ISSN 00329452, Journal of Ichthyology, 2015, Vol. 55, No. 4, pp. 497–526. © Pleiades Publishing, Ltd., 2015.

Species Structure of Pelagic Ichthyocenes in Russian Waters of Far Eastern Seas and the Pacific Ocean in 1980–2009 O. A. Ivanova and V. V. Sukhanovb a

Pacific Research Fisheries Center (TINRO center), ul. Shevchenko 4, Vladivostok, 690950 Russia b Institute of Marine Biology, Far Eastern Branch, Russian Academy of Sciences, ul. Pal'chevskogo 17, Vladivostok, 690059 Russia email: [email protected] Received November 28, 2014

Abstract—Based on the results of pelagic trawl catches in 1980–2009 a general list of ichthyofauna in Russian waters of the Far Eastern seas and the Pacific Ocean comprise 450 species, among which 114 species are iden tified in the Sea of Japan, 258 species in the Sea of Okhotsk, 170 species in the Bering Sea, and 319 species in the Russian waters of the Pacific Ocean. The species composition of the ichthyofauna was classified according to taxonomical, biotopical, and zoogeographical status of the species. The Simson’ index of dom inance varies in a narrow range from 1.58 to 2.66, and the Pielou’s evenness index has low values: 0.17 (Sea of Japan and Sea of Okhotsk), 0.19 (Russian waters of the Pacific Ocean), and 0.27 (Bering Sea). Values of the indexes are confirmed by the conclusion about the dominance of one or two to three species in the sur veyed ichthyocenoses made according to the analysis of profiles of rank curves. The highest species diversity (Margalef’s index) is observed in Russian waters of the Pacific Ocean and the lowest diversity is in the Sea of Japan. When comparing qualitative and quantitative composition of ichthyofauna in respect to regions, the lowest index of diversity is found for the fauna in the Sea of Japan. The dynamics of biomass (tons/km2) of all components of pelagic ichthyocenoses is observed according to the periods of years and their species structure is determined. The average longterm density of pelagic fauna concentrations in the area under survey con stitute 16.8 tons/km2 and its total resources are 70–80 million tons. Keywords: pelagial, ichthyocene, composition, structure, biomass, diversity, rank curves, Far Eastern seas, Pacific waters of Russia DOI: 10.1134/S0032945215040037

INTRODUCTION Since the beginning of the 1980s, comprehensive ecosystem studies, largescale monitoring of the state of natural communities in the Far Eastern seas and adjacent waters of the Pacific Ocean have been con ducted at the level of macrosystems at the Pacific Research Fisheries Center (TINRO center). Over a 30year period of studies conducted in this vast area (more than 5 mln km2), the unique data (which were obtaining according to the uniform standard method) on the composition and structure of pelagic commu nities of hydrobionts was obtained. The applied char acter of studies made it possible to obtain such a great volume of data for a longterm period. On the one hand, these data were efficiently used by the specialists of TINRO for prediction of catches of commercial fishes, thus fulfilling the task for fish industry. On the other hand, they were used as the basis of fundamental generalizations that predicted ecosystem reorganiza tion (Shuntov et al., 1993, 2007; Volkov, 1996; Borets, 1997; Shuntov, 2000, 2001, 2004, 2012; Dulepova, 2002; Ivanov and Sukhanov, 2002; Chuchukalo, 2006; Volvenko, 2009; Sukhanov and Ivanov, 2009; Shuntov and Temnykh, 2011, 2013). These generalizations

added much and changed our conception of biota and functioning of hydrobiont communities in the Far Eastern seas of Russia and adjacent waters of the Pacific Ocean. In particular, their bio and fish pro ductivity was reconsidered towards their increasing values. The main generalizing conclusion in the work of Shuntov and Temnykh (2013) is that marine macro ecosystems of the main fishing basin of Russia in a normal functional state will preserve their dominant fishing importance in the predictable prospect. Thus, we have a large amount of unique materials on the composition and abundance of ichthyofauna of the Far Eastern seas of Russia and adjacent waters of the Pacific Ocean. The aim of this work is to system atize the materials with regard to taxonomical, zoo geographical, and biotopical status of the species to detect features of the species structure of ichthy ocenes. MATERIALS AND METHODS In our study, we used the data on the species com position and abundance of ichthyofauna in catches by pelagic trawling that were obtained during expeditions

497

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(a)

(b) 60° 58°

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44° 128°

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62°

46°

60°

44° 58° 42° 55° 40° 54° 165°

170°

175°

180° 175° E

170° W

144°

148°

168° E

Fig. 1. Schematic map of the surveyed region with points of trawling (⋅): (a) the northwestern part of the Sea of Japan, (b) the Sea of Okhotsk, (c) the western part of the Bering Sea, and (d) Pacific waters of Kamchatka and Kuril Islands.

in the Russian economic zone in the northwestern part of the Pacific Ocean (Sea of Japan, Sea of Okhotsk, and Bering Sea, Pacific waters of Kam chatka and Kuril Islands) in 1980–2009 (Fig. 1). Dur ing trawl surveys, a complex of oceanological and planktonic studies was carried out. This study is based on materials of 273 marine expeditions during which more than 23 000 trawlings were performed (Table 1).

Trawling was performed outside territorial waters (a 12mile zone) with 13 types of variabledepth rope trawls; their technical features and parameters are described in a series of publications (Nekton…, 2003– 2006). All trawls have a smallmeshed (10–12 mm) liner inserted in the trawl bag along the last 12–15 m. Trawling (59.7%) was mainly performed in the upper layer of the epipelagial (0–70 m). The portion of epi JOURNAL OF ICHTHYOLOGY

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Table 1. Regions and dates of sampling Number Region

Period of observation

Northwestern part of the Sea of Japan Sea of Okhotsk Western part of the Bering Sea Pacific waters of Kamchatka and Kuril Islands

10.02.1981–26.11.2003 27.02.1980–20.11.2009 20.05.1982–11.10.2009 24.12.1979–20.10.2009

of surveys

of trawlings

32 76 40 124

2110 8754 3756 8770

pelagial trawling was 79.2% and mesopelagial trawling constituted 20.8%. The vertical opening of the trawl averaged 43.26 ± 0.07 m, and the horizontal opening was 41.11 ± 0.08 m. The depth of the movement of the upper edge of the trawl ranged from 0 to 1230 m. The taxonomical status of fishes and fishlike animals was determined when catches were sorted on board of the vessel, however, when the species identification was too difficult, qualified experts identified animals in laboratory conditions on the coast. Some species or taxonomical ranks identified in field conditions were considered dubious and, after critical analysis, the species were dropped from the list or attached to another taxonomical group (if the collection material was available). Higher taxa of fishes and fishlike ani mals correspond to the Nelson system (Nelson, 2006), and their species names are presented according to Eschmeyer (Eshmeyer, 1998), taking into account the latest upgrading (with the exception of walleye pol lock; its species name Theragra chalcogramma remained unchanged). The species composition of the ichthyofauna was systematized according to biotopical (in respect to vertical zonation of pelagic and benthal biotopes) and zoogeographical (range type) status of the species. Latitudinal typization of the ranges was made by subdivision of planetary latitudinal zones of the northern hemisphere into arctic, boreal, and trop ical zones (Perestenko, 1982) and by subdivision of each of these zones into two subzones (high and low Arctic, high and low boreal, subtropical and tropical subzones). The correspondence of the meridional component to the range type is determined according to the published works (Willis et al., 1988; Fedorov and Parin, 1998; Fedorov, 2000; Sheiko and Fedorov, 2000; Tokranov, 2010).

and stabilization of fish productivity due to the increase in walleye pollock abundance.

The subdivision of the primary materials into three periods is based on the concept of reorganization in the ecosystems of the Far Eastern seas (Shuntov et al., 2007). The first period, 1980–1990, corresponds to the maximal productivity of fish caused by the peak of the abundance of Japanese sardine Sardinops melanos tictus and a high abundance of walleye pollock. The second period, 1991–1995, is marked by a sharp decrease in fish productivity when walleye pollock in part and Japanese sardine completely lost their influ ence on the structure of pelagic ichthyocenes. The third period, 1996–2009, is marked by the increase

Standard statistical parameters, such as an average value, mode, standard error, and standard deviation, were used for description of quantitative characteris tics. Classification of the Far Eastern seas and adjacent waters of the Pacific Ocean on the basis of the similar ity of the species composition and the species structure of the ichthyofauna was made by the method of multi dimensional scaling (Guttman, 1968; Ivanov and Sukhanov, 2002). The Pearson linear correlation coef ficient is a measure of a quantitative relationship between the variables. The coefficient of Sörensen Chekanovsky was used as a measure of similarity for

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The biomass of each component of the ichthy ocene was determined by the aerial method (Aksyutina, 1968). All estimates of the abundance of species or groups of species are given with account for the coefficients of catching efficiency for adult speci mens without differentiation of the coefficients according to sizeage groups of fishes as was done for some species in the published works (Nekton…, 2003– 2006). The appropriateness of the use of the coeffi cients of catching efficiency has been repeatedly dis cussed in publications (Volvenko, 1998, 1999; Ivanov and Sukhanov, 2002; Lapshin, 2009). In our opinion, the use of the coefficients is necessary when we try to approach the concept of “the species structure of the community” instead of the concept of “the species structure of catches.” The coefficient of catching effi ciency is an integral parameter. Its value depends on the efficiency of fishing gears, biological features of caught organisms (size, physiological state, velocity of movement, response to fishing gears, etc.), and spatial and temporal features of the effect of different envi ronmental factors on organisms. The rank curves were built as follows. The species list was ranked according to descending biomass. The number of the species in the list (rank) was plotted on the axis of abscissa and the logarithm of its abundance was plotted on the axis of ordinates. In publications in English, such constructions are called “curves of the species significance” (Pianka, 1981) and, in Russian publications, they are called “curves of relative domi nance” (Fedorov and Gilmanov, 1980). For short, we call them rank curves.

