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Pacific herring
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Pacific herring
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Clupeiformes
Family: Clupeidae
Genus: Clupea
Species:
C. pallasii
Binomial name
Clupea pallasii
Valenciennes in Cuvier and Valenciennes, 1847

The Pacific herring (Clupea pallasii) is a species of the herring family associated with the Pacific Ocean environment of North America and northeast Asia. It is a silvery fish with unspined fins and a deeply forked caudal fin. The distribution is widely along the California coast from Baja California north to Alaska and the Bering Sea; in Asia, the distribution is south to Japan, Korea, and China. Clupea pallasii is considered a keystone species because of its very high productivity and interactions with many predators and prey. Pacific herring spawn in variable seasons, but often in the early part of the year in intertidal and sub-tidal environments, commonly on eelgrass, seaweed[2] or other submerged vegetation. They do not die after spawning and can breed in successive years. According to government sources, the Pacific herring fishery collapsed in the year 1993 and is slowly recovering to commercial viability in several North American stock areas.[3] The species is named for Peter Simon Pallas, a noted German naturalist and explorer.

There are disjunct populations of Clupea pallasii in North-East Europe, which are often attributed to separate subspecies Clupea pallasii marisalbi (White Sea herring) and Clupea pallasii suworowi (Chosha herring).

Morphology

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Pacific herring have a bluish-green back and silver-white sides and bellies; they are otherwise unmarked. The silvery color derives from guanine crystals embedded in their laterals, leading to an effective camouflage phenomenon. There is a single dorsal fin located mid-body and a deeply forked tail-fin. Their bodies are compressed laterally, and ventral scales protrude in a somewhat serrated fashion. Unlike other genus members, they have no scales on heads or gills;[4] moreover, their scales are large and easy to extract. This species of fish may attain a length of 45 centimetres (18 in) in exceptional cases and weigh up to 550 grams (19 oz), but a typical adult size is closer to 33 centimetres (13 in). The fish interior is quite bony with oily flesh.

This species has no teeth on the jawline, but some are exhibited on the vomer. Pacific herring have an unusual retinal morphology that allows filter feeding in extremely dim lighting environments. This species is capable of rapid vertical motion, due to the existence of a complex nerve receptor system design that connects to the gas bladder.[5]

Life cycle

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Juvenile fish

Pacific herring prefer spawning locations in sheltered bays and estuaries.[6] Along the American Pacific Coast, some of the principal areas are San Francisco Bay, Richardson Bay, Tomales Bay and Humboldt Bay. Adult males and females make their way from the open ocean to bays and coves around November or December, although in the far north of the range, these dates may be somewhat later. Conditions that trigger spawning are not altogether clear, but after spending weeks congregating in the deeper channels, both males and females will begin to enter shallower inter-tidal or sub-tidal waters. Submerged vegetation, especially eelgrass, is a preferred substrate for oviposition. A single female may lay as many as 20,000 eggs in one spawn following ventral contact with submerged substrates. However, the juvenile survival rate is only about one resultant adult per ten thousand eggs, due to high predation by numerous other species.

The precise staging of spawning is not understood, although some researchers suggest the male initiates the process by release of milt, which has a pheromone that stimulates the female to begin oviposition. The behavior seems to be collective so that an entire school may spawn in the period of a few hours, producing an egg density of up to 6,000,000 eggs per square meter.[7] The fertilized spherical eggs, measuring 1.2 to 1.5 millimeters in diameter, incubate for approximately ten days in estuarine waters that are about 10 degrees Celsius. Eggs and juveniles are subject to heavy predation.[8]

Fisheries

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Global capture production of Pacific herring (Clupea pallasii) in thousand tonnes from 1950 to 2022, as reported by the FAO[9]

Pacific herring fisheries (fishing grounds) had been sustainably exploited by indigenous people for millennia, not only on the Pacific Coasts of North America, but in Japan and the Russian Far East. In all these cases, industrial fishing for herring oil and fertilizer has encroached or seized these fishing areas, leading to collapses in the fish stock.[10]

The Ainu of Ezo (now Hokkaido) caught herring using basic dip nets (hand nets),[a] but Japanese fishermen during the late Edo Period into Meiji Era began to operate increasingly large-scaled capture of herring in these grounds, first using gillnets and later "pound nets" (or traps).[11][12][14] Intensive fishing resulted in the so-called "Million-Ton Era" of the late nineteenth century onward.[16] Herring fishery near Hokkaido collapsed in the late 1950s.[11][17]

Museum diorama of pacific herring being caught with traditional nets in Hokkaido, Japan

Much like Japan, commercial herring fisheries in Alaska, US, and British Columbia underwent the phase of § Reduction fishery (for fertilizer and oil), and when Japanese herring fleets suffered scarcity in the late 1950s, North American fisheries began to cater to the Japanese market especially for the herring roe (§ Roe fishery; § Spawn on kelp fishery), known in Japan as § kazunoko. Alaska Department of Fish and Game has managed Alaskan resources and issues quota has released their biomass estimate figures since 1975, but the figures remain highly volatile.[18]

Pacific herring as sushi

Herring has long been fished by First Nations on the Central Coast of British Columbia, and elsewhere. In 1997 the Supreme Court of Canada rendered its decision in the Gladstone decision (R. v. Gladstone)- recognizing a pre-existing aboriginal right to herring that includes a commercial component to the Heiltsuk Nation.

