805
Zhineshkesu RiverZhy47°38′09″85°06′31″476
Table 2. Fish diversity and species distribution in the basin of the Black Irtysh.
Table 2. Fish diversity and species distribution in the basin of the Black Irtysh.
SpeciesCommon NameAbbreviationIrtysh and ZaisanRight Side InflowsLeft Side
Inflows
Indigenous:
Lethenteron sp.Siberian brook lamprey or ArcticLet0+0
Hucho taimenSiberian TaimenHuc0+0
Brachymystax lenokLenokBrl++0
Brachymystax savinoviMarkakol LenokBrs0+0
Thymallus arcticusArctic GraylingThy0+0
Esox luciusNorthern PikeEso+++
Leuciscus baicalensisSiberian DaceLba+++
Leuciscus idusIdeLid++0
Leuciscus aspiusAspLas+00
Rutilus lacustrisRoachRla+++
Carassius carassiusCrucian CarpCar+0+
Rhynchocypris percnurusLake MinnowPho00+
Phoxinus poljakowiBalkhash MinnowRhy00+
Phoxinus ujmonensis Phu+++
Tinca tincaTenchTin+0+
Gobio sibiricusSiberian gudgeonGos+++
Gobio acutipinnatus Goa0+0
Gymnodiptychus dybowskiiNaked osmanGym00+
Triplophysa dorsalisGray loachTdo00+
Triplophysa strauchiiSpotted thicklip loachTst+++
Triplophysa stolickaiTibetan stone loachTsz00+
Triplophysa sewerzowiiSevertsov’s loachTsw00+
Triplophysa sp. Tsp00+
Barbatula toni Bar0++
Cobitis melanoleucaSpined loachCob+++
Lota lotaBurbotLot++0
Perca fluviatilisEuropean perchPer+++
Gymnocephalus cernuusRuffeCer++0
Cottus sibiricusSiberian sculpinCot0+0
Cottus dzungaricus Cot0+0
Alien:
Coregonus albulaVendaceCoa+0+
Coregonus lavaretusEuropean whitefishCor+0+
Carassius gibelioPrussian carpCag+++
Abramis bramaFreshwater breamAbr+++
Alburnus alburnusBleakAlb+++
Leucaspius delinetausSunbleak (=belica)Leu00+
Cyprinus carpioCommon carpCyp+++
Pseudorasbora parvaTopmouth gudgeonPse0++
Abbottina rivularisChinese false gudgeonAbb00+
Misgurnus sp.Pond loachMis0++
Sander luciopercaPike-perchSan+0+
+ the species were revealed; 0 the species were not revealed.
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MDPI and ACS Style

Mamilov, N.S.; Sapargaliyeva, N.S.; Kegenov, E.; Kegenova, G.B.; Magda, I.N.; Lopatin, O.E.; Barinova, G.; Sharakhmetov, S.E.; Gabdullina, A.; Bolbotov, G.; et al. Asymmetric Distribution of Fish Diversity in Inflows of the Black Irtysh River (Central Asia, Kazakhstan). Diversity 2025, 17, 648. https://doi.org/10.3390/d17090648

AMA Style

Mamilov NS, Sapargaliyeva NS, Kegenov E, Kegenova GB, Magda IN, Lopatin OE, Barinova G, Sharakhmetov SE, Gabdullina A, Bolbotov G, et al. Asymmetric Distribution of Fish Diversity in Inflows of the Black Irtysh River (Central Asia, Kazakhstan). Diversity. 2025; 17(9):648. https://doi.org/10.3390/d17090648

Chicago/Turabian Style

Mamilov, Nadir Shamilevich, Nazym Sapargaliyevna Sapargaliyeva, Erlan Kegenov, Gulnar Bolatovna Kegenova, Igor Nikolaevich Magda, Oleg Efimovich Lopatin, Gulnaz Barinova, Sayat Ermukhanbetovich Sharakhmetov, Aliya Gabdullina, Gleb Bolbotov, and et al. 2025. "Asymmetric Distribution of Fish Diversity in Inflows of the Black Irtysh River (Central Asia, Kazakhstan)" Diversity 17, no. 9: 648. https://doi.org/10.3390/d17090648

APA Style

Mamilov, N. S., Sapargaliyeva, N. S., Kegenov, E., Kegenova, G. B., Magda, I. N., Lopatin, O. E., Barinova, G., Sharakhmetov, S. E., Gabdullina, A., Bolbotov, G., Rudoi, V., & Vorobyov, V. (2025). Asymmetric Distribution of Fish Diversity in Inflows of the Black Irtysh River (Central Asia, Kazakhstan). Diversity, 17(9), 648. https://doi.org/10.3390/d17090648

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