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Clupeiformes (Herrings)

Clupeiformes

(Herrings)

Class Actinopterygii

Order Clupeiformes

Number of families 5


Evolution and systematics

Fishes in the order Clupeiformes are teleosts, a group of fishes characterized evolutionarily by the presence of true bone forming their skeletons and by specific bone structures in the tail and skull. Teleosts arose during the early Mesozoic era (approximately 200 million years ago). Four subsequent radiations gave rise to the current major groups of fishes, with one of these radiations producing the Clupeomorpha. A major evolutionary feature that distinguishes clupeomorphs is the extension of the gas bladder into the brain case so that it contacts the inner ear, and thereby increasing the hearing ability of the fish. Modern Clupeiformes also possess other evolutionary advances over their closest ancestors: a modified joint in the jaw caused by fusion of the angular to the articular and a reduction of the caudal skeleton.

Two suborders, Clupeoidei and Denticipitoidei, are recognized in the Clupeiformes. The Clupeiodei includes the families Chirocentridae (wolf herrings; 1 genus, 2 species), Clupeidae (herrings, menhadens, pilchards, sardines, shads, and sprats; 5 subfamilies, 56 genera, 214 species), Engraulidae (anchovies; 2 subfamilies, 16 genera, 145 species), and Pristigasteridae (sawbelly herrings; 2 subfamilies, 9 genera, 36 species). The Denticipitoidei includes the family Denticipitidae (denticle herring; 1 genus, 1 species).

Physical characteristics

Clupeoids are small fusiform (tapering toward each end) fishes with streamlined bodies that facilitate fast swimming in open water. They have dark shading on their backs and bright silvery sides. Except for the head, their bodies are completely covered in large scales. Most clupeoids lack a lateral line, and only in the deticipitoid herring does this line extend along the body. The fins of clupeoids lack spines. A single dorsal fin is located near the middle of the body, and the tail is forked. Many clupeoids have a row of scutes, modified scales that usually have sharp points towards the rear, along the medial line of the belly. The smallest cluepoid is the Sanaga pygmy herring (Thrattidion noctivagus), measuring only 0.83 in (2.1 cm) in standard length; male wolf herrings (Chirocentrus spp.) are the largest herring, attaining standard lengths of 39 in (100 cm).

Distribution

Clupeiformes are widely distributed worldwide between 70°N to 60°S latitude. They primarily live in oceans, but some species inhabit coastal margins and fresh water for at least a portion of their lives.

Habitat

Nearly all Clupeiformes are open-water, pelagic species. Four-fifths of all species are marine, with habitats ranging from nearshore littoral zones to nearly 100 mi (160 km) offshore. Many are found near the surface at times but often move to deeper waters during the day. Some Clupeiformes live in inland waters or are anadromous, moving inland to spawn. These species utilize bays, estuaries, marshes, rivers, and freshwater streams as habitats. Landlocked populations have formed as shads, alewives, and herrings moved into lakes or rivers and became trapped between dams.

Behavior

Clupeiformes are perhaps best recognized for the large schools they form. Schools may include hundreds or thousands of individuals ranging from the young to adults, but individuals in a school are usually of similar size. Schooling is

a form of organization in which behavior is synchronized; large numbers of fish may swim parallel to each other in the same direction with fairly uniform spacing. These synchronized aggregations are believed to confer swimming efficiency and, most importantly, to enable fishes to avoid or deter predators. Clupeiformes also congregate in smaller, less-organized shoals, particularly during spawning seasons. In addition to schooling, many Clupeiformes undertake some type of migration. Some clupeoid fishes are anadromous, migrating from the ocean to streams and rivers for spawning. They also may migrate inshore or latitudinally on a seasonal basis. Many clupeoids migrate in the water column on a diel basis, staying at deep depths during the day and moving to shallow depths at night.

Feeding ecology and diet

Most Clupeiformes filter feed by straining water through their long and numerous gill rakers. They consume plankton, particularly small crustaceans and the larval stages of larger crustaceans and fishes. Some herrings visually locate and target food particles. Clupeoid fishes are important prey for larger fishes, seabirds, and marine mammals.

Reproductive biology

Clupeiformes produce large numbers of offspring, either through a single seasonal spawning event or by spawning in seasonal peaks throughout the year. Most Clupeiformes spawn in shoals by broadcasting large numbers of small, buoyant eggs in waters near the surface. The eggs and larvae drift passively in currents as they develop. Herrings, on the other hand, produce demersal eggs that sink to the bottom, where they often adhere to the substrate until they hatch. After hatching, larvae become pelagic.