Due to overfishing,[19] the total North American Pacific herring fishery collapsed in 1993 and is slowly recovering with active management by North American resource managers. In various sub-areas, the Pacific herring fishery collapsed at slightly differing times; for example, the Pacific herring fishery in Richardson Bay collapsed in 1983.[20] The species has been re-appearing in harvestable numbers in a number of North American fisheries including San Francisco Bay, Richardson Bay, Tomales Bay, Half Moon Bay, Humboldt Bay all in California, and Sitka Sound, Alaska. In other areas, such as Auke Bay, Alaska, which in the late 1970s was the largest harvestable stock of herring in Alaska, the species remains severely depleted.[21]

Pacific herring are currently harvested commercially for bait and for roe. Past commercial uses included fish oil and fish meal.[21]

Reduction fishery

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Processing pacific herring in Alaska

The Alaskan herring industry began in 1880s as "reduction" plants which processed herrings into fish meal and oil, with the meal utilized mostly as animal fodder or fertilizer,[22] and the oil mostly for soap.[23] Since it began the reduction in 1882 until around 1917, the business was a practical monopoly of the North West Trading Company which established its processing plant at Killisnoo, Alaska.[24] The use of "Norwegian method" of catching using oar-propelled seine boats did continue until 1923 here, but was being supplanted by the purse seine (purse seiner [de]) introduced into herring fishery after around 1900. .[25]

Concerns had developed regarding this practice as early as the 1900s, regarding localized fish stock depletion, adverse food chain effects on commercially valuable fish types that prey on herring, and the ethics of taking fish for purposes other than human food or bait.[26] However, the industry persisted in Alaska until it ceased operations in 1966.[27]

In Canada, the earliest recorded catches were for the purpose of producing dry-salted herring, starting around 1904, peaking around the 1920s,[b] but declining to initial catch tonnages by 1934 due to sagging demand.[28] Reduction (fertilizer) fishing operated in Canada during the years 1935–1967. The end was due to the collapse of the fish population.[28]

Roe fishery

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Just as the reduction industry was phasing out in Alaska in the 1960s, there emerged an alternate industry to exploit herring in another way, i.e., harvesting only the "roe sacs" ("egg skein") inside the females, to meet the Japanese demand for "kazunoko".[c][27] A similar shift took from the defunct reduction fishing took place in Canada: after the herring population recovered somewhat, a Canadian roe fishery industry sprang up in 1971 to cater to the Japanese market.[32][d]).

A commercially viable product demands the eggs to be "ripe", or swollen to the right size, which only occurs within a few days of spawning, and there is a narrow window for the catch.[32][36] Accordingly, the season is very short, a matter of days: it lasted all of 90 minutes in the April 1975 season.[36][37]

These egg skeins need to retain perfection of shape to fetch highest value, and to that end, the fish are frozen or brine-frozen then rethawed in freshwater before extracting the egg skeins.[37]

Spawn on kelp fishery

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Shoals of herring during the reproductive season lay clusters of eggs on kelp and other seaweed,[e][f] and the seasonal collection has been a time-honored traditional practice among the natives of Pacific Coast of Alaska and Canada,[41][42] witnessed and recorded in the 18th and 19th centuries,[46] and has been traded [41] and a trade item.[44] The natives traditionally foraged wild-grown eggs on various seaweed, or laid on introduced hemlock branches,.[45][47]

The Japanese market for kazunoko kombu (数の子コンブ; 'herring roe kelp') or (子持ちコンブ; 'child holding kelp') is best served, so it has been claimed, by preferably using products laid on giant kelp (Macrocystis pyrifera), which only grew in Southeast Alaska [48][g] or down in Canada.[h][i]

[j] commercial harvest of wild-caught roe began in that region at Craig/ Craig/ Klawock ,[k] in 1959[l][54] Export to Japan began 1962.[m] So that in wild foraging surged at Craig/Klawock 1963, burgeoned in Sitka in 1964, and at a third site at Hydaburg in 1966 were harvesting in southeast Alaska:[55] overfilling their 250 tons quota in 1966.[57] The season had to be drastically shortened or canceled due to depletion from the following year.[58]

Transplanting and impounding kelp

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In 1960 and 1961 "open-pounds", stocked with kelp to lure herring egg-laying, were operated in the town of Craig, on Prince of Wales Island, probably for the first time in Alaska.[55] But afterwards, intensified harvest led to closure of season, and it was not until 1992 that harvest of semi-farmed eggs on kelp in closed-pounds resumed.[59]

The shortage of spawn led to seeking new harvesting grounds in areas where giant kelp do not naturally grow, and demand and harvest developed for eggs on alternate seaweeds, such as Desmarestia sp. or "hair seaweed".[e][59] Amidst the 1968 shortage, commercial collection of spawn of Fucus began,[f] in Bristol Bay, east of Togiak.[60] And in 1959 spawn from various algae began to be commercially collect from Prince William Sound, peaking in 1975, ending with the depletion of the "kelp".[60][i]

During the shortage, an enterprising operator experimented with transplanting "unused" kelp from remoter areas into kelp-depleted spawning grounds, or into eelgrass territory. He sometimes attached kelp cut elsewhere to barges he owned.[56]

In Canada, "impoundments" began to be used, whereby floating enclosures at sea are stocked with kelp, mature herring are introduced, and the egg-deposited kelp to be later harvest. Canada issued their first licenses in 1975, initially about half to indigenous operators, in Northern British Columbia.[32] The enclosure ("closed pounds") technique was subsequently copied by Alaskans.[61] The "impoundments" or "closed ponds" consisted of a square (wooden) frame holding a pocket of "suspended webbing" as enclosure space. Inside, rows of kelp are hung on strings. [32][61][63]

Decline

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Alaska's principal areas for roe fishery, according to the 2022 season allotted tonnage were: Sitka Sound (late March) 45,164 short tons (90,000,000 lb), Kodiak Island (April 1) 8,075 short tons (16,000,000 lb), and Togiak[n] (May) 65,107 short tons (130,000,000 lb). However the allowed quotas were hardly expected to be filled, given the drastic downturn in Japanese demand. During the heydays of the 1990s, the pre-spawn herring commanded $1000 per ton, yielding a gross $60 million to fisherman, but by 2020 the tally fell to a $5 million figure.[65] In 2023, the last roe processing plant in Togiak indicated it would not be purchasing herring, and the season was cancelled.[66]

Conservation

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On April 2, 2007, the Juneau group of the Sierra Club submitted a petition to list Pacific herring in the Lynn Canal, Alaska, area as a threatened or endangered distinct population segment under the criteria of the U.S. Endangered Species Act (ESA).[67] On April 11, 2008, that petition was denied because the Lynn Canal population was not found to qualify as a distinct population segment. However, the National Marine Fisheries Service did announce would be initiating a status review for a wider Southeast Alaska distinct population segment of Pacific herring that includes the Lynn Canal population.[68] The Southeast Alaska DPS of Pacific herring extends from Dixon Entrance northward to Cape Fairweather and Icy Point and includes all Pacific herring stocks in Southeast Alaska.

