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Caplin Research At Bryant's Cove   (DRAFT) - Taxonomy Species capelin Species Id M010068 Date 27 AUG 96 TAXONOMY NAME - capelin OTHER COMMON NAMES - Caplin, Lodde, Whitefish, Angmagssat and Fr: Capelan E LEMENT CODE - CATEGORY - Fish PHYLUM AND SUBPHYLUM   CLASS AND SUBCLASS - Osteichthyes   ORDER AND SUBORDER - Salmoniformes   FAMILY AND SUBFAMILY - Osmeridae,   GENUS AND SUBGENUS - Mallotus,   SPECIES AND SSP - villosus,   SCIENTIFIC NAME - Mallotus villosus   AUTHORITY - Muller, 1777   TAXONOMY REFERENCES - 258 and 232 STATUS Coded Status   Migrant Commercial/consumption Commercial/industrial Commercial/bait See Comments   REFERENCES FOR STATUS - 258 and 232   COMMENTS ON STATUS - An intensive migration inshore by coastal populations takes place prior to spawning activities on beaches *258*.   Although used as human food when sun-dried by the Newfoundland people for many years, abundance of capelin led to its use in the early part of century as a fertilizer, as bait for the cod line fishery, and as dogfood. Although most of the world's supply is now being processed into fish meal and oil, capelin and capelin products are considered high in nutritional value. Expolitation of previously underutilized stocks resulted in extensive experimentation with numberous methods of processing for use as human food, especially in Canadian laboratories and Norway *258*.   Capelin with high fat content, i.e., prespawning fish, are preferred for all types of processing and consumption. The fat content can be as high as 23 percent in fish in prime condition *258*.   When dried and smoked, capelin are relished by the Japanese and served with their native 'sake'. The Greenland people value the product and, when sun-dried, store it for human consumption during the winter months when food supplies are short *258*.   As food for cage-reared fishes, such as trout and salmon, its use is growing in significance *258*.   The decline in stocks of capelin is of great concern to the cod fishery because capelin is such an important prey of the Atlantic cod. Exploitation by foreign fleets aggravates the problem of continuing food supply for this important species. An account of status stocks on the Southeast Shoal of the Grand Bank in 1946 was published. Causes of annual variations in year-class strength are discussed *258*.   Capelin stocks in the Canadian Atlantic area have received increased attention since about 1980. Landings in 1982 and 1983 totaled 31,893 t and 31,041 t respectively, with landed value of $6,691,000 (1982) and $5,958,000 (1983) *258*.   About a month before spawning, capelin migrate to shallow water and segregate into schools by sex *232*.   The ADF&G has managerial authority over caplein, extending to 3 mi offshore.   Status - 1 (DRAFT) - Status Species capelin Species Id M010068 Date 27 AUG 96   Because harvesting of caplein is minimal, no management plan has been formulated. A policy statement, however, has been established that defines the relationship between herring and capelin fisheries that could occur simultaneously *232*. Status - 2   (DRAFT) - Distribution Species capelin Species Id M010068 Date 27 AUG 96 DISTRIBUTION Distribution - 1 HABITAT ASSOCIATIONS HABITAT - MARINE REFERENCES FOR HABITAT - 258 and 232 LAND USE - Water Beaches REFERENCES FOR LAND USE - 258 COMMENTS ON HABITAT ASSOCIATIONS - The capelin is a marine fish of cold, deep water, found in the Atlantic Ocean on the offshore banks and in coastal areas, occasionally spending the winter and early spring months in deep bays off the east coast of Newfoundland. It is a plentiful species off the north shore of the Gulf of St. Lawrence but the largest concentrations in Canadian waters are found off Newfoundland and the Labrador coast. The presence of a spawning population on the Southeast Shoal of the Grand Bank was noted. This population may remain in deep water in this region throughout life, moving close to the bank to spawn *258*. An intensive migration inshore by coastal population takes place prior to spawning activities on beaches. This inshore movement