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0.6

PO

0.4

0.2 JS 0

OS

–0.2 BS

–0.4

–0.6 –0.8

–0.4

0

0.4

Fig. 2. Classification of the Far Eastern seas and adjacent waters of the Pacific Ocean according to similarity of the species composition of the ichthyofauna (explanations are given in the text). Regions: JS—northwestern part of the Sea of Japan, OS—Sea of Okhotsk, BS—western part of the Bering Sea, PO—waters of the Pacific Ocean near Kuril Islands and Kamchatka.

the qualitative comparison of the species composition (Odum, 1971). The elementary parameter, the species richness (number of species in the sample) was a measure of the ichthyofauna diversity. The Simpson dominance index, the Margalef diversity index, and the Pielou evenness index were used when assessing the species diversity of the ichthyocene (Odum, 1971; Magurran, 1991). RESULTS AND DISCUSSION According to the results of trawl catches, 450 spe cies are included in the general list of the ichthyofauna (Table 2): two species of lampreys, 15 species of carti laginous fishes (Chondrichthyes), and 433 of bony fishes (Actinopterigii). The class of bony fishes exceeds other classes of fishes and fishlike animals in the number of species; its portion is 96.2%. The list of species, of course, does not comprise all representa tives of the ichthyofauna of Russian marine and oce anic waters in the Far Eastern region. According to preliminary estimates, the complete list should com prise more than 1000 species of fishes and fishlike animals (Fedorov, 1973, 2000; Fedorov and Parin, 2004; Sokolovskii et al., 2007; Tokranov, 2010; Izmy atinskii, 2014; Parin et al., 2014). Tables 3 and 4 present the summarized data on the taxonomical diversity of the ichthyofauna. On the

whole, the highest number of taxa is recorded in the water area of the Kuril Islands and Kamchatka in the Pacific Ocean. The Sea of Okhotsk, the Bering Sea, and the Sea of Japan follow this area in descending order. The same tendency exists for the taxonomic diversity within classes (except lamprey) and for the species richness in the regions. Pacific waters near Kamchatka and the Kuril Islands are characterized by the highest species richness (319 species). The Sea of Okhotsk (258 species) and the Bering Sea (170 spe cies) follow in the descending order, and the lowest species richness was recorded in the Sea of Japan. In total, when ranking 96 families by the number of species, the families Liparidae (39 species), Myc tophidae (38), and Cottidae (34) take the first three places. It is typical that, according to the number of species, only these three families participate in distri bution of the first rank position among regional subdi visions in the surveyed water area: Cottidae in the Sea of Japan and in the Bering Sea, Liparidae in the Sea of Okhotsk, and Myctophidae in Russian waters of the Pacific Ocean. The families represented in Table 4 (their portion is 33.3% of the total number of families) constitute 80.0% of all species detected in catches. The portion of monotypical families was 42.7% in all regions, 54.5% in the Sea of Japan, 50.9% in the Sea of Okhotsk, 41.5% in the Bering Sea, and 38.1% in the Pacific waters of Russia. The comparison of the qualitative composition of the ichthyofauna has demonstrated that the Sea of Okhotsk and the Bering Sea are the most similar in the species composition, and the Sea of Japan and Pacific waters near Kamchatka and the Kuril Islands are less similar in the species composition. The Sörensen Chekanovsky coefficient of similarity (equal to 1 at the complete coincidence of faunas) was 0.63 in the first case and 0.35 in second case. It should be mentioned that the comparative coefficient marks out the Sea of Japan: in all compared cases, the coefficient of simi larity of the species composition is low and varies in the range 0.35–0.40 (Table 5, Fig. 2). Of the total number of species (450) in the surveyed region, only 42 species (9.3%) were detected in catches in each region. This small list (scaleyey plaice Acanthopsetta nadeshnyi, Pacific sand launce Ammodytes hexapterus, Bering wolfish Anarhichas ori entalis, smooth lumpsucker Aptocyclus ventricosus, Alaska skate Bathyraja parmifera, crested sculpin Blepsias bilobus, salmon snailfish Careproctus rastri nus, Pacific herring Clupea pallasii, Pacific saury Cololabis saira, Pacific navaga Eleginus gracilis, Japa nese anchovy Engraulis japonicus, Pacific cod Gadus microcephalus, threespine stickleback Gasterosteus aculeatus, purplegray sculpin Gymnacanthus detrisus, threaded sculpin G. pistilliger, banded Irish lord Hemilepidotus gilberti, shaggy sea ravon Hemitripterus villosus, flathead sole Hippoglossoides elassodon, salmon shark Lamna ditropis, Pacific daubed shanny Leptoclinus maculatus diaphanocarus, Arctic lamprey JOURNAL OF ICHTHYOLOGY

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Table 2. Taxonomical composition of ichthyofauna in the northwestern part of the Sea of Japan, Sea of Okhotsk, western part of the Bering Sea and Pacific waters of Russia according to the data of pelagic trawl surveys of TINRO center in 1980–2009 Region Number

Taxon

1. 1. 2.

2. 3. 4. 3. 5. 6. 4. 7. 5. 8. 6. 9. 7. 10. 8. 11. 12. 13. 14. 15. 16. 17.

9. 18. 10. 19. 11. 20. 21.

CLASS PETROMYZONTIDA Order Petromyzontiformes Family Petromyzontidae Lethenteron camtschaticum (Tilesius, 1811) Entosphenus tridentatus (Richardson, 1836) CLASS CHONDRICHTHYES Order Lamniformes Family Alopiidae Alopias pelagicus Nakamura, 1935 A. vulpinus (Bonnaterre, 1788) Family Lamnidae Isurus oxyrinchus Rafinesque, 1810 Lamna ditropis Hubbs et Follett, 1947 Order Carchariniformes Family Carcharchinidae Prionace glauca (Linnaeus, 1758) Order Squaliformes Family Squalidae Squalus suckleyi (Girard, 1854) Family Etmopteridae Etmopterus pusillus (Lowe, 1839) Family Somniosidae Somniosus pacificus Bigelow et Schroeder, 1944 Order Rajiformes Family Rajidae Bathyraja aleutica (Gilbert, 1896) B. isotrachys Günther, 1877 B. maculata Ishiyama et Ishihara, 1977 B. parmifera (Bean, 1881) B. tzinovskii Dolganov, 1983 B. violacea (Suvorov, 1935) Rhinoraja taranetzi Dolganov, 1983 CLASS ACTINOPTERYGII Order Albuliformes Family Albulidae Pterothrissus gissu Hilgendorf, 1877 Order Anguilliformes Family Synaphobranchidae Histiobranchus bathybius (Günther, 1877) Family Nemichthydae Avocettina infans (Günther, 1878) Nemichthys scolopaceus Richardson, 1848

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JS

OS

BS

PO

+

+ +

+ +

+ +

+ + + +

+

+

+

+

+

+

+ +

+

+

+ +

+

+

+

+

+

+

+ +

+

+ + +

+ + + + +

+

+

+ +

+

+ +

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Table 2. (Contd.) Region Number

Taxon JS

12. 22. 23. 13. 24. 14. 25. 15. 26. 27. 28. 29. 16. 30. 17. 31. 32. 33. 34. 35. 18. 36. 37. 38. 39. 40. 41. 19. 42. 43. 44. 20. 45. 21. 46. 47. 48. 49. 22. 50.

Family Congridae Conger myriaster (Brevoort, 1856) Gnathophis heterognathos (Bleeker, 1858) Family Serrivomeridae Serrivomer sector Garman, 1899 Order Clupeiformes Family Engraulidae Engraulis japonicus (Temminck et Schlegel, 1846) Family Clupeidae Clupea pallasii Valenciennes, 1847 Etrumeus micropus Temminck et Schlegel, 1846 Konosirus punctatus (Temminck et Schlegel, 1846) Sardinops melanostictus (Temminck et Schlegel, 1846) Order Argentiniformes Family Argentinidae Glossanodon semifasciatus (Kishinouye, 1904) Family Opisthoproctidae Bathylychnops exilis Cohen, 1958 Dolichopteryx longipes (Vaillant, 1888) D. parini Kobyliansky et Fedorov, 2001) Macropinna microstoma Chapman, 1939 Winteria telescopa Brauer, 1901 Family Microstomatidae Bathylagus pacificus Gilbert, 1890 Leuroglossus schmidti Rass, 1955 Lipolagus ochotensis (Schmidt, 1938) Melanolagus bericoides (Borodin, 1929) Nansenia candida Cohen, 1958 Pseudobathylagus milleri (Jorden et Gilbert, 1898) Family Platytroctidae Holtbyrnia innesi (Fowler, 1934) Maulisia argipalla Matsui et Rosenblatt, 1979 Sagamichthys abei Parr, 1953 Family Alepocephalidae Bajacalifornia megalops (Lütken, 1898) Order Osmeriformes Family Osmeridae Hypomesus japonicus (Brevoort, 1856) H. olidus (Pallas, [1814]) Mallotus villosus catervarius Pennant, 1784) Osmerus mordax dentex Steindachner, 1870 Family Salangidae Salangichthys microdon (Bleeker, 1860)

OS

BS

PO + + +

+

+

+

+

+ + + +

+

+

+

+

+

+

+

+

+ +

+

+ +

+ + +

+ + +

+ +

+ +

+ + + + + +

+

+ + +

+ + +

+ + +

+ +

+ +

+

+ +

+ + + +

+

+

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Table 2. (Contd.) Region Number

23. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 24. 61. 62. 63. 64. 65. 66. 67. 25. 68. 69. 70. 71. 72. 73. 26. 74. 75. 27. 76. 77. 78. 79. 80. 81. 82. 83. 84. 85. 86.

Taxon Order Salmoniformes Family Salmonidae Oncorhynchus gorbuscha (Walbaum, 1792) O. keta (Walbaum, 1792) O. kisutch (Walbaum, 1792) O. masou (Brevoort, 1856) O. nerka (Walbaum, 1792) O. tshawytscha (Walbaum, 1792) Parasalmo mykiss (Walbaum, 1792) Salvelinus leucomaenis (Pallas, [1814]) S. malma (Walbaum, 1792) S. kraschennikovi Taranetz, 1933 Order Stomiiformes Family Gonostomatidae Cyclothone alba Brauer, 1906 C. atraria Gilbert, 1905 C. pseudopallida Mukhacheva, 1964 Diplophos orientalis Matsubara, 1940 Gonostoma elongatum Günther, 1878 Echiostoma barbatum Lowe, 1843 Sigmops gracilis (Günther, 1878) Family Sternopthychidae Argyropelecus aculeatus Valenciennes, 1850 A. sladeni Regan, 1908 Maurolicus japonicus Ishikawa, 1915 Polyipnus matsubarai Schultz, 1961 Sternoptyx diaphana Hermann, 1781 S. pseudobscura Baird, 1971 Family Phosichthyidae Ichthyococcus elongatus Imai, 1941 Vinciguerria nimbaria (Jordan et Williams, 1895) Family Stomiidae Aristomias scintillans (Gilbert, 1915) Astronesthes fedorovi Parin et Borodulina,1994 A. indicus Brauer, 1902 Chauliodus macouni Bean, 1890 Ch. sloani Bloch et Schneider, 1801 Flagellostomias boureei (Zugmayer, 1913) Idiacanthus antrostomus Gilbert, 1890 Leptostomias gladiator (Zugmayer, 1911) L. robustus Imai, 1941 Malacosteus niger Ayers, 1848 Melanostomias pauciradius Matsubara, 1938

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+ + + + + +

+ + + + + +

+ + + + + + +

+ +

+

+

+

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+

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+

+ + + + + + +

+ +

+ + + + + + +

+ +

+

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Table 2. (Contd.) Region Number

Taxon JS

87. 88. 89. 90. 28. 91. 92. 29. 93. 94. 30. 95. 96. 97. 98. 99. 100. 101. 102. 31. 103. 32. 104. 33. 105. 34. 106. 107. 108. 109. 110. 111. 112. 113. 114. 115. 116. 117. 118. 119. 120.

