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Contribution to the biology of the Pacific herring, Clupea pallasii, and the condition of the fishery in Alaska

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... Herring have not been taken for food purposes in Alaska in recent years. Rounsefell (1930) compiled Alaska herring catch statistics through 1928 relying on various sources of information. For herring harvests prior to 1904, RounsefeU (1930 relied on catches reported in Moser (1899 and1902) and Cobb (1906). ...
... Beginning in 1904, the Bureau of Fisheries required every individual or company fishing in Alaska to make a sworn annual statement of the total amounts and kinds of fshery products prepared, and of the amounts, kinds, and value of fishing gear, boats, and other apparatus used. Rounsefell (1930) consulted these annual sworn statements as well as the annual statistical review and monthly numbers of the trade Journal "Pacific Fisherman", individual herring company production records, and company receipt books in his compilation of Alaska herring catch statistics. Rounsefell (1930) listed herring catch statistics from Southeast Alaska for 19 10-1928, Prince William Sound for 19 17-1928, Cook Inlet for 1914-1928, and Kodiak for 1912-1928, Rounsefell(1935 extended the Prince William Sound catch series to 1930. ...
... Rounsefell (1930) consulted these annual sworn statements as well as the annual statistical review and monthly numbers of the trade Journal "Pacific Fisherman", individual herring company production records, and company receipt books in his compilation of Alaska herring catch statistics. Rounsefell (1930) listed herring catch statistics from Southeast Alaska for 19 10-1928, Prince William Sound for 19 17-1928, Cook Inlet for 1914-1928, and Kodiak for 1912-1928, Rounsefell(1935 extended the Prince William Sound catch series to 1930. ...
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Historical and contemporary fisheries for herring bait in Alaska are reviewed and summarized.
... Beluga distribution within Cook Inlet has been shifting to the upper Inlet from a broader distribution that more regularly included the lower (Kachemak Bay) and the mid-inlet . Belugas used to feed on herring in Kachemak Bay prior to the late 1920's when herring declined there (Rounsefell, 1930). In April, hunters from Nanwalek would harvest belugas when they fed on herring in Halibut Cove within Kachemak Bay (Rounsefell, 1930;Stanek, 1994). ...
... Belugas used to feed on herring in Kachemak Bay prior to the late 1920's when herring declined there (Rounsefell, 1930). In April, hunters from Nanwalek would harvest belugas when they fed on herring in Halibut Cove within Kachemak Bay (Rounsefell, 1930;Stanek, 1994). The distributional shift to the upper Inlet could be due to fewer belugas available to fi ll the historic habitat, the upper Inlet becoming better habitat, or the lower Inlet becoming poorer habitat. ...
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At least five stocks of beluga whales, Delphinapterus leucas, are found in Alaska waters: Beaufort Sea, eastern Chukchi Sea, eastern Bering Sea, Bristol Bay, and Cook Inlet. The two northernmost stocks (Beaufort Sea and eastern Chukchi Sea) are highly migratory; the two southernmost stocks (Bristol Bay and Cook Inlet) are nonmigratory. Little is known about the seasonal movements and distribution of the eastern Bering Sea stock. Beluga populations in Alaska are thought to be stable or increasing, except for the Cook Inlet stock which is listed as endangered under the Endangered Species Act. We analyzed stomach contents from beluga whales collected between the months of March and November taken in subsistence harvests, from belugas found dead, and from belugas collected for research. We describe prey species and their percent frequency of occurrence (% FO) as well as potential biases from the seasonality of prey relative to the timing of sampling, and differential feeding and digestion. Diet was highly variable among stocks. The predominant fish species of the Beaufort Sea stock was Arctic cod, Boreogadus saida (21% FO), although shrimp (60% FO) and smoothskin octopus, Benthoctopus leioderma (42% FO) were found more frequently. Although the eastern Chukchi Sea stock ate more saffron cod, Eleginus gracilis (7% FO) than Arctic cod (3% FO), shrimp (73% FO) and echiurids (27% FO) were more prevalent than fish. The eastern Bering Sea stock had the most diverse diet, and dominant fish species included saffron cod (95% FO), rainbow smelt, Osmerus mordax (62% FO), several species of sculpin (Family Cottidae) and flatfish (Family Pleuronectidae), both at 48% FO, and Arctic cod at 43%. Dominant invertebrates included shrimp (86% FO), with polychaetes, isopods, bivalves, amphipods, and echiurids ranging from 29 to 38% FO. Pacific salmon, Onchorhyncus spp., predominated over cod in Bristol Bay (81% FO) and Cook Inlet (67% FO) beluga stocks, and invertebrates appeared to be less prevalent prey. In Bristol Bay, smelt were also eaten more often (43% FO) than cod (3% FO), while in Cook Inlet cod were eaten more often (39% FO) than smelt (11% FO). Invertebrates were common in the diet of all Alaska beluga stocks and shrimp (mostly from the family Crangonidae) were the most prevalent.
... From 1995 through 1997, very few herring were observed in region 5 compared to regions 1, 3, 6, and 8 (Stokesbury et al. 2000). Earlier this century, fisheries biologists grappled with the issue of delineating herring populations between the OK and PWS (Rounsefell 1930). Schroeder (1989) speculated that the OK was an " overflow " juvenile nursery area of the PWS stock and should be managed as part of the PWS population. ...
... Respondents observed mixed-age and size-structured schools of herring as well as spawning areas not reported by ADF&G. Mixed-age schools (with juveniles ) in PWS previously had been reported to occur after spawning (Rounsefell 1930). The mixed schools observed by respondents were in regions or seasons outside of those studied by ADF&G. ...