Conservation status

Two Clupeiformes are listed as Endangered by the IUCN: the Alabama shad (Alosa alabamae) and the Laotian shad (Tenualosa thibaudeaui). The Alabama shad is found in the northern portion of the Gulf of Mexico, from the Mississippi delta eastward to the Choctawhatchee River in Florida. It also occurs in inland rivers from Iowa to Arkansas and eastward to West Virginia. The Laotian shad occurs in the Mekong River basin, including inland waters of Thailand, Laos, and Cambodia. Most Clupeiformes are not threatened by severe population disruptions, but populations do show natural variability due to fluctuations in reproductive success. This natural variability is exacerbated by fishing pressure and global climate patterns.

Significance to humans

Clupeiformes are some of the most economically important fishes in the world's oceans. They have been widely exploited throughout human history, primarily for food but also as a source of oil, fertilizer, and animal feed. Herring fishing was one of the earliest occupations of coastal peoples, as first described in England in a chronicle that dates back to a.d. 709. The first commercial fishing establishment opened in Heligoland, a small island in the North Sea off the coast of Germany, in 1425.

Clupeiformes continue to constitute a large portion of world's commercial fisheries. Although 186 species are exploited by pelagic fisheries worldwide, 50% of the total landings in 1997 were represented by only seven species. Among these seven, four are Clupeiformes: the anchoveta (Engraulis ringens), Atlantic herring (Clupea harengus), Japanese pilchard (Sardinops melanostictus), and South American pilchard (Sardinops sagax). Herrings and anchovies constitute approximately 25% of the total fisheries harvest worldwide.

In addition to being heavily utilized by humans, Clupeiformes are an important component of the broader marine ecosystem. They serve as food items for larger predatory fishes, sea birds, and marine mammals. Thus, clupeoids sustain other organisms of importance to humans through ecosystem interactions.

Species accounts

List of Species

Dorab wolf herring
Alewife
American shad
Atlantic menhaden
Atlantic herring
Pacific herring
Gizzard shad
European pilchard
South American pilchard
Bay anchovy
Northern anchovy
Anchoveta

Dorab wolf herring

Chirocentrus dorab

family

Chirocentridae

taxonomy

Chirocentrus dorab Forsskål, 1775, Jiddah, Saudi Arabia.

other common names

English: Blackfin wolf herring, dorab, silver barfish, wolf herring; French: Chirocentre dorab, sabre; German: Wolfshering; Spanish: Arencón dorab, arenque lobo de la India, sabre; Arabic: Gairi, kerli, dorab, samak abu sayf.

physical characteristics

Reaches lengths of approximately 3.3 ft (1 m). Elongate, highly compressed body. Silvery with a bright blue-gray back. Unlike species of the Clupeidae, the wolf herring has no scutes along the belly. Two distinctive fang-like canine teeth point forward in the upper jaw, and a series of canine teeth is present in the lower jaw.

distribution

Warm coastal waters of the Indo-Pacific, from the Red Sea and East Africa, to the Solomon Island, north to Japan, and south to Australia.

habitat

Nearshore portions of oceans and inshore brackish areas.

behavior

Does not form large schools, but a small number of individuals may be found together. Known for its leaping powers.

feeding ecology and diet

Feeds on other small schooling fishes, such as herrings and anchovies. To a lesser extent, it also eats fish eggs, larvae, and crustaceans.

reproductive biology

Researchers believe that Dorab wolf herrings broadcast eggs into the water column, from which pelagic larvae hatch.

conservation status

Not listed by the IUCN.

significance to humans

Minor commercial fishery in the Indo-Pacific, with products marketed fresh, frozen, dried, and salted.


Alewife

Alosa pseudoharengus

family

Clupeidae

taxonomy

Alosa pseudoharengus Wilson, 1811, Delaware River at Philadelphia, Pennsylvania, United States.

other common names

English: Bigeye herring, branch herring, freshwater herring, golden shad, grayback, gray herring, green shad, kyack, mulhaden, sawbelly, spring herring, white herring; French: Alose gaspareau, gaspareau, gasparot, gasperot; German: Maifisch; Spanish: Alosa, pinchagua.

physical characteristics

Sea-run alewives reach a maximum length of 15 in (38.1 cm); landlocked alewives are typically around 6 in (15.2 cm). Small bodies. Strongly compressed laterally. Silvery bodies with grayish green backs. A row of scutes runs along the ventral edge of the belly. A single black spot is present behind the head.

