ICES Marine Science Symposia, 215: 315-322. 2002 Long-term discussion on the stocks of the White Sea herring: historical perspective and present state

Dmitry L. Lajus

Lajus, D. L. 2002. Long-term discussion on the stocks of the White Sea herring: his­ torical perspective and present state. - ICES Marine Science Symposia, 215:315-322.

There are a number of White Sea herring (Clupea pallasi marisalbi) stocks which dif­ fer with respect to features such as growth rate, spawning time, migrations, and exter­ nal morphology. Two questions concerning this herring provoke discussion even today: 1) taxonomic status and origin, and 2) population structure. Research on the popula­ tion structure of White Sea herring started in the 1920s under the strong influence of herring studies by Friedrich Heincke and Johan Hjort, key figures at that time in activ­ ities of the International Council for the Exploration of Sea. The first description of White Sea herring was made earlier by Heincke himself. The first concept of White Sea herring population structure developed in the 1920s with the use of methods developed under the ICES umbrella. During subsequent decades, the White Sea her­ ring’s amazing diversity was explained by several concepts ranging from the model of a single population to the concept of polyphiletic origin. However, these concepts have not been confirmed by recent data. Data on chromosomal sets and variation of meris- tic characters obtained recently make it possible to propose a new concept of White Sea herring population structure.

Keywords: herring, historical perspective, population structure. White Sea.

Dmitry L. Lajus: Zoological Institute, Russian Academy o f Sciences, Universitetskaya nab. 1, 199034 St. Petersburg, Russia; tel: + 7 812 114 0097; fax: +7 812 328 2941; e-mail: [email protected]. ras.spb. ru.

Introduction Taxonomic status of the White Sea herring White Sea herring (Clupea pallasi marisalbi) (Berg, 1923) is considered to be a subspecies of the Pacific The first researcher to study the taxonomic status of herring (Clupea pallasi). It has occupied the White Sea White Sea herring was the German ichthyologist Fried­ for about 10 000 years (Derjugin, 1928; Andriiashev, rich Heincke (1898). By the end of the 19th century, 1957), formed many stocks during this time, and repre­ he had published several papers establishing a new ap­ sents an excellent example of high intra-species diversi­ proach in fisheries biology. The essence of this new ty within a comparatively small body of water. Slow- approach was the shift from species to population as the and fast-growing herring are distinguishable; at 4-5 appropriate unit of study (Sinclair and Solemdal, 1988). years of age, they differ 3-5 times in weight. The spawn­ Among many herring samples from different loca­ ing period lasts from April to June for different stocks tions, Heincke studied a sample collected near the Solo­ (Dmitriev, 1946; Tambovtsev, 1957) (Figure 1). vetsky Islands in the central part of the White Sea. Herring is the most important commercial fish in the Heincke had found that the White Sea herring was mor­ White Sea and has great significance for the local econ­ phologically closer to the Pacific herring than to the omy. In addition to Pacific herring, the White Sea also Atlantic herring. He had also reported several speci­ includes Atlantic herring, which comes from the mens with higher numbers of vertebrae than was typical Barents Sea for feeding and never spawns in the White for Pacific herring, but did not draw particular attention Sea. In some years, the Atlantic herring contributes to them. Not until the later 1920s did Sergei Averintsev greatly to the total herring catch (Tambovtsev, 1957). (1927, 1928) describe them as Atlantic herring coming Problems and even heated debates have surrounded from the Barents Sea. the taxonomic status and population structure of White The particular location of the White Sea, with this Sea herring during the whole period of its study. Even mixture of two herring species, led to a long-standing now when the overall situation is becoming clearer, con­ error in the taxonomy of the genus Clupea. At the begin­ siderable work is still needed before the relationship be­ ning of the 20th century, the Russian ichthyologist Petr tween different stocks can be fully understood. Shmidt (1904) compared Atlantic and Pacific herring 316 D. L. Lajus