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We documented qualitative ecological information about non-harvested fish age classes and spe- cies from resource users and area residents. Our primary objective was to compile local and traditional eco- logical knowledge about the distribution, abundance, ecology, and associated changes over time of Pacific herring Clupea pallasiand other forage fish species in Prince William Sound (PWS) and the Outer Kenai Pen- insula (OK) in Southcentral Alaska. A secondary objective was to provide ecological information to aid in de- veloping study or management plans concerning herring and other forage fish. Both objectives were met by developing an oral interview protocol, selecting and interviewing key informants in 5 Alaskan communities, and developing a geographic database. Researchers tape-recorded and mapped,respondents’ observations. Survey questions fell into 6 categories: 1) life history stage and species of the fish observed, 2) fish behav- ior and school characteristics,3) presence and behavior of co-occurring predators, 4) seasonal spatial distri- butions observed, 5) decadal shifts observed, and 6) observation and method activity. Forty-eight interviews were conducted. The earliest observation was from 1934. Thirty-seven respondents were commercial,fisher- men and 17 were pilots. Respondents made most observations of juvenile herring schools from planes. Other observations came from net catches, visual sightings, and sonar output. Most observations were made dur- ing summer (June through August), probably due to both shallow distribution of schools and an increase in human,activity during this season. In PWS the spring spatial distribution of herring was significantly differ- ent from summer and fall-winter, but the latter 2 were not significantly different. Spatial distributions of her- ring in the OK were significantly different from one another in all 3 seasons, and the differences were more highly significant than in PWS. Most observations concerned juvenile herring, but locations of herring spawn- ing overlap with adult herring, Pacific sand lance Ammodytes hexapterus, capelin Mallotus villosus, capelin spawning, and eulachon Thaleichthys pacificuswere also documented. Most respondents were able to dis- tinguish herring from other species by their school shape, school color, behavior, and location within a bay. Some pilots believed sunny days were better than overcast days for distinguishing herring from forage fish schools because herring schools “flash silver” and forage fish (mainly sand lance), also called “feed fish” or
... Capelin were also captured in three of eight seine sets in December. Although herring were present throughout the summer (Fig. 3), almost all of them (99%) were captured in five June and four August sets at one area (Halibut Cove) historically known for its aggregations of herring (Rounsefell, 1930). Species abundance, diversity (H'), and richness (D) in the nearshore Kachemak Bay fish community increased steadily from April to June, peaked in July, and declined rapidly in September. ...
... Unequivocal declines in seabird populations (predominantly murres and black-legged kittiwakes) at Chisik Island (Slater et al., 1994) may be related to declines of locally abundant forage fish, particularly sand lance. It is possible that historically larger numbers of capelin (Piatt and Anderson, 1996 ) or herring (or both) (Rounsefell, 1930; Reid, 1971) may have inhabited this region when stocks of pelagic seabirds were higher prior to the mid-1970s. Colder than average temperatures prior to the late 1970s would have favored both of these fish species (Ware, 1995; Frank et al., 1996). ...
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Nearshore and shelf fish communities were studied in three areas of lower Cook Inlet, Alaska: the Barren Islands (oceanic and well-mixed waters), Kachemak Bay (mixed oceanic waters with significant freshwater runoff), and Chisik Island (estuarine waters). Fish were sampled with beach seines (n=413 sets) and midwater trawls (n=39 sets). We found that lower Cook Inlet supported a diverse nearshore fish community of at least 52 species. Fifty of these species were caught in Kachemak Bay, 24 at Chisik Island, and 12 at the Barren Islands. Pacific sand lance dominated Barren Islands and Kachemak Bay nearshore habitats, comprising 99% and 71% of total individuals, respectively. The nearshore Chisik Island fish community was not dominated by any one species; instead it exhibited higher diversity. These spatial differences appeared linked to local oceanographic regimes and sediment influx. Analysis of historical data revealed that the nearshore Kachemak Bay fish community changed significantly between 1976 and 1996, showing increased diversity and abundance in several taxa, notably gadids, salmonids, pleuronectids, and sculpins. Decadal differences appeared to be related to large-scale climate changes in the North Pacific. Catches of most taxa peaked in May-August, and were low during other months of the year. Several species were present for only part of the summer. Species composition of seine catches differed significantly between consecutive high and low tides, but not between consecutive sets or years. Midwater trawls took 26 species, 14 of which were present in Kachemak Bay, 19 near Chisik Island, and 7 at the Barren Islands. Community structures in shelf and nearshore waters were similar: diversity was high and abundance low at Chisik Island, whereas a few abundant species dominated at both Kachemak Bay and the Barren Islands. In addition, the low fish abundance near Chisik Island appeared to be related to declining seabird numbers at this colony.
... For instance, it is possible that exceptional feeding conditions in Southeast Alaska may allow fish to grow both gonadally and somatically, whereas that may not occur for NSS Atlantic herring. It is also possible that the shorter migrations of Sitka Sound Pacific herring (based on limited studies of Southeast Alaska stocks; Rounsefell, 1930;Rounsefell and Dahlgren, 1933;Dahlgren, 1936;Carlson, 1980;Csepp et al., 2011) allow for a smaller reduction in somatic growth compared to reductions in somatic growth from energetically costly migrations for NSS Atlantic herring (Slotte, 1999b). As scales for this study were collected in a single year, it is possible that unusual environmental influences may have limited the reduction in growth for maturing Sitka Sound Pacific herring in 2017 (such as small first year gonads in response to poor feeding conditions), whereas NSS Atlantic herring scale collections made over many years moderated year-specific environmental influences thereby revealing differences in growth between immature and mature fish. ...
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Age at maturation is linked to population productivity and directly impacts forecasts of population biomass and resultant harvest limits set by management agencies. Based on the original concept that scale growth can be used as an index of body growth (length) from Johan Hjort and his collaborators, the current study examined a new way of estimating maturity empirically using scales for Pacific herring. We hypothesized that Pacific herring that will spawn in a particular year (spring) will have reached a sufficient size and have reduced growth (length) the summer prior, compared to immature herring, as energy will be allocated to reproduction rather than somatic length. Model results suggested that there was no difference in the measurement of all scale growth prior to the last increment (growth up until the summer prior) nor in the outer ring measurement (growth the summer prior to spawning) of immature and mature female herring; using scales to estimate maturity empirically was not successful in this study. It is possible that the sample size in this study may have been too low to detect differences in growth or differences in growth may be better represented by changes in length-specific mass.
... Also important are other energy-rich fishes, such as Pacific herring Clupea pallasii, capelin Mallotus villosus, Pacific sand lance Ammodytes hexapterus, northern lampfish Stenobrachius leucopsarus, and eulachon Thaleichthys pacificus. All these fish species occur seasonally in the diet of Steller sea lions in Alaska (Pitcher 1981, Sinclair & Zeppelin 2002, but little is known about their life history, spawning patterns, and distribution in Alaska, with the exception of herring (Rounsefell 1930, Rounsefell & Dalgreen 1935, Carlson 1980. These seasonally abundant, energy-rich forage fish are critical to the biology of many predators (Springer 1992, Skov et al. 2000, Bishop & Green 2001, Litzow et al. 2002, Marston et al. 2002, Rodway et al. 2003. ...