was noted in a study of the Atlantic cod off Newfoundland, which reported that tagged Atlantic cod migrated inshore following their preferred prey, the capelin *258*. Capelin in the Canadian Arctic have been reported spawning in warm, brackishh water. Capelin spawn either offshore in deep water or on sand and gravel beaches *232*. Adult and larval capelin are pelagic fish *232*. ANIMAL/PLANT SPECIES ASSOCIATIONS - Fish - 10 sp. Marine mammals - 5 sp. Marine birds - 9 sp. Atlantic cod (Gadus morhua) Haddock (Melanogrammus aeglefinus) Atlantic salmon (Salmo salar) Atlantic herring (Clupea harengus harengus) Flounders Dogfish Sculpins Eelpout Minke whales (Balaenoptera acutorostrata) Fin whales (Balaenoptera physalus) Harp seal (Phoca groenlandica) Ringed Seal Seabirds Thick-billed murres Winter flounder Cestode (Eubothrium parvum) Nematode (Anisakis sp.) Nematode (Contracaecum sp.) Protozoan (Glugea sp.) Cod Salmon Saffron cod   Habitat Associations - 1   Starry flounder Harbor seal Sea lion Ribbon seal Spotted seal Belukha whale Artic cod Squid   REFERENCES FOR SPECIES ASSOCIATIONS - 258 and 232   COMMENTS ON SPECIES ASSOCIATIONS - Predation and Competition: The role of capelin as food for other fishes, birds, and marine mammals is most significant and has been estimated to be in the order of 5 million t annually at virgin population levels. A study listed 10 fish species, 5 species of marine mammals, and 9 marine bird species known to forage extensively on capelin *258*.   Capelin are preyed upon heavily by the Atlantic cod, Gadus morhua, probably their chief predator. They are followed inshore during the spawning migration. During the period from June to early August 98 percent of the food of the Atlantic cod was capelin. Offshore Atlantic cod also feed heavily on this species. Capelin and their eggs were eaten extensively by haddock, Melanogrammus aeglefinus, on the Southeast Shoal of the Grand Bank during times when haddock were plentiful in the area. They are also consumed in large quantities by haddock on the spawning grounds of Norway and are the chief food of the Atlantic salmon, Salmo salar, in the northwest Atlantic *258*.   Juveniles and larvae often form part of the food of smaller fishes such as Atlantic herring, Clupea harengus harengus. Many other fishes, such as flounders, dogfish, sculpins, and eelpout, prey on them *258*.   Several species of whales, notably minke whales (Balaenoptera acutorostrata), feed intensively on capelin and they are frequently food of fin whales (B. physalus). Their role as food for other species extends to seals, especially the harp seal (Phoca groenlandica) but including ringed seal, and seabirds. A study reported the occurrence of capelin in the stomachs of thick-billed murres captured at Digges Sound, Hudson Bay. Winter flounder feed heavily and systematically on capelin eggs, supporting up to 25 percent of their annual somatic growth from this food source *258*.   Parasites and Disease: The capelin in Atlantic waters appears to be only lighly parasitized. A study listed one species of cestode, Eubothrium parvum; two species of nematodes, Anisakis sp. larva and Contracaecum sp.; and the protozoan Glugea sp., for which the capelin was described as a new host *258*.   A study redescribed and published new locality records of monogenean trematodes taken from the gills of capelin in the northwest Atlantic and Icelandic waters *258*.   Haddock and winter flounder consume large quantities of Atlantic capelin eggs during the spawning period. Egg mortality from predation may be significantly greater at deep-water dermersal spawning sites and may have a significant impact on recruitment of demersal spawners. Many fish are predators of adult and juvenile capelin, including cod and salmon in the Atlantic and arctic char, saffron cod, and starry flounder in the Canadian arctic. Marine mammals also consume capelin as a primary part of their diet. In the Gulf of Alaska, harbor seals and Steller sea lions are predators, especially in nearshore waters in the spring and summer. Other capelin Habitat Associations - 2 predators include ribbon seals in the Pribilof Islands area, spotted seals in the southeastern Bering Sea in the spring, and belukha whales in the coastal northern Bering and Chukchi seas in summer. Competition between capelin and other zooplankton-feeding species may. In the Newfoundland area, the diets of capelin, arctic cod, and squid overlap. In years when capelin have reduced their zooplankton consumption, the abundance of arctic cod and squid has increased *232*.   