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The fishes and fishery in Baikal A. N. Matveyev* and V. P. Samusenok Faculty of Biology and Soil, State University, Irkutsk, *Corresponding author: [email protected]

This article reviews the fish fauna of Lake Baikal, the world’s deepest and largest lake in volume. The nucleus of the fish fauna of Lake Baikal is a unique complex of endemic cottoid fishes (56% of the total number of species and 41% of the total number of genera) and endemic species and subspecies of the coregonid and thymallid families. Abundance and composition of the fish fauna have changed during the 20th Century in each of the three main parts of the lake: north, central and south. Historical and present environmental conditions, such as main habitats, quality and anthropogenic impacts, are discussed. Key objectives of fishery management on Lake Baikal include: (i) protection of fishes, especially commercial species, during periods of preparation for spawning, on spawning runs and in spawning areas; (ii) progressive restriction of the industrial fisheries and rapid expansion of sports and recreational fisheries; (iii) reallocation of priorities in the artificial breeding of fishes to restore the natural population structure of and to restore the abundance of other valuable species that are important for sport fishing; (iv) prevention of the introduction into the Baikal basin of any new non- indigenous fish species; (v) conservation of habitats used by different ecological groups of fishes.

Keywords: , fishery goals, introductions, conservation

Introduction their effectiveness, and outline the future prospects for fisheries development. Addition of Lake Baikal to the World Heritage List necessitates a critical review of human activi- ties on the shores of the lake and development of Discussion new environmental management strategies. This review describes changes in the fish community of A brief physical-geographical review the lake, resulting from human activities, and anal- of Lake Baikal yses the status of fisheries in Lake Baikal. For sev- eral hundred years human impacts on this Lake Baikal is by volume (23,015 km3) and by ecosystem were mainly expressed in the intensifi- depth (over 1630 m) the earth’s largest freshwater cation of fisheries. Only since the mid-20th body (Ravens et al., 2000). It holds about 20% of Century has the pace of industrial development world freshwater reserves, and, at the same time, increased in the lake basin, leading to a significant is an important Russian inland fishery . increase in adverse impacts on the ecosystem of Lake Baikal is located in Eastern at 454 Lake Baikal, including the fish stocks. The authors meters above sea level and fills the biggest central review the actions taken in recent decades to pre- depression among a series of tectonic breaks of the vent these effects primarily on the fishes, evaluate Baikal zone. The lake is 636 km long and 25

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Aquatic Ecosystem Health & Management, 18(2):134–148, 2015. Copyright Ó 2015 AEHMS. ISSN: 1463-4988 print / 1539-4077 online DOI: 10.1080/14634988.2015.1028868

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to 80 (averaging 48) km wide with a water surface about half of river inflow (some 30 km3). Other big area of 31,500 km2, a catchment area of tributaries are the River Verkhnyaya 540,000 km2, a shoreline length of 1800 km, a (640 km long) with an annual inflow of 9.42 km3, mean depth of 731 m and a maximum depth of and River (700 km long) with an annual 1631 m (Efimov, 1970). influx of 3.54 km3. A single river, the Angara, The lake bed consists of three depressions flows out of the lake. Water level has a seasonal (Figure 1). The most ancient is the southern depres- fluctuation within limits of 1.4–2.7 m with the min- sion with the maximum depth, by different sources, imum level observed in April and the maximum in of 1394–1432 m; the maximum depth of the mid- September. The water’s mineral content is less than dle depression reaches, by different sources, 1632– 100 mg l¡1 and the calcium content is less than 1642 m; the most recently formed and shallow 15 mg l¡1. One of the unique features of Lake Bai- northern depression is 889 m deep. More than 300 kal is the high concentration of oxygen in the water rivers and small streams flow into the lake; the big- at all depths. Oxygen saturation varies from 75 gest one is the River which accounts for (at the bottom) to 115%. Transparency by Secchi

Figure 1. A map of Lake Baikal showing the main basins and their deepest points (m), notable bays, sors and rivers, along with the sites of the main fish hatcheries: (1) Bol’sherechenskiy; (2) Selenginskiy; (3) Barguzinskiy.

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disk is 5–8 m in summer and 30–40 m in winter most abundant and productive (0.25–0.64 g (Domysheva, 2009). m¡3) species of mesoplankton is a small The thermal regime of the lake is harsh and endemic crustacean Epischura baicalensis only exposed to seasonal fluctuations in the sur- Sars, 1900. The main species of macroplankton face 200–250 metres. In summer, the temperatures (up to 3 g m¡3) is an endemic pelagic amphi- of the layer are 14–16C, at pod Macrohectopus branickii (Dybowski, 10 metres 10–12C, and in some shallow bays sur- 1874). The pelagic endemic fish community face values may reach 20–24C. There are two consisting of five or six pelagic and benthopela- periods of the temperature stratification – direct in gic species is discussed below; summer and inverse in winter. Also, there are two 2. The presence in slope zones of a macroplank- periods of homothermy, in spring (May–June) and ton community, that possesses a very high bio- in autumn (October–November), when the water logical productivity, analogous to the temperature of the surface 0¡250 m water layer upwelling zone in the ocean rich in nutrients; equals to about 4C. Below those depths, there is a 3. The benthic community is characterized by permanently stratified deep water mass with tem- high species diversity and high productivity perature throughout the year equaling to 3.1– (up to 30 g m¡2 y¡1 in the open littoral zone 3.8C (Shimaraev et al., 1993; Ravens et al., and up to 80 g m¡2 y¡1 in the bays). Key spe- 2000). The lake is covered with ice from January– cies of the benthic community are oligochaetes, February until early June. Maximum ice thickness mollusks, amphipods, and chironomids; ranges from 0.6–0.8 to 1.2–1.5 m. Bottom sedi- 4. The presence of the fauna at a great ments vary in different parts of the lake from depth (more than 100 m) analogous to that stony-pebble and sandy bottoms in the shore zone known only in oceans and seas; and, to silty sand, silt, and clay in the deeper zones. 5. The terminal species in the food chain is an Sediment accumulation is 4.2 cm per 1000 years endemic Pusa sibirica (Gmelin, (Efimov, 1970; Domysheva, 2009). The recent 1788), the world’s only completely freshwater state of hydrology of Lake Baikal was reviewed seal with a population size between 70,000 and by Shimarev and Domysheva (2012). 120,000.