carried out investigations of the main Russian fisheries (Borisov, 1964). This was among the earliest organized scientific investigation of marine fisheries (Smith, 1994). In 1859, the Expedition worked in the White Sea where its scientists studied the herring fishery. Like Heincke, Danilevsky paid attention to fishermen who claimed that herring caught in different years were not exactly alike. In the report of the Expedition, Danilevsky raised a question which set the direction of further studies, particularly "whether or not the White Sea herring is a single stock" (Danilevsky, 1862, p. 16). Raising such a question, Danilevsky obviously reject­ ed the Dodd-Anderson theory, although it was finally rejected by the scientific community only after Figure 1. Growth rate and spawning time of the White Sea her­ Heincke’s studies at the end of the 19th century. Dani­ ring (A - April, M - May, J - June). levsky based his ideas on the herring studies in the Bohuslän area (Sweden) by Nilsson, who found that a stock which once disappeared was not recovered from for taxonomic purposes. Shmidt took one sample of her­ neighbouring stocks (Danilevsky, 1862). ring from the Pacific area and another, as a representa­ Danilevsky described differences in morphology and tive of Atlantic herring, from the White Sea. His appro­ spawning behaviour of several stocks of herring, con­ priate conclusion was that these forms differed at the sidering stocks from Onega and Kandalaksha Bays to be subspecies level. different stocks, but the fast- and slow-growing herring This conclusion was repeated by Berg (1923) and from Kandalaksha Bay to be different age groups of the consequently by Svetovidov in his well-known book same stock. on the taxonomy of Clupeidae (Svetovidov, 1952). Although both Berg and Svetovidov classified the White Sea herring as Pacific herring, they merely repeated First systematic studies: races and populations Shmidt’s conclusion without a critical analysis of the primary data. Following Svetovidov, for several decades The new period of studies of the White Sea herring pop­ many researchers considered Atlantic and Pacific her­ ulation structure began in the 1920s with work by ring to be subspecies. Only quite recently has this mis­ Aleksandr Rabinerson (1925) and Sergei Averintsev take been recognized (Rass, 1985; Rass and Wheeler, (1926, 1927, 1928, 1934). Their studies were based on 1991). the most modern ichthyological methods of the time. Both researchers used techniques for age determination by scale reading and analysis of age composition of Population structure of the White Sea samples developed by Johan Hjort (1910, 1914) and herring Einar Lea (1910) as well as methods for statistical analysis of morphological characters. Interestingly, in The beginning of studies in the 19th and 20th the introduction to his book, Rabinerson acknowledged centuries: formulation of the problem "help with application of variation-statistical method" from Theodosii Dobzhansky, who soon emigrated from The first data on White Sea herring were reported by Russia and became one of the founders of evolutionary Ivan Lepekhin (1805). He supported the theory, devel­ synthesis. oped by Dodd and Anderson in the middle of the 18th It should be noted that, in the 1920s, Russian scien­ century, that herring shoals, belonging to one aggrega­ tists, in spite of their more-or-less political and econom­ tion with no local varieties, migrated each spring from ic isolation, considered themselves part of the broad sci­ Arctic waters to the North Atlantic and returned in the entific community. Links established earlier with their autumn (Sinclair and Solemdal, 1988). Lepekhin, how­ western colleagues were not forgotten. Rabinerson and ever, thought that the White Sea herring was not identi­ Averintsev were very aware of the herring research be­ cal to the parental herring stock inhabiting polar waters ing done abroad. Lev Berg, one of Rabinerson’s super­ because "it is re-born in several generations during its visors, attended oceanographic courses in Bergen in journey to the White Sea" (Lepekhin, 1805, p. 343). 1903 where he met Hjort and other Norwegian marine Also, Lepekhin indicated the existence of two White biologists. Averintsev, who was much older than Rabi­ Sea herring groups differing in growth rate. nerson, visited Bergen in 1904 as a director of the From 1851 to 1870, the Expedition for Studies of Murman Biological Station of the St. Petersburg Na­ Fishery led by Karl von Baer and Nikolai Danilevsky turalist Society. Long-term discussion on the stocks of the White Sea herring: historical perspective and present state 317