distribution

Atlantic coast of North America, from the St. Lawrence River and Nova Scotia, Canada, to North Carolina. Numerous translocations of this species have resulted in the establishment of landlocked populations in many inland water bodies, including all of the Great Lakes.

habitat

Spends a large portion of its life at sea and migrates into inland freshwater streams for spawning. Landlocked populations live entirely in inland lakes or river systems.

behavior

Forms schools in open water. Sea-run alewives migrate from the sea to freshwater streams to spawn. Populations in lakes migrate on a diel basis, moving inshore at night and retreating to deeper offshore waters during the day.

feeding ecology and diet

Feeds on zooplankton, primarily copepods, cladocerans, mysids, and ostracods. Inshore adults eat insect larvae as a large part of their diet.

reproductive biology

Moves from the sea to rivers for spawning, with the timing of migration dependent on water temperature. In lakes, alewives move onto shallow beaches and into ponds to spawn. Males and females mature at one and two years of age, respectively. Females usually move to the spawning site before males. Groups of 2–3 fish spawn at night over gravel or rocky substrates. Eggs are broadcast by females and fertilized in the water column by males. Immediately after spawning, the eggs sink and adhere to the substrate. Upon hatching, the young remain at the spawning grounds until the late larval stage and then move slowly into deeper water or the sea.

conservation status

Not listed by the IUCN, but overfishing, pollution, and impassable dams contribute to stock declines.

significance to humans

Harvested commercially and used as fresh, dried, or salted meat, bait for crab or lobsters, and sometimes as animal feed. Also serves as a forage base for other commercial or recreational fish stocks. In lakes, alewives can be a nuisance to humans. They may become so abundant that they clog industrial water intake pipes, and they often die off in massive events related to fluctuations in water temperature and dissolved oxygen.


American shad

Alosa sapidissima

family

Clupeidae

taxonomy

Alosa sapidissima Wilson, 1811, Delaware River at Philadelphia, Pennsylvania, United States.

other common names

English: Atlantic shad, common shad, Connecticut river shad, herring jack, North River shad, Potomac shad, Susquehanna shad, white shad; French: Alose, alose canadienne, alose savoureuse; German: Amerikanische Finte, Amerikanischer Maifisch; Spanish: Sábalo americano.

physical characteristics

Average size 15 in (38 cm) long. Silvery with blue or blue-green metallic shading on its back. Moderately compressed body with a distinct keel of ventral scutes along the belly. A large black spot is located behind the rear edge of the gill cover, followed by several smaller dark spots.

distribution

Occurs off the Atlantic coast of North America, from the St. Lawrence River and Nova Scotia to central Florida. Introduced to the Sacramento River in California by the U.S. Fish Commission in 1971. Since then, its range has expanded greatly, and it can now be found from Cook Inlet, Alaska, to Baja, Mexico.

habitat

Occurs in the ocean at depths to 820 ft (250 m). As an anadromous fish, it spends most of its life at sea but migrates into estuaries and freshwater streams to breed.

behavior

Forms schools after reaching a juvenile size of approximately 8–12 in (20–30 cm). Anadromous species, migrating into freshwater areas for spawning.

feeding ecology and diet

Planktonic feeder that consumes primarily copepods and mysids (shrimp-like crustaceans), but occasionally eats small fishes.

reproductive biology

Ascends freshwater rivers and streams in the spring to spawn. Males mature at four years of age; females first spawn between five and seven years. Spawning commences at water temperatures of 53.6°F (12°C) and peaks at 65°F (18.3°C). The males arrive at the spawning location before the females. During spawning, several males and a female swim close to the surface during the evening. Females release eggs in the open water, where they are fertilized by males. Eggs are 0.1–0.15 in(2.5–3.5 mm) in diameter after fertilization, and after 8–10 days, small larvae of 0.35–0.40 after spawning, but juveniles spend their first summer in the river and reach the sea by autumn. Most American shad spawn more than once, but some die after spawning at southern latitudes of the Atlantic coast.

conservation status

Not listed by the IUCN, but the presence of dams in rivers and streams impedes spawning migrations and has contributed to the decline of some populations.

significance to humans

Commercially fished in rivers and estuaries during spawning migrations. The flesh is eaten fresh, salted, or smoked, and this species is prized for its tasty roe. American shad play a central role in shad planking parties, where the fish are slow-cooked on oak boards over campfires. Shad planking parties are central to political rallies and community gatherings, particularly in the mid-Atlantic region of the United States.