OS

Opostomias mitsuii Imai, 1941 Pachystomias microdon (Günther, 1878) Photonectes albipennis (Döderlein, 1882) Tactostoma macropus Bolin, 1939 Order Aulopiformes Family Notosudidae Scopelosaurus adleri (Fedodrov, 1967) S. harryi (Mead et Teylor, 1953) Family Scopelarchridae Benthalbella dentata (Chapman, 1939) B. linguidens (Mead et Böhlke, 1953) Family Paralepididae Arctozenus risso (Bonaparte, 1840) Lestidiops jayakari jayakari (Boulenger, 1889) L. ringens (Jordan et Gilbert, 1880) L. sphyraenopsis Hubbs, 1916 Lestrolepis intermedia (Poey, 1868) L. japonica (Tanaka, 1908) Magnisudis atlantica (Kröyer, 1868) Stemonosudis rothschildi Richards, 1967 Family Alepisauridae Alepisaurus ferox Lowe, 1833 Family Anotopteridae Anotopterus nikparini Kukuev, 1998 Order Myctophiiformes Family Neoscopelidae Scopelengys tristis Alcock, 1890 Family Myctophidae Benthosema suborbitale (Gilbert, 1913) Ceratoscopelus warmingii (Lütken, 1892) Diaphus chrysorhynchus Gilbert et Gramer, 1897 D. coeruleus (Klunzinger, 1871) D. effulgens (Goode et Bean, 1896) D. gigas Gilbert, 1913 D. kuroshio Kawaguchi et Nafpaktitis, 1978 D. metopoclampus (Cocco, 1829) D. perspicillatus (Ogilby, 1898) D. richardsoni Tåning, 1932 D. schmidti Tåning, 1932 D. theta Eigenmann et Eigenmann, 1890 Electrona rissoi (Cocco, 1829) Lampadena luminosa (Garman, 1899) L. urophaos (Coleman et Nafpaktitis, 1972) JOURNAL OF ICHTHYOLOGY

BS

PO +

+ +

+

+ +

+ +

+ +

+ +

+

+

+ +

+

+

+ + + + + + + +

+

+

+

+

+

+

+

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+

+

+

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Table 2. (Contd.) Region Number

Taxon JS

121. 122. 123. 124. 125. 126. 127. 128. 129. 130. 131. 132. 133. 134. 135. 136. 137. 138. 139. 140. 141. 142. 143. 35. 144. 36. 145. 146. 37. 147. 148. 149. 150. 151. 38. 152. 153. 154. 155.

L. yaquinae Paxton, 1963 Lampanyctus festivus Tåning, 1928 L. jordani Gibert, 1913 L. nobilis Tåning, 1928 L. steinbecki Bolin, 1939 L. tenuiformis (Brauer, 1906) L. turneri (Fowler, 1934) Myctophum asperum Richardson, 1845 M. nitidulum Garman, 1899 M. selenops Tåning, 1928 M. spinosum (Steindachner, 1867) Nannobrachium fernae (Wisner, 1971) N. nigrum Günther, 1887 N. regale (Gibert, 1892) Notoscopelus caudispinosus (Johnson, 1863) N. japonicus (Tanaka, 1908) N. resplendens Richardson, 1845 Protomyctophum thompsoni (Chapman, 1944) Stenobrachius leucopsarus (Eigenmann et Eigenmann, 1890) S. nannochir ( Gibert, 1892) Symbolophorus californiensis (Eigenmann et Eigenmann, 1890) S. evermanni (Gilbert, 1905) Tarletonbeania taylori Mead, 1953 Order Lampridiformes Family Lampridae Lampris guttatus (Brünnich, 1788) Family Trachipteridae Desmodema lorum Rosenblatt et Butler, 1977 Trachipterus ishikawai Jordan et Snyder, 1901 Order Gadiformes Family Macrouridae Albatrossia pectoralis (Gibert, 1892) Coelorinchus macrochir Günther, 1877 Coryphaenoides acrolepis (Bean, 1884) C. cinereus (Gibert, 1896) C. longifilis Günther, 1877 Family Moridae Antimora microlepis Bean, 1890 Halargireus johnsonii Günther, 1862 Laemonema longipes Schmidt, 1935 Lepidion schmidti Svetovidov, 1936

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+ + +

+ +

+ +

+

+

+

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PO + + + + + + + + + + + + + + + + + + + + + + +

+

+

+

+ +

+

+

+ + +

+ +

+ +

+ +

+ + + + + + + + +

506

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Table 2. (Contd.) Region Number

Taxon JS

39. 156. 157. 158. 159. 40. 160. 41. 161. 42. 162. 163. 43. 164. 165. 166. 167. 44. 168. 45. 169. 46. 170. 47. 171. 172. 173. 48. 174. 49. 175. 50. 176. 51. 177. 178.

Family Gadidae Boreogadus saida (Lepechin, 1774) Eleginus gracilis (Tilesius, 1810) Gadus macrocephalus Tilesius, 1810 Theragra chalcogramma (Pallas, 1814) Order Ophidiiformes Family Ophidiidae Bassozetus zenkevitchi Rass, 1955 Family Bythitidae Thalassobathia pelagica Cohen, 1963 Order Lophiiformes Family Ceratiidae Ceratias holboelli Kröyer, 1845 Cryptopsaras couesii Gill, 1883 Family Oneirodidae Bertella idiomorpha Pietsch, 1973 Chaenophryne draco Beebe, 1932 Oneirodes bulbosus Chapman, 1939 O. thompsoni (Schultz, 1934) Family Gigantactinidae Gigantactis elsmani Bertelsen, Pietsch et Lavenberg, 1981 Order Beloniformes Family Exocoetidae Cheilopogon pinnatibarbatus japonicus (Franz, 1910) Family Scomberesocidae Cololabis saira (Brevoort, 1856) Order Stephanoberyciformes Family Melamphaidae Melamphaes lugubris Gilbert, 1891 M. suborbitalis (Gill, 1883) Poromitra crassiceps (Günther, 1878) Family Rondeletiidae Rondeletia loricata Abe et Hotta, 1963 Order Beryciformes Family Anoplogastridae Anoplogaster cornuta (Valenciennes, 1833) Family (50) Berycidae Beryx splendens Lowe, 1934 Order Zeiformes Family Zeidae Zenopsis nebulosa (Temminck et Schlegel, 1845) Zeus faber Linnaeus, 1758

OS

BS

PO

+ + +

+ + + +

+ + +

+ + +

+

+

+

+

+ + +

+

+ +

+

+ + + +

+ +

+

+ +

JOURNAL OF ICHTHYOLOGY

+

+

+

+ + +

+

+

+

+ +

+

+ +

+ + Vol. 55

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SPECIES STRUCTURE OF PELAGIC ICHTHYOCENES

507

Table 2. (Contd.) Region Number

52. 179. 180. 53. 181. 182. 54. 183. 184. 185. 186. 187. 188. 189. 190. 191. 192. 193. 194. 195. 196. 197. 55. 198. 56. 199. 200. 201. 202. 203. 57. 204. 205. 206. 207. 208. 209. 210. 211. 212.

Taxon Order Gasterosteiformes Family Gasterosteidae Gasterosteus cf. aculeatus Linnaeus, 1758 Pungitius pungitius (Linnaeus, 1758) Order Sygnathiformes Family Syngnathidae Hippocampus mohnikei Bleeker, 1854 Syngnathus schlegeli Kaup, 1856 Order Scorpaeniformes Family Scorpaenidae Sebastes aleutianus (Jordn et Evermann, 1898) S. alutus (Gilbert, 1890) S. borealis Barsukov, 1970 S. ciliatus (Tilesius, 1813) S. glaucus Hilgendorf, 1880 S. iracundus (Jordan et Starks, 1904) S. minor Barsukov, 1972 S. owstoni (Jordan et Thompson, 1914) S. polyspinis (Taranetz et Moiseev, 1933) S. steindachneri Hilgendorf, 1880 S. taczanowskii Steindachner, 1880 S. trivittatus Hilgendorf, 1880 Sebastolobus alascanus Bean, 1890 S. altivelis (Gilbert, 1896) S. macrochir (Günther, 1877) Family Anoplopomatidae Anoplopoma fimbria, (Pallas, 1814) Family Hexagrammidae Hexagrammos lagocephalus (Pallas, 1810) H. octogrammus (Pallas, 1810) H. stelleri Tilesius, 1810 Pleurogrammus azonus Jordan et Metz, 1913 P. monopterygius (Pallas, 1810) Family Cottidae Alcichthys elongatus (Steindachner, 1881) Artediellus aporosus Soldatov, 1922 A. dydymovi Soldatov, 1915 A. ochotensis Gilbert et Burke, 1912 A. pacificus Gilbert, 1896 A. schmidti Soldatov, 1915 Enophrys diceraus (Pallas, 1788) Gymnacanthus detrisus Gilbert et Burke, 1912 G. galeatus Bean, 1881

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JS

OS

BS

PO

+ +

+ +

+ +

+

+ +

+ + +

+ +

+ +

+ + + +

+

+ + + + + +

+ + +

+

+ +

+ + + +

+ +

+

+ + + + +

+ + +

+ + + + + +

+ + + +

+ +

+

+ + + +

+ +

+ + +

508

IVANOV, SUKHANOV

Table 2. (Contd.) Region Number

Taxon JS

213. 214. 215. 216. 217. 218. 219. 220. 221. 222. 223. 224. 225. 226. 227. 228. 229. 230. 231. 232. 233. 234. 235. 236. 237. 58. 238. 239. 240. 241. 242. 59. 243. 244. 245. 246. 247. 248. 249. 250. 251. 252. 253.

G. herzensteini Jordan et Starks, 1904 G. pistilliger (Pallas, 1814) G. tricuspis (Reinchardt, 1631) Hemilepidotus gilberti Jordan et Starks, 1904 H. hemilepidotus (Tilesius, 1811) H. jordani Bean, 1881 H. papilio Bean, 1880 Icelus cataphractus (Pavlenko, 1910) I. spatula Gilbert et Burke, 1912 I. spiniger Gilbert, 1896 I. stenosomus Andriashev, 1937 Megalocottus platycephalus (Pallas, 1814) Myoxocephalus jaok (Cuvier, 1829) M. ochotensis Schmidt, 1927 M. polyacanthocephalus (Pallas, 1814) M. scorpioides (Linnaeus, 1758) M. stelleri Tilesius, 1811 M. tuberculatus Soldatov et Pavlenko,1922 M. verrucosus (Bean, 1881) Stelgistrum beringianum Gilbert et Burke, 1912 Trichocottus brashnikovi Soldatov et Pavlenko, 1915 Triglops forficatus (Gilbert, 1896) T. jordani (Schmidt, 1903) T. pingelii Reinchardt, 1831 T. scepticus Gilbert, 1896 Family Hemitripteridae Blepsias bilobus Cuvier et Valenciennes, 1830 B. cirrhosus (Pallas, 1814) Hemitripterus bolini (Myers, 1934) H. villosus (Pallas, 1814) Nautichthys pribilovius (Jordan et Gilbert, 1898) Family Agonidae Agonomalus jordani Schmidt, 1904 . A. proboscidalis (Valenciennes, 1858) Anoplagonus occidentalis Lindberg, 1950 Aspidophoroides bartoni Gilbert, 1896 Freemanichthys thompsoni (Jordan et Gilbert, 1898) Leptagonus decagonus (Bloch et Schneider, 1801) Occella dodecaedron (Tilesius, 1813) O. iburia (Jordan et Starks, 1904) Pallasina barbata (Steindachner, 1876) Percis japonica (Pallas, 1769) Podothecus accipenserinus (Tilesius, 1813)

OS

+ + +

+

BS

+

+ + +

+ + + + + + +

+ +

PO + + + + +

+ +

+ +

+ +

+

+ + + +

+ + +

+ + + +

+ + +

+ + +

+ + + + +

+ + + +

+

+ +

+ +

+ +

+

+ + + + + + +

+ + + + +

+

+ + + + +

+ + + +

+

+ + + +

+

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+ + + Vol. 55

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Table 2. (Contd.) Region Number

Taxon JS

254. 255. 256. 257. 258. 259. 260. 261. 60. 262. 263. 264. 265. 61. 266. 267. 268. 269. 270. 271. 272. 273. 274. 275. 276. 277. 278. 279. 62. 280. 281. 282. 283. 284. 285. 286. 287. 288. 289. 290. 291. 292. 293.