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Energetic demands are high for Steller sea lions Eumetopias jubatus during spring, when females are pregnant and lactating and males are preparing for extended fasts on breeding territories. Therefore, we predicted that the distribution of sea lions in SE Alaska in spring would be influenced by the distribution of spring spawning aggregations of high-energy prey species (Pacific herring Clupea pallasii and eulachon Thaleichthys pacificus). The spatial distribution of sea lions during spring reflected the distribution of spawning eulachon in northern Southeast Alaska, particularly in Lynn Canal and along the Yakutat forelands. Haulouts with peak numbers of sea lions in spring were located significantly closer to eulachon spawning sites than haulouts that peaked at other times of year. Some haulouts were occupied only during the eulachon spawning period. The maximum number of sea lions at haulouts in spring was inversely correlated with the distance to the closest eulachon aggregation and was positively associated with the number of eulachon within 20 km. Aerial surveys conducted every 7 to 10 d during March through May in 2002 and 2003 revealed large numbers of sea lions in the water at herring spawning sites in 2002 and 2003; however, there were no significant relationships between the number of herring spawning sites and number of sea lions (except at distances > 60 km). The number of sea lions was greater at herring spawning sites in 2003, corresponding to higher herring biomass. Seasonally aggregated, high-energy prey species influence the seasonal distribution of sea lions and may be critical to their reproductive success.
... Adults of this species are extremely abundant and widespread in bays and inlets throughout the Salish Sea. (Lord, 1866;Günther, 1868;Haist and Stocker, 1985;Hammond, 1887;Rounsefell, 1930;Fitch and Lavenberg, 1975;Miller and Borton, 1980;Morrow, 1980;Whitehead, 1985;Swift et al., 1993;Penttila 7 ; Gaydos and Brown, 2009) ...
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As part of a current effort to restore the Salish Sea, a 16,925-km2 inland waterway shared by Washington State and British Columbia, a definitive, up-to-date list of the fishes that inhabit this marine ecosystem has been badly needed. The last such effort was published more than three decades ago. In response to this deficiency, we compiled information from various sources and identified 253 fish species observed in marine or brackish waters of the Salish Sea ecosystem, an increase of nearly 14% since the last published checklist. These 253 species, encompassing 1 myxinid, 2 petromyzontids, 18 chondrichthyans, 2 chondrosteans, and 230 teleosts, are contained within 78 families and 31 orders. This comprehensive list of the Salish Sea ichthyofauna will serve as a foundation for determining the occurrence of new species and perhaps the disappearance of others, enabling the selection of species as indicators of ecosystem health, and will provide a basis for identifying the mechanisms responsible for marine animal declines.
... In spring, between March and May in SE Alaska, herring move to spawning areas (Carlson 1980). During summer, herring typically move to feeding areas, many of which are located along the outer coast of SE Alaska (Rounsefell 1930). ...
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Steller sea lions Eumetopias jubatus are central-place foragers that forage in the marine environment while using terrestrial sites to rest and care for young. Some terrestrial sites are used seasonally; however, the reasons for doing so are not fully understood. We addressed the hypothesis that seasonal availability of prey influences seasonal abundance and diet of sea lions. We quantified monthly prey availability and sea lion abundance and quarterly diet composition at Benjamin Island in SE Alaska (2001-2004). Large numbers of sea lions occupied Benjamin Island during the non-breeding season from October to April when Pacific herring Clupea pallasii biomass was highest. Herring was the most common species in sea lion diet (frequency of occurrence [FO] = 90%) and comprised over 81 % of the available pelagic prey biomass and 96 % of the energy encountered during pelagic surveys. Walleye pollock Theragra chalcogramma accounted for 19 % of the available prey biomass but was only slightly less common in sea lion diet (FO = 88 %) than herring. Herring biomass was correlated with the number of sea lions; in contrast, there was no relationship between pollock biomass and number of sea lions. Several fish species were found in nearshore areas, but were uncommon in sea lion diet. Sea lions consumed the available pelagic prey but little of the available nearshore prey. The FO of herring and pollock in sea lion diet did not differ significantly between seasons; however, the FO of other seasonal prey species differed between seasons. Seasonal occupation of Benjamin Island by sea lions is influenced by seasonally available, densely aggregated, energy-rich prey.
... Thus, although estuarine utilization is more limited for Pacific than Atlanticcoastfishes (McHugh, 1967;Pearcy &Myers, 1974), the problemofestuarine entry may be particularly acute for Pacific coast fishes whose young inhabit these nursery areas. Pacific estuaries are important to several fish species, including Pacific herring, Clupea harengus pallasi, which migrate to, and spawn in, estuaries (Rounsefell, 1930), and anadromous salmon which utilize estuaries as nurseries during their migration to the sea (Healey, 1980). Few offshore Pacific species; however, use estuaries as nurseries after metamorphosis (McHugh, 1967). ...
Article
English sole, Parophrys vetulus, spawn in shelf waters off the west coast of North America and early development occurs in coastal waters. Near metamorphosis, however, larvae recruit to nearshore and estuarine nursery areas, an uncommon life-history feature for a species in this region. Recruitment of larval P. vetulus to Yaquina Bay, Oregon, was sampled with moored nets on a weekly basis. Recruitment began during night flood tides in late February and was characterized by three peaks during the season. Planktonic recruitment continued into June, whereas benthic juveniles began to emigrate from the estuary in late May.The data suggest that two developmental stages of recruits immigrated to Yaquina Bay. The first are newly transforming larvae which enter the bay earliest within each peak of recruitment; peaks of abundance are related to onshore Ekman transport. This transport also brings larvae to shallow areas along the open coast where they settle. These transforming stages are still capable of pelagic swimming activity and continue to recruit to the estuary, typically swimming deeper in the water column than the early stages. Estuarine factors, particularly bottom salinity at the end of ebb tide, are most strongly correlated with recruitment of these stages. It seems that tidal stream transport is the primary mechanism used by English sole to recruit to the estuary, but the mechanism of locating the estuary and timing of entry is relatively complex.
... The geographic structure of Pacific herring (Clllpea p a l h i ) stocks has been intensively studied, especially along the Pacific coast of Canada, because of its commercial importance. Geographic variation in morphological traits, particularly vertebral number and scale patterns (Rounsefell 1930; Rounsefell and Dahlgren 1935; Tester 1949; Rowel1 1980 ), and tagrecovery methods (Stevenson 1955; Hourston 1982) have been used to describe its stock structure. The results of these studies show that Pacific herring are subdivided into several more or less distinct populations with limited migration between spawning areas. ...