Atlantic menhaden

Brevoortia tyrannus

family

Clupeidae

taxonomy

Brevoortia tyrannus Latrobe, 1802, Chesapeake Bay, United States.

other common names

English: Bony fish, bugfish, bunker, fatback, menhaden, moss-bunker, pogy, whitefish; French: Alose tyran, menhaden tyran, menhaden; German: Menhaden; Spanish: Lacha tirana.

physical characteristics

Adults typically 12–14 in (30.5–35.6 cm) in length. Deep bodied and laterally compressed. Silvery in color, with brassy sides and a dark blue-green back. A row of sharp scutes extends along the ventral edge of the belly. A large dark spot is located behind the gill cover, followed by several smaller spots.

distribution

Western Atlantic from Nova Scotia, Canada, to Indian River, Florida.

habitat

Primarily pelagic fish. Found in waters over the continental shelf. Moves inshore to bays, inlets, and estuaries in the summer.

behavior

Forms large, compact schools of juveniles and adults. Stratifies by size along the Atlantic seaboard during annual north-south migrations. Fish of all ages congregate near Cape Hatteras, North Carolina, during the winter months. Most adults move northward after March, with the largest fish migrating as far as the Gulf of Maine; some adults move southward as far as to waters off of Florida. Fish of all ages and sizes then return to Cape Hatteras in late autumn. Atlantic menhaden also move in and out of inshore habitats with the tides, the season, and the weather.

feeding ecology and diet

Feeds by filtering phytoplankton and zooplankton, including diatoms, copepods, and euphausids.

reproductive biology

Spawns in the open ocean throughout the year. Eggs are buoyant and hatch at sea. Over one to three months, larvae are transported to estuaries by ocean currents, where they develop into juveniles. Most Atlantic menhaden first spawn during their third year of life. Females exhibit high fecundity levels, producing about 38,000–50,000 eggs per female.

conservation status

Not listed by the IUCN.

significance to humans

Previously used by Native Americans and early European colonists as fertilizer. A fishery developed for menhaden during the late 1800s and early 1900s. Today used primarily to produce fish meal and oil, although some are marketed fresh, salted, canned, or smoked. Due to the large biomass represented by this species, menhaden constituted around 30% of commercial fisheries landings along the U.S. Atlantic coast in 2000.


Atlantic herring

Clupea harengus

family

Clupeidae

taxonomy

Clupea harengus Linnaeus, 1758, European seas. This species was previously considered a subspecies, Clupea harengus harengus, but recent taxonomic classifications distinguish it as a separate species.

other common names

English: Bank herring, brit, fall herring, hern, herning, herron, labrador herring, mesh herring, murman hering, Norwegian herring, sardine, sea Atlantic herring, sea stick, shore herring, split, spring herring, summer herring, yawling; French: Hareng atlantique, hareng de l'Atlantique; German: Allec, Hering, Silling; Spanish: Arenque, arenque del Atlántico.

physical characteristics

Maximum size 17.7 in (45 cm), but most fish captured in fish-eries are 11.8–13.8 in (30–35 cm). Elongate and slender. Back is dark blue-green (or bluish green), sides and belly are silvery, and snout is blackish blue. There are no distinct dark spots on the body or fins. The belly is rounded with scutes, but has no prominent keel. The lower jaw of the Atlantic herring is slightly longer than the upper jaw.

distribution

Eastern Atlantic Ocean, from the northern Bay of Biscay northward to Spitzbergen and Novaya Zemlya, and around Iceland and southern Greenland. Also found in the western Atlantic, from southwestern Greenland, around Labrador, and south to South Carolina.

habitat

Coastal pelagic species. Found at depths ranging from near the surface down to 656 ft (200 m).

behavior

Schools in coastal waters. Exhibits complex feeding and spawning migrations, but the timing and extent of migrations varies by morphological race of the fish. Stays in deep water during the day but moves to the surface at night. Most migrate to coastal spawning grounds at the onset of spawning seasons. Also migrates north and south seasonally to feeding areas and for over wintering.

feeding ecology and diet

During their first year of life, Atlantic herring feed on small planktonic copepods. Thereafter, they consume mostly copepods, but also amphipods, mysids, small fishes, and ctenophores. Atlantic herring locate food particles visually, but they can switch to filter feeding if they encounter a high density of small food particles. This species is important as prey in the marine food chain, and it is consumed by larger fishes, squids, skates, whales, and seabirds.