P. sachi (Jordan et Snyder, 1901) P. sturioides (Guichenot, 1869) P. veternus Jordan et Starks, 1895 Sarritor frenatus (Gilbert, 1896) S. leptorhynchus (Gilbert, 1896) S. knipowitschi Lindberg et Andriasev, 1937 Tilesina gibbosa Schmidt, 1904 Ulcina olriki (Lütken, 1877) Family Psychrolutidae Dasycottus setiger Bean, 1890 Malacocottus gibber Sakamoto, 1930 M. zonurus Bean, 1890 Eurymen gyrinus Gilbert et Burke, 1912 Family Cyclopteridae Aptocyclus ventricosus (Pallas, 1769) Cyclopsis tentacularis Popov, 1930 Cyclopteropsis bergi Popov, 1929 C. lindbergi Soldatov, 1930 C. popovi Soldatov, 1929 Eumicrotremus andriashevi Perminov, 1936 E. asperrimus (Tanaka, 1912) E. derjugini Popov, 1926 E. orbis (Günther, 1861) E. pacificus Schmidt, 1904 E. schmidti Lindberg et Legeza, 1955 E. soldatovi Popov, 1930 E. taranetzi Perminov, 1936 E. tartaricus Lindberg et Legeza, 1955 Family Liparidae Careproctus bathycoetus Gilbert et Burke, 1912 C. colletti Gilbert, 1896 C. cyclocephalus Kido, 1983 C. cypselurus (Jordan et Gilbert, 1898) C. furcellus Gilbert et Burke, 1912 C. homopterus Gilbert et Burke, 1912 C. macrodiscus Schmidt, 1950 C. mederi Schmidt, 1916 C. ostentum Gilbert, 1896 C. ovigerus (Gilbert, 1896) C. pycnosoma Gilbert et Burke, 1912 C. rastrinus Gilbert et Burke, 1912 C. rhodomelas Gilbert et Burke, 1912 C. rosseofuscus Gilbert et Burke, 1912

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+ + +

OS

BS

PO

+ + + +

+ + +

+ + + +

+ +

+ +

+

+ + +

+

+

+

+ +

+ + +

+ + + + + + + + + + + + + + + + + + + + + + + + + + +

+ + +

+ + +

+

+ +

+ + + +

+ + + + +

+ +

+ +

+ +

+

+

+

510

IVANOV, SUKHANOV

Table 2. (Contd.) Region Number

Taxon JS

294. 295. 296. 297. 298. 299. 300. 301. 302. 303. 304. 305. 306. 307. 308. 309. 310. 311. 312. 313. 314. 315. 316. 317. 318. 63. 319. 64. 320. 65. 321. 322. 323. 324. 325. 326. 327. 66. 328. 67. 329. 330.

C. sinensis Gilbert et Burke, 1912 C. trachysoma Gilbert et Burke, 1912 Crystallias matsushimae Jordan et Snyder, 1902 Crystallichthys mirabilis Jordan et Gilbert, 1898 Elassodiscus obscures Pitruk et Fedorov, 1993 E. tremebundus Gilbert et Burke, 1912 Liparis agassizii Putnam, 1874 L. callyodon (Pallas, 1814) L. curilensis (Gilbert et Burke, 1912) L. cyclopus Günther, 1861 L. frenatus (Gilbert et Burke, 1912) L. gibbus Bean, 1881 L. latifrons Schmidt, 1950 L. ochotensis Schmidt, 1904 L. pravdini Schmidt, 1951 L. pulchellus Ayres, 1855 L. punctatus Schmidt, 1950 L. tanakai (Gilbert et Burke, 1912) L. tessellates (Gilbert et Burke, 1912) Nectoliparis pelagicus (Gilbert et Burke, 1912) Paraliparis dactylosus Gilbert, 1896 P. entochloris Gilbert et Burke, 1912 P. melanobranchus Gilbert et Burke, 1912 P. rosaceus Gilbert, 1890 Squaloliparis dentatus (Kido, 1988) Order Perciformes Family Percichthyidae Howella parini Fedoryako, 1976 Family Echeneidae Remora remora (Linnaeus, 1758) Family Carangidae Caranx sexfasciatus Quoy et Gaimard, 1825 Decapterus kurroides Bleeker, 1855 D. muroadsi (Temminck et Schlegel, 1844) D. russelli (Rüppell, 1830) Seriola lalandi Valenciennes, 1833 S. quinqueradiata Temminck et Schlegel, 1845 Trachurus japonicus (Temminck et Schlegel, 1844) Family Coryphaenidae Coryphaena hippurus Linnaeus, 1758 Family Bramidae Brama japonica Hilgendorf, 1878 Taractes asper Lowe, 1843

OS

+ + +

BS

PO +

+ + + + + +

+ +

+ +

+ +

+ + + + + + +

+ + +

+ +

+ + + + +

+

+ + +

+ + + + + + +

+ + +

+

+ +

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511

Table 2. (Contd.) Region Number

Taxon JS

68. 331. 69. 332. 333. 334. 70. 335. 71. 336. 337. 338. 339. 340. 341. 342. 343. 344. 345. 346. 347. 348. 349. 350. 351. 352. 353. 354. 355. 356. 357. 358. 359. 360. 361. 362. 363. 72. 364. 365. 366. 367. 368.

Family Caristidae Caristius macropus (Bellotti, 1903) Family Bathymasteridae Bathymaster caeruleofasciatus Gilbert et Burke, 1912 B. derjugini Lindberg et Soldatov, 1930 . B. signatus Cope, 1873 Family Oplegnathidae Oplegnathus fasciatus (Temminck et Schlegel, 1844) Family Zoarcidae Bothrocara c.f. brunneum (Bean, 1890) B. hollandi (Jordan et Hubbs, 1925) B. tanakae (Jordan et Hubbs, 1925) B. zestum Jordan et Fowler, 1902 Bothrocarina microcephala (Schmidt, 1938) B. nigrocaudata Suvorov, 1935 Gymnelus hemifasciatus Andriashev, 1937 Lycodapus fierasfer Gilbert, 1890 Lycodes albolineatus Andriashev, 1955 L. brevipes Bean, 1890 L. brunneofasciatus Suvorov, 1935 L. concolor Gill et Townsend, 1897 L. diapterus Gilbert, 1892 L. fasciatus (Schmidt, 1904) L. japonicus Matsubara et Iwai, 1951 L. macrochir Schmidt, 1937 L. mucosus (Richardson, 1855) L. palearis Gilbert, 1896 L. polaris (Sabine, 1824) L. raridens Taranetz et Andriashev, 1937 L. sigmatoides Lindberg et Krasyukova, 1975 L. soldatovi Taranetz et Andriashev, 1935 L. tanakae Jordan et Thompson, 1914 L. uschakovi Popov, 1931 L. ygreknotatus Schmidt, 1950 Lycogrammoides schmidti Soldatov et Lindberg, 1928 Lycozoarces regani Popov, 1933 Zoarces elongatus Kner, 1868 Family Stichaeidae Acantholumpenus mackayi (Gilbert, 1896) Leptoclinus maculatus diaphanocarus (Scmidt, 1904) Lumpenella longirostris (Evermann et Goldsborough, 1907) Lumpenus fabricii Reinhardt, 1836 L. sagitta Wilimovsky, 1956

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BS

PO

+

+

+ + + +

+

+ +

+

+ + + +

+

+ + + + + +

+ + +

+ +

+ + + + + + + + + + + + + + + +

+

+

+ + + + +

+ +

+ + +

+ +

+

+ +

+ +

+

+

512

IVANOV, SUKHANOV

Table 2. (Contd.) Region Number

Taxon JS

73. 74.

75. 76. 77. 78.

79.

80. 81. 82. 83.

84.

85

369. Stichaeus grigorjewi Herzenstein, 1890 370. Stichaeopsis nevelskoi (Schmidt, 1904) 371. S. punctatus (Fabricius, 1780) Family Cryptacanthodidae 372. Cryptacanthoides bergi (Lindberg, 1930) Family Anarhichadidae 373. Anarhichas orientalis Pallas, 1814 374. Anarrhichthys ocellatus Ayres, 1855 Family Ptilichthydae 375. Ptilichthys goodei Bean, 1881 Family Mugilidae 376. Mugil cephalus Linnaeus, 1758 Family Zaproridae 377. Zaprora silenus Jordan, 1896 Family Chiasmodontidae 378. Chiasmodon niger Johnson, 1864 379. Kali indica Lloyd, 1909 380. Pseudoscopelus scriptus Lütken, 1892 Family Trichodontidae 381. Trichodon trichodon (Tilesius, 1813) 382. Arctoscopus japonicus (Steindachner, 1881) Family Pholidae 383. Rhodymenichthys dolichogaster (Pallas, 1814) Family (81) Ammodytidae 384. Ammodytes hexapterus Pallas, 1814 Family Icosteidae 385. Icosteus aenigmaticus Lokington, 1880 Family Gempylidae 386. Diplospinus multistriatus Maul, 1948 387. Gempylus serpens Cuvier, 1829 388. Lepidocybium flavobrunneum (Smith, 1843) 389. Nealotus tripes Johnson, 1865 390. Nesiarchus nasutus Johnson, 1862 Family Trichiuridae 391. Aphanopus arigato Parin, 1994 392. Benthodesmus elongatus (Clarke, 1879) 393. B. tenuis (Günther, 1872) 394. Trichiurus lepturus Linnaeus, 1758 Family Scombridae 395. Auxis rochei rochei (Riss, 1810) 396. A. thazard thazard (Lacepede, 1800) 397. Katsuwonus pelamis (Linnaeus, 1758) 398. Sarda orientalis (Temminck et Schlegel, 1844)

OS

BS

PO

+

+

+ + + + +

+

+ +

+ +

+

+

+

+

+

+

+

+

+ + +

+

+ +

+

+ +

+

+

+

+

+

+

+ +

+ + + + + + + + +

+

+ + + +

+

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Table 2. (Contd.) Region Number

Taxon JS

399. 400. 401. 402. 403. 86. 404. 87. 405. 406. 88. 407. 408. 89. 409. 410. 411. 90. 412. 91. 413. 92. 414. 415. 416. 417. 418. 419. 420. 421. 422. 423. 424. 425. 426. 427. 428. 429. 430. 431. 432. 433.