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Studies of stock structure in Pacific herring (Chpea p a l h i) using geographic variation in morphology and growth, and tagging methods, have shown that Pacific herring are subdivided into numerous more or less distinct populations or stocks having limited migration between stocks. Homing of adults to previous spawning areas and larval retention mechanisms may enhance reproductive isolation between spawning areas and have been postulated to produce genetic differences between stocks. In this study the geographic distributions of inherited biochemical markers were used to measure the genetic component of stock structure in Pacific herring. The gene products of 40 protein-coding loci were examined by starch-gel electrophoresis in 2% samples collected from locations extending over most of the range of Pacific herring in the North Pacific Ocean and Bering Sea. These results show that there is a very small genetic component to the stock structure described by nongenetic methods. No significant allele-frequency differences were detected among three samples of Asian herring. In the eastern Bering Sea, the northern populations were genetically distinct from southern populations on the Seward Peninsula and in Bristol Bay, but no genetic structure was detected within these areas. In the eastern North Pacific Ocean, the Gulf of Alaska populations were genetically distinct from the remaining southern populations. Some genetic structure was detected in the Gulf of Alaska but not among the remaining southern populations. Using the stepping-stone model of migration and the observed amount of genetic divergence among populations, we show that small-scale genetic differences would not be expected, given the amount of migration observed between spawning areas. Another important result of this study is the discovery of two genetic races, Asian-Bering Sea herring and eastern North Pacific herring, having an average Nei genetic distance (D) between samples of the two races of 0.839; B between populations within each race averaged 0.0009. We postulate that the two North Pacific races of herring arose as a result of repeated Pleistocene glaciation on the southern coast of Alaska, which created a barrier to gene flow and permitted the two oceanic groups to diverge genetically. Des etudes sur la structure des stocks de harengs du Pacifique (Clupea pa88asi) basees sur la variation geographique de la morphologie et de la croissance, ainsi que des marquages, indiquent que cette espece se subdivise en un grand nombre de populations ou stocks plus ou moins distincts, avec echanges limit& entre stocks. Le retour des adultes a des frayeres qu'ils ont deja frequentees, de meme que des mecanismes de retention des larves, peuvent favoriser I'isolement reproducteur entre zones de ponte, et on a suppose qu'ils causaient des differences genetiques entre stocks. Dans la presente etude, nous avons utilise la distribution geographique de marques biocherniqrres hereditaires pour mesurer la composante genetique de la structure des stocks. Sur 2% echantillons recueillis 2 des endroits repartis sur la plus grande partie de
... Pacific Herring (Clupea pallasi) is a pelagic forage fish species widely distributed in the northeast Pacific, from California to the Bering Sea (Rounsefell, 1930;Miller and Schmidtke, 1956;Wespestad and Fried, 1983). There is a long-standing interest in Pacific Herring biology and ecology as this species generates large economic benefits for commercial fisheries and is a key species in First Nations communities and cultures in the northeast Pacific (Taylor, 1964;Schweigert et al., 2009;DFO, 2018). ...
Article
Determining how fish respond to variation in biotic and abiotic conditions is a crucial prerequisite to forecasting changes in productivity and spatial distribution of fish stocks and designing sustainable marine resource management strategies. In the present study, we investigated the physical and biological drivers of the spatio-temporal dynamics of Pacific Herring (Clupea pallasi), which is a marine forage fish species important for commercial fisheries and First Nations in the northeastern Pacific. We fit multivariate spatio-temporal models to fisheries-independent trawl- and acoustics-based data collected off the West Coast of Vancouver Island (WCVI), Canada, during summers over the period 2006–2014. We evaluated the effects of the main ocean environmental drivers of WCVI lower trophic level productivity, including sea surface temperature (SST), chlorophyll a, fluorescence, salinity, oxygen, transmissivity and zooplankton density on variation in Pacific Herring biomass. Models were also used to measure spatio-temporal covariation with other pelagic, semi-pelagic, and bottom-associated fish species occurring off the WCVI to address potential competitive and predation interactions. Through application of these spatio-temporal models we found: (i) Pacific Herring biomass off the WCVI increased during 2006–2014; the highest Pacific Herring biomass was repeatedly found on the continental shelf (depths < 185 m) while low densities were usually observed along the shelf break (depths > 185 m), where Euphausiids, Pacific Hake, Sablefish and Arrowtooth Flounder were more abundant, which could reflect predation avoidance behaviour; (ii), the local biomass of Pacific Herring was related quadratically to the average SST in May; (iii) a positive covariation in spatio-temporal densities between Pacific Herring and its common zooplankton prey, supporting a potential bottom-up control hypothesis; (iv) a negative covariation in spatio-temporal densities between Pacific Herring and both Pacific Hake and Pacific Sardine, which could reflect predation and competitive interactions, respectively; and (v) a positive covariation in spatio-temporal densities between Pacific Herring and several groundfish species (i.e., Arrowtooth Flounder, Sablefish, Pacific Halibut, Pacific Cod), which highlights the need for an accurate assessment of the relative contribution of those species to the total summer predation pressure experienced by Pacific Herring off the WCVI. The findings of this study contribute to a better understanding of the WCVI marine ecosystem.
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Pacific herring Clupea harengus pallasi are winter-spring spawners which exhibit a S-N latitudinal cline in spawning time. Atlantic herring Clupea h. harengus consist of both winter-spring and summer-autumn spawning groups characterized in the NE Atlantic by oceanic, shelf, and coastal populations. The oceanic group are large migratory fish spawning off the coasts of Norway and Iceland. The shelf group includes the various locally migratory North Sea populations adjacent to the British Isles. The coastal groups consist of smaller fish restricted to the Baltic and White seas. In the NW Atlantic, spawning occurs from N Labrador to Virginia with spring spawners predominating in the north and fall spawners in the south. Temperature is one of the factors that determine when spawning occurs. The Atlantic herring exhibits sexual dimorphism in the spawning act with only the female interacting with the spawning substrate. Both sexes of the Pacific herring make physical contact with the substrate on which the adhesive eggs are deposited. Spawning grounds are located in high-energy environments, either nearshore for spring spawners or in tidally active areas for fall spawners. Spawn is deposited on marine vegetation or on bottom substrate, such as gravel, which is free from silting. The eggs are tolerant to temperatures in the range of 5-14oC and salinities in the range of 3-33per mille. Egg mortality results mostly from suffocation due to high egg densities and silting, predation, and, in intertidal spawn, from stresses imposed by exposure to air and from egg loss by wave action.-from Authors
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Allozyme markers have been used to investigate the genetic structure of stocks of herring. Results from these studies show little genetic differentiation between the recognised stocks. Much of the traditional evidence for discrete stocks, based on morphometrics, spawning times and tag returns, is shown to be weak and based on a typological concept. In fact this evidence indicates the occurrence of gene flow between neighbouring spawning aggregations. It is suggested that the herring stock is a transient sub-division and has no taxonomic or evolutionary status.