In his paper, Rabinerson even presented his data in a tence of self-dependent stocks in different parts of the format used previously by Lea to facilitate an easier White Sea. Those who agreed described the new stocks comparison of results. To the White Sea herring, he of Pacific herring in northern parts of the White Sea and applied the same techniques of age composition of sam­ in the eastern part of the Barents Sea. In a monograph ples for studying migrations which Hjort had success­ on White Sea herring, 10 self-dependent stocks were fully applied to herring in the Norwegian Sea. listed (Dmitriev, 1946). Rabinerson, however, failed to find any correlation between the age composition of samples from different bays. Based on these results, he concluded that different stocks did not migrate from the bays where they were Heated debates: a single population or different born. species? Rabinerson had systematically distinguished two forms of the White Sea herring and gave them the status In the 1950s, the problem of methodology for herring of varietas, emphasizing that the growth rate was the population studies arose and became the subject of main difference between them. He used the term vari­ serious discussions. In 1956, ICES organized a special etas instead of race because the forms he had distin­ meeting devoted to the problem of "herring races". guished did not differ in their geographical distribution. By this time, it had become obvious that morpho­ He even described the mixing between these two forms logical characters, on which analysis of the popu­ in the spawning stocks in different bays. However, the lation studies had been mainly based since Heincke, absence of morphological differences prevented him exhibit plasticity under the influence of environmen­ from describing them as different races. tal factors (Bückmann and Parrish, 1958). This problem Averintsev emphasized the insufficiency of the mor­ was soon overcome by the application of methods of phological approach for discriminating races because biochemical genetics (Ridgway, 1958; Sindermann and morphological characters were not adequately sensitive Mairs, 1959; Sindermann, 1962). However, the new and could be affected directly by environmental condi­ genetic methods were not applied to the White Sea her­ tions. "To solve the question whether we deal with one ring for a long time. or two races, one needs to investigate conditions and The period from the end of the 1940s to the middle of area of their spawning, conditions and area of distribu­ the 1960s was a difficult time for population biology tion and development of larvae, fluctuation of race char­ and genetics in the Soviet Union. It was a period of pre­ acters etc." (Averintsev, 1934, p. 67). He stressed the dominance of Lysenkoism (Joravsky, 1970). Lysenko importance of data on life history and even advocated argued that phenotypic variation had no genetical basis using "fishermen’s knowledge". and, therefore, phenotypes could vary to an unlimited Although Rabinerson and Averintsev discussed their extent. Most of the recognized Russian zoologists re­ results from rather different positions and hotly debated jected Lysenko’s views and did not follow them when with each other, it is apparent even from their papers his period of dominance ended. that their conclusions were rather similar. Both consid­ Perhaps the only well-known zoologist who basically ered the White Sea as Pacific herring, forming two large expressed support for the ideas of Lysenko concerning groups, slow- and fast-growing. Each of these groups problems of spéciation was the ichthyologist Georgy formed self-dependent stocks in different locations Nikolsky (Vorontsov and Yablokov, 