Areas with a depth up to 20 m (only about 5% of Review of fish fauna composition the total lake area) are the main fishing zones and history adjacent to the mouths of main rivers such as Selenga, Verkhnyaya Angara, Kichera, Barguzin , taxonomic contents, and endemism or located in bays, relatively larger (Barguzin Lake Baikal Lake has a unique and rich Bay, Chivyrkuy Bay, Maloye Morye [Small Sea] endemic flora and fauna. Specialists predict there Strait) or smaller, so-called sors (North Baikal could be as many as 3500 animal species and 1500 Sor, Proval Sor, Posolskiy Sor, Dubininskiy Sor). plant species (Timoshkin, 2001). The most diverse A scheme of zoogeographic was proposed are infusorians (350 free-living and parasitic spe- by Taliev (1955) for Lake Baikal with the lake cies), flatworms (more than 290 species), divided into two main bathymetric zones, the (more than 200 species), oligochaetes (more than shallow zone of 0–400 m depth and the deep 200 species), and crustaceans (more than 690 spe- zone (400 m down to the maximum depth). The cies). The Chironomidae contain more than 120 shallow zone is subdivided into the northern part species, mollusks (gastropods and bivalves) more and the central-southern part with nine open than 180 species, and fishes 61 species. Besides a water districts within both, and shallow isolated great number of endemic species, there are also areas (sors) neighboring river mouths. The deep some endemic genera and families. zone is subdivided into north-central and southern In its structure and characteristics, the Baikal parts. Based on modern data on fish distribution ecosystem is similar to oceanic ecosystems with in the lake, the scheme has been modified and no the following characteristic features: subdivisions are accepted now as it was shown that there are no distributional barriers for most 1. The presence of a pelagic community with low deep-lake species (Matveyev and Samusenok, taxonomic diversity and high productivity. The 2006).

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There have been a number of fundamental non-indigenous species – Peled peled investigations of the and ecology of Gmelin, 1789, Amur Catfish Parasilurus asotus Baikal fishes (Taliev, 1955; Skryabin, 1969, 1979; Linnaeus, 1758, and Amur Sleeper Percottus Smirnov and Shumilov, 1974; Tugarina, 1981; glenii Dybowski 1877, are also established there. Sideleva, 1982, 1994, 2003; Matveyev et al., The Baikal littoral zone is an area of interaction 2008). The Baikal fish fauna includes 61 species between ‘endemic Baikal complex’ and ‘common and subspecies (Sideleva, 1994, 1998, 2001, 2003; Siberian complex’ and is a transit zone for energy Knizhin et al., 2004; Matveyev and Samusenok, and nutrient exchange to and from the deep zone. 2006; Matveyev et al., 2008) from 15 families; a At the same time, the littoral zone is of great sig- list of taxa is given in Table 1. nificance for fishery as it is inhabited by valuable Significant differences of the fish fauna of Lake species and, thus, is the main zone of traditional Baikal from those in adjacent basins is the reason fishing. for establishing it as a separate zoogeographic sub- Species abundances and the composition of the region belonging to the Holarctic Region (Berg, fish fauna in each of the three main parts of the 1909). The nucleus of the fish fauna of Lake Bai- lake have undergone considerable changes during kal is a unique complex of endemic cottoid fishes the 20th Century (Table 1). According to our 20- (56% of the total number of species and 41% of year-long investigations and literature (Taliev, the total number of genera). 1955; Bazikalova and Vilisova, 1959; Ustyuzha- Using the species endemism and distributions nina, 1971), the fish composition of the Baikal litto- in the lake, three groups of fishes can be distin- ral zone and the slope zone up to the depth of 200– guished. The first group comprises endemic spe- 300 m consists, at present, of 46 species and sub- cies of the families Cottidae, Comephoridae and species inhabiting it permanently or sporadically. (the latter two families are consid- Species that mainly inhabit the littoral zone are as ered to be within Cottidae by some authors) (34 follows (in decreasing order with regard to their bio- species from 12 genera) which are widely distrib- mass and abundance): Sand Leocottus kes- uted at all depths all over Lake Baikal. The second slerii (Dybowski, 1874), Stone Sculpin Paracottus group was defined as a ‘Baikal-Siberian complex’ knerii (Dybowski, 1874), Black Baikal Grayling Thy- by Vereshchagin (1935) and consists of five fishes mallus baicalensis (Dybowski, 1874), Baikal Omul endemic at the species or subspecies level: Baikal C. migratorius migratorius (Georgi, 1775), Bighead Acipenser baerii baicalensis Nikolski, Sculpin baicalensis (Dybowski, 1896; Baikal Omul, or just Omul Coregonus 1874), White Baikal Grayling Thymallus brevipinnis migratorius migratorius Georgi, 1775; Baikal Svetovidov, 1935, Baikal Lake Whitefish C. pid- Lake Whitefish C. pidschian baicalensis Dybow- schian baicalensis Dybowski, 1874, and Siberian ski, 1874; Black Baikal Grayling Thymallus baica- Whitefish C. pidschian (Gmelin, 1788). In the sublit- lensis Dybowski, 1874; and White Baikal Grayling toral zone the main species are Baikal Omul and Yel- T. brevipinnis Svetovidov, 1935. The third group lowfin Baikal Sculpin grewingkii is a complex of non-endemic ‘common Siberian’ (Dybowski, 1874) and other cottoid species are also species inhabiting the lake’s littoral zone and present. mouths of rivers, which consists of 16 indigenous Depths of more than 300 m are inhabited by species and 6 introduced species. Many species only endemic cottoid fishes. This is a very specific from this third group are very rare or occasional in autochthonous lineage that evolved in the lake the lake (Table 1). itself (Mamontov, 2001). At depths of more than Fishes of the ‘Baikal-Siberian complex’ inhabit 1000 m, there are only six species, and at the max- the lake up to depths of 250–300 m; however their imum depth only four species occur: Cottinella maximum abundance occurs at depths of 20– boulengeri (Berg, 1906), Neocottus werestscha- 50 m. The same zone is also inhabited by several gini Taliev, 1935, Abyssocottus gibbosus Berg, species of the ‘endemic Baikal complex’ and of 1906 and Abyssocottus korotneffi Berg, 1906 the ‘common Siberian complex’. The latter group (Sideleva, 1994, 1998, 2003). is represented there by so-called ‘Baikal forms’ The pelagic ichthyocenosis is more productive, specialized for inhabiting the Baikal littoral zone though composed by a limited number of species: and having increased growth rate (Kozhov and two pelagic viviparous cottoid species (Big Oilfish Misharin, 1958; Kartushin, 1958). A few of the baicalensis [Pallas, 1776]) and Small

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Table 1. Freshwater fishes occurring in three main sub–basins of Lake Baikal, their Red Book status, and changes noted in their distribution and abundance2 over three periods in the 20th Century: 1900¡1954, 1955¡1973 and 1974¡1999.