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A spawning pheromone in the milt (semen) and testes of the Pacific herring,Clupea harengus pallasi, triggers spawning in sexually mature fish of both sexes and is thought to facilitate school spawning of this species. We found the response to the pheromone to be a stereotyped behavioral sequence consisting of a graded extension of the gonadal papilla, release of gametes, and spawn deposition behavior. The response is triggered by an olfactory stimulus, as demonstrated by the elimination of the response by occlusion of the nares. Stimulus concentrations of an approximate 1:500 dilution of fresh milt or the equivalent of 0.02 g of fully mature testes per milliliter were required to elicit a response in 50% of ripe herring that are responsive to the pheromone. Female fish appeared to be less sensitive to the pheromone in milt than males early in the spawning season, but not thereafter. The average duration of responses of male fish was longer after exposure to concentrated milt than to testes extracts, but no consistent difference in response times of the two sexes was detected. Factors other than the spawning pheromone, maturity of the fish, and stress also were found to influence the spawning response. For example, exposure to shallow (3 cm) water in a small tank induced “spontaneous” papilla extension and spawning approximately 20 min after refilling the tank; occluding the nares prevented this response. Also, the presence of floating kelp (Macrocystis) resulted in prolonged spawning in a large tank after pheromonal stimulation.
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In wenigen Sätzen will ich hier die aus der Rassenanalyse des Herings gewonnenen Hauptergebnisse zusammenfassen.
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Fecundity relationships of 156 female herring (Clupea harengus pallasi) from Prince William Sound were studied. Upon comparison with other North American populations of Pacific herring, we found a progressive decrease in fecundity at a particular body size with increasing latitude. We also found an increase in mean length and mean fecundity of spawner with latitude.
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This paper uses GIS and spatiotemporal analysis of a historically and culturally modified marine ecosystem to evaluate Pacific herring abundance, declines, vulnerabilities, and future prospects, about which a Native Tribe and state fisheries managers disagree. In 2008, the Sitka Tribe of Alaska (STA) requested that an area within its traditional waters be closed to commercial sac roe fishing to protect spawning Pacific herring (Clupea pallasi), a key species for Native subsistence and marine ecosystem health. This proposal was opposed by the Alaska Department of Fish and Game (ADF&G), which estimated that adequate biomass was available to accommodate all herring users' needs. The disagreement exposes divisions between the Tribe's and the State's conceptualizations of the status, health, and management priorities for fisheries and marine ecosystems. The Tribe's model is one of cultivated abundance, wherein herring eggs are harvested conservatively and habitat is enhanced to make coastal spawning areas more productive, stable, and resilient. The State's paradigm, in contrast, follows a constitutional mandate to manage fisheries for Maximum Sustained Yield (MSY). A single-species biomass model is used to estimate "surplus" herring for commercial roe harvesting within management areas. This work analyses and compares the spatiotemporal prescriptions of State and Indigenous models of herring fisheries management as they are used within debates over a closed area (Proposal 239), and assesses their relative potential for improving herring fisheries and marine ecosystem management using a combination of GIS spatial and scientific analysis and traditional ecological knowledge.
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The life of Frederic Morton Chamberlain (1867–1921) was marked by his many contributions to fishery science. As an assistant with the old U.S. Bureau of Fisheries, he pioneered life history studies of salmon in California and Alaska, and assisted in the collection of hundreds of thousands of natural history specimens from the Pacific Ocean as resident naturalist on the U.S.S. Albatross. He was innovative in the use of field photography in fishery work, and the design and construction of capture nets and collecting gear. His detailed reports were important in documenting the conditions of the native ichthyofauna in Alaska, Arizona, and California at the turn of the century and have been heavily relied upon by subsequent investigators. His ability to design and conduct scientifically justifiable studies to solve fishery problems while under many kinds of hardships is a testament to his background and dedication to the field. He represents the prototype of the modern fishery biologist.