On February 5, 2018, researchers at Western Washington University began researching causes for the decline in Pacific herring populations in the Puget Sound; a prominent speculated reason is the loss of eelgrass, an important spawning substrate for the herring.[69]

Kazunoko

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The herring egg roe or "egg skein", called kazunoko had traditionally commanded a good price in Japanese markets, and the herring roe fishery and processing industry (especially in Alaska), geared towards export to that country, has been described above under § Roe fishery.

As for the culinary aspects, the kazunoko merchandized in Japan primarily fall into either hoshi kazunoko (塩数の子; 'dried kazunko') or shio kazunoko (干し数の子; 'dried kazunko').[70] There is also a lower-grade substitute[73] called shio kazunoko (味付け数の子; 'dried kazunko'),[34] made from Atlantic herring roe (which is considered a softer or "less crunchy" in texture).[34][35][71] [o]

The roe is eaten mostly as the New Year's fare,[74] called osechi, consisting of an assortment of symbolically propitious foods, with herring representing fertility (production of many children).[75][76]

Notes

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References

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Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
The Pacific herring (Clupea pallasii) is a small, silvery clupeid characterized by unspined and a deeply forked caudal fin, inhabiting the coastal and neritic zones of the temperate North . Its range extends from northward along the eastern Pacific to and the , and across to the western Pacific including the , , and Korea. Adults typically measure 25-38 cm in length, form dense schools, and undertake seasonal migrations to shallow coastal spawning grounds where females broadcast adhesive eggs onto substrates such as , eelgrass, and rocks in intertidal or subtidal areas. As a foundational species, Pacific herring plays a pivotal role in marine food webs, serving as primary prey for piscivorous fishes like and groundfish, marine mammals, seabirds, and even transient predators such as orcas. Its high content enhances its nutritional value, supporting the growth and reproduction of dependent predators and contributing to . The supports commercially and culturally significant fisheries, particularly for roe-on-kelp in and , where indigenous communities have harvested eggs for millennia as a subsistence staple. Population dynamics vary regionally, with some stocks exhibiting cyclic fluctuations influenced by environmental conditions, predation, and fishing; while overall North Pacific herring abundance remains managed through quotas and monitoring, certain localized groups, such as those in or Cherry Point, have faced decline and prompted petitions for distinct population segment status under the U.S. Endangered Species Act, though none have been listed to date.

Taxonomy and Distribution

Scientific Classification

The Pacific herring is classified in the genus Clupea within the family Clupeidae, order Clupeiformes, class Actinopterygii, phylum Chordata, and kingdom Animalia. Its binomial name is Clupea pallasii, originally described by Valenciennes in 1847 based on specimens from the North Pacific. Clupea pallasii is recognized as a distinct species from the Atlantic herring (Clupea harengus), with which it was formerly considered conspecific; separation is supported by biochemical genetic analyses demonstrating consistent differences in allozyme loci and other markers. Phylogenetic studies using mitochondrial DNA indicate that the two species diverged from a common ancestor approximately 2 million years ago, reflecting vicariant speciation across the Arctic following Pleistocene glacial cycles and trans-Arctic migrations. Nuclear and mitochondrial genomic data further reveal adaptations in C. pallasii to North Pacific salinity, temperature, and prey dynamics, including loci associated with osmoregulation and lipid metabolism absent or divergent in C. harengus. Taxonomic debates persist regarding subspecies status for certain populations, such as the herring (Clupea pallasii marisalbi) in the northeastern Atlantic, which exhibits morphological and genetic isolation but limited with Pacific stocks. In the North Pacific, regional variants like those in isolated fjords (e.g., stocks) show minor allozyme and divergences suggestive of local adaptation, though insufficient for formal subspecific elevation under current criteria emphasizing . Ongoing genomic sequencing reinforces C. pallasii as a cohesive with clinal variation rather than discrete , prioritizing empirical genetic clustering over historical morphological designations.

Geographic Range

The Pacific herring (Clupea pallasii) occupies coastal and nearshore waters across the North Pacific Ocean. In the eastern Pacific, its range spans from northern Baja California, Mexico, northward through the coastal United States and Canada to Alaska, encompassing the Bering Sea and extending into the Beaufort Sea. In the western Pacific, populations inhabit regions from Anadyr Bay and the southward along the eastern coasts of Kamchatka and the to and the western coast of Korea. The species maintains a primarily coastal distribution, with schools seasonally migrating into enclosed bays, sounds, and estuaries, including in Washington and in . Populations exhibit genetic differentiation, reflecting localized adaptations and limited . For instance, the stock is recognized as a distinct population segment, genetically distinguished from adjacent populations, and underwent a comprehensive status review under the U.S. Endangered Species Act in 2014. Such delineations highlight core areas of abundance in the and peripheral extents toward southern latitudes, where occurrences are less frequent.

Physical Characteristics

Morphology and Anatomy

The Pacific herring (Clupea pallasii) has a , laterally compressed body covered in large, thin, scales that are easily detached. Ventral scales protrude in a serrated , forming a keel-like structure. The dorsal surface is blue-green to olive, transitioning to silvery sides and belly, which enhances in open water. Adult specimens typically measure 25-38 cm in length, with maximum sizes varying by population up to 46 cm and weights reaching approximately 0.5 kg. Fins are unspined, with a single positioned mid-body and a deeply forked caudal fin aiding propulsion. The mouth is terminal, featuring small teeth and a moderately protrusible upper suited for filter-feeding. The lateral line system, though not prominently visible as a scaled row, includes sensory neuromasts for detecting hydrodynamic cues. The eyes are relatively large relative to body , supporting in dimly lit coastal habitats. Sexual dimorphism is minimal in external morphology, though females often exceed males in , especially during spawning when abdominal distension from maturing ovaries occurs.