North Baikal Middle Baikal South Baikal 1900– 1955– 1974– 1900– 1955– 1974– 1900– 1955– 1981– Status1 Family, species 1954 1973 1999 1954 1973 1999 1954 1980 1999 Family Acipenseridae R Baikal sturgeon Acipenser C !!CCCCC !! baerii baicalensis (Nikolski, 1896) Family Cyprinidae A Bream Abramis brama ––––! !–– (Linnaeus, 1758) Common Ide Leuciscus idus !!!!!!––– (Linnaeus, 1758) Siberian Dace L. baicalensis C C C CC CC CC C C C (Dybowski, 1874) Common Minnow Phoxinus CCCCCCCCC phoxinus (Linnaeus, 1758) Lake Minnow Rhynchocypris ––––––––! percnurus (Pallas, 1814) Roach Rutilus rutilus (Linnaeus, CC CC CC CC CC CC C CC CC 1758) Siberian Gudgeon Gobio –––––!!!! cynocephalus (Dybowski, 1869) Crucian Carp Carassius gibelio !!!–!!–!! (Bloch, 1782) A Amur Carp Cyprinus –––!CC –!! rubrofuscus (Lacepede, 1803) Family Nemacheilidae Siberian Stone Loach Barbatula ––!–!!!!! toni (Dybowski, 1869) Family Cobitidae Siberian Spiny Loach Cobitis ––!–!!!!! melanoleuca (Nichols, 1925) Family Siluridae A Amur Catfish Parasilurus –!!–!C –!C asotus (Linnaeus, 1758) Family Salmonidae R Brachymystax lenok CCC CCC CC CC C C CCC CC C (Pallas, 1773) R Taimen Hucho taimen (Pallas, CCC CC C CC C ! CC – 1773) R Frolikha Char Salvelinus !?–––––––– alpinus erythrinus (Georgi, 1775)

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Table 1. Freshwater fishes occurring in three main sub–basins of Lake Baikal, their Red Book status, and changes noted in their distribution and abundance2 over three periods in the 20th Century: 1900¡1954, 1955¡1973 and 1974¡1999. (Continued) North Baikal Middle Baikal South Baikal 1900– 1955– 1974– 1900– 1955– 1974– 1900– 1955– 1981– Status1 Family, species 1954 1973 1999 1954 1973 1999 1954 1980 1999 Family Coregonidae A Vendace Coregonus albula ––––!?!?––– (Linnaeus, 1758) E Baikal Omul C. migratorius CCC CCC CCC CCC CCC CCC CCC CC CCC migratorius (Georgi, 1775) E Baikal Lake Whitefish C. C C C CCC CC CC C C ! pidschian baicalensis (Dybowski, 1874) Siberian Whitefish C. pidschian CCC CCC CC C C C ––– (Gmelin, 1788) A Peled C. peled (Gmelin, 1789) ––––! !–! ! Family Thymallidae E Black Baikal Grayling CCC CCC CCC CCC CCC CCC CCC CC CC (Dybowski, 1874) ER White Baikal Grayling T. CCC CC CC CCC CC CC CC C C brevipinnis (Svetovidov, 1935) Family Esocidae Northern Pike Esox lucius C C C CC CC CC C C C (Linnaeus, 1758) Family Lotidae Burbot Lota lota (Linnaeus, CCCCCCCCCCCCCC 1758) Family Percidae Perch Perca fluviatilis C C C CC CC CC C C C (Linnaeus, 1758) Family Eleotrididae A Amur Sleeper Percottus glenii ––!–!CC –!C (Dybowski, 1877) Family Cottidae E Bighead Sculpin C C CC CCC CCC CCC CCC CCC CCC Batrachocottus baicalensis (Dybowski, 1874) E Mottlefin Sculpin B. CC CC CC CC CC CC CC CC CC multiradiatus (Berg, 1907) E Fat Sculpin B. nikolskii (Berg, CC CC CC CC CC CC CC CC CC 1900) E Taliev’s Sculpin B. talievi ––CC ––CC ––CC (Sideleva, 1999)

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Table 1. Freshwater fishes occurring in three main sub–basins of Lake Baikal, their Red Book status, and changes noted in their distribution and abundance2 over three periods in the 20th Century: 1900¡1954, 1955¡1973 and 1974¡1999. (Continued) North Baikal Middle Baikal South Baikal 1900– 1955– 1974– 1900– 1955– 1974– 1900– 1955– 1981– Status1 Family, species 1954 1973 1999 1954 1973 1999 1954 1980 1999 E Baikal Yellowfin CC CC CC CCC CC CCC CCC CC CCC Cottocomephorus grewingkii (Dybowski, 1874) E Longfin Baikal Sculpin C. CCC CCC CCC CCC CCC CCC CCC CCC CCC inermis (Jakovlev, 1890) E North Baikal Yellowfin C. CCC CCC CCC CCC CCC CCC CC CC CC alexandrae (Taliev, 1935) Sand Sculpin Leocottus CCC CC CCC CCC CC CCC CCC CCC CCC kesslerii (Dybowski, 1874) Stone Sculpin Paracottus knerii CC CC CCC CCC CCC CCC CCC CCC CCC (Dybowski, 1874) Family Comephoridae E Big Baikal Oilfish Comephorus CCC CCC CCC CCC CCC CCC CCC CCC CCC baicalensis (Pallas, 1776) E Little Baikal Oilfish C. CCC CCC CCC CCC CCC CCC CCC CCC CCC dybowski (Korotneff, 1905) Family Abyssocottidae E Elokhin’s Sculpin Abyssocottus !–––––––– elochini (Taliev, 1955) E White Sculpin A. gibbosus CCCCCCCCC (Berg, 1906) E Smalleye Sculpin A.korotneffi !!!!!!!!! (Berg, 1906) E Deepwater Baikal Sculpin ––––––CCC Asprocottus abyssalis (Taliev, 1955) E Herzenstein’s Rough Sculpin A. CC CC CC CC CC CC CC CC CC herzensteini (Berg, 1906) E Koryakov’s Sculpin A. ––C ––C ––C korjakovi (Sideleva, 2001) E Half-Naked Sculpin A. CC CC CC CC CC CC C C C intermedius (Taliev, 1955) E Armored Sculpin A. parmiferus CC CC CC C C C !!! (Taliev, 1955) E Flathead Sculpin A. CCCCCCCCCCCC platycephalus (Taliev, 1955) E Narrow-Snouted Sculpin A. CCCCCC ––– pulcher (Taliev, 1955) E Short–Headed Sculpin CCCCCCCCC Cottinella boulengeri (Berg, 1906)