Scomber australasicus Cuvier, 1832 S. japonicus Houttuyn, 1782 Thunnus alalunga (Bonnaterre, 1788) T. obesus (Lowe, 1834) T. thynnus (Linnaeus, 1758) Family Xiphiidae Xiphias gladius Linnaeus, 1758 Family Istiophoridae Makaira mazara (Jordan et Snyder, 1901) Kajikia audax (Philippi, 1887) Family Centrolophidae Hyperoglyphe japonica (Döderlein, 1884) Icichthys lockingtoni Jordan et Gilbert, 1880 Family Nomeidae Nomeus gronovii (Gmelin, 1789) Psenes pellucidus Lütken, 1880 Psenopsis anomala (Temminck et Schlegel, 1844) Family Tetragonuridae Tetragonurus cuvieri Risso, 1810 Family Stromateidae Pampus argenteus (Euphrasen, 1788) Order Pleuronectiformes Family Pleuronectidae Acanthopsetta nadeshnyi Schmidt, 1904 Atheresthes evermanni Jordan et Starks, 1904 A. stomias Jordan et Gilbert, 1880 Cleisthenes herzensteini Schmidt, 1904 Clidoderma asperrimum (Temminck et Schlegel, 1846) Glyptocephalus stelleri Schmidt, 1904 G. zachirus Lockington, 1879 Hippoglossoides dubius Schmidt, 1904 H. elassodon Jordan et Gilbert, 1880 H. robustus Gill et Townsend, 1897 Hippoglossus stenolepis Schmidt, 1904 Lepidopsetta mochigarei Snyder, 1911 L. polyxystra Orr et Matarese, 2000 Limanda aspera (Pallas, 1894) L. punctatissimus (Steindachner, 1879) L. sakhalinensis Hubbs, 1915 Liopsetta glacialis (Pallas, 1776) Myzopsetta proboscidea (Gilbert, 1896) Paralichthys olivaceus (Temminck et Schlegel, 1848) Platichthys stellatus (Pallas, 1788)

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+

OS

BS

PO

+

+ + + + +

+

+

+

+ + +

+

+ + + + +

+

+

+

+

+ + +

+ + +

+ + +

+ +

+ +

+ +

+ +

+ + + +

+ + +

+ + +

+ + + +

+ +

+ +

+

+

+

+ + +

+

+

+

+ + + +

+ +

514

IVANOV, SUKHANOV

Table 2. (Contd.) Region Number

93.

94. 95.

96.

Taxon

434. Pleuronectes quadrituberculatus Pallas, 1814 435. Pseudopleuronectes schrenki (Schmidt, 1904) 436. Reinhardtius hippoglossoides matsuurae Jordan et Snyder, 1901 Order Tetraodontiformes Family Monacanthidae 437. Paramonacanthus japonicus (Tilesius, 1809) 438. Stephanolepis cirrhifer (Temminck et Schlegel, 1850) 439. Thamnaconus modestus (Gunther) 440. T. multilineatus (Tanaka, 1918) Family Ostraciidae 441. Tetrosomus concatenatus (Bloch, 1785) Family Tetraodontidae 442. Lagocephalus lagocephalus (Linnaeus, 1758) 443. L. wheeleri Abe, Tabeta et Kitahama, 1984 444. Takifugu chinensis (Abe, 1949) 445. T. niphobles (Jordan et Snyder, 1901) 446. T. porphyreus (Temminck et Schlegel, 1850) 447. T. rubripes (Temminck et Schlegel, 1850) 448. T. vermicularis (Temminck et Schlegel, 1850) 449. T. xanthopterus (Temminck et Schlegel, 1850) Family Molidae 450. Mola mola (Linneus, 1758)

JS

OS

BS

PO

+

+ + +

+

+

+

+

+ + +

+ + + + + +

+ + + + +

+

+ +

+

Regions (here and in Tables 3–11): JS—northwestern part of the Sea of Japan; OM—Sea of Okhotsk; BS—western part of the Bering Sea; PO—Pacific waters near Kuril Islands and Kamchatka; “+” means that the species is recorded in catches.

Table 3. Distribution of taxonomic categories in three classes of fishes and fishlike animals in the Far Eastern seas and adjacent waters of the Pacific Ocean Number of orders/families/genera/species Region JS OS BS PO All regions

Petromyzontida

Chondrichthyes

Actinopterigii

all classes

1/1/1/1 1/1/2/2 1/1/2/2 1/1/2/2 1/1/2/2

4/4/5/5 4/5/6/10 3/4/4/5 4/7/8/13 4/7/9/15

12/39/77/108 21/58/138/246 17/48/108/163 22/76/205/304 22/88/235/433

17/44/83/114 26/64/146/258 21/53/114/170 27/84/215/319 27/96/246/450

Lethenteron camtschaticum, yellowfin sole Limanda aspera, Sakhalin flounder L. sakhalinensis, snake prickleback Lumpenus sagitta, darkfin sculpin Mala cocottus zonurus, Pacific capelin Mallotus villosus catervarius, plain sculpin Myoxocephalu jaok, great sculpin M. polyacanthocephalus, pink salmon Onco rhynchus gorbuscha, chum salmon O. keta, coho salmon O. kisutch, masu salmon O. masou, Asiatic

smelt Osmerus mordax dentex, starry flounder Platich thys stellatus, Alaska plaice Pleuronectes quadrituber culatus, hawk poacher Podothecus sturioides, veteran poacher P. veternus, quillfish Ptilichthys goodie, Pacific spiny dogfish Squalus suckleyi, walleye pollock Theragra chalcogramma, ribbed sculpin Triglops pin gelii, spectacled sculpin T. scepticus) also includes nektonbenthic species that occasionally penetrate JOURNAL OF ICHTHYOLOGY

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the pelagial from the bottom biotopes. Twentyone species from this list (4.7% of the total list) can be characterized as nonrandom elements of pelagic com munities (they are given in semibold type). The pres ence of species of benthal biotopes is certainly tempo rary. Some species adapted themselves to use the pela gial resources at a certain time of day (e.g., Pacific sand lance is a elittoral species at night and neritope lagic at daytime), the other species feed and grow in the pelagial at different stages of their life cycle (e.g., Bering wolfish at the juvenile stage is a neritopelagial species and at the adult stage it is an elittoral species. Such species as smooth lumpsucker, crested sculpin, and Sakhalin flounder are not occasionally recorded in catches (both at night and at daytime); they pur posefully use nutritive resources of the pelagial. In the strict sense, there are no true dwellers in the upper lay ers of the pelagial (holoepipelagic species according to Parin, 1968) that constantly inhabit the biotope (under conditions of climatic zones of Far Eastern seas and adjacent waters of the Pacific Ocean) at all stages of the life cycle (salmon shark is the first candidate to be a true dweller). In addition to the species that were detected in catches in all surveyed areas, we present statistics of species that were recorded in only one of the regions (conventional endemics). Such species amounted to 25 species (21.9%) in the Sea of Japan, 57 species (22.2%) in the Sea of Okhotsk, 23 species (13.5%) in the Bering Sea, and 119 species (37.3%) in the adja cent waters of the Pacific Ocean. A higher portion of such species in the Pacific waters is, probably, due to the physicogeographical position of the region. As a part of the subarctic region of the northern Pacific Ocean, it extends more than 2000 km in a latitudinal meridional direction and is completely occupied by a western variety of waters of the subarctic structure (Shuntov, 2001). The mixing of the population of the pelagial of moderate and low latitudes occurs in the region as a result of interaction of water masses of dif ferent origin, which are biotopes of pelagic biocenoses (Beklemishev, 1969). The boreal fauna in the region becomes enriched in a warm period of the year (sum mer–autumn) due to subtropical and tropical migrants from low latitudes. Among the ichthyofauna species caught with a pelagic trawl, the dwellers both of the pelagial and benthal were detected and the portion of the benthal species (55.5%) in the general list exceeds the portion of true dwellers of the pelagial (Table 6). Findings of nektobenthic species in the water column are due to either their biological features (change of their mode of life at a certain stage of development and temporary vertical migrations) or accidental reasons (transport from the coastal regions by currents, setting of the trawl on the ground when catching in the neritic pela gial). This phenomenon is well known and is typical for pelagic communities of seas and coastal and island regions of oceanic waters (Parin, 1968; Radchenko, JOURNAL OF ICHTHYOLOGY

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Table 4. Distribution of the number of species fishes and fishlike animals according to families in the Far Eastern seas and adjacent waters of the Pacific Ocean Region Family Liparidae Myctophidae Cottidae Zoarcidae Pleuronectidae Agonidae Stomiidae Scorpaenidae Cyclopteridae Salmonidae Scombridae Paralepididae Stichaeidae Tetraodontidae Carangidae Gonostomatidae Rajidae Microstomatidae Sternopthychidae Gempylidae Hexagrammidae Macrouridae Opisthoproctidae Hemitripteridae Trichiuridae Oneirodidae Moridae Monacanthidae Gadidae Psychrolutidae Osmeridae Clupeidae

Total 39 38 34 28 23 19 15 15 14 10 9 8 8 8 7 7 7 6 6 5 5 5 5 5 4 4 4 4 4 4 4 4

JS

OS

BS

PO

5 – 13 1 12 10 – 4 6 5 2 – 4 5 3 – 1 – 1 – 1 – – 3 1 – – 2 3 1 2 4

33 10 26 23 17 12 4 10 14 8 2 2 6 2 – 3 6 5 – – 5 4 2 4 – 3 2 1 3 3 4 2

7 7 21 9 14 8 4 6 5 7 – 3 4 – – 3 2 5 1 – 5 3 2 5 – 3 2 – 4 3 3 1

14 38 18 11 16 11 13 9 9 8 9 8 4 2 6 6 5 6 5 5 5 5 4 2 4 4 4 3 3 3 2 2

Families with a total number of species less than four are not included in the table.

1994; Lapko, 1996; Fedorov, 2000). It should be noted that some conventionalities and difficulties exist in ascertaining the species to a certain biotope (and dis tinguishing the boundaries between vertical zones) because of a deficiency of knowledge about environ mental and biological features of the species. Some fish species not only successfully settled in several ver tical zones of the pelagic and benthal but were also able

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IVANOV, SUKHANOV

the species from pelagic biotopes prevail over species from the benthal biotopes by 53.9 and 46.1%, respec tively). The penetration of nectobenthic species into the pelagial of open Pacific waters is restricted by small areas of the shelf macrostructures and the slope of the region.