reproductive biology

Exhibits a wide range of spawning behaviors and strategies that vary by stock and race. At least one population spawns in any month of the year, but each race spawns at a different time and place. Off Greenland, these fishes spawn in up to 16 ft (5 m) of water; autumn-spawning herrings in the North Sea spawn at depths of 656 ft (200 m). The eggs are adhesive and are laid over rocks, stones, gravel, sand, algae, or vegetation on the seabed. Atlantic herring reach their sexual maturity at three to nine years of age, and females may produce 20,000–40,000 eggs.

conservation status

Populations fluctuate widely. Some stocks are considered over-fished, while others have shown recent increases in abundance. Not listed by the IUCN.

significance to humans

The commercial fishery for Atlantic herring ranks among the most important in the world in terms of biomass and value. Atlantic herring were a major item of trade between Scandinavian countries and western European countries as early as the twelfth century. The western Atlantic fishery developed in the 1800s and supplied fish that were sold as bait or sardines. In the United States and Canada, most Atlantic herring are sold as sardines or are converted to fish meal or oil. Smoked, salted, pickled, fresh, and frozen herring are more common in Europe. Herring form an important part of the cuisine of certain cultures; it is particularly central to the Jewish cuisine due to dietary and cooking restrictions observed by many Jews.


Pacific herring

Clupea pallasii

family

Clupeidae

taxonomy

Clupea pallasii Valenciennes, 1847, Kamchatka, Russia. This species was previously considered a subspecies, Clupea harengus pallasii, but recent taxonomic classifications distinguish it as a separate species. One subspecies: Clupea pallasii marisalbi Berg, 1923.

other common names

English: Herring, North Pacific herring, Oriental herring; French: Hareng du Pacifique, hareng pacifique; German: Pazi-fischer Hering; Spanish: Arenque del Pacifico.

physical characteristics

Maximum size 18 in (46 cm), but most are less than 11.8 in (30 cm). Elongate and slender, with a rounded belly and scutes. Back is dark blue to olive, the sides and belly are silvery. There are no distinct dark spots on the body or fins. Distinguished from the Atlantic herring by having fewer vertebrae and fewer postpelvic scutes.

distribution

Arctic Sea from the White Sea eastward to the Ob Inlet. They range in the Western Pacific along the coast of Russia, south to Japan and Korea. In the eastern Pacific, they are found from the Kent Peninsula and Beaufort Sea, around Alaska, and south to Baja California, Mexico.

habitat

Coastal pelagic species that uses estuaries and bays for spawning.

behavior

Begins schooling as a juvenile and is found in inshore waters during this life history stage. Adults form schools at sea, but they migrate to inshore waters, including bays and estuaries, to spawn. They migrate from the sea to inshore waters, entering bays and estuaries to spawn. They also migrate daily from deep waters during the day to shallower portions of the water column at night. Pacific herring do not undertake extensive latitudinal migrations along the coast.

feeding ecology and diet

Feeds on zooplankton, including euphausids, copepods, mysids, and amphipods. Serves as food for a wide variety of predatory fishes and seabirds.

reproductive biology

Breeds from December to July, with the precise timing of spawning varying with latitude. Adults congregate near inshore spawning grounds several weeks before spawning. Herring spawn over a variety of substrates, including rocks, algae, vegetation, and flat surfaces. The female deposits rows of eggs along the substrate; the eggs are then fertilized by the males. Spawning occurs several days at a time, with events separated by a day to several weeks. In California, these herring spawn at age two to three, but members of the species that inhabit waters at higher latitudes reach maturity later.

conservation status

Not listed by the IUCN, but populations are sensitive to shoreline development and fishing pressures.

significance to humans

Previously used by Native Americans as fresh or salted food and for bait. In the early 1900s, commercial fisheries developed along the west coast of the United States and Canada for salted and canned herring and for the reduction of the fish to meal and oil. These fishes are valued in Russia as food and are utilized fresh, dried, smoked, canned, and frozen. In the eastern Pacific, the roe and eggs are taken to supply Asian markets. Pacific herring also are used in Chinese medicine.


Gizzard shad

Dorosoma cepedianum

family

Clupeidae

taxonomy

Dorosoma cepedianum Lesueur, 1818, Baltimore, Maryland and Philadelphia, Pennsylvania, United States.

other common names

English: American gizzard shad, eastern gizzard shad, gizzard mud, gizzard nanny shad, hickory shad, mud shad, nanny shad, skipjack winter shad; French: Alose à gésier, alose a gésier américaine, aloser noyer; Spanish: Sábalo molleja.

physical characteristics

Maximum size 22.4 in (57 cm). Silver to brassy body with a bluish back and white belly. There is a large dark spot behind the operculum and above the pectoral fin. Snout is rounded and blunt. The last dorsal fin ray extends to a long filament.