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Table 1. Freshwater fishes occurring in three main sub–basins of Lake Baikal, their Red Book status, and changes noted in their distribution and abundance2 over three periods in the 20th Century: 1900¡1954, 1955¡1973 and 1974¡1999. (Continued) North Baikal Middle Baikal South Baikal 1900– 1955– 1974– 1900– 1955– 1974– 1900– 1955– 1981– Status1 Family, species 1954 1973 1999 1954 1973 1999 1954 1980 1999 E Thick–Snouted Sculpin C C C CC CC CC CC CC CC Cyphocottus eurystomus (Taliev, 1955) E Sculpin C. megalops CC CC CC CC CC CC C C C (Gratzianov, 1902) E Flat Baikal Sculpin Limnocottus CC CC CC CC CC CC CC CC CC bergianus (Taliev, 1935) E Guttate Sculpin L. godlewskii –––C C C CC CC CC (Dybowski, 1874) E Dark Sculpin L. griseus (Taliev, CCCCCCCCCCCC 1955) E Narrow Sculpin L. pallidus C C C CC CC CC CC CC CC (Taliev, 1955) E Loose Sculpin Neocottus ––––––CCC werestschagini (Taliev, 1935) E Thermal water Sculpin N. ––C –––––– termalis (Sideleva, 2002) E Red Sculpin Procottus jeittelesii C C C CC CC CC CC CC CC (Dybowski, 1874) E Goto’s Sculpin P. gotoi –––––C ––CC (Sideleva, 2001) E Big Red Baikal Sculpin P. CC CC CC CCC CCC CCC CC CC CC major (Taliev, 1949) E Dwarf Sculpin P. gurwicii –––! ! !CCC (Taliev, 1946)

1Status: R: rare species listed in Red Data Book of Russia; A: alien species; E: endemic species. 2Distribution and abundance: CCC: numerous species; CC: common species; C: species of rare occurrence; !: sporadic occur- rence; ?: doubtful occurrence.

Oilfish Comephorus dybowski Korotneff, 1905) History of the fish fauna formation and three benthopelagic fishes (Yellowfin Baikal Sculpin, Longfin Baikal Sculpin (Cottocomepho- The formation of the unique present-day fish rus inermis [Jakovlev, 1890]), and Baikal Omul. fauna in Lake Baikal is the result of a long evolu- The of Big Oilfish was estimated by our tion and adaptation to the diversity of existing bio- observations as 73,300–112,000 tonnes (t); Small topes. Prior to the existence of the modern Baikal Oilfish 61,500–103,000 t, Yellowfin Baikal Scul- there were several pre-historic water bodies. Their pin 5000 t, Longfin Baikal Sculpin 3000 t, and shape and number changed through time Omul 23,000–30,000 t. (reviewed by Timoshkin, 2001). The first Endemic species are absent from shallow appeared between 25–30 and 60–70 million years bays and sors where the most abundant species ago. The most ancient (Oligocene–Miocene) deep- are cyprinids, in particular Roach Rutilus rutilus water lake appeared first in the South Baikal (Linnaeus, 1758), and also Perch Perca fluviatilis depression and it was inhabited by , Linnaeus, 1758 and Pike Esox lucius Linnaeus, , and mollusks that were closely related or 1758. identical to the present Baikal species. The most

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‘recent’ part is North Baikal which appeared only catches decreased during the same period from some 6 million years ago. The process of mountain 8000 t to 1500–2000 t (Popov, 1958) but stabi- uplift started around Baikal only 3.5 million years lized at levels of 5000–6000 t in 1930s. During ago. This orogenesis led to significant water level the Second World War, Omul catches increased fluctuation in the lake and changes of its drainage to 8000–9000 t due to increased fishing activity. pattern. It is considered that Lake Baikal was sub- During that period, the catches of other fishes sequently connected with three great Siberian also increased, and fishing of Baikal cottoid fishes drainage basins: , Yenissey and Amur (Korz- began. In 1945 to 1955, Omul catch was 4000– huyev, 1977). In general, the border of the 5000 t, but by the early 1960s fell to 1700 t. In and Pacific Ocean watersheds were displaced to 1969–1975, Omul fishing was banned, but the the south. At that time, in whole Siberia, and par- Omul stock did not recover and its main popula- ticularly in the Baikal Basin, the warm-water Ter- tion characteristics continued to deteriorate. The tiary ichthyofauna became extinct and was restrictive measures led to increased fishing of replaced by a complex of cold-water species of other valuable fish species, in the first place, sal- Arctic and Boreal origin. The only species that has monids, whitefishes, and Burbot, during their pre- persisted from the Tertiary period up to present is spawning and spawning periods at mouths of riv- the Baikal subspecies of . Cold- ers. Such practices quickly destroyed the fishery water species such as Taimen Hucho taimen stocks of these species and from mid-1980s on (Pallas, 1773) and Lenok Brachymystax lenok they did not play any significant role in the fish- (Pallas, 1773) probably appeared in Lake Baikal ing industry. At present, the most important com- as late as the Pliocene; the most probable time of mercial fish species are still Omul, as well as the appearance of Burbot Lota lota (Linnaeus, Black Baikal Grayling, Pike, Roach, Burbot, 1758), graylings and whitefishes is Pliocene– Perch, and some cyprinids that are rare or spo- Pleistocene. radic in the lake but caught in lower reaches of The most uncertain timing is that of the pene- rivers. In recent years, introduced Amur Catfish tration of the ancestors of modern cottoids and for- and Bream Abramis brama (Linnaeus, 1758) mation of their diversity in the Baikal Lake. Berg have appeared in commercial catches in small (1922) and Sideleva (1994) hypothesized that the amounts. Commercial fishing for the following Baikal cottoids are ancient autochthonous forms. species is forbidden: Baikal Sturgeon, Taimen, Another theory was proposed by Dorogostayskiy Lenok, White Baikal Grayling, and whitefishes. (1923) and worked out in details by Taliev (1955). These species are included in Red Book data at It emphasizes the evolutionary youth of the various administrative levels. endemic cottoid fishes and their rapid in The total official catch fluctuates from 2700 to the lake. New genetic and molecular data (Kiril- 3450 t, of which up to 65–70% is Omul. Another chik et al., 1995) support the latter hypothesis con- 20–25% is Roach. The total catch of all other spe- firming the age of cottoid diversification as cies does not exceed several tens of tonnes. Expert recently as about 2 million years only (Timoshkin, estimates of illegal catches, at the same time, are 2001). put at about 700 t. Hatchery activities are in place for Baikal Stur- geon, Omul, Baikal Lake Whitefish, Black Baikal Fishery and fish-farming Grayling and White Baikal Grayling. There are three still functioning fish farms located on a tribu- Fishing in Lake Baikal has had a long history tary to Posolskiy Sor (Bol’sherechenskiy hatchery beginning in the 17th Century. The primary farm), on River Selenga (Selenginskiy hatchery objects of the commercial fishery were Baikal farm), and River Barguzin (Barguzinskiy hatchery Sturgeon and Omul. The first fishing regulations farm) (Figure 1). The main species that is artifi- appearing at the end of 19th Century limited net cially reared is Omul. In 1920s, K. N. Panteleyev fishing of Omul in the pre-spawning period; these first experimented with keeping breeders in cages, regulations had very little effect. At the beginning artificial spawning and egg incubation at a fish of the 19th Century, the sturgeon catch was as farm in Ulan-Ude; since then, number of individu- high as 200 t in some years, and declined to only als released increased to 120 million in 1930s and 10 t in the 1920s (Egorov, 1961, 1985). Omul 2.53 billions in 1998 (50% of this being produced