Size, Growth, and Variation

Pacific (Clupea pallasii) exhibit rapid growth in their early , with juveniles reaching lengths of approximately 11-15 cm by the end of the first year in various populations, though precise measurements vary by location and environmental conditions. Growth rates decelerate after the initial phase, particularly following the typical attainment of maturity at 2-3 years, leading to slower annual increments thereafter. Maximum lifespan ranges from 9 to 20 years, with longevity increasing latitudinally; southern populations rarely exceed 9-14 years, while northern stocks in the and can reach 18-20 years. Intraspecific variation is pronounced regionally, with eastern Pacific herring generally smaller and slower-growing than western Pacific counterparts, attaining lower asymptotic weights (e.g., 143 g in versus 278 g in ). Adult body lengths increase with latitude, averaging 10-24 cm in compared to maxima of 43-46 cm in northern areas like . These differences reflect a combination of and local ecological factors, including prey availability influencing condition rather than strict .

Habitat and Ecology

Environmental Preferences

Pacific herring inhabit temperate coastal waters of the North , predominantly in nearshore environments such as bays, estuaries, and sheltered inlets, while avoiding deep oceanic pelagic zones beyond the continental shelf. These are eurythermal but exhibit preferences for cooler temperatures, with optimal ranges for growth and distribution typically between 0.2°C and 9.7°C, averaging around 2.8°C based on occurrence data across their range. Embryonic development proceeds optimally at approximately 7°C, with hatching times inversely related to temperature in spawning habitats. Salinity tolerance varies by life stage; adults thrive in fully marine conditions of 30-35 ppt, but juveniles readily enter brackish estuarine waters, associating with lower salinities in inner bays during early rearing. Larvae demonstrate tolerance to reduced salinities as low as 0-6 ppt during initial development, though prolonged exposure below optimal marine levels can impair survival. Pacific herring require well-oxygenated waters, showing avoidance and reduced abundance when dissolved oxygen falls below 2 mg/L, a threshold associated with hypoxia in coastal estuaries. Spawning favors structured substrates in intertidal and shallow subtidal zones, including macroalgae like , seagrasses such as eelgrass, and occasionally rock or other hard surfaces, which provide adhesion sites for adhesive eggs. These vegetated or structured habitats in sheltered coastal areas support egg deposition and initial larval survival by offering protection from wave action and predation.

Diet, Feeding, and Behavior

Pacific herring ( pallasii) exhibit a planktivorous diet primarily composed of , including copepods, , and amphipods, as revealed by stomach content analyses of juveniles and adults across various populations. Juveniles additionally consume microplankton such as diatoms, protozoans, bivalve veligers, and smaller copepods, transitioning to larger crustaceans and occasional small as they mature. Seasonal variations in prey selection reflect availability rather than rigid preferences, with higher consumption of lipid-rich northern copepods during periods of peak abundance to support fat storage for overwintering. Feeding occurs mainly through visual predation and particulate selection in low densities, supplemented by filter-feeding via elongated rakers that strain from water currents, particularly effective during schooling in phytoplankton-rich zones. Diel patterns show intensified feeding at night, with ascending from near-bottom positions during daylight to surface or shallow waters, aligning with vertical distributions and reduced predation risk under low light. Schooling behavior enhances foraging efficiency by concentrating prey encounters and enables predator evasion through the confusion , with schools tightening during threats and dispersing slightly at dusk for individual feeding bouts. These formations, often numbering thousands, facilitate resource partitioning among co-occurring , as stomach content studies indicate minimal dietary overlap with species like despite shared habitats.

Role in Food Web and Predators

Pacific herring (Clupea pallasii) serves as a foundational in the Northeast Pacific , facilitating transfer from planktonic to higher trophic levels through its high abundance and schooling behavior. As a mid-trophic species, it supports piscivorous fishes, marine mammals, and seabirds by providing a concentrated, predictable prey resource that enhances predator efficiency and energy flow. flow models indicate that herring consumption by predators constitutes a significant proportion of total herring in regions like the and , underscoring its role in sustaining predator populations without implying disproportionate dependence. Key predators of Pacific herring encompass a range of taxa, including Pacific salmon (Oncorhynchus spp.), Pacific cod (Gadus macrocephalus), and other groundfishes, which prey on herring juveniles and adults to meet substantial dietary needs. Marine mammals such as harbor seals (Phoca vitulina) and California sea lions (Zalophus californianus) also consume herring, particularly during spawning aggregations, contributing to observed predation pressures. Seabirds, including gulls (Larus spp.) and cormorants (Phalacrocorax spp.), target herring schools near the surface, with consumption rates reflecting herring availability as a buffer against competition for shared prey. Humpback whales (Megaptera novaeangliae) represent a dominant predator in recent decades, with escalating consumption linked to their population recovery following historical depletion. A 2024 analysis of natural mortality trends in Pacific stocks demonstrated that predation, predominantly by humpback whales, accounts for the observed increase in herring natural mortality rates since the early 2010s, as quantified through diet reconstruction and abundance estimates in areas like . Empirical consumption estimates from these models prioritize bottom-up herring availability as a driver of whale shifts, rather than invoking unsubstantiated top-down control mechanisms. Regional data further show humpback whales removing up to a notable fraction of available herring biomass in overwintering sites like , , though variability underscores context-specific predation intensity.