Habitat Associations - 3 (DRAFT) - Food Habits Species capelin Species Id M010068 Date 27 AUG 96 FOOD HABITS TROPHIC LEVEL - CARNIVORE REFERENCES FOR TROPHIC LEVEL - 258 and 232 LIFESTAGE FOOD FOOD PART General Plankton Not Specified General Crustaceans Not Specified General Copepods Not Specified General Crustaceans See Comments General Copepods See Comments General Invertebrate Cordates Not Specified Adult Copepods Not Specified Adult Crustaceans Not Specified Juvenile Copepods Not Specified Juvenile Crustaceans Not Specified Larva Copepods Egg/Fetus stage Larva Crustaceans Larva stage Larva Invertebrate Cordates Egg/Fetus stage Larva Bacillariophyceae Not Specified Juvenile Crustaceans Adult stage Adult Crustaceans Adult stage   REFERENCES FOR GENERAL FOOD - 258 and 232   REFERENCES FOR ADULT FOOD - 232   REFERENCES FOR JUVENILE FOOD - 232   REFERENCES FOR LARVAE FOOD - 232   COMMENTS ON FOOD - Food of capelin consists of planktonic organisms. Consumption of euphausiids dominates by weight the items consumed, although copepods, especially Calanus, occur in greater numbers and are more frequently found in stomachs. Amphipods and other planktonic invertebrates are also taken. Feeding is seasonal, intensifying in later winter and early spring in the prespawning period, declining as the spawning season approaches, and virtually ceasing during spawning. Several weeks after the spawning period, surviving capelin commence feeding and continue until cessation in early winter *258*.   Capelin are filter feeders, consuming euphausiids, copepods, amphipods, and a variety of planktonic invertebrates *232*.   Factors Limiting Availability of Food: The diet of capelin is dominated by zooplankton, which, in turn, feed on phytoplankton. In polar waters, there is no productivity throughout most of the year, and an outburst of exceedingly high productivity occurs in the short summer season. Primary production in arctic waters is usually limited by available sunlight and, Food Habits - 1 (DRAFT) - Food Habits   Species capelin Species Id M010068 Date 27 AUG 96   less often, by the availability of nutrients for phytoplankton *232*   Feeding Behavior: Capelin's feeding is highly seasonal. Feeding intensity increases before spawning in late winter and early spring and declines with the spawning migration. Feeding ceases during spawning and proceeds again at high intensity until early winter, when zooplankton becomes less abundant. Daily feeding peaks occur in mornings and evenings between May and August in eastern Canada, and a single midday feeding occurs in October and November *232*.   COMMENTS ON ADULT FOOD - Adult capelin north of Norway feed mainly on copepods and euphausiids, and adults in eastern Canada feed primarily on copepods, euphausiids, and appendicularians. The diets of juvenile and adult capelin overlapped, with a shift from copepods to adult euphausiids as the capelin reached a total length of 140 mm *232*.   COMMENTS ON JUVENILE FOOD - Juveniles in eastern Canada feed primarily on copepods, euphausiids, and appendicularians. The diets of juvenile and adult capelin overlapped, with a shift from small copepods to adult euphausiids as the capelin reached a total length of 140 mm *232*.   COMMENTS ON LARVAE FOOD - The diet of first-feeding capeline larvae consists mainly of copepod eggs and nauplii. A shift in diet occurred during growth, with early larval capelin consuming invertebrate eggs, copepods, and diatoms *232*.   