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at Bol’sherechenskiy, 43% at Selenginskiy and 7% monoxide, 2000 t of nitrogen oxides). In 1990s, at Barguzinskiy hatchery). In the 2000s, the aver- the release of specific substances (dimethyl sul- age number of fry released was 1.26 billions per fide, methyl mercaptan, hydrogen sulfide, , year. Artificial spawning of Baikal Sturgeon was etc.) decreased by 10–15% (State Report on Envi- started in the middle 1980s when a dramatic ronmental Conditions in Irkutsk Province in 1998, decline of this species occurred. It is based at the 2000). In general, since then a clear trend toward experimental hatchery station on River Selenga reducing atmospheric emissions of pollutants has (Selenginskiy), the main spawning river for stur- been seen in the region. The most prominent sour- geon in the Baikal basin. From 1996 to 2002, ces of liquid sewage are the major pulp and paper 4.36 million sturgeon fingerlings were released production plants in the south of Lake Baikal and into the lake. Artificial rearing of Baikal white- in the lower reaches of the Selenga River. For fishes and White Baikal Grayling was also begun example, in 1998, dumping partially-treated waste at the Selenga hatchery. It is still in the experimen- water from sewage treatment plants at BPPM tal stages and the releases are only 15 million amounted to 33.7 106.m3, and the lake received whitefishes and 250,000 Grayling per year. 146 t of lignin, 12 t of nitrates, 4215 t of sulfates, and 1596 t of chlorides. As with air pollution in recent years, there has been a significant reduction State of environment: of liquid pollutants as a result of equipment upgrades at treatment facilities. Main habitats, water quality and Increased coastal erosion caused by the higher anthropogenic changes water level on Lake Baikal, necessitated large- scale bank protection works at the south-east coast During the second half of the 20th Century, the where the Trans-Siberian Railway passes. These pace of economic development activities in the measures had negative impacts on the coastal basin of Lake Baikal grew dramatically. In the spawning sites of Baikal Yellowfin Sculpin. Also, early 1960s, habitats of many Baikal aquatic beginning in the mid-1970s, the Baikal ecosystem organisms were seriously impacted by the rise in was strongly negatively impacted by the construc- the water level (about 1 m) and changes in the nat- tion of the Baikal-Amur Railway close to the shore seasonal fluctuations due to construction of of the northern part of the lake, due to large-scaled the Irkutsk Hydroelectric Power Station on the straightening of the shoreline, deforestation, tim- Angara River, 70 km downstream from its outflow ber floating, and a sharp rise in water freight trans- from the lake. Based on this source of cheap port with concomitant release of oil products into power, an industrial area was developed (the the lake. The population growth was associated Irkutsk-Cheremhovsky Industrial Region), which with increased illegal harvesting of valuable fish then became a source of significant pollution from species to levels thought to be as great as the offi- aluminum, pulp and paper and chemical industries, cial (legal) catch. as well as from further power plants that burn solid All these categories of human activity had the fuels. However, because of regional topography greatest impact on biota of the littoral zone, where and prevailing wind directions, contamination the highest number of endemic species occurs, from the industrial region have had less impact especially at the higher levels of the food chain in than that from industrial enterprises located in the the ecosystem of Lake Baikal – the fish and seal Baikal Depression proper. The greatest negative populations. For example, one of the consequences impacts have arisen from the air emissions of sul- of the lake water level increase was a drastic drop fur dioxide, nitrogen oxides, sulfur-and hydrocar- in numbers of Baikal Yellowfin Sculpin, an impor- bon, methyl mercaptan, formaldehyde, and tant part in the Baikal food chain and an important phenol. The biggest source of air pollution of the food source of Omul, a species that had long been environment located directly on the shores of Lake a dominant component of the fish fauna and the Baikal is the Baikal Pulp and Paper Mill (BPPM). fishery in Lake Baikal. Due to the rise in water The total annual emissions of air pollutants from level, spawning sites of Baikal Yellowfin Sculpin stationary and mobile sources at Baikalsk, where were placed beyond the favored surf zone. In addi- the plant is located, are 8100 t (2100 t of solid sub- tion, southern lake shores along the Trans-Siberian stances, 2000 t of sulfur dioxide, 200 t of carbon Railway were reinforced with solid concrete