Life History

Reproduction and Spawning

Pacific herring (Clupea pallasii) exhibit an iteroparous reproductive strategy, spawning annually after reaching at 3–4 years of age. Females typically produce a single batch of eggs per spawning event, with ranging from approximately 20,000 eggs on average, though this varies by body size, age, condition, and geographic —southern populations like those in often show higher egg production per female length compared to northern stocks in . Spawning occurs in synchronized group events during winter to spring, with timing varying by : October to April in waters and primarily to May in Alaskan regions. These events involve dense aggregations of adults in shallow, nearshore areas, where females broadcast demersal, adhesive eggs onto submerged vegetation such as , eelgrass, or rocky substrates, while males simultaneously release for . The resulting high-density egg layers—often multiple batches over days—enhance fertilization success through sheer volume, though exact densities depend on local and spawner biomass, as observed in 20th-century surveys of sites like . Stocks demonstrate strong site fidelity, with adults returning to traditional natal spawning grounds annually, a behavior documented across North Pacific populations and contributing to localized reproductive cycles. This supports stock-specific management, as disruptions to preferred sites can affect deposition strategies, though empirical data from fisheries monitoring emphasize the role of synchronous timing in maximizing overlap for viable fertilization rates.

Development and Life Stages

Pacific herring (Clupea pallasii) eggs are demersal and adhesive, typically deposited in large clusters on subtidal or substrates, where they undergo embryogenesis influenced primarily by water temperature. Incubation duration ranges from 10 to 20 days, with occurring in 10-14 days at 11.8-13.5°C and extending to 12-15 days at 9-10°C or up to three weeks at cooler temperatures. Embryos develop external features such as eyes and sacs, with yielding larvae approximately 5-7 mm in length that rely initially on residual reserves. Newly hatched larvae enter a pelagic phase, dispersing offshore while feeding exogenously on zooplankton after 5-6 days of yolk utilization. This larval stage lasts 2-3 months (or 30-60 days pre-metamorphosis), during which growth is rapid under favorable conditions, reaching sizes conducive to transformation. Metamorphosis to the juvenile form occurs at 25-38 mm total length, marked by fin development, scale formation, and shift to adult-like morphology and pigmentation. Post-metamorphosis, age-0 juveniles school in protected nearshore nurseries such as bays, inlets, and fjords, seeking refuge from open-water predators while continuing growth. This settlement phase, spanning 5-9 months in estuarine habitats, supports higher survival through reduced exposure compared to pelagic larvae. Early life stages exhibit pronounced survival bottlenecks, with cohort analyses revealing that larval and juvenile mortality—driven by predation, starvation, and environmental stressors—exerts the dominant influence on year-class strength, far exceeding embryo-stage losses. Field and mesocosm studies quantify these vulnerabilities, showing interannual variability tied to hatching timing, prey availability, and advection, where unfavorable conditions can reduce cohort survival by orders of magnitude.

Natural Mortality and Longevity

Natural mortality in Pacific herring (Clupea pallasii) is characterized by elevated rates during early life stages, primarily driven by predation and environmental variability, with lower rates among adults until advanced age. Juvenile mortality exceeds 90% annually, reflecting density-independent factors such as predation by piscivorous , seabirds, and marine mammals, alongside stochastic environmental conditions like fluctuations and food scarcity. Acoustic-trawl surveys in , , estimated instantaneous natural mortality rates for young-of-the-year herring at 0.009 (SD = 0.002) for the 1995 cohort and 0.016 (SD = 0.012) for the 1996 cohort, while rates for 1-year-olds averaged 0.003 (SD = 0.007) and 0.008 (SD = 0.005), respectively, indicating persistent high vulnerability post-larval settlement. These estimates, derived from cohort-specific abundance tracking, underscore predation as the dominant cause, with limited evidence of strong density-dependent regulation in juveniles due to schooling diluting per-capita at high densities. Adult natural mortality remains low, typically 0.1–0.2 annually, increasing with age due to and cumulative predation exposure, as modeled from age-structured catch data spanning 1951–1998 in southern . Tagging studies and population models distinguish these rates from density-dependent effects, revealing minimal compensation at low abundances, where predation pressure scales with biomass but is modulated by predator switching. Recent analyses attribute upward trends in adult mortality—rising from historical baselines to 0.3+ in some stocks since the 2010s—to surging populations of marine mammals, particularly humpback whales (Megaptera novaeangliae), whose consumption accounts for 20–50% of losses in predator hotspots like the . This predator-driven dynamic, validated through bioenergetic models integrating whale sighting data and diet composition, challenges assumptions of stable low adult M and highlights trophic feedbacks absent in earlier fisheries-independent estimates. Pacific herring longevity varies latitudinally, with southern populations rarely exceeding 9 years due to compounded annual mortality and faster in warmer waters, while northern stocks in attain 12–16 years under cooler conditions favoring slower metabolism and reduced predation. Age validation from readings confirms maximum observed ages of 19 years in exceptional cases, though median lifespan hovers at 6–8 years, reflecting selective pressures that prioritize early over extended . Modeling exercises incorporating size-dependent M further parse these patterns, showing longevity inversely tied to growth rates, with slower-maturing northern cohorts evidencing lower cumulative mortality.