1973), who wrote a (Figure 2a). "It is not important for the fishery how to book, Species and Spéciation, which has appeared in name these self-dependent stocks - races, local forms or several editions (Nikolsky, 1953, 1962a, 1972). In this unmixed shoals. It is important that they are self- book, he "does not recognise, does not understand and, dependent, and therefore the samples of herring from as the main point, does not want to recognise and to the Onega Bay say nothing to us about the structure of understand, the paramount methodical importance of the stock from Kandalaksha Bay and vice versa" genetics for the problem of spéciation" (Vorontsov and (Borodatova, 1928). Yablokov, 1973, pp. 116-118). Nikolsky was a very It was becoming more and more clear that morpho­ influential person in both Soviet and world-wide ichthy­ logical differences and population differentiation were ology, and a Vice-President of ICES in 1969-1972 not the same. Hjort proposed differentiating the term (Went, 1972). Though Nikolsky himself did not study "races" as only morphologically distinct groups, while White Sea herring, he considered it a very convenient "populations" and "stocks" meant groups which could case for applying his views. be distinguished both geographically and biologically In the early 1960s, Yury Lapin posed a new concept of (Hjort, 1945). the White Sea herring population structure. He consid­ Averintsev (1934), summing up the results of studies ered White Sea herring to be a single population, "a sin­ conducted in the 1920— 1930s, wrote that the occurrence gle biological entity" (Lapin, 1966) (Figure 2b). Lapin’s of different races of herring in such a small area as the theoretical views can be seen from his epilogue to the White Sea was so strange that, even despite strong evi­ book by Nikolsky, where he wrote: "Within-species dence, not all ichthyologists could agree on the exis­ groups are reversible, inter-transitory, although they are 318 D. L. Lajus

Although many ichthyologists opposed Lapin views, the delay in applying genetic methods postponed the solution of the problem. Studies were continued by means of traditional techniques, such as analysis of age and size composition, growth rate, fecundity (Ano­ khina, 1963; Altukhov, 1975), migrations (Gosheva, 1967), salinity resistance (Galkina, 1962; Dushkina, 1973), analysis of life cycles (Mukhomediarov, 1975), larvae pigmentation (Dushkina, 1975, 1988), composi­ tion of parasites (Kulachkova, 1977), and anatomical features (Korovina and Zhuk, 1985). Data from many of the studies were used as arguments for the reproductive isolation of the White Sea herring stocks. Attempts to model herring population dynamics primarily using data on catch statistics, fecundity, and size were based on the concept of isolated stocks (Krixunov, 1991). Some opponents of Lapin developed a concept of polyphiletic origin of the White Sea herring, of which a consequence would be reproductive isolation. Ladda Dushkina (1975), who had found similarities between some stocks of the White Sea herring and Atlantic her­ ring and between other stocks and Pacific herring, argued that this showed their different origins. On the basis of these data, Dushkina argued that different stocks of the White Sea herring "have a systematic Figure 2. Schemes of concepts of the White Sea herring pop­ ulation structure: a) isolated populations of Pacific origin range not lower than that of a subspecies, or even divided into groups of fast- and slow-growing stocks species" (Dushkina, 1975, p. 249) (Figure 2c). (Rabinerson, 1925; Averintsev, 1927); b) one population, "sin­ Taisia Skvortsova (1975), in a book edited by gle biological entity", inhabiting different parts of the sea at Dushkina, reported data on consistent chromosomal dif­ different periods of the life cycle (Lapin, 1966); c) populations ferences between the White Sea herring stocks. One of having different origin from Atlantic and Pacific ancestors the populations, namely the spring-spawning herring of (Dushkina, 1975). Kandalaksha Bay (known as "St George herring" because it spawns near St George’s Day), had a diploid chromosomal number (2n) equal to 52, and several relatively reproductively isolated. However, fluctuations other populations