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constructions, and local people started collecting Currently, the coordinating role is provided by Yellowfin eggs for consumption. The joint impact the Scientific Fishery Council affiliated at FGU of these factors led to the disappearance of several “Baykalrybvod,” which draws representatives generations of Baikal Yellowfin Sculpin. As a from these organisations and a number of research result, yellowfin catches decreases in 1970s to 10 t institutions. The membership of the Council is comparing to 1400 t in the 1950s. By the middle approved by the Main Department on Protection, 1980s, the numbers of yellowfin had Reproduction of Fish Resources and Fishery Reg- decreased dramatically—by a factor of 300 (Mat- ulation (“Glavrybvod”) of the Committee on Fish- veyev et al., 2008). In turn, this caused a decline eries of Russian Federation. of Omul numbers of at least two times and of its General coordination of scientific research and biological characteristics as well because Baikal consultations regarding strategic decisions on the Yellowfin Sculpin was the main food item for use of Baikal biological resources were previously Omul. In Omul diet, Yellowfin contributed up to implemented by East-Siberian Branch of Ichthyo- 38% in the 1950s, 12% at the end of the 1960s, logical Commission of the Ministry Fisheries of and only 0.2% in the 1970s (Toporkov, 1987). the USSR (now the Interdepartmental Ichthyologi- Beginning in the 1980s, there was some evidence cal Commission of Russian Academy of Science, of improvements in the biological characteristics Ministry of Agriculture, and State Fishery Com- of the stocks with increasing of numbers of eggs mittee). However, the activities of the Commission laid and of Yellowfin young-of-the-year in the have not been effective in recent years. upper water layers. It appears that the Yellowfin Fishing in Lake Baikal is governed by the Rus- stock is slowly coming out of the depression state sian Federation Law “Protection of Lake Baikal,” that began at the end of 1960s. In the late 1980s, which was adopted in May 1999. The State Reso- the contribution of Yellowfin increased in the lution “On peculiarities of protection and harvest- Omul diet. Omul again became more abundant in ing of endemic water animals and gathering of the southern part of Baikal where it had been only endemic water plants in Lake Baikal” (No. 67, 28 occasionally caught in the 1970s. In 1995–2004, January 2002) is also in effect. Annual TACs are concentrations of foraging Omul become compa- adopted on the base of a scientific forecast devel- rable to those in the 1950s, and Baikal Yellowfin oped by the “Vostsibrybcenter” using the State’s Sculpin young-of-the-year comprised up to 30%– ecological expertise. Recreational fishing is 90% of Omul’s ration by mass. licensed; the licenses are distributed by the local fishery authorities. Data from “Vostsibrybcenter” shows that, for example, in 2003, the total recrea- Fishery management tional fishing catch was 170 t, including 115 t of Omul, 17 t of Roach, 12 t of Perch, and 10 t of Management of the Baikal fishing industry is Siberian Dace. To ensure of the sustainability now implemented by three independent organiza- of the fisheries, a special law of the Government tions: (1) Baikal Basin Department on Protection, of the Russian Federation (No. 234, 26 March Reproduction of Fish Resources and Fishery Reg- 2001) established allowable limits of the water ulation (Federal State Institution FGU level fluctuations in Lake Baikal of 1 m between “Baikalrybvod”) whose main duties are protection, 457 and 456 m (according to the Pacific System of regulation of fishing and inspection of biological Elevations). In 2011, for example, the water level resources use; (2) East-Siberian Scientific-Produc- was at 456.28 m (Pacific system). tion Center of Fishery (FGUP “Vostsibrybcenter”) The main direction of work to maintain and which assesses the size of fish stocks, develops restore the fish stocks of Lake Baikal was to max- fishery forecasts (i.e. the total allowable catch, imize the Omul fishery. This fisheries manage- TAC) and provides research support for fishery ment strategy has had both positive and negative industry and fish farming; (3) Union of Fishing consequences. Persistent decreases in Omul com- Organisations (ZAO “Baikalrybhoz”) that per- mercial catches in the first half of the 20th Cen- forms fishing and processing of fishes and Baikal tury became the main driving force for the Seal. intensification of efforts to create technologies for These organisations were part of the Ministry artificial reproduction. The main idea was to of Fisheries of the RSFSR and the USSR. maintain a high abundance of natural spawning

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populations through contracted hatcheries at all waterbodies of the Baikal basin including Lake spawning rivers. Hatcheries on tributaries of Pos- Baikal. At present, Amur Sleeper inhabits the delta olskiy Sor, Chivyrkuy Bay, Maloye Morye Strait, of River Selenga, all shallow-water bays along the Selenga and Barguzin rivers were built while that lake eastern coast, coastal areas of the southern on the Angara was only planned. The size of the part of the lake, and some locations along the Omul populations in the Chivyrkuyskiy Bay and western coast. In some locations at the eastern the Maloye Morye was small and could not pro- coast its abundance, in 1990s, reached 75–90 vide enough breeders to sustain the hatcheries; specimens per m2 (Pronin et al., 1998), and it so, those hatcheries were closed. Similar situa- became a competitor of native shallow water tions were repeated at the Selenginskiy and Bar- fishes, especially roach and dace (Litvinov and guzinskiy fish farms. As a result only half of their Gorman, 1996). At the same time, Amur Sleeper facilities are used for re-stocking population of became a common part of the food diet of Pike, the rivers where they are located. Other parts of Perch, sometimes White Baikal Grayling, and their facilities are used to incubate eggs from the piscivorous birds. In recent years, a marked down- Posolskiy Sor population. Hence, the Posolskiy ward trend in the number of Amur Sleeper and a Sor Omul became the main reared population reduction of its distribution in the lake has been though the fingerlings are released, in part, into recorded. As to deep water habitats in Baikal, little the lake areas non-native for this population. The is known. Posolskiy Sor Omul is a benthic deepwater Modes of fishing, fishing gear, protective meas- dweller inhabiting mainly depths from 200 to ures, and hatchery activity were mainly focused 400 m. Thus, re-stocking of this population has on Omul. The young of other species of fishes, been ineffective as it cannot be fished at those similar in size to Omul, were caught alongside depths. As a result, there is now an imbalance in Omul with no special consideration of additional size of the different Omul populations and the pressures on populations of these species. The reproductive state of their spawning stocks. long-term depression of species other than Omul Another direction for “improvement” of com- has given rise to the viewpoint that those species mercial fish stocks was the introduction of new have no commercial value in the lake (Mamontov, exotic species as it had been thought that native 1996). Thus, Omul fishing in Lake Baikal has fishes did not use all the food resources of the acquired the characteristics of a monoculture. lake. Introduction of new species, mainly of com- mercial value, began in the early 1930s. Up to 1970s, introductions were made without any effort Nature protection and to account for specific features of the Baikal fish conservation communities or the Baikal ecosystem in general, and without any long-term predictions. Altogether, The present state of fisheries and fish stocks thirty fish species were proposed for introduction, described above has arisen from the negative and sixteen were introduced (Mamontov, 2001). impacts of economic activity in the Baikal basin. Most of these introductions failed. Only five non- In the 1980s, these changes become obvious to all native fish species (Peled, Amur Carp, Amur Cat- those involved in research on the Baikal ecosys- fish, Bream and Amur Sleeper) became estab- tem. This situation made necessary specific efforts lished; all of them are caught periodically in the to make improvements. In the late 1980s and early lake. The Amur Carp stock reached a commer- 1990s due to the political changes occurring in cially valuable level up to the end of the 1970s, Russia, a movement formed to protect Lake though later its catches declined to a level so that Baikal, which brought together scientists and a official records are not kept. Although the present wide range of the general public. As a result, Lake numbers of Peled, Bream, and Amur Catfish are Baikal was added to the UNESCO list of World also very low, these fishes are still included in the Heritage sites in 1996. This act along with ratifica- TACs for Lake Baikal. tion by Russia of the Convention on Amur Sleeper was unintentionally introduced became the basis for the “Strategy on Biodiversity along with Amur Carp into a lake in the Selenga Conservation in the Baikal ” river drainage. From there it expanded its range which was supported by the World Bank. A full rapidly and became established in a number of range of activities to preserve the Baikal