Population Dynamics

Historical Fluctuations

Archaeological evidence from coastal sites in the Northeast Pacific indicates that Pacific herring (Clupea pallasii) populations were historically superabundant, supporting dense indigenous settlements and forming a key component of human diets for millennia prior to intensive commercial exploitation. Indigenous oral histories and place names along the and coasts further attest to periods of exceptional abundance, with accounts describing as so plentiful that they could be harvested en masse from the surface, reflecting long-term reliance on these fish as a cultural and ecological staple. These records suggest inherent boom-bust cycles driven by environmental variability, rather than solely human impacts, consistent with the species' sensitivity to oceanic conditions. In the early , following the collapse of Pacific stocks in the late 1940s, herring fisheries expanded rapidly along Canada's , with annual catches peaking at approximately 100,000 tonnes in the early . This boom was followed by widespread stock collapses attributed primarily to , leading to fishery closures in by 1967, where reductions in catch mirrored sharp declines in spawning biomass across multiple regions. Similar patterns emerged in , where early commercial harvesting intensified in the amid global demand, contributing to localized busts amid strong year-class variability. Natural oceanographic regimes, such as the (PDO), have modulated these fluctuations independently of fishing pressure, with herring spawn abundance exhibiting a bowl-shaped response to PDO phases—higher during neutral periods and lower at extremes—over multi-decadal scales. In the , proxy records from sediment cores reveal sustained high abundance from AD 1417 to 1870 during cooler climatic conditions akin to the , followed by declines linked to warming and circulation shifts, underscoring climate's role in long-term cycles. Post-collapse rebounds in areas like parts of demonstrated partial recovery in biomass during favorable environmental windows in the late , highlighting resilience when anthropogenic pressures eased alongside supportive marine conditions. In the , the primary Canadian stock of Pacific herring, the forecasted spawning for 2025 stands at 65,894 short tons (range: 36,824–118,078 short tons), with a 0% probability of exceeding the limit reference point. For the West Coast stock, spawning has remained low for nearly 20 years, with a 2025 forecast of 8,329 short tons (range: 3,677–20,355 short tons). In , the total spawning for the southern stock in 2024 was 11,404 metric tons, reflecting a 36% decline from 2023 levels. The four-year average spawning for other stock complexes decreased to approximately 12,700 metric tons in 2024. Pacific herring stocks in have exhibited high variability in spawning since 1992, with populations supporting commercial , bait, and fresh fish fisheries as well as recreational into 2025, though specific biomass estimates for the year remain tied to ongoing annual spawn surveys. In , particularly , herring stocks are assessed annually via spawn surveys and remain above management thresholds, enabling sac fisheries without indications of . Across core North American populations, monitoring incorporates aerial and ground-based spawn surveys alongside acoustic-trawl methodologies, with no listings under the U.S. Endangered Act as of 2025. A 2014 NOAA Fisheries status review affirmed that Southeast Alaska's distinct population segment does not warrant endangered status, consistent with ongoing dynamic but non-critical trends in regional assessments.

Drivers of Abundance Changes

has been the predominant driver of major historical declines in Pacific herring abundance, particularly evident in the coastwide stock collapse along the coast in the early 1960s, which prompted the closure of the commercial reduction fishery in 1967. This event followed decades of intensive harvesting that exceeded sustainable levels, reducing spawning across multiple and illustrating how age-selective exploitation can truncate population age structures, impairing recovery. In contrast, recent abundance changes in certain populations reflect heightened natural mortality from predation, rather than harvest pressure alone. predation, which has intensified with population recoveries, accounts for much of the observed increase in natural mortality rates, as documented in the Ecosystem and , where whales consumed 3-13% of available in 2021. Variations in prey quality for , linked to shifts in euphausiid and ocean productivity, have also contributed to reduced growth and success in regions like the west coast of , underscoring predation and bottom-up trophic effects as complementary causes beyond . Climate-driven environmental variability further modulates abundance through impacts on spawning and early life survival. Sea surface temperature (SST) shifts tied to North Pacific atmospheric patterns have advanced spawning timing in some stocks by weeks over multi-decadal scales, with models forecasting an average 9-day earlier onset by 2100 under warming scenarios, potentially misaligning larval development with peak availability and reducing recruitment. Oscillations in indices like the (PDO) and North Pacific Gyre Oscillation (NPGO) exhibit nonlinear, "bowl-shaped" relationships with spawn abundance across coastal regions, amplifying natural fluctuations independent of . As a short-lived clupeid, Pacific herring exhibits inherently high variability as the primary engine of , where oceanographic conditions during early life stages dominate over consistent anthropogenic signals. This stochasticity, evidenced in age-3 models incorporating environmental covariates, challenges monocausal explanations of declines, as predation surges and climatic regime shifts can override harvest controls in driving observed trends. Empirical stock assessments integrating these factors reveal that while precipitated past crashes, contemporary abundance variance integrates biological interactions and physical forcing, necessitating multifaceted causal analysis.

Exploitation and Fisheries

Historical Harvesting

along the , including Coastal First Nations, Native Americans, and , harvested Pacific herring (Clupea pallasii) and its for subsistence over thousands of years prior to European contact. Archaeological evidence from sites in indicates herring use dating back more than 10,000 years, with bones frequently found in middens reflecting consistent exploitation for food. In the region, herring remains represent the most abundant fish in 55% of analyzed sites, comprising a dietary staple for at least 12,500 years. Traditional methods emphasized collection on and sustainable practices to avoid depletion, as documented in oral histories and archaeological records. Commercial harvesting of Pacific herring commenced in the late , initially for bait and local food markets in regions such as , , and . In , the fishery began supplying dry-salted products by the early 1900s, with annual catches reaching approximately 30,000 tons by 1904 for export to . Southeast Alaska's reduction fishery, processing herring into oil and meal, started around the same period, while California's operations in focused on similar early uses. By the 1870s, bait fisheries had expanded from Washington to Alaska, transitioning to larger-scale reduction processes. Mid-20th-century exploitation peaked during reduction fisheries for oil and meal, driven by industrial demand, with landings exceeding 200,000 metric tons annually in the early and reaching 237,600 metric tons in the 1962-63 . These highs, part of broader North Pacific trends including catches of 146,000 metric tons in 1970, led to stock collapses, such as 's fisheries closing in 1967 after sustained high harvests. Post-1967 moratoriums shifted focus to sac- harvesting from the early onward, reopening fisheries with quotas to rebuild populations, as roe products gained market value in . Regulations intensified following these peaks to curb evident in declining yields.