had 2n equal to 54 chromosomes. of environment disrupts nascent deviations from the ini­ Although most of these data were obtained by a rather tial form and the differentiation appears to be rever­ old technique which does not allow the accurate count­ sible" (Nikolsky, 1980, p. 153). ing of chromosomes (Vasil’ev, 1985), they were According to Lapin, White Sea herring inhabits dif­ nevertheless considered a very strong support for ferent parts of the sea at different stages of its life cycle Dushkina’s concept of polyphiletic origin of the White which are most appropriate for the particular stages Sea herring. of ontogenesis (Lapin et al., 1962; Lapin, 1971, 1978; These data seemingly disproved Lapins concept. Lapin and Pokhiliuk, 1993). Nikolsky immediately re­ However, at least theoretically, there was the opportuni­ acted to Lapin’s concept, noting that "studies by Lapin ty to consider the existence of chromosomal differences and his colleagues force us to revise a concept of isolat­ among the White Sea herring stocks reported by ed stocks of the White Sea herring. Most likely, what Skvortsova within the framework of the concept of a was considered earlier as the separated stocks (on the single population. It was possible, if the White Sea her­ base of formal biometrical analysis) represent ... differ­ ring were a chromosomally polymorphous form, i.e., ent age groups of a single population" (Nikolsky, 1962b, if individuals had different chromosome numbers, that p. 6). 52 and 54 could exist within the same population. Lapin’s concept quickly became popular, probably Such data were soon published in a book edited by because of its convenience for the fishery. The main Lapin (Krysanov, 1978). It was reported that, in one gear for fishing herring in the White Sea is the bottom population from Onega Bay, most of the individuals trawl, but its use is limited because of the rocky bottom. had 52 chromosomes, whereas some individuals had Thus, acceptance of the concept of a single population 54 chromosomes. Under such circumstances, data by allowed fishermen to take the whole herring quota in the Skvortsova could not be considered as indispu­ few locations which were most convenient for trawling table proof of the reproductive isolation of the herring without danger of overfishing. populations. Long-term discussion on the stocks of the White Sea herring: historical perspective and present state 319

The present state: hierarchical population structure

Chromosomal differences between the White Sea her­ iiMiniHiH.i. 2n- = 52 - - ring stocks reported in the 1970s suggested that chro­ 'lU P tl tltOM M 'l# »»MX*«« K mosomal analysis could be a potential method for pop­ i>nonnni\Afionnn« J. II = ^1 'v ulation studies. However, data on the existence of her­ ring with 54 chromosomes and, consequently, data on 2n = 50 ) chromosomal differentiation of the White Sea herring populations, were not confirmed. All studied specimens had 52 chromosomes (Lajus, 1987; Klinkhardt, 1993). Continuation of research in this direction seemed to be non-productive. However, further chromosomal stud­ ies led to surprising results. Robertsonian polymor­ phism was described in the White Sea herring. Indivi­ duals with 52, 51, and 50 chromosomes, with an equal number of chromosomal arms (NF=60), were found to occur in the same population (Lajus, 1989, 1996a) (Figure 3a). Robertsonian polymorphism, a rather rare phenomenon in fish, is described for only 5% or less of all species (Vasil’ev, 1985). Moreover, it became appar­ ent that different populations of the White Sea herring differ in the frequency of individuals with different numbers of chromosomes, which permits the use of chromosomal analysis as a convenient tool in White Sea herring population studies (Figure 3b). Figure 3. Chromosomal variation in the White Sea herring and On the basis of chromosomal studies and data on corresponding concept of the population structure (based on variation of meristic characters, a new concept of the Lajus, 1996a, 1996b): a) Robertsonian polymorphism: kary­ White Sea herring population structure was proposed otypes of specimens with different diploid number of chro­ (Lajus, 1996b) (Figure 3c). According to this concept, it mosomes (50, 51. and 52); the