distribution

Atlantic coast of the United States from New York, southward into the Gulf of Mexico and to the basin of the Rio Pánuco in Mexico. Found inland in the Great Lakes, St. Lawrence, Mississippi, Atlantic, and Gulf Slope drainages, with its east-west range extending from Quebec to North Dakota in the north and from Florida to New Mexico in the south.

habitat

Can tolerate salinities from fresh water to 33 or 34 parts per 1,000, but is most common in freshwater lakes, reservoirs, swamps, and slow-moving rivers. Adults are found in estuaries and protected bays; the young are sometimes found far upstream in small streams.

behavior

Forms schools. Individuals may be seen leaping out of the water and skipping along the surface on their sides, giving rise to one common name for the species, the skipjack.

feeding ecology and diet

Young gizzard shad feed on zooplankton. As they mature, gizzard shad become herbivorous filter feeders, consuming phytoplankton and algae on the bottom of the water body.

reproductive biology

Spawns from late winter through most of the summer; the exact spawning dates relate to latitude and water temperature. Adults spawn near the surface, usually in groups of two males and one female. The eggs, which are very small (0.03 in [0.75 mm] in diameter) sink and adhere to the bottom. They hatch in three days to one week, depending on temperature.

conservation status

Not listed by the IUCN.

significance to humans

Young gizzard shad are important forage fishes that sustain other game and commercial fish species. In some areas, the abundance of gizzard shad creates a nuisance for humans when low temperatures, low oxygen, or disease trigger the death of a large number of fish that often wash onto shores and decay. Gizzard shad also create problems for fisheries; they entangle the nets of commercial fishermen and out-compete many preferred recreational species. The gizzard shad is used for fertilizer and as a component of cattle and hog food.


European pilchard

Sardina pilchardus

family

Clupeidae

taxonomy

Sardina pilchardus Walbaum, 1792, Cornwall, England.

other common names

English: Pilchard, sardine, true sardine; French: Sardine, sardine commune, sardine d'Europe; German: Pilchard, Sardine; Spanish: Majuga, parrocha, sardina, sardiña, xouba.

physical characteristics

Can attain lengths of 9.8 in (25 cm). Body elongate with a rounded belly that has scutes but not a defined keel. Back dark green or olive in color, with golden flanks and a silvery white belly. There is a series of dark spots along the upper flanks.

distribution

Northeast Atlantic from Iceland and the North Sea south to Senegal. Also found in the western Mediterranean and Adriatic Seas.

habitat

Coastal, pelagic species that is found at water depths of 82–180 ft (25–55 m) during the day and 49–115 ft (15–35 m) at night.

behavior

Forms schools and undertakes diel migrations in the water column, staying at deeper depths during the day and rising to shallower waters at night.

feeding ecology and diet

Feeds mainly on planktonic crustaceans, but may also consume larger planktonic organisms.

reproductive biology

Breeds at 66–82 ft (20–25 m) depth up to 62 mi (100 km) offshore. Spawning time varies by geographic location, taking place in April in the English Channel, from June to August in the North Sea, from September to May along European coasts in the Mediterranean, and from November to June along

African coasts in the Mediterranean. The eggs are buoyant and develop in the water column.

conservation status

Not listed by the IUCN.

significance to humans

Constitutes an important fishery throughout its range. Catches have steadily risen since 1950, and in 1999, over 900,000 tons (816,466 tonnes) was harvested. The fish is marketed fresh, frozen or canned but may also be utilized in dried or smoked forms.


South American pilchard

Sardinops sagax

family

Clupeidae

taxonomy

Sardinops sagax Jenyns, 1842, Lima, Peru.

other common names

English: Chilean pilchard, Chilean sardine, Pacific American sardine, pilchard, sardina, sardine; French: Hareng, Pilchard sudaméricain, Sardinops du Chili; German: Chilenishe Sardine, Südeamerikanische Sardine; Spanish: Pilchard chileña, Sardina, Sardina española.

physical characteristics

May grow to 12 in (30 cm), but sizes of 8 in (20 cm) are more common. Bluish on the back and silvery on the sides. Roundbodied and has a keel with scutes along the belly.

distribution

Eastern Pacific along the coasts of Peru and Chile; also found near the Galápagos Islands.

habitat

Coastal, pelagic species that is found to depths of around 131 ft (40 m).

behavior

Forms large schools and migrates up and down in the water column on a diel cycle.

feeding ecology and diet

Feeds mainly on planktonic crustaceans but may also consume phytolankton. Bethnic organisms, including ostracods and polychaete worms make up a minor portion of the diet of adults.

reproductive biology

Spawns twice each year by broadcasting eggs. Eggs and larvae are pelagic and develop in the water column.

conservation status

Not listed by the IUCN.

significance to humans

Forms the basis of a substantial fishery off Peru and Chile. The fishery reached a peak of 6.5 million tons (5.9 million tonnes) in 1985, corresponding to the crash of the Peruvian anchoveta fishery caused by El Niño. However, catches have declined since then, and the total harvest in 1999 was less than 450,000 tons (408,233 tonnes).