Commercial Methods and Yields

Purse seining constitutes the primary commercial method for harvesting Pacific herring, involving the deployment of large encircling nets to capture dense schools during spawning aggregations near coastal areas, which facilitates high-volume extraction when are predictably aggregated. Gillnets are employed as a secondary technique, using vertical panels to entangle by gills, often targeting similar aggregations for efficiency in shallower waters. These methods exploit herring's schooling behavior but face challenges such as depredation, which can disrupt net sets. Advancements in acoustic technologies, including and fish locators, have optimized yields by enabling precise detection and estimation of schools prior to net deployment, allowing operators to select optimal sets and reduce search time. However, gear selectivity remains constrained, as purse seines and gillnets capture a broad range of sizes with limited differentiation by age or maturity, potentially impacting population structure despite technological aids. Commercial yields fluctuate annually based on stock assessments and environmental factors, with total allowable catches varying by region; for instance, Canada's 2024-2025 Integrated Fisheries Management Plan sets a food and bait quota of 12,787 tonnes in the Strait of Georgia at a 14% harvest rate of forecasted spawning biomass. In Puget Sound, bait-directed fisheries produced 74 short tons in 2024, reflecting conservative exploitation primarily on juvenile fish.

Specific Fishery Types

Pacific herring fisheries are categorized by end-use, including reduction for meal and oil, roe production via sac-roe and spawn-on-kelp methods, bait for recreational fishing, and special uses such as kelp impounding. Historically, reduction fisheries dominated, processing herring into meal and oil; a reduction plant was established in British Columbia in 1937, targeting bulk volumes for industrial applications. These operations reduced large catches to byproducts, but their scale diminished as roe markets grew, with recent emphasis shifting toward targeted quotas to align with stock assessments. Roe fisheries, prominent since the late , focus on high-value eggs for export, particularly to . Sac-roe harvesting involves pre-spawn capture of gravid females, while spawn-on-kelp entails suspending fronds or hemlock branches in spawning grounds to collect eggs naturally. In , these fisheries allocate significant portions of total allowable catch, though volumes have declined post-1990s due to reduced Japanese demand and price drops, leading to fewer active permits. Quotas have been adjusted downward, such as a total allowable catch reduction to 10% of in some areas by 2022, reflecting responses to pressures. Food and bait fisheries target whole , with bait uses prominent in regions like , where juvenile are harvested for sport fishing bait, averaging around 270 short tons annually in the south Sound. In 2024, bait catches totaled 74 short tons, involving a small number of vessels focused on central and south-central areas. These operations remain limited compared to roe harvests, serving regional recreational markets. Special use fisheries include techniques like kelp impounding, where enclosures stock vegetation to concentrate spawning, and kelp or branches laden with eggs to enhance local production or support indigenous practices. These methods, sometimes integrated with spawn-on-kelp, allow for controlled harvests in specific locales, such as Alaska's coastal communities, prioritizing smaller-scale, site-specific volumes over mass reduction.

Management and Controversies

Regulatory Approaches

In Canada, (DFO) manages Pacific herring fisheries through Integrated Fisheries Management Plans (IFMPs), which outline strategies, quotas, and monitoring for specific stocks such as , Prince Rupert District, Central Coast, West Coast Vancouver Island, and . The 2024-2025 IFMP, effective from November 7, 2024, to November 6, 2025, incorporates annual stock assessments using statistical catch-age models to forecast mature biomass and determine options, ensuring quotas align with biomass estimates above limit reference points. Management Strategy Evaluations (MSEs), initiated in 2018, test control rules via computer simulations to evaluate stock sustainability under varying scenarios. In the United States, the Alaska Department of Fish and Game (ADFG) oversees Pacific herring fisheries in state waters on a sustained yield basis, setting total allowable catches (TACs) annually for distinct management areas including , , and . TACs are derived from hydroacoustic surveys, spawn timing observations, and age-structured models projecting and , with allocations divided among sac roe, bait, and food fisheries. Federal oversight by NOAA Fisheries applies in the , emphasizing bycatch limits in groundfish trawl fisheries through measures like Herring Savings Areas in the . Stock-specific limits predominate in both nations to account for regional spawning and migration patterns, with DFO applying coast-wide harvest rates adjusted per stock (e.g., lower rates for rebuilding stocks like under a 2024-approved plan) and ADFG tailoring TACs to local abundance indices. Scientific integration includes biomass forecasting and uncertainty quantification in decision rules, as in Bayesian assessments for stocks. For transboundary areas like the , DFO treats northern stocks as a single migratory unit for quota-setting, while U.S. portions fall under separate state-federal assessments without formal bilateral herring agreements, unlike treaties for or .

Conservation Efforts

In response to the collapse of the Pacific herring reduction fishery in 1967, driven by decades of intensive harvesting, Fisheries and Oceans Canada (DFO) implemented a coast-wide moratorium on commercial fishing from 1968 to 1971. This closure allowed spawning biomass to recover, with post-moratorium assessments indicating partial stock rebuilding that supported the transition to a roe-on-kelp fishery starting in 1972. Biomass levels in areas like the Strait of Georgia subsequently stabilized at levels permitting limited harvests, though long-term trends varied by region. Habitat protection efforts target spawning substrates such as eelgrass beds and giant kelp, which provide attachment sites for eggs comprising up to 20,000 per female. In , the California Department of Fish and Wildlife (CDFW) integrates spawn ground surveys into its management framework, emphasizing preservation of nearshore bays and estuaries where herring deposit eggs in dense mats. Ongoing initiatives, including exploratory habitat mapping in the , have documented spawn deposition to inform site-specific protections, with intensities used to estimate local biomass. Monitoring programs form a core conservation tool, with CDFW's public reporting system—active as of January 2025—collecting observations of spawning events via submissions including location, date, and photos to track distribution and abundance. Similar DFO-led spawn surveys in , such as those in Barkley Sound during the 2025 season, provide real-time data on deposition and larval , enabling adaptive quota adjustments. These efforts have contributed to data-driven status reviews by NOAA Fisheries, which in 2023 determined that the Georgia Basin distinct population segment does not warrant Act listing, citing sufficient resilience and regulatory responsiveness. Stock enhancement trials, though limited, include managed roe-on-kelp practices in and , where kelp lines are deployed to concentrate spawning and harvest eggs while releasing adults, potentially bolstering local without genetic dilution of wild stocks. Evaluations indicate these methods maintain productivity when fishing pressure on enhanced areas avoids impacts on wild diversity, with 's 2023 roe-on-kelp yields reflecting sustained participation. Overall, such interventions have averted federal endangered status for multiple segments through empirical tracking, though regional variability persists.