specimens have the same num­ ber of chromosomal arms (number fundamental, NF=60); b) is possible to differentiate at least three levels. First, her­ proportion of specimens with different number of chromo­ ring from different parts of the Sea differ in the fre­ somes in different locations of the White Sea; c) scheme of a quency of chromosomal rearrangement. Second, herring concept of the population structure: reproductively isolated from the same location (Kandalaksha Bay) spawn at dif­ populations of Pacific origin with poor gene flow differing in ferent times. They have the same frequency of chromo­ spawning time and growth rate. somal rearrangement, but differ in the mean number of meristic characters, in developmental stability, meas­ ured by fluctuating asymmetry, and also in salinity resistance of embryos and larvae. Third, no difference studies clearly showed that two forms of herring from (except growth rate) was found between slow- and fast- Kandalaksha Bay are. genetically speaking, more close­ growing herring from the same location. ly related to each other than to populations from other In discussions about population structure of the White parts of the White Sea. This indicates that the location Sea herring, those from Kandalaksha Bay occupied a where herring live, not growth rate, is the principal fac­ distinctive place. The Bay includes two forms of her­ tor accounting for their differentiation. ring: St John herring, a fast-growing herring which As for St George, or spring-spawning herring, a spawns in June (near St John’s Day) at temperatures of hypothesis which explains its origin has been proposed 8-12°C, and St George herring, a slow-growing herring (Lajus, 1996b). Owing to early spawning, the critical which is much more numerous than St John herring and period of larval development of springspawners, i.e., the which spawns in April beneath the ice under the most first month after spawning (Ivanchencko, 1983), coin­ severe environmental conditions experienced by the cides with maximum plankton abundance needed for other Pacific herring. Because of this clear differentia­ larval feeding, which usually takes place in June tion between St John and St George herring, Rabinerson (Prygunkova, 1982). Favourable feeding conditions for and Averintsev and all later researchers probably as­ larvae result in considerably higher numbers of spring sumed slow- as opposed to fast-growing herring is the spawners relative to summer spawners. On the other principal differentiation among the forms of the White hand, development stability of spring spawners, meas­ Sea herring. Dushkina (1975) set St George herring of ured by fluctuating asymmetry (Lajus, 1991, 2001 ) and Kandalaksha Bay against all other populations with salinity resistance (Dushkina, 1973; Ivanchenko and respect to polyphiletic origin. However, chromosomal Lajus, 1985), is lower. Therefore, decreased develop- 320 D. L. Lajus mental stability can be considered as compensation for navaga and low vertebrate herring). Materialy po komplek- the higher abundance. snomu izucheniiu Belogo moria, 1. Moscow-Leningrad, Izdatelstvo Akademii Nauk SSSR (Materials on complex Chromosomal data are still the only genetic data on the investigation of the White Sea, 1. Moscow-Leningrad, White Sea herring. Although attempts to analyse protein USSR Academy of Sciences Press): 117-125. (In Russian). variation were undertaken, results have not yet been fully Anokhina, L. E. 1963. Some aspects of the fecundity of the published (Andreeva et a l., 1995). In the general context herring in the White Sea. Rapports et Procès-Verbaux des of herring biology and systematics, it would be of inter­ Réunions du Conseil International pour l’Exploration de la est to study the relationship between White Sea herring Mer, 154: 123-127. Averintsev, S. V. 1926. Herring of the White Sea. Wissen­ and other forms of Pacific herring in the northern seas. schaftlichen Meeresuntersuchungen, 15: 1-24. It was determined long ago that the Pacific herring Averintsev, S. V 1927. Sel'di Belogo moria ch. 1 ( Herring of the occurs in the eastern part of the Barents Sea (Esipov, White Sea. part 1 ). Trudy Nauchnogo instituta rybnogo kho- 1938; Shut ova-Korzh, 1958) and in the mouths of the ziaistva (Proceedings of the Research Institute for Fish­ great Siberian rivers (Galkin, 1940; Svetovidov, 1952), eries), 2: 41-77. (In Russian). but no one has applied genetic methods to this herring. Averintsev, S. V 1928. Sel’di Belogo moria ch. 2, 3 (Herring of the White Sea, parts 2 and 3). Trudy Nauchnogo instituta The use of a complex of methods including protein elec­ rybnogo khoziaistva (Proceedings of the Research Institute trophoresis made it possible to describe the stocks of her­ for Fisheries), 3-4: 73-142. (In Russian). ring in the eastern Norwegian ljords as being related to Averintsev, S. V. 1934. Seldi severa SSSR: Vazhneishie cherty Pacific herring (Jorstad e t a t., 1991). ikh biologii i promysla (Herring of the north of the USSR: Genetic methods such as protein electrophoresis and the most important features of its biology and fishery). variation in structure of nucleic acids were widely ap­ Priroda, 10: 64-68. (In Russian). Berg, L. S. 1923. Ryby presnykh vod Rossii (Freshwater Fish plied to herring from the Atlantic and Pacific (Anderson of Russia). 2nd Edition. Gosudarstvennoe izdatel’stvo, Mos­ e t a l., 1981; Kornfield et a t., 1982; Grant, 1984; Grant cow. 536 pp. (In Russian). and Utter, 1984; Ryman e ta t., 1984; Kobayashi, 1993). Borisov, P. G. 1964. Nauchno-promyslovye issledovania na These studies indicated that, in general, the herring morskikh i presnykh vodoemakh (Marine and Freshwater represents an example of a species with a very low Fishery Research). 2nd Edition. Pischevaia promyshlen- degree of population differentiation (Kirpichnikov, nost’, Moscow. 260 pp. (In Russian). Borodatova, Z. A. 1928. Materialy k poznaniu belomorskikh 1987). Contrary to this conclusion, the chromosomal sel’dei. I. Sel’d’ vesennego promysla v raione Soroki v 1927 analysis of the White Sea herring shows a rather clear godu. 2. Sel’di Solovetskikh ostrovov (Materials to knowl­ genetic differentiation between herring populations. edge of the White Sea herring. 1. Herring of spring fishery near the town of Soroka in 1927. 2. Herring of Solovetsky Islands). Trudy Nauchnogo instituta rybnogo khoziaistva Acknowledgements (Proceedings of the Research Institute for Fisheries), 3(2): 37-82. (In Russian). Bückmann, A„ and Parrish, B. B. 1958. Conclusions reached 1 would like to thank Julia Lajus for her advice on his­ at Special Scientific Meeting on "Herring Races" held at torical matters and help in preparing the manuscript, and Charlottenlund on 25-27 September 1956. Rapports et Pro­ Emory Anderson and an anonymous referee for their cès-Verbaux des Réunions du Conseil International pour valuable comments on the manuscript and improve­ l’Exploration de la Mer, 143(2): 8-9. ments to the English. Danilevsky, N. J. 1862. Rybnye i zverinye promysly v Belom i Ledovitom moriakh (Fisheries and marine hunting in the White and Arctic Seas). Issledovania o sostoianii rybolovst- va v Rossii (Investigations on the Status of Fisheries in Rus­ References sia), Vol. 6, St. Petersburg. 257 pp. (In Russian). Derjugin, K. M. 1928. Fauna Belogo moria i usloviia ee Altukhov, K. A. 1975. Sel'd' Solovetskikh ostrovov (Herring of suschestvovaniia (Fauna of the White Sea and environmen­ Solovetsky Islands). Issledovaniia fauny morei (Exploration tal conditions of its existence). Issledovania morei SSSR of Fauna of Seas), 16: 53-91. (In Russian). (Exploration of the USSR seas), Vol. 7/8, Leningrad. 445 Anderson, L., Ryman, N., Rosenberg, R., and Stahl, G. 1981. pp. (In Russian). Genetic variability in Atlantic herring (Clupea harengus Dmitriev, N. A. 1946. Biologiia i promysel belomorskoi sel’di harengus): description of protein loci and population data. (Biology and Fishery of the White Sea Herring). Pische- Hereditas, 95: 69-78. promisdat, Moscow. 88 pp. (In Russian). Andreeva, A. P., Jorstad K., Karpov, A. K.. and Novikov, G. G. Dushkina, L. A. 1973. Influence of salinity on eggs, sperm and 1995. Belkovyi polimorfizm sel'di Belogo moria (Protein larvae of low-vertebrate herring reproducing in the coastal polymorphism of the White Sea herring). Problemy waters of the Soviet Union. Marine Biology, 19: 210-223. izucheniia, ratsional’nogo ispol’zovaniia i okhrany pripod- Dushkina, L. 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