Thesis
Sebastes schlegelii and Sebastiscus marmoratus are commercially important marine ovoviviparous fish in the Northwestern Pacific. In the present study, morphological, otolith morphological, mitochondrial DNA sequence, AFLP markers and microsatellite DNA markers were used to estimate their gene diversity, population structure and phylogeographic pattern. The main results are as follows: (A)Molecular phylogeography of Sebastes schlegelii (1)In order to study the morphological variations among nine populations of S. schlegelii (244 individuals) collected from Yantai, Lianyungang, Rizhao, Dandong, Qingdao, Dalian, Weihai, Jiaonan and Aomori, eleven meristic characters and fifteen measureable characters were measured and the data were standardized by using log-transformed method and then analyzed by using one-way ANOVA, discriminant analysis method and principal component analysis method. One-way ANOVA results showed that there were no significant differences between eight Chinese populations, but significant differences were found between Aomori population and eight Chinese populations. Discriminant analysis results showed that the correct classification ratio was 96.0% in Aomori population, and the cumulative contribution ratio was 80.7%. In the result of principal component analysis, four principal components explained 72.3% of the total variance; the first two principal components reflected the morphological difference of fish’s head and trunk. The results of multivariate statistical analysis methods showed that there were significant differences between Aomori population and eight Chinese populations. (2)Multivariate statistical analysis that includes one way analysis of variance, discriminant analysis and principle analysis was conducted for 9 otolith shape indices of 228 individuals of Chinese S. schlegelii. Fourier analysis was used to be compared with shape indices method. One way analysis of variance showed significant differences were found between Dandong population and Rizhao population in 9 otolith shape indices; Rongcheng population showed significant differences in 9 otolith shape indices from Jiaonan, Qingdao, and Rushan population, respectively. While insignificant differences were found between Rushan and Jiaonan in their 9 otolith shape indices (P>0.05); the first two principle components explained 71.953% of the total variance; the discriminant analysis of the 9 shape indices showed that the correct classification was 64.9% among the 8 populations. Fourier analysis showed that the correct classification was 43.8%, which was 21.1% lower than that of shape indices analysis; the first 9 principle components explained 89.2% of the total variance. Three processes of multivariate statistical analysis and Fourier analysis showed that there was insignificant difference between eight populations of Chinese S. schlegelii. (3)The present study deals with the structure of mitochondrial DNA control region of the black rockfish, S. schlegelii. Two TASs and two cTASs were detected in the species as well as CSB-F, CSB-E, CSB-D, CSB1, CSB2 and CSB3. The results indicated that the structures of these blocks are similar with most marine fishes, but it is special that there are two TASs and two cTASs in the control region of S. schlegelii. One conserved region was found from 450bp to the end of the control region, which is also a special feature of S. schlegelii. All sequences of CSB1, CSB2 and CSB3 blocks were the consensus among different individuals, which is quite different from most vertebrates. In addition, the complete mtDNA control region sequences and the first 449bp of the control region are used to analyze the phylogenetic relationships of S. schlegelii. The phylogenetic trees show a lack of genetic structure among individuals. This study also indicated that the genetic diversity is similar between the wild and cultured individuals, which are important to the fisheries management. (4)S. schlegelii is one of the representative fish that aggregate around drifting seaweed during its early developmental period. Transport of drifting seaweed by ocean currents probably gives larval and juvenile S. schlegelii fish an opportunity to expand their distribution area and to exchange genetic material between widely separated populations. To examine the population genetic structure of S. schlegelii, a 452-bp fragment of the hypervariable portion of the mtDNA control region was sequenced and used to interpret life history characteristics and larval dispersal strategy as well as its historical demography. Two hundred and twenty one individuals from 13 sites along the coasts of China, Japan and Korea were analyzed. Both the levels of haplotype diversity and nucleotide diversity were not very high. The neighbor-joining tree and the network showed that no significant genealogical structures corresponding to sampling locations existed. AMOVA, pair-wise FS T and exact-P test revealed no significant genetic differentiation among populations. The migration rate among populations was strong based on the result of LAMARC. Both mismatch distribution analysis and neutrality tests showed S. schlegelii have experienced a recent population expansion in the late Pleistocene. It is suggested larval dispersal with drifting seaweed, the current environmental factors and late Pleistocene glaciations play an important role in shaping the contemporary phylo-geographic pattern of S. schlegelii. (5)AFLP marker was used to investigate the population genetic structure of S. schlegelii among populations from China and Japan. The results indicated S. schlegelii showed high Nei’s gene diversity. No significant genealogical branches or clusters corresponding to sampling localities were detected by UPGMA tree. The results of AMOVA analysis and pairwise FS T values showed no genetic divergence among different geographic populations and high gene flow was existed. The result of STRUCTURE showed all the populations of S. schlegelii shared one gene pool. The results of AFLP were in accordance with those of mtDNA. (6)Eight microsatellite DNA loci were used to investigate the genetic diversity and population structure of S. schlegelii. The results indicated polymorphism information content (PIC) and observed heterozygosity were high, indicating a high level in gene diversity of S. schlegelii. No significant genealogical branches or clusters corresponding to sampling localities were detected by UPGMA tree. The results of AMOVA analysis and pairwise FS T values showed no genetic divergence among different geographic populations and high gene flow was existed. The result of STRUCTURE showed all the populations of S. schlegelii shared one gene pool. The results of AFLP were in accordance with the results of mtDNA and AFLP. (B)Molecular phylogeography of Sebastiscus marmoratus (1)In order to study the morphological variations among seven populations of S. marmoratus (114 individuals) collected from China (Huizhou and Haikou ) and Japan (Hakata Island and Yokosuka), eleven meristic characters and fourteen morphological characteristics were measured and the data were analyzed by using one-way ANOVA, discriminant analysis method, principal component analysis method and cluster analysis method. One-way ANOVA results showed that there were no significant differences between Hakata Island and Yokosuka in nine indices, but significant differences were found between Hakata Island and Huizhou in twelve indices. Discriminant analysis results showed that the correct classification ratio was 100% in Huizhou, and the cumulative contribution ratio was 90.35%. In the result of principal component analysis, four principal components explained 53.977% of the total variance, the first principal component reflected the feature of fish’s trunk, and the second principal component reflected the feature of fish’s head and rump. Cluster analysis results showed that there were significant differences among populations in China and Japan, Chinese and Japanese populations were pooled in one group separately. The results of multivariate statistical analysis methods showed that there were significant differences among populations in China and Japan. (2)In order to study the otolith morphological variations among four populations of S. marmoratus (101 individuals) collected from China (Huizhou and Haikou) and Japan (Hakata Island and Yokosuka), nine otolith shape index and seventy-seven elliptic Fourier parameters were measured. The data were analyzed by using discriminant analysis method, principal component analysis method and cluster analysis method. The results of discriminant analysis showed that the correct classification ratio ranged from 75% to 100%, and the average classification ratio was 91.1%. The results of cluster analysis showed that there was significant difference among populations in China and Japan, and Chinese and Japanese populations were pooled in one group respectively. According to the result of principal component analysis, the divergence of populations of S. marmoratus could be measured by fewer otolith morphological variables. The results of multivariate statistical analysis methods showed that there was significant difference in otolith morphology among populations in China and Japan. (3)To examine the population genetic structure of S. marmoratus, a 458-bp fragment of the hypervariable portion of the mtDNA control region was sequenced and used to interpret life history characteristics and larval dispersal strategy as well as its historical demography. Two hundred and forty four individuals from 16 sites along the coasts of China and Japan were analyzed. Both the levels of haplotype diversity and nucleotide diversity were high. The neighbor-joining tree and the network showed that no significant genealogical clades corresponding to sampling locations existed. AMOVA, pair-wise Fst and exact-P test revealed no significant genetic differentiation among populations. Both mismatch distribution analysis and neutrality tests showed S. marmoratus have experienced a recent population expansion in the late Pleistocene. (4)AFLP was used to investigate the population genetic structure of S. marmoratus among populations from China and Japan. The results indicated S. schlegelii showed high Nei’s gene diversity. No significant genealogical branches or clusters corresponding to sampling localities were detected by UPGMA tree. The results of AMOVA analysis and pairwise Fst values showed no genetic divergence among different geographic populations and high gene flow was existed. The result of STRUCTURE showed all the populations of S. marmoratus shared one gene pool. The results of AFLP were in accordance with the results of mtDNA. (5)S. marmoratus is a typical sedentary fish and usually lives in the bottom of the coastal zone from Japan to the East China Sea. Next-generation sequencing platform was used to obtain microsatellite loci sequences of S. marmoratus, and forty-eight perfect microsatellite repeats were suitable for primer design. Nine polymorphic microsatellite markers were exploited and characterized for S. marmoratus. The allele number of all polymorphic microsatellite markers ranged from 4 to 12. Expected and observed heterozygosity varied from 0.302 to 0.909 and 0.190 to 0.625, respectively. Six of the nine markers deviated obviously from Hardy– Weinberg equilibrium (P<0.05). In further studies, the genetic variation and population structure of this species would be investigated using these microsatellite loci. (C)DNA barcodes of eight species in genus Sebastes DNA barcode is effective for biological taxonomy and is able to identifying species from any life-history stage. In the present study, eight species which belong to four different subgenera of genus Sebastes found in China sea waters were identified by cytochrome c oxidase I (COI) barcode. The results indicated that the intra-species variation in DNA barcode was less than inter-species variation. When the phylogenetic trees were reconstructed by neighbor-joining (NJ), maximum parsimony (MP), maximum likelihood (ML) and Bayesian methods, all the species clustered in their groups distinguishable by high bootstrap values, which proved that COI barcode is a powerful means to differentiate between species of Sebastes and supports their identification. When the molecular tree was compared to the morphological tree, only S. trivittatus in subgenus Sebastocles settled in the different positions. It is suggested that S. trivittatus is one of the shallowest occurring species in the Northwest Pacific due to its life characters.