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ecosystem was elaborated in the sub-programme However, it is clear that besides the develop- “Protection of the Lake Baikal and Baikal Natural ment priorities for Lake Baikal uses such as Territory” of the National Programme “Ecology increased tourism and recreation, or as a source of and Natural Resources of Russia (2002–2010), hydropower and drinking water, fisheries are which was approved by the Government of Rus- becoming less significant. Traditionally extensive sian Federation in 2001. fishing methods, their low efficiency, large-scale According to the national law “On Protection of poaching, along with the prospects for the reorien- Lake Baikal,” the Baikal natural territory is selected tation of economic activities in the Central zone of with an area of 30.7 million hectares. This area is the Baikal Natural Territory in accordance with subdivided into the following zones: (1) The Cen- the principles of the Convention “On the Global tral zone that includes Lake Baikal, all its islands Natural and Cultural Heritage,” make it necessary and adjacent protected areas, where the main pro- to implement several urgent actions including the tective activity is conservation of rare and endemic reorganization of fisheries and measures to species and their habitats; (2) the Buffer zone that improve the lake’s ecosystem. overlaps the whole Baikal basin (except the lake Reorganization of fisheries proposals include: proper, or the Central zone) in the Russian Federa- (1) changes in fishing strategy aimed at managing tion, where the main tasks here are protection and the intensity of fishing instead of catch volume regulation of the traditional uses of the natural limitations; (2) more effective security measures resources (grassland farming and fishing), elimina- on the part of fishing and local authorities; (3) opti- tion of timber logging, and reduction of other indus- mization of the technology and scale of artificial trial impacts; (3) the Zone of Atmosphere influence propagation of Omul and other valuable fish spe- that includes the territory out of the Baikal basin cies. Today’s excessive emphasis on artificial within the range of 200 km from the lake towards reproduction of Omul does not provide the neces- the west and the north-west from the lake encom- sary economic and environmental benefits while, passing those areas where some industrial plants at the same time, significantly altering the natural are located that could have negative impacts on the structure of Omul populations and their ecological Baikal ecosystem, where the main tasks are to mini- relationships in the lake. It appears that preserving mize influences of the hydro-construction on the or expanding artificial reproduction, aimed at max- water level in Baikal, decrease the air pollution imizing the catch on the one hand, and maintaining from industrial plants, and prevent negative impacts the natural population structure of Omul on the from the planned industrial activities (e.g. the con- other hand, is compatible. It is necessary to limit struction of the power transmission lines to other the scale of artificial Omul reproduction to levels countries). At present, the new legislation has been sufficient to maintain the rate of natural reproduc- passed framing these basic strategies, but imple- tion, and to significantly extend the capacity of mentation has been very slow. hatcheries for other valuable fish rearing, in partic- There are also some special protected areas in ular, of Baikal Sturgeon, Taimen, Lenok, white- the Central zone. There are three state nature fishes, and graylings. Such measures will not only reserves (Barguzinskiy, Baikalskiy, Baikalo-Len- restore the natural structure of fish communities, skiy), two national parks (Pribaikalskiy, Zabaikal- but will provide an opportunity to greatly intensify skiy), and also four wildlife sanctuaries. All are sport fishing. There is evidence, that recreational located on the shores of the Baikal, but aquatic fishing as part of the tourism industry can have a areas are not included in any of them. All special greater economic impact than traditional fishing. protected areas currently only serve to protect ter- Any introduction of non-indigenous fish species restrial ecosystems around the lake. to the basin of Lake Baikal is entirely forbidden at present, and a prohibition of incubation of eggs of non-native species in Baikal hatcheries has been Conclusions recommended. Protection of the environment should be con- Fishing has been the traditional focus of man- sidered as another important activity. The priori- agement on Lake Baikal and its modernization ties in this area include: (1) stabilization of the needs to be considered as part of the economic lake level regime and its seasonal variation within development plans for the Baikal region. the limits that are close to natural regime; (2)