Debates on Overfishing and Sustainability

Critics of Pacific fisheries, particularly in 's sac-roe sector, contend that mismanagement has precipitated local stock crashes, citing correlations between elevated harvest rates and subsequent declines. For instance, in the , advocacy groups have highlighted a proposed quota increase to 11,000 tonnes in despite spawning projections of around 20,000 tonnes, warning that exceeding historical harvest rates of 20% of risks irreversible affecting and orcas. Historical precedents, such as concerns over behavioral changes and population reductions from intensive , underscore arguments that unchecked exploitation disrupts spawning aggregations, with meta-analyses of showing overfished Pacific populations more likely to close fisheries at low thresholds compared to Atlantic counterparts. These views often draw on catch- models indicating that pre-1970s reductions in followed peak harvests exceeding 100,000 tonnes annually, attributing causation to regulatory delays rather than environmental factors alone. Counterarguments emphasize empirical evidence of natural variability and predation as dominant drivers, challenging claims of harvest-induced inevitability. A 2024 analysis found that predation, primarily by recovering populations consuming up to 70% of in some areas, explained rising natural mortality rates from 2010 onward, with abundance correlating more strongly to trends than pressure. Similarly, a evaluating historical cetacean impacts quantified predation rates sufficient to account for observed declines without invoking , noting 's short lifespan (rarely exceeding 9 years) and boom-bust cycles tied to oceanographic conditions like . Proponents of continued harvest argue sustainable yields remain viable for this dynamic, high-fecundity species under adaptive rules limiting exploitation to 20% of mature , as demonstrated by fisheries balancing industrial and subsistence needs amid fluctuating recruitment. These perspectives critique conservation models for underweighting economic data, such as 's role in supporting 10,000 jobs in , and overemphasizing static targets that ignore predator-prey feedbacks. Debates also reveal tensions between indigenous subsistence rights and industrial operations, with First Nations groups like the WSÁNEĆ asserting that roe-on-kelp fisheries undermine traditional priorities by prioritizing export markets over . In , indigenous representatives have disputed sac-roe impacts, arguing they subvert communal access amid neoliberal allocations favoring corporate seiners, though empirical stock assessments show no broad collapse when harvests align with age-structured models. Such conflicts highlight potential biases in regulatory frameworks, where advocacy-driven calls for moratoriums may overlook data on resilience, as evidenced by post-closure rebounds in closed stocks without corresponding ecosystem recovery. Overall, while local mismanagement critiques warrant scrutiny, causal realism favors multifactor explanations integrating predation and cycles over singular harvest blame, supporting targeted reforms over blanket restrictions.

Human Significance

Economic Contributions

The Pacific herring (Clupea pallasii) supports commercial fisheries valued in the tens of millions of dollars annually, primarily through sac harvests in and , with additional contributions from and food markets. In , the 2022 sac roe fishery generated $12.7 million in ex-vessel value, the highest since 2013, though recent annual figures have hovered around $5 million amid market fluctuations. In British Columbia, the fishery yielded approximately $50 million for harvesters and processors combined in 2017 and 2018, reflecting its role in roe-on-kelp and spawn products. Historical peaks exceeded $55 million in Alaska during high-demand periods for Japanese markets, underscoring the species' fiscal importance despite variability tied to stock abundance and global prices. These operations bolster employment in harvesting, roe processing, and logistics within remote coastal economies, particularly in where spawning concentrations drive seasonal activity. While herring-specific job data is sparse, the fishery integrates into Alaska's broader seafood sector, which contributed $5.7 billion in statewide economic output in 2019 and supported localized processing infrastructure. Ex-vessel revenues from Pacific herring, averaging $28.9 million annually in the with landings of 47,100 metric tons, historically amplified regional GDP through multiplier effects in supply chains. Portions of the catch also supply bait for groundfish and , as well as reduction to meal and oil for aquaculture feeds, indirectly enhancing aquaculture economics in the North Pacific where herring-derived proteins reduce feed costs. Empirical trends indicate that quota-managed harvests sustain yields without collapsing stocks, as seen in persistent revenues post-1990s adjustments, favoring long-term community benefits over unchecked exploitation despite periodic market downturns.

Cultural and Culinary Uses

Pacific herring (Clupea pallasii) holds profound cultural significance for of the , including the Haida, , and other coastal First Nations, who have harvested it for subsistence and ceremonial purposes for thousands of years. These communities traditionally preserved whole herring through smoking over open fires or air-drying to create storable foods essential for winter survival and social gatherings. Archaeological evidence and oral histories indicate that such practices sustained populations by leveraging the fish's seasonal spawning aggregations, with often collected directly from natural substrates like or cultivated on placed hemlock branches in "herring gardens" to enhance yields without depleting stocks. In and , herring remains a integral to , prepared fresh during spring spawns or fried on with minimal seasonings to highlight its natural . The Haida term iinang encompasses this deep ecological knowledge, where sustainable harvesting protocols, such as timing collections to respect spawning cycles, reflect principles of reciprocity with marine ecosystems. Culinary applications extend to East Asian traditions, where Pacific herring , processed as kazunoko, features in Japanese New Year's dishes for its crunchy texture and symbolic abundance. Salted intact roe skeins, often paired with kelp, provide a preserved form tied to spawning seasonality, with modern preparations maintaining historical methods of and . These uses underscore the fish's role in bridging Indigenous stewardship and global palates, though contemporary access varies due to regulatory limits on roe harvests.

References

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