Chapter
Publisher Summary This chapter discusses the behavior and physiology of herring and other clupeids. The various characteristics of clupeids are discussed. The Clupeidae are found in all habitats from fresh water, for example some Pomolobus and Harengula species, to sea water. Many of them are anadromous such as Alosa, Caspiolosa and Pomolobus. The family contains species that lay demersal eggs offshore (herring), demersal eggs in shallow water (Clupea pallasii) and pelagic eggs (Sardinops). The basic structure and composition of the gametes is discussed. The herring sperm is of major importance. In the body of the parent herring the gametes are maintained in a relatively favorable environment. The embryonic development of the herring is reviewed. The rate of development is closely linked with temperature, for example at 14° C gastrulation begins 22 hr after fertilization, whereas at 11.5°C the eggs after 22 hr are still in the blastula stage. The anatomy and development of the skeleton and organ systems after hatching is studied in herring.
Article
Meristic characters are enumerable morphological features of fishes. Strictly, meristic features are those that corresponded evolutionarily with body segmentation; however, today, the term is used more broadly. The most commonly enumerated features have been external, including fin spines and fin rays, gill rakers, and scales (along several possible vectors). Internal meristic features that have been analyzed include pterygiophores, vertebrae, branchiostegal rays, and pyloric caeca. Meristic analysis remains a technically simple, inexpensive alternative for the stock identification of fishes. Drawbacks include that in almost all instances, specimens must be sacrificed-it is difficult (although not impossible) to make accurate counts of meristic features on living fish. Also, variation among year-classes as a consequence of environmental effects must be considered. However, there may be instances in which ecophenotypic variation of meristic traits is more effective toward stock identification than are genetic approaches. One instance is where genetic differences are limited because some or all populations to be analyzed are very recent in origin due to natural recolonization or stocking. Another instance is where there is sufficient gene flow among populations to preclude or erode genetic differentiation.
Article
The transformation of Pacific herring (Clupea pallasii) fisheries from communal to commons to neoliberal regulation has had significant impacts on the health and sustainability of marine ecosystems on the Northwest Coast of North America. Due to their abundance, seasonality, and sensitivity in disturbance, herring were carefully cultivated and protected by coastal Tlingit, Haida, and Tsimshian communities. The early industrial fishing era undermined this communalist approach in favor of an unregulated commons for bait and reduction fisheries, attracting non-local fleets and leading to conflicts with local Natives and tragedy of the commons style overexploitation of herring stocks by the mid-twentieth century. Since the 1970s, a re-regulated neoliberal sac roe fishery for Japanese markets has provided new opportunities for limited commercial permit holders, but with further depredations on local spawning populations. This paper uses frame theory and historical and political ecology to show how this transformation was justified by three critical but dubious (re)framings of Southeast herring populations under modern scientific management: (1) a reductionist framing of single species productivity models, expressed as herring “biomass,” within space and time (baseline scale framing); (2) the selective framing and privileging of human industrial predation under maximum sustainable yield (MSY) within a dynamic ecosystem of multiple predator populations (actor relations framing); and (3) the strategic framing of spawning failure events and policy responses to those events by professional fisheries managers (event–response framing). Finally, the paper argues for a new social–ecological systems approach, based on aboriginal models of herring cultivation, to sustain a commercial, subsistence, and restoration economy for the fishery.
Chapter
Abundance of Pacific herring (Clupea harengus pallasi) populations may be limited by (1) the available area of summer feeding habitat, mainly on coastal shelf waters with depths of 200 m or less, and (2) the area of suitable spawning habitat. Shelf areas may serve as summer feeding areas that account for most of the production of herring biomass. In offshore waters, adult herring rarely are found in waters exceeding 200 m depth and herring schools often are seen close (< 5 m) to the bottom. Throughout the N Pacific, the largest herring populations occur in association with large continental shelf areas. Spawning and nursery habitat consists of shallow, near-shore areas protected from the open sea. The relative proportion of shelf and near-shore habitat varies among areas and populations. The Sea of Okhotsk has a broad shelf area, but maximal herring production often may not be realized because of limited available near-shore habitat suitable for spawning and rearing of the larval and juvenile stages. In contrast, in British Columbia there is a relative abundance of near-shore spawn and nursery habitat and a relative scarcity of suitable shelf areas.