Bay anchovy

Anchoa mitchilli

family

Engraulidae

taxonomy

Anchoa mitchilli Valenciennes, 1848, New York, United States.

other common names

English: Common anchovy; French: Anchois américain; Spanish: Anchoa de caleta.

physical characteristics

Small species, typically 3–4 in (7.6–10.2 cm) total length. Nearly transparent and greenish in color, with a silvery band along the side of the body. Snout overhangs the mouth and low jawbone extends well beyond the eye.

distribution

Atlantic coast of North America from Casco Bay, Maine, to the Florida Keys, and westward around the Gulf of Mexico south to the Yucatán peninsula.

habitat

Primarily an estuarine and inshore coastal species. Utilizes a wide variety of habitats including bays, sandy beaches, marshes, islands, and spoil banks. Typically found over muddy bottoms or in vegetation. Tolerates a wide range of salinities but is often found in brackish water.

behavior

Swims in schools. Migrates seasonally from deep waters in winter to shallow shores and wetlands in summer.

feeding ecology and diet

Plankton feeder; primarily consumes mysids and copepods. Small fishes, gastropods, and isopods are occasionally taken.

reproductive biology

Spawns from late winter to early fall when water temperatures are around 68°F (20°C). Spawning takes place during the evening hours in shallow waters near barrier islands, in bays, and in estuaries. Spawning usually occurs in large schools. Females broadcast eggs, which are fertilized in the water column by males. Eggs float near the water surface for approximately 24 hours after fertilization before they hatch. Bay anchovies mature to adults in two and one-half months.

conservation status

Not listed by the IUCN.

significance to humans

Used as bait and to make anchovy paste. Important as forage fishes in food chains that sustain other commercial and recreational fishery species in estuaries and coastal areas.


Northern anchovy

Engraulis mordax

family

Engraulidae

taxonomy

Engraulis mordax Girard, 1854, San Francisco, California, United States.

other common names

English: Anchovy, Californian anchovy, North Pacific anchovy, pinhead; French: Anchois de California, anchois du nord, anchois du Pacifique, anchois du Pacifique nord; German: Amerikanische Nordpazifische, Amerikanische Sardelle; Spanish: Anchoa de California, anchoa del Pacifico, anchoveta, anchoveta de California, anchoveta norteña.

physical characteristics

Grows to 4 in (10 cm). Body slender and elongate but round in cross section. Back is green and sides are silvery, including a stripe along the flank in young individuals. Large head and mouth with a pointed snout.

distribution

Northern Pacific from Vancouver Island, Canada, to Baja California, Mexico.

habitat

Pelagic marine species found to depths of 984 ft (300 m). Usually stays in coastal waters within 18.6 mi (30 km) of shore but may range as far as 298 mi (480 km) offshore. Enters estuaries, bays, and inlets during the spring and summer.

behavior

Forms schools during most of the year, although these become smaller or break up in late spring, typically around the end of spawning in April or May. Moves to inshore waters during spring and summer and migrates offshore in the fall and winter. Diel migrations also occur, with the northern anchovy remaining at depths during the day and approaching the surface in low-density schools at night.

feeding ecology and diet

Obtains food both by filter feeding and particulate biting. Feeds on plankton, primarily euphausids, copepods, and decapod larvae. Northern anchovies are an important forage species for other fishes, birds, and marine mammals.

reproductive biology

Spawns in inlets and offshore, with spawning activity occurring at night. Two major spawning areas exist in coastal waters off southern California and Baja California, Mexico. Eggs are broadcast and fertilized in the water column, then float and incubate for two to four days before hatching. Spawning takes place throughout the year, and individual females may spawn several times each year. However, as a whole, the species exhibits a clear peak in spawning during the winter and early spring.

conservation status

Not listed by the IUCN.

significance to humans

Supports a commercial and bait fishery, which developed after the collapse of the Pacific sardine fishery in the 1940s. Approximately 25 million pounds were landed in 2000, with most used for fish meal, fertilizer, and animal feed. A small portion is consumed by humans in pickled or salted forms.