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relocation of enterprises now operating directly on Litvinov, A.G., O’Gorman, R., 1996. Biology of Amur Sleeper the shores of Lake Baikal; (3) reduction of the neg- (Perccottus glehni) in the Delta of the Selenga River, Bury- ative impacts coming from industrial, agricultural atia, Russia. Journal of Research 22(2), and household pollution in the basin; (4) in the 370–378. Mamontov, A.;. 1996. Reproduction, hybridization, and artifi- future, complete elimination of pollution loads cial stocking of Baikal whitefishes. In: O.M. Kozhova, P. into the lake; (5) reduction of the impacts of con- Ya. Tugarina (Eds.), Ichthyological investigations of Baikal struction and upgrading of shore facilities and Lake and its basin at the end of the 20th century, pp. transport routes, of which some level of develop- 41¡48. (In Russian). Irkutst State University, Irkutsk. ment is inevitable. Mamontov, A.M., 2001. Acclimatisation of fishes in the Baikal. Trudy Kafedry Zoologii Pozvonochnykh, Irkutsk State Uni- versity 1, 134–239. (In Russian). References Matveyev, A.N., Samusenok, V.P., 2006. Biodiversity and structure of fish community in mountainous water bodies of Bazikalova, A.Ya., Vilisova, I.K., 1959. Feeding of benthofa- the . Bulleten’ VSNTs SO RAMN, 2, 84– gous fishes of Maloye Morye. Trudy ’skoy Limnolo- 91. (In Russian). gicheskoy Stantsii AN SSSR 17, 382–497. (In Russian). Matveyev, A.N., Pronin N.M., Samusenok, V.P., Bobkov, A.I., Berg, L.S., 1909. Fishes of Amur Basin. Zapiski Zoologiche- Sokolov, A.V., 2008. Fauna, atlas with identification skogo Instituta Imperatorskoy Akademii Nauk, Seriya 8, keys, and fish resources of Lake Baikal. BNTs SO RAN, 24, 9, 1–270. (In Russian). Ulan-Ude. (In Russian). Berg, L.S., 1922. Fauna of the Baikal and its origin. In: Climate Popov, P.F., 1958. Historical review of fisheries in the Baikal. and Life, pp. 28–53. Gosizdat, Moscow. (In Russian). In: Fish and Fisheries in the Lake Baikal Basin, pp. 33–42. Domysheva, V.M., 2009. Hydrochemistry. In: A. K. Tulokho- OGIZ, Irkutsk. (In Russian). nov (Ed.), Baikal: nature and people. Encyclopedia, pp. Pronin, N.M., Selgebi, D.Ch., Litvinov, A.G., Pronina, S.V., 45¡70. Ekos, Ulan-Ude. (In Russian). 1998. Comparative ecology and parasitic fauna of exotic Dorogostayskiy, V.Ch., 1923. Vertical and horizontal distiribu- species introduce in Great Lakes of the World: Perccottus tion of the fauna of Lake Baikal. Trudy Profesorskogo i Pre- glehni in Lake Baikal and Gymnocephalus cernuus in Lake podavatel’skogo Sostava Irkutskago Gosudarstvennogo Superior. Sibirskiy Ecologicheskiy Zhurnal, 5, 397–406. (In Universiteta 4, 1–31. (Works by professors and lecturers of Russian). the Irkutsk State University. In Russian). Ravens, T.M., Kocsis, O., Wuest,€ A., Granin, N., 2000. Small- Efimov, A.I. (Ed.), 1970. Hydrogeology of the USSR. Volume scale turbulence and vertical mixing in Lake Baikal. Lim- 22. Buryatskaya ASSR. Nedra, Moscow. (In Russian). nology and Oceanography 45(1), 159¡173. Egorov, A.G., 1961. Baikal Sturgeon Acipenser baeri stenor- Shimaraev, M.N., Domysheva, V.M., 2012. Trends in hydro- hynchus natio baicalensis Nikolski. Buryatskoye Knizh- logical and hydrochemical processes in Lake Baikal under noye Izdatel’stovo, Ulan-Ude.(In Russian). conditions of modern climate change. In: C.R. Goldman, Egorov, A.G., 1985. Fishes of water bodies of the southern M. Kumagai, R.D. Robarts (Eds.), Climatic Change and West Siberia (Petromyzontidae, Acipernseridae, Salmoni- Global Warming of Inland , pp. 43–66. John Wiley dae, Coregonidae, Thymallidae, Esocidae). Irkutsk State & Sons, Ltd., Chichester, UK. University, Irkutsk. (In Russian). Shimaraev, M.N., Granin, N.G., Zhdanov, A.A., 1993. Deep Kartushin, A.I., 1958. Biology of Siberian roach, dace, ide and ventilation of Lake Baikal waters due to spring thermal crucian carp in the system of Lake Baikal. In: Fish and bars. and Oceanography 38(5), 1068¡1072. Fisheries in the Lake Baikal Basin, pp. 334–380. OGIZ, Sideleva, V.G., 1982. The lateral-line system and ecology of the Irkutsk. (In Russian). Baikal sculpins (Cottoidei). Nauka, Novosibirsk (In Knizhin, I.B., Bogdanov, B.E., Matveyev, A.N., Samusenok, V. Russian). P., 2004. Fishes of Lake Baikal and its basin. Irkutskiy Uni- Sideleva, V.G., 1994. Speciation of endemic Cottoidei in Lake versitet, Irkutsk. (In Russian). Baikal. Archiv fur€ Hydrobiologie–Beiheft Ergebnisse der Kiril’chik, S.V., Slobodyanyuk, S.Ya., Belikov, S.I., Pavlova, Limnologie 44, 441–450. M.E., 1995. Phylogenetic relationships of 16 species of cot- Sideleva, V.G., 1998. Cottidae, Comephoridae, Abyssocottidae. toid fishes of Lake Baikal based on the analysis of nucleo- In: Annotated check-list of Cyclostomata and Fishes of the tide sequence of a fragment of the cytochrome b gene of continental waters of Russia, pp. 149–158. Nauka, Moscow. mitochondrial DNA. Molekulyarnaya Biologiya 29 (4), (In Russian). 817–825. (In Russian). Sideleva, V.G., 2001. List of fishes from Lake Baikal with Korzhuyev, S.S., 1977. Morpho-structural analysis of river val- descriptions of new taxa of cottoid fishes. In: A.V. Neelov leys and hydroelectric construction. Nauka, Moscow. (In (Ed.), New contributions to freshwater fish research, pp. Russian). 45–79. Zoological Institute of RAS, St. Peterburg. Kozhov, M.M., Misharin, K.I., 1958. Systematic composition Sideleva, V.G., 2003. The endemic fishes of Lake Baikal. Back- of the fish fauna of Lake Baikal and its basin. In: Fish and huys Publishers, Leiden. Fisheries in the Lake Baikal Basin, pp. 91–100. OGIZ, Skryabin, A.G., 1969. Biology of Baikal whitefishes. Nauka, Irkutsk. (In Russian). Moscow (In Russian).

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Skryabin, A.G., 1979. Coregonid fishes of southern Siberia. object of Baikal omul and other fishes. In: A.G. Egorov Nauka, Novosibirsk. (In Russian). (Ed.), Investigations of fishes of East Siberia, pp. 18–26. Smirnov, V.V., Shumilov, I.P., 1974. of the Baikal. Irkutskiy Universitet, Irkutsk. (In Russian). Nauka, Novosibirsk. (In Russian). Tugarina, P.Ya, 1981. Graylings of Baikal. Nauka, Novosi- State Report on Environmental Conditions in Irkutsk Province birsk. (In Russian). in 1998. 2000. Irkutsk. (In Russian). Ustyuzhanina, L.A., 1971. Feeding and feeding relationships Taliev, D.N., 1955. Sculpins of Baikal (Cottoidei). Izdatel’stvo between benthophagous fishes. In: Proceedings of Limno- Akademii Nauk SSSR, Moscow-Leningrad. (In Russian). logical Institute, Siberian Branch of the Russian Academy Timoshkin, O.A., 2001. Index of animal species inhabiting of Sciences. Limnology of deltaic areas of Baikal (Selenga Lake Baikal and its catchment area. Vol. 1. Lake Baikal. region) 12, 132–157. Nauka, Leningrad. (In Russian). Book 1. Nauka, Novosibirsk. (In Russian). Vereshchagin, G.Yu, 1935. Two types of biological complexes Toporkov, I.G., 1987. Influence of the Baikal water level in the Baikal. Trudy Baykal’skoy Limnologicheskoy Stant- increase on resources of yellowfin sculpin as the main food sii AN SSSR 6, 199–212. (In Russian).

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