Table 5. Matrix of similarity (according to Sörensen–Che kanovsky) of the qualitative composition of icthyofauna in the Far Eastern seas and adjacent waters of the Pacific Ocean Region (species richness) Region JS OS BS PO

JS (114) 0.40 0.35 0.35

OS (258)

BS (170)

PO (319)

0.40

0.35 0.63

0.35 0.61 0.53

0.63 0.61

The analysis of the data on the icthyofauna species distribution according to biotopical groups (Table 6) has revealed the typical features for each of the regions. Thus, the fauna in the Sea of Japan compared to other surveyed areas has a small number of fishes in deepwater biotopes. Species poverty of the deep water ichthyofauna has been known for a long time (Suvorov, 1948; Andriashev, 1953). This can be explained by historical reasons and modern feature of deepwater biotopes. In the opinion of Dolganov and Saveliev (2010), poverty of the deepwater ichthyo fauna in the Sea of Japan is caused by its geographical isolation from deep Pacific waters, which occurred at the end of Pliocene (rise of the JapanKurils Cordille ras) simultaneously with continuous anomalous cool ing at the end of Pliocene–at the beginning of Pleis tocene. During that period, subtropicallowboreal and lowboreal fauna was shifted to the south and became extinct. In this geological period, only high boreal and borealarctic species of fish had a chance to survive under conditions of global climate cooling. Shallow waters of straits did not limit migrations of some deepwater fauna, because they can overcome them at early stages of development. The main factor that prevents migration of the deepwater fauna from

0.53

to adapt to habitation in the conditions of these two large biotopes. They are not ontogenetic transloca tions (widely spread phenomenon) from pelagic larvae of juveniles to the bottom mode of life of adult speci mens. This concerns the cases when adult specimens acclimatized successfully in biotopes of the pelagial and bottom (e.g., walleye pollock, smooth lump sucker, Sakhalin flounder, etc.). Sometimes, in such situations, we should follow the path of enlargement of a biotope or, with a certain portion of conventionality, (depending on preferences of researches and profun dity of the knowledge of the species biology) to include such species either in a pelagial or benthal biotope. In all Far Eastern seas, the ratio of species from benthal and pelagic biotopes is similar (Table 6) and this ratio is reverse only in waters of the Pacific Ocean;

Table 6. Number (N) and portion (%) of the ichthyofauna species in biotopical groups of the Far Eastern seas and adjacent waters of the Pacific Ocean Total

JS

OS

BS

PO

Biotopic groups Pelagic: ⎯neritic marine ⎯neritic anadromous ⎯epipelagic marine ⎯epipelagic anadromous ⎯epimesopelagic ⎯mesopelagic ⎯bathypelagic ⎯abyssopelagic Benthal: ⎯littoral ⎯sublittoral ⎯elittoral ⎯upper bathyal ⎯lower bathyal ⎯abyssobenthal

N

%

N

%

N

%

N

%

N

%

200 23 6 28 11 7 43 81 1 250 2 28 123 72 24 1

44.4 5.1 1.3 6.2 2.4 1.6 9.5 18.0 0.2 55.6 0.4 6.2 27.3 16.0 5.3 0.2

40 18 3 9 6 2 2 – – 74 – 10 50 14 – –

35.1 15.8 2.6 7.9 5.3 1.8 1.8 – – 64.9 – 8.8 43.9 12.3 – –

80 8 6 5 9 4 29 18 1 178 1 20 89 48 20 –

31.0 3.1 2.3 1.9 3.5 1.6 11.2 7.0 0.4 69.0 0.4 7.8 34.4 18.6 7.8 –

65 5 5 3 8 1 22 21 – 105 1 15 53 26 10 –

38.2 2.9 2.9 1.8 4.7 0.6 12.9 12.4 – 61.8 0.6 8.8 31.2 15.3 5.9 –

172 11 3 27 9 7 40 74 1 147 1 12 73 46 14 1

53.9 3.4 0.9 8.5 2.8 2.2 12.5 23.2 0.3 46.1 0.3 3.8 22.9 14.4 4.4 0.3

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the adjacent areas of the Pacific Ocean to depths of the Sea of Japan is a special regime of deep waters. This is the isolation of the Sea of Japan from inflow of Pacific deep water masses and severe temperature conditions for existence of hydrobionts (water temperature is 0.1–0.2°C in winter and 0.3–0.5°C in summer (Shuntov, 2001; Zuenko, 2008)), under which larvae and juveniles of many deepwater fishes do not survive. As opposed to a poor deepwater fauna in the Sea of Japan, the deepwater fauna in Pacific waters of Russia (Table 6) demonstrates the highest (compared to all surveyed regions) species richness, 182 species or 57.4% of the total number of recorded species in the region (38.2% pelagic biotopes and 19.1% benthal biotopes). The similar results were obtained by Fedorov and Parin (1998) in pelagic and benthope lagic biotopes in Pacific waters of Russia where the portion of deepwater ichthyofauna was approxi mately 60%. According to the total number of such species, these authors distinguish the water area of Kuril Islands with the highest species richness (132 species) because of the presence numerous warmwater species from neighboring subtropical latitudinal regions. The similar distribution of the ichthyofauna species according to biotopical groups is observed in the Sea of Okhotsk and in the Bering Sea. The most significant differences in this parameter were found in the Sea of Japan and in Pacific waters of Russia (Table 6). The portion of species of the deepwater ichthyofauna in the Sea of Okhotsk and the Bering Sea is high and is a little bit smaller than in Pacific waters of Russia: 46.6 and 47.1 vs. 57.3%). Among species of pelagic biotopes in the Sea of Okhotsk and the Bering Sea, the highest species richness is recorded in a mesopelagic biotopical group (29 and 22 species, respectively); ner itic species has the highest species richness (21 species or 18.4%) in the Sea of Japan and bathypelagic species (74 species or 23.2%) in Pacific waters of Russia. The elittoral biotopic group dominates among species from benthal biotopes in all regions. Elittoral fishes take the first place by the number of species and by generalized data among all species from benthal and pelagic biotopes, 122 species or 27.2%. When considering the species composition of the ichthyofauna as a whole (450 species) from the point of view of the type of the geographical range of the spe cies in the system of latitudinal zones (Table 7), it should be noted that the contribution of high and low (lowborealsubtropical and subtropicaltropical) lat itudes corresponds to the ratio 3 : 2 (61.1 : 38.9%). This ratio differs between regions: in the Sea of Japan, one species from a low latitude range is verses 2.3 spe cies from high latitude ranges (70.3, 29.7%), 5.4 in the Sea of Okhotsk (84.4 : 15.6%), 5.5 in the Bering Sea (84.7 : 15.3%), and 1.1 in Pacific waters of Russia (53.3 : 46.7%). In the ichthyofauna structure, the sufficient differ ences in the zoogeographical status of species are dis tinguished between biotopes of the pelagial and JOURNAL OF ICHTHYOLOGY

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benthal. Thus, in total in all regions, most species of the pelagic icthyofauna (74.5%) have low latitudinal ranges, whereas most species of the benthal fauna (89.6%) dwell in coldwater or moderately coldwater regions of latitudinal zones (Table 7). Such significant discrepancy between ranges of the pelagial and benthal ichthyofauna can be explained by higher adaptation of the pelagial fauna to seasonal latitudinal migrations. On a regional scale, the minimal portion of species of the benthal fauna with low latitude ranges dwells in the Bering Sea (0.9%), and the maximal portion of species inhabits Pacific waters of Kamchatka and Kuril Islands (14.3%) (Table 7). The same situation is observed among species of the pelagic fauna: the min imum number of species with low latitude ranges is recorded in the Bering Sea (38.5%) and the maximum number of species is recorded in Pacific waters of Kamchatka and Kuril Islands (74.4%). Regional dif ferences in the zoogeographical composition of the ichthyofauna in biotopes of the pelagial and benthal correspond to the location of the surveyed regions in the system of natural latitudinal zones. An important factor of such differences is the degree of formation of the continental shoal in this or that region (the pene tration of the benthal fauna into the pelagial increases with the increase of its area). And, of course, high mobility of the pelagial biotope and the absence of sta ble substrate do not promote the settled mode of life of the pelagic ichthyofauna in contrast to the mode of life of the population of bottom ichthyocenes. The fact of the considerable latitudinal extension of the surveyed area from the Arctic Circle to subtropical latitude supposes the existence of different abiotic environmental conditions (the law of latitudinal zon ality). The diversity of biotopes causes differences in the structural organization of marine biocenoses (in particular, pelagic ichthyocenes), because the com munities and their components are associated with particular biotopes (Beklemishev et al., 1973; Levush kin, 1982). The comparison of the qualitative composition of the ichthyofauna in the studied region (Fig. 2) does not make it possible to point to its synecological orga nization. Such considerations require the comparison of the list with the indexes of abundance. None of the species in the community are equally abundant and, as a rule, most of them are represented by few specimens. The distribution of the species abundance occurs according to a certain system (Whittaker, 1965; Vai levich, 1969; Pielou, 1975). The general idea of the features of the species struc ture of the ichthyocene in a particular region is repre sented in Fig. 3, where the biomass of the species (kg/m2) is a parameter of abundance. Instead of the cumulative percentage curve, the figure represents rank curves approximated by lognormal mathematical models, whose statistical parameters are given in Table 8. It is known that a small number of abundant species and a large portion of rare species (when the large

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10 5 0 –5 –10 JS

BS

OS PO

–15 0

100

200

300

Fig. 3. Curves of dominance of the ichthyofauna of Rus sian Far Eastern seas and adjacent waters of the Pacific Ocean. The rank of species is plotted along the axis of abscissa, and the natural logarithm of biomass is on the axis of ordinates; see other designations in Fig. 2.

number of samples collected in large territories with diverse biotopes) suggest that such distribution is approximated by the model of lognormal distribution. When the number of samples is large, which is typical for communities with high species richness, the distri bution of species by abundance corresponds to the lognormal distribution (Whittaker, 1980; Shitikov et al., 2011). S forms of distribution of actual data (Fig. 3) testify to a lognormal distribution of species by abundance (Magurran, 1991; Gilyarov, 2007). Such distribution is often associated with biocenotic rela tions in communities (through the concept of niche and competition) and is even considered as a feature of undisturbed communities (natural background state). But the opinions exist that the lognormal distribution of species in respect to their abundance is a statistical property of large samples or is the result of their ran dom mixing (May, 1975; Sugihara, 1980; Hughes, 1986). It is apparent that the shorter a rank curve, the less species diversity of the community. The shortest rank curve characterizes the species structure of the pelagic ichthyocene in the Sea of Japan and the longest rank curve characterizes the species structure of the pelagic ichthyocene in Pacific waters near Kamchatka and Kuril Islands. According to the profiles of rank curves, we can conclude that the dominance of one or several species in the pelagial of all Far Eastern seas of Russia and adjacent waters of the Pacific Ocean is well expressed. This is testified by sharp descending of all four curves to the axis of abscissa. Some criteria widely applied in synecology were used to estimate the degree of biodiversity of ichthy ocenes of the Far Eastern seas and adjacent waters of the Pacific Ocean (Table 9). One of such criteria is the

polydominance index, which shows the number of species in a hypothetical collection where all species have equal abundance if the collection has the same diversity as the given collection (Pesenko, 1982). In extreme cases, the value of the index tends to zero (one species community) or becomes equal to the number of species in the community if all species have equal abundance (Sukhanov and Ivanov, 2009). In other words, this index estimates the species diversity in the sample as if it was simply the number of detected spe cies and all these species were equal, i.e., had similar abundance in the sample. According to our materials, the polydominance index in ichthyocenes in the Far Eastern seas and adja cent waters of the Pacific Ocean vary in a small range from 1.58 to 2.66 (Table 9). Such values of the poly dominance index and low evenness of the species structure (0.17–0.27) indicate a wellexpressed domi nance of 2–3 species. If all species in each studied ich thyocenes were equally abundant, according to the observed species structure (Fig. 3), the species rich ness (the number of species) would constitute from 1.6 to 2.7. The common idea of the species structure of the nucleus of the ichthyocenes for each region does not contradict this conclusion (the dominance of 2–3 spe cies). This is as follows: the Sea of Japan (four spe cies)—Japanese sardine (78.0%), walleye pollock (14.3%), Pacific herring (3.3%), Japanese anchovy (2.0%); Sea of Okhotsk (five species)—walleye pol lock (75.3%), Pacific herring (16.7%), northern smoothtongue (2.2%), Japanese sardine (1.2%), cape lin (1.2%); the Bering Sea (six species)—walleye pol lock (57.7%), capelin (17.4%), Pacific herring (9.8%), polar cod (9.1%), Northern lampfish (2.6%), chum salmon (1.1%); Pacific waters of Russia (three spe cies)—walleye pollock (51.5%), Japanese sardine (37, 1%), chub mackerel (7.9%). When comparing the number of species (species richness) in the compared communities in order to exclude “the sampling effect,” we use some indexes of diversity in which the number of species is normalized anyhow (Levich, 1980; Protasov, 2002). The Margalef index is one the indexes that is simple in use. The index is applied as a comparative measure of the spe cies richness in respect to the surveyed area (the more surveyed the area, the more the abundance of detected species). The highest values of the Margalef index (the concentration of the species richness or the species density) are recorded in Pacific waters of Russia, and the lowest Margalef index characterizes the species struc ture of the ichthyocene in the Sea of Japan (Table 9). On the whole, the index does not completely correlate with the latitude, though it shows a decreasing trend with latitude northwards in three of four surveyed areas of water, except the Sea of Japan. It is apparent, that, in addition to the latitudinal zonality, other factors that affect the species diversity of the community become important. They are generally known factors of productivity and severity of environment: low spe JOURNAL OF ICHTHYOLOGY