Chapter
Pacific herring (Clupea harengus p.11asi) spawn synchronously in large schools of several million fish in the near-shore environment, markedly discolouring the surrounding water with suspended milt. This spawning activity has a rapid onset as a school moves inshore. Males appear to initiate spawning in small pockets from which the activity spreads to the rest of the school (Hay, 1985). Spawning continues for several hours in small schools (Hourston and Rosenthal, 1976) and several days in large schools, after wh:.ch the fish move back into deeper waters (Haegele and Schweigert,:985). Without overt behavioural interaction, the spawning female and ma:.e herring deposit gametes on submerged vegetation or other surfaces as trails of sticky eggs or viscous milt (Schaeffer, 1937). Over 20 layers of eggs result in some cases (Haegele and Schweigert, 1985). The deposited milt, together with milt released mid-water, gradually diss:.pates, resulting in the milkiness typical of herring spawning s:.tes. Hay (1985) suggests that high concentrations of milt may inhibit spawning and hence regulate the density of egg deposition.
Chapter
Marine fishery research had its origin in man’s reactions to a universal natural phenomenon, fluctuations in abundance. From the earliest days of fishing it has been known that catches of particular species are variable. We now know that these fluctuations were caused by annual changes in the success of spawning, or by changes in environmental conditions which altered patterns of migration or geographic distribution, or by both. The natural inclination then, as today, was to blame decreased catches upon one or more activities of man. When fishing was responsible for the decline, the nature of the effect was misunderstood, as it often is today. In a fully developed, but still healthy fishery, the catch-per-unit-of-effort is much less than it was when the virgin stock was first fished, the average size and age of the fish are considerably less, and the stock and its biology may be altered in various other ways. Fishery research began because people wanted to know the reasons for these changes, and if possible how to correct the situation.
Article
Full-text available
Department s f Fisheries and Oceans, Fisheries Research Branch, Pacific BisBogical Station, Nanairno, B.6. V9R 5K6 Haegele, C. W., and J. F. Schweigert. 1985. Distribution and characteristics of herring spawning grounds and description of spawning behavior. Can. J. Fish. Aquat. Sci. 42 (Suppl. I) : 39-55. Pacific herring (Clupea harengus pallasi) are winter-spring spawners which exhibit a south t o north latitudinal cline i n spawning time. I n the eastern Pacific, major spawning populations are concentrated near San Francisco, i n Puget Sound, and along the coasts of British Columbia and southeastern Alaska. l n the Bering Sea, herring spawn in Alaska and along the Kamchatka Peninsula. In the western Pacific, herring spawn in the Sea of Bkhotsk and the Japan Sea. Atlantic herring (Clupea harengus harengus) consist of both winter-spring and summer-autumn spawning groups characterized i n the northeast Atlantic by oceanic, shelf, and coastal populations. The oceanic group are large migratory fish spawning off the coasts of Norway and Iceland. The shelf group includes the various locally migratory North Sea populations adjacent t o the British Isles. The coastal groups consist of smaller fish restricted t o the Baltic and White seas. Bn the northwest Atlantic, spawning occurs from northern Labrador t o Virginia with spring spawners predominating i n the north and fall spawners in the south. Herring typically congregate near their spawning grounds for several weeks t o months prior t o spawning. Temperature is one of the factors that determine when spawning occurs. The Atlantic herring exhibits sexual dimorphism i n the spawning act with only the female interacting with the spawning substrate. Both sexes of the Pacific herring make physical contact with the substrate o n which the adhesive eggs are deposited. Spawning grounds are located in high-energy environments, either nearshore for spring spawners or i n tidally active areas for fall spawners. Spawn is deposited o n marine vegetation or o n bottom substrate, such as gravel, which is free from silting. The eggs are tolerant t o temperatures i n the range of 5-14°C and salinities i n the range of 3-33%. Egg mortality results mostly from suffocation due t o high egg densities and silting, predation, and, i n intertidal spawn, from stresses imposed by exposure to air and from egg loss by wave action. Le hareng d u Pacifique (Clupea harengers pa!lasi), qui fraye en hiver et au printemps, exhibe u n cline latitudinal d u sud au nord au moment de la fraie. Dans le Pacifique oriental, les principales populations reproductrices sont concentr6ies pres de San Francisco, dans le Puget Sound et le long des c6tes de la Colombie-Britannique et d u sud-est de IfAlaska. Dans la mer de Bering, le hareng fraye en Alaska et Be long de %a peninsule de Mamchatka tandis que, dans le Pacifique oriental, il se reproduit dans la mer d'Okhotsk et la mer d u Japon. kes harengs de I'Atlantique (Clupea harengus harengus) se composent de groups qui frayent pendant la periode hives-printemps et efe-automne, groupes caracterises dans I'Atlantique nord-est par des populations peuplant ['ocean, la plate-forme o u les c8tes. Le groupe oceanique est compos6 de gros poissons migrateurs qui se reproduisent au large de la NowGge et de I'lslande, tandis que celui de la piate-forme compte les diverses populations qui migrent Iocalernent dans la mer d u Nord pres des iles Britanniques. Par contre, le groupe c6tier consiste en de plus petits poissons restreints aux mers Baltique et Blanche. Dans IfAt!antique nord-ouest, la fraie a lieu d u Labrador septentrional a la Virginie; les reproducteurs printaniers se concentrent au nord et les geniteurs d'autornne, au sud. Typiquernent, le hareng se regroupe pres des frayeres pendant plusieurs semaines o u mois avant la fraie. ha temperature represente u n des facteurs qui d6terminent le moment de la fraie. hors de l'accouplement, le hareng de I'Atlantique exhibe u n dimorphisme sexuel : seulement la femelle interagit avec le substrat de la frayere. Par contre, chez le hareng d u Pacifique, les deux sexes entrent en contact physique avec le subtrat, o u les oeufs adherents sont deposes. hes frayeres sont situees dans des environnements a haut potentiel energetique, soit pres des c6tes pour les reproducteurs printaniers o u dans Bes zones a maree active pour les geniteurs automnacmx. he frai est depose sur la his paper was g ~ s e n t e d at the International Symposium on the Biological Characteristics of Herring and Their Implication for Management, convened at Nanaimo, B .C., June 20-22. 1983, commemorating the 75th anniversary of the Pacific Biological Station. '~sder of authorship alphabetical.
Article
In a Baltic herring (Clupea harengus membras L.) spawning ground study, carried out in summer 1982 in the northern Baltic proper, spawning herring were encountered on two line transects on June 14. The spawning behaviour of minor groups as well as of a large school was studied in situ by four divers for approximately three hours. No behavioural differences or systematic coordinated behaviour between the sexes could be detected. Most of the spawning took place in a limited depth interval, 0.5 to 4 m, and Ceramium tenuicorne was the substrate most used.
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