Anchoveta

Engraulis ringens

family

Engraulidae

taxonomy

Engraulis ringens Jenyns, 1842, Iquique, Chile.

other common names

English: Anchovy, Peruvian anchoveta; French: Anchois du pérou, anchois péruvien; German: Perusardelle, Südamerikanische Sardelle; Spanish: Anchoa, anchoa bocona, anchoveta, anchoveta peruana, atunera, chicora, manchuma, manchumilla, peladilla, sardina bocona.

physical characteristics

Grows to 7 in (18 cm) and has a slender, elongated body that is round in cross section. Its large snout and mouth are similar to those of other anchovies. Silvery in color; juveniles have a stripe along the flank.

distribution

West coast of South America along the Peru Current, typically from around Aguja Point, Peru, south to Chiloë, Chile.

habitat

Pelagic species. Occurs in surface waters, usually within 49 mi (80 km) of the shore, but occasionally offshore to 99 mi(160 km).

behavior

Forms huge schools. Descends to deeper waters during the day but rises to the surface at night.

feeding ecology and diet

Filter feeder. Depends entirely on the plankton of the Peru Current for food. Diet consists largely of diatoms, which make up as much as 98% of its consumption according to some studies. It also consumes copepods, euphausids, fish eggs, and dinoflagellates. Seabirds prey heavily on schools of anchoveta.

reproductive biology

Matures at around one year of age. Breeds throughout the year, but two peaks of spawning occur in late winter and early autumn. Their buoyant ellipsoidal eggs are broadcast into the water column and fertilized.

conservation status

Not listed by the IUCN, but populations vary greatly with climatic conditions. El Niño, the oceanographic condition that results in warmer water in the Pacific Ocean, slows and may stop the upwelling of nutrient rich waters in the Peru Current. This can have devastating effects on the anchoveta by dramatically reducing its planktonic food base.

significance to humans

Previously the world's largest fishery in terms of biomass in 1971, with over 13 million tons (11.7 million tonnes) harvested. Populations declined during the 1970s and 1980s due to overfishing and the occurrence of severe El Niño events in 1972–1973 and 1982–1983. Since that time, anchoveta populations have recovered and the species again constitutes a major fishery. In 2000 the anchoveta comprised over 12 million tons (11 million tonnes) of a total harvest in the southeastern Pacific of around 16.5 million tons (15 million tonnes) for all fishes. Peru and Chile rely upon this fishery as a major export and source of income. It is utilized primarily as fish meal and oil.


Resources

Books

Laws, Edward A. El Niño and the Peruvian Anchovy Fishery. Sausalito, CA: University Science Books, 1997.

Lecointre, G., and G. Nelson. "Clupeomorpha, Sister-Group of Ostariophysi." In Interrelationships of Fishes, edited by Melanie L. J. Stiassney, Lynne R. Parenti, and G. David Johnson. San Diego: Academic Press, 1996.

Periodicals

Petitgas, P., D. Reid, P. Carrera, M. Iglesias, S. Georgakarakos, B. Liorzou, and J. Masse. "On the Relation Between Schools, Clusters of Schools, and Abundance in Pelagic Fish Stocks." ICES Journal of Marine Science 58(2001): 1,150–1,160.

Whitehead, P. J. P. "Clupeoid Fishes of the World (Suborder Clupeoidei). An Annotated and Illustrated Catalogue of the Herrings, Sardines, Pilchards, Sprats, Shads, Anchovies, and Wolf-Herrings. Part 1—Chirocentridae, Clupeidae, and Pristigasteridae." FAO Fisheries Synopsis 125, no. 7(1985): 303.

Whitehead, P. J. P, G. J. Nelson, and T. Wongratana. "Clupeoid Fishes of the World (Suborder Clupeoidei). An Annotated and Illustrated Catalogue of the Herrings, Sardines, Pilchards, Sprats, Shads, Anchovies, and Wolf-Herrings." Part 2—Engraulidae." FAO Fisheries Synopsis 125, no. 7 (1988): 274.

Organizations

Menhaden Resource Council. 1901 N. Fort Myer Drive, Suite 700, Arlington, VA 22209 USA. Phone: (703) 796-1793. E-mail: [email protected] Web site: <http://www.menhaden.org>

Other

FAO Fisheries Department. (13 Nov. 2002) <http://www.fao.org/fi/default.asp>

FishBase. 8 Aug. 2002 (13 Nov. 2002). <http://www.fishbase.org/search.cfm>

Katherine E. Mills, MS

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