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– 28 11 43 106

⎯highboreal

⎯boreal

⎯lowboreal

⎯lowborealsubtropical

⎯subtropicaltropical

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29 156 51 51 124

⎯highboreal

⎯boreal

⎯lowboreal

⎯lowborealsubtropical

⎯subtropicaltropical

%

27.6

11.3

11.3

34.7

6.4

7.6

0.2

0.9

100

4.0 (7.2)

1.8 (3.2)

8.9 (16.0)

28.5 (51.2)

6.4 (11.6)

5.1 (9.2)

0.2 (0.4)

0.7 (1.2)

55.6 (100)

23.6 (53.0)

9.6 (21.5)

2.4 (5.5)

6.2 (14.0)

–

2.4 (5.5)

0.2 (0.5)

44.4 (100)

Total

14

20

19

39

2

20

–

–

114

4

3

17

37

2

11

–

–

74

10

17

2

2

–

9

–

40

N

%

12.2

17.5

16.7

34.3

1.8

17.5

100 –

–

3.5 (5.4)

2.6 (4.1)

14.9 (22.9)

32.5 (50.0)

1.8 (2.7)

9.6 (14.9)

–

–

64.9 (100)

8.7 (25.0)

14.9 (42.5)

1.8 (5.0)

1.8 (5.0)

–

7.9 (22.5)

–

35.1 (100)

JS

23

17

39

123

24

31

1

–

258

1

2

31

99

24

20

1

–

178

22

15

8

24

–

11

–

80

N

%

–

8.9

6.6

15.1

47.7

9.3

12.0

0.4

100

0.4 (0.6)

0.8 (1.1)

12.0 (17.4)

38.3 (55.6)

9.3 (13.5)

7.8 (11.2)

0.4 (0.6)

–

69.0 (100)

8.5 (27.5)

5.8 (18.8)

3.1 (10.0)

9.3 (30.0)

–

4.3 (13.7)

–

31.0 (100)

OS

14

12

11

86

14

29

–

4

170

1

–

3

66

14

18

–

3

105

13

12

8

20

–

11

1

65

N

%

8.2

7.1

6.5

50.5

8.2

17.1

2.4

100 –

0.6 (0.9)

–

1.8 (2.9)

38.8 (62.9)

8.2 (13.3)

10.6 (17.1)

–

1.8 (2.9)

61.8 (100)

7.6 (20.0)

7.1 (18.5)

4.7 (12.3)

11.7 (30.8)

–

6.5 (16.9)

0.6 (1.5)

38.2 (100)

BS

113

36

32

95

17

26

–

–

319

17

4

24

68

17

17

–

–

147

96

32

8

27

–

9

–

172

N

The portion of species: their total number is out of brackets and the number of species in a particular faunistic complex of the pelagial or benthal is in the brackets.

34

⎯arcticboreal

1

18

⎯tropicalsubtropical 4

8

⎯lowborealsubtropical

⎯arctichighboreal

40

⎯lowboreal

⎯arctic

128

⎯boreal

450

29

Total:

23

⎯highboreal

1

⎯arctichighboreal

⎯arcticboreal

3

⎯arctic

250

11

⎯arcticboreal

benthal:

1

200

N

⎯arctic

Pelagic:

Zoogeographical groups

%

–

–

35.4

11.3

10.0

29.9

5.3

8.1

100

5.3 (11.6)

1.3 (2.7)

7.5 (16.3)

21.4 (46.2)

5.3 (11.6)

5.3 (11.6)

–

–

46.1 (100)

30.1 (55.8)

10.0 (18.6)

2.5 (4.7)

8.5 (15.7)

–

2.8 (5.2)

–

53.9 (100)

PO

Table 7. Number (N) and portion (%) of the ichthyofauna species in the composition of zoogeographical groups of the Far Eastern seas and adjacent waters of the Pacific Ocean

SPECIES STRUCTURE OF PELAGIC ICHTHYOCENES 519

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IVANOV, SUKHANOV

Table 8. Statistical parameters of models of lognormal curves of the ichthyofauna dominance in Russian Far Eastern seas and adjacent waters of the Pacific Ocean Region

Mode

Σ

R2

Error of the model, %

JS OS BS PO

–4.890 ± 0.029 –5.363 ± 0.094 –4.315 ± 0.077 –6.824 ± 0.484

4.384 ± 0.026 4.813 ± 0.050 4.924 ± 0.052 5.677 ± 0.202

0.9995 0.9995 0.9995 0.9982

2.1 2.9 4.2 8.6

Table 9. Average longterm estimate of the fish abundance in the pelagic ichthyocenes and parameters of their ecological diversity (1980–2009) Region Parameter Biomass, tons/km2 Species richness Nucleus the community** ⎯number of species ⎯biomass, % Margalef index impson dominance index Pielou evenness index

JS

OS

BS

PO

11.7* 114

17.9 258

15.5 170

22.1 319

4 97.6 9.60 1.58 0.17

5 96.6 21.10 1.68 0.17

6 97.2 12.78 2.66 0.27

3 96.7 25.30 2.43 0.19

* Chapanese pilchard constituted 77% of biomass in 1980–1990; ** species with biomass ≥1% formed the nucleus of the community.

Table 10. Average longterm estimate of the fish abundance in Russian Far Eastern seas and adjacent waters of the Pacific Ocean in different periods Biomass, tons/km2 Region 1980–1990 JS OS BS PO On average for regions

1991–1995

1996–2009

17.98 21.29 21.40 31.52

1.08 7.43 5.80 1.66

0.11 23.82 12.36 6.85

23.05

3.99

10.79

cies diversity of the pelagic ichthyocene in the Sea of Japan is the result of poverty of the deepwater ichthy ofauna and narrowness of the shelf zone (fishes of the bottom and near bottom biotopes have a large effect on coastal pelagic ichthyocenes). Figure 4 represents the results of the comparison of all four seas according to similarity of the list of fish species with their biomasses. The comparison was made using the Pierson correlation coefficient and was based on the method of multidimensional scaling. The following features should be mentioned. The Sea of Okhotsk is close both to the Bering Sea and to the Pacific region. At the same time, the regions of the

Bering Sea and Pacific Ocean are maximally remote from each other. The Sea of Japan is at equally large distances from other regions. Coordinates of the two dimensional plane where points of compared seas are plotted cannot be interpreted in a meaningful sense, and, thus, x and y axis on the plot in Fig. 4 (as in Fig. 2) are not named. Despite this fact, proximity and remoteness of objects both in an initial space with the size more than 400 (the length of the general list of species in the region) and in an abstract twodimensional plane approximating this space convenient for perception are estimated and depicted on the plot rather objec tively and visually. The basic characteristic of communities is the bio mass of its components. According to this parameter (average longterm estimates), the maximal density of fish aggregations was recorded in Pacific waters of the Russian special economic zone and then the Sea of Okhotsk, the Bering Sea, and the Sea of Japan follow in a descending order (Table 9). According to the concept of reorganization in eco systems of the Far Eastern seas (Shuntov et al., 2007), the dynamics of the biomass of ichthyocenes during three periods (1980–1990, 1991–1995 and 1996– 2009) had synchronous trends during the first two periods of survey (the periods of the maximum and a sharp decrease in fish productivity) in all regions (Table 10). The third period of increase and stabiliza tion of fish productivity in the Far Eastern seas was JOURNAL OF ICHTHYOLOGY

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observed in all regions except the Sea of Japan. Of course, it was not expected that the place of Japanese sardine, which it took during the phenomenal peak of its abundance in 1980–1990 in the pelagial ichthy ocene in the Russian zone of the Sea of Japan, will be completely occupied by other species of pelagic fishes (Shuntov, 1999); first of all, this is because Japanese sardine has lower trophic status (its diet consists of phytoplankton) compared to most other pelagic spe cies (Chuchukalo, 2006). Is it is known that the integ rity of communities and biocenoses is provided by the principle of energy conductivity according to which the flow of energy (biomass) damps gradually when passing through trophic levels (producers–consum ers–decomposers) (the law of the pyramid of energy). The transition from one trophic level of the pyramid of energy to another higher level is accompanied by mul tiple losses of energy (Reimers, 1994). It should be taken into account that dynamics of abundance and productivity of generations of the main pelagic competitors of Japanese sardine, such as Pacific saury, Japanese anchovy, and chub mackerel, is formed at the ichthyoplankton stage of development of the species outside the Russian zone of the Sea of Japan. Thus, the main production factors of the envi ronment (nutrients, hydrochemical composition of water, dynamics of water, etc.) in the northwestern part of the Sea of Japan do not have a direct effect on dynamics of the species abundance. Species of fishes formally comprising the faunistic complex of subtrop ical (seasonal) migrants together with Japanese sar dine (Japanese anchovy, mackerel, and Pacific saury) can replace alternatively Japanese sardine. In fact, this did not happen; probably none of these fish species demonstrated efficient reproduction to occupy the vacant place of the dominant species (Japanese sar dine). Many species forming the nucleus of the com munity during this or that period of observations can cardinally change the situation of the present decline in fish productivity in the northwestern part of the Sea of Japan. However, this has not yet happened and, to date, it is apparent that the bioproductional potential of the Russian zone in the Sea of Japan is explored inefficiently by fishes (Shuntov et al., 1998; Dudarev et al., 2004). This fact is confirmed by the available data. Based on the data of the summer pelagic trawl survey of 1997, Shuntov and other researches (1998) observed that the epipelagial in the Russian zone of the Sea of Japan was poorly inhabited by fishes. The abso lute biomass of nekton (only fishes and squids) in the epipelagial of the Russian economic zone in the Sea of Japan during that period constituted 0.33 million tons (Shuntov and Temnykh, 2011), whereas the absolute biomass was 2.57 million tons on average in the period of the maximum fish productivity (Shuntov, 2009). Thus, only in the Russian zone of the Sea of Japan during the last of the considered periods (1996–2009), the fish productivity did not increase, but its decrease continued with the following stabilization at a low JOURNAL OF ICHTHYOLOGY

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521

0.2 BS JS 0.1

0 OS –0.1

–0.2 –0.8

PO –0.4

0

0.4

0.8

Fig. 4. Classification of the Far Eastern seas and adjacent waters of the Pacific Ocean according to the species struc ture of pelagic ichthyocenes (explanations are given in the text; see designations in Fig. 2).

level, which was confirmed by the survey of 2012 when the nekton biomass was evaluated as 0.28 million tons. Changes in the species structure of ichthyocenes in the Far Eastern seas and adjacent waters of the Pacific Ocean during the distinguished periods are presented in Table 11. When considering the results of evalua tions of abundance of some components of ichthy ocenes, it should be taken into consideration that they are average statistical data and, in some cases, cannot be used for assessment of an actual dynamics of spe cies. It is well known that resources of mesopelagic fishes in the Bering Sea were rather considerable (Shuntov, 2012) and the dominant species was North ern lampfish with biomass of 9 tons/km2 (Balanov and Il’inskii, 1992; Beamish et al., 1999). Our estimate of the biomass of the species for the period 1980–1990 was an order lower (0.7 tons/km2) because of underes timate of the mesopelagic ichthyofauna in the epipela gial (where we conducted 80% of the surveys). Not all aboriginal dwellers of the mesopelagial perform night migrations to the epipelagial, and interzonal species of fishes were also underestimated during surveys in the surface layer of the epipelagial (the trawl opening cov ers a layer of 30–50 m). According to evaluations made by Radchenko (1994), during that period in the epipelagial of the Bering Sea, up to 600000 tons of mesopelagic species of fish were taken into account (Northern lampfish as a dominant species, whereas the population in the mesopelagial constituted more than 6 million tons (Radchenko, 2007). In addition, when analyzing the obtained estimates of the abun dance of some species (e.g., capelin), it should be

PO

BS

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1996–2009

6.85

1.66

1991–1995

12.36

1996–2009 31.52

5.80

1991–1995

1980–1990

21.40

23.82

1996–2009

1980–1990

7.43

1991–1995

1996–2003

21.29

0.11

1991–1995

1980–1990

1.08

1980–1990

JS

OS

17.98

Period, years

Region

Biomass, tons/km2 subdominant

Japanese anchovy (31.7)

Chub mackerel (8.7)

Pacific herring (12.4), capelin (12.4), chum salmon (3.6), sand lance (1.4)

Capelin (2.2), Northern lampfish (1.9), chum salmon (1.5), pink salmon (1.3)

Polar cod (8.4), Pacific herring (8.3), North ern lampfish (3.1)

Capelin (1.3), Sakhalin flounder (1.0)

Pink salmon (12.9), northern smoothtongue (3.3), chum salmon (1.0)

Northern smoothtongue (5.3), Japanese sar dine (3.2)

Pacific saury (8.9), arabesque greenling (8.8), salmon shark (6.3), masu salmon (4.7), pur ple puffer (fugu) (1.3)

Capelin (5.0), chub mackerel (2.0), threespine stickleback (1.5)

Pacific herring (3.4), Japanese anchovy (1.9)

the remaining species

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Walleye pollock (52.1) Pacific saury (112.5)

Japanese anchovy (11.1), pink salmon (5.9), California headlightfish (4.9), Northern lampfish (2.9), chum salmon (2.4), Japanese lanternfish (1.4)

Walleye pollock (43.0) California headlightfish (11.2) Pink salmon (9.8), Northern lampfish (7.7), Japanese anchovy (6.0), chum salmon (4.7), lowsail ribbonfish (3.1), northern smooth tongue (2.5), Japanese lanternfish (1.7), blue lanternfish (1.5), Pacific pomfret (1.4)

Walleye pollock (53.0) Japanese sardine (36.4)

Walleye pollock (50.6) Polar cod(15.9)

Walleye pollock (69.4) Pacific herring (21.5)

Walleye pollock (58.6) capelin (20.1)

Walleye pollock (85.5) Pacific herring (9.1)

Walleye pollock (58.1) Pacific herring (20.7)

Walleye pollock (57.9) Pacific herring (30.8)

Pink salmon (37.0)

Walleye pollock (78.1) Japanese anchovy (9.5)

Japanese sardine (80.3) Walleye pollock (12.6)

dominant

Nucleus of icthyocene⎯species (portion of biomass, %)

Table 11. Species structure of pelagic ichthyocenes in Far Eastern seas and Pacific waters of Russia in different periods

233

167

260

130

106

171

215

170

194

27

91

94

N

6.6

7.4

1.9

3.7

2.2

1.5

3.1

4.0

2.8

1.3

3.9

1.8

%

Other species

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Table 12. Some parameters of ecological diversity of the ichthyofauna in the Far Eastern seas of Russia and adjacent waters of the Pacific Ocean in different periods 1980–1990

1991–1995

1996–2009

Region JS OS BS PO

1/D

E

S

1/D

E

S

1/D

E

S

1.51 2.31 2.51 2.37

0.16 0.21 0.24 0.19

98 198 176 263

1.61 2.51 1.89 4.52

0.21 0.25 0.22 0.42

96 175 112 178

3.86 1.35 3.18 3.26

0.46 0.12 0.22 0.36

33 219 136 241

I/D is the polydominance index (species diversity according to Simpson in interpretation of Gibson (1966), E is evenness according to Pielou, and S is the species richness.

taken into account that our surveys were performed outside the territorial waters and the coefficients of catching efficiency are not 100% accurate. In the Sea of Japan and in waters near Kuril Islands, the main resource components of the ichthy ocene in the epipelagial during the period of the max imum fish productivity in the Far Eastern seas (1980– 1990) were two species (a dominant and subdomi nant)—Japanese sardine and walleye pollock—and the nucleus of the ichthyocene in the first region was formed by four species and that in the second region was by three species. The main contribution to the total fish productivity in the Sea of Okhotsk was made by walleye pollock and herring during all periods of survey (Table 11), and the structure of the ichthyocene nucleus was formed by four to five species. In the Ber ing Sea, walleye pollock and capelin made the greatest contribution to fish productivity, and the nucleus of the community was formed by five species. In the following period (1991–1995), a sharp decline in fish productivity was observed in two regions: in the Sea of Japan and in Pacific waters near Kuril Islands. The stock of Japanese sardine and wall eye pollock decreased simultaneously. In the Sea of Japan, walleye pollock was a dominant species and Japanese anchovy was a subdominant species during this period. walleye pollock dominated in Pacific waters (in the region of shelf and underwater slope), and theta lanternfish dominated in open ocean waters. In the Bering Sea, the pair of a dominant and subdom inant species was formed by walleye pollock and Pacific herring. During the period of the increase and stabilization of fish productivity in the Far Eastern seas (1996– 2009), the main increase in the ichthyocene biomass was due to walleye pollock in all regions, including the Sea of Japan. It is typical that this period in the Sea of Okhotsk was more productive by general parameters of the abundance of the pelagic fauna than the period of the maximum fish productivity. This fact is directly related to extremely high abundance of walleye pol lock in the 2000s (Ovsyannikov, 2011; Ovsyannikov et al., 2013). A pair of dominant and subdominant species was formed by pink salmon and Japanese JOURNAL OF ICHTHYOLOGY

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anchovy in the Sea of Japan, walleye pollock and Pacific herring in the Sea of Okhotsk, walleye pollock and Polar cod in the Bering Sea, and walleye pollock (shelf and slope) and Pacific saury (neriticoceanic waters) in Russian waters of the Pacific Ocean. Changes in the species structure of the pelagic ich thyocenes in Russian Far Eastern seas and adjacent waters of the Pacific Ocean can be traced by changes in some parameters of the ecological diversity (Table 12). In the Sea of Japan, the dynamics of these parameters was as follows depending on the period of surveys. The species richness was at one level during the first two periods of studies and decreased notably in the third period. This fact can be explained by a smaller number of surveys during the last period (only two surveys). The indexes of dominance (Tables 11 and 12) of the pelagic ichthyocene in the Sea of Japan demonstrate that the polydominance of the community increases when fish productivity declines. This correlation is observed in other regions except the Bering Sea, where one of the indexes of dominance, the index of polydominance, decreased (it increased in other regions) during the period of a sharp decrease in fish productivity. The probable reason for this is a sharp weakening of advection of Pacific waters in the Bering Sea and the following increase in the current intensity by 1995 (Verkhunov and Tkachenko, 1992; Cokelet et al., 1996; Balykin, 2008). According to the biotopic principle of Beklemishev (Beklemishev et al., 1973), feeding migrations of warmloving species of fish from the ocean decreased, which resulted in the decrease of the index of polydominance. Highly expressed polydominant properties of the pelagic community of fishes were observed in the period of a sharp decline in fish productivity (1991– 1995) in the Pacific region (the index of polydomi nance was 4.52), and the lowest polydominance was during the period of increase and stabilization of fish productivity in the Sea of Okhotsk (1.35), which indi cates a monodominant type of the community. The higher the index of polydominance, the more even the species structure of the community, which is equiva lent to higher species diversity (Magurran, 1992). The minimal evenness of the species structure of the

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pelagic fish community was recorded in 1996–2009 in the Sea of Okhotsk, where the portion of walleye pol lock by biomass exceeded 85% and it was the highest value for the period of surveys in all regions (Tables 11 and 12). The species evenness in respect to abundance increased when the fish productivity decreased. It is of interest that the period of increase and stabilization of fish productivity was manifested in the Sea of Japan and was more expressed in the Sea of Okhotsk. As a result, the species evenness in respect to abundance was the highest in the Sea of Japan (0.46) and the low est (0.12) in the Sea of Okhotsk. The reliability of these simple and evident relations between integral charac teristics of communities was confirmed statistically by the example of nekton from Pacific waters near Kuril Islands and the Sea of Japan (Ivanov and Sukhanov, 2002; Sukhanov and Ivanov, 2009) and all macrofauna in the northwestern part of the Pacific Ocean (Vol venko, 2009). Table 9 presents the average, longterm estimate of the fish biomass in the pelagial of Russian waters of the Far Eastern seas and adjacent regions of the Pacific Ocean. The biomass of pelagic fishes was, on average, 16.8 tons/km2 in all regions. If we know the total area of the region under study, which is 3.74 million km2 within the Russian economic zone (Volvenko, 2003), and the value of the average longterm biomass of pelagic fishes (16.8 tons/km2), we can calculate the total value of their biomass (the total stock of fishes in the pelagial), 63 million tons. This estimate is compa rable with the estimate of nektonic resources (fishes and cephalopods) made by Shuntov (2009, 2012) in the Russian Far Eastern seas in the pelagial layer of 1 km, 81.3 million tons (for the same period). Taking into consideration that the mesopelagial ichthyofauna was not properly studied during surveys (only 20% of trawling was performed in the mesopelagial layer), the total resources of all pelagic fishes in the Russian waters of the Far Eastern seas in the epi and mesope lagial is 70–80 million tons for an average period of longterm observations. ACKNOWLEDGMENTS We are grateful to all participants of research cruises, who performed highly qualified work on sam pling the materials. O.A. Ivanov expresses his special appreciation to the staff (former and present) of labo ratories of Applied Biocenology, Hydrobiology, and other laboratories of TINRO center for the miles we travelled together and for their support and assistance on land and sea in our common activities aimed at the study of marine biota. REFERENCES Aksyutina, Z.M., Elementy matematicheskoi otsenki rezul’tatov nablyudenii v biologicheskikh i rybokhozyaistven nykh issledovanii (Elements of Mathematical Analysis of

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Translated by N. Ruban

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