Food Habits - 2 (DRAFT) - Environment Associations Species capelin Species Id M010068 Date 27 AUG 96 ENVIRONMENTAL ASSOCIATIONS G = General A = Adult LIM = Limiting RA = Resting Adult J = Juvenile FA = Feeding Adult RJ = Resting Juvenile BA = Breeding Adult FJ = Feeding Juvenile P = Pupae L = Larvae E = Egg RL = Resting Larvae FL = Feeding Larvae   LIFESTAGE ENVIRONMENTAL ASSOCIATIONS   A Water Depth Preference: 200-500 ft. G Water Temperature: Specified in Comments G Water Depth Preference: Specified in Comments BA Water Temperature: Specified in Comments BA Tidal inundation frequency/ duration: specified in comments BA Bottom Type [Aquatic]: Sand BA Bottom Type [Aquatic]: Gravel E Bottom Type [Aquatic]: Sand E Bottom Type [Aquatic]: Gravel BA Bottom Type [Aquatic]: Pebble E Bottom Type [Aquatic]: Pebble E Wave intensity: specified in comments BA Water Temperature: Below 15 degrees C E Water Temperature: Below 15 degrees C E Water Temperature: Specified in Comments E Water Depth Preference: 1-5 ft. E Water Depth Preference: 5-10 ft. E Water Depth Preference: 100-200 ft. E Water Depth Preference: 200-500 ft. E Water Depth Preference: Specified in Comments BA Water Depth Preference: 1-5 ft. BA Water Depth Preference: 5-10 ft. BA Water Depth Preference: 100-200 ft. BA Water Depth Preference: 200-500 ft. BA Water Depth Preference: Specified in Comments E Dissolved Oxygen: Specified in Comments E Air Temperature: Specified in Comments E Relation to Substrate: Specified in Comments E Water Temperature: Between 15-21 degrees C L Water Temperature: Specified in Comments BA Coastal Features: Specified in Comments BA Wave intensity: specified in comments BA Coastal Features: Sand beaches BA Substrate: Specified in Comments BA Bottom Type [Aquatic]: Specified in Comments L Tidal inundation frequency/ duration: specified in comments L Coastal Features: Specified in Comments L Substrate: Specified in Comments L Bottom Type [Aquatic]: Gravel L Bottom Type [Aquatic]: Specified in Comments   Environment Associations - 1 (DRAFT) - Environment Associations Species capelin Species Id M010068 Date 27 AUG 96   LIFESTAGE ENVIRONMENTAL ASSOCIATIONS   E Substrate: Specified in Comments E Substrate: Sand E Bottom Type [Aquatic]: Specified in Comments E Soil Depth: Less than 10 inches E Soil Depth: Specified in Comments E Coastal Features: Specified in Comments E Relation to Substrate: Occurs in substrate [penetrating] E Relation to Substrate: Attached - normally sessile BA Water Temperature: Specified in Comments L Water Temperature: Specified in Comments G Water Depth Preference: 100-200 ft. G Water Depth Preference: 200-500 ft. G Water Depth Preference: Specified in Comments G Water Depth Preference: 500-1000 ft. G Water Depth Preference: 50-100 ft. BA Water Depth Preference: 500-1000 ft. BA Water Depth Preference: Specified in Comments BA Water Depth Preference: 200-500 ft. L Water Depth Preference: 100-200 ft. L Water Depth Preference: Specified in Comments L Water Depth Preference: 50-100 ft. BA Water Temperature: Specified in Comments BA Water Temperature: Between 15-21 degrees C BA Water Temperature: Below 15 degrees C A Water Temperature: Below 15 degrees C A Water Temperature: Specified in Comments L Water Temperature: Below 15 degrees C L Water Temperature: Specified in Comments A Aquatic Habitat Zonation: Open water [pelagic zone] L Aquatic Habitat Zonation: Open water [pelagic zone] A Aquatic Habitat Zonation: Specified in Comments L Aquatic Habitat Zonation: Specified in Comments BA Coastal Features: Sand beaches BA Coastal Features: Specified in Comments E Coastal Features: Sand beaches E Coastal Features: Specified in Comments E Substrate: Sand E Substrate: Rocks E Substrate: Specified in Comments E Relation to Substrate: Attached - normally sessile E Relation to Substrate: Specified in Comments E Water Temperature: Specified in Comments E Water Temperature: Below 15 degrees C   Environment Associations - 2 (DRAFT) - Environment Associations Species capelin Species Id M010068 Date 27 AUG 96   REFERENCES FOR ENVIRONMENTAL ASSOC_ - 258 and 232   REFERENCES FOR ADULT ENVIRONMENTAL ASSOC_ - 232   REFERENCES FOR BREEDING ADULT ENVIRONMENTAL ASSOC_ - 258 and 232   REFERENCES FOR LARVAE ENVIRONMENTAL ASSOC_ - 258 and 232   REFERENCES FOR EGG ENVIRONMENTAL ASSOC_ - 258 and 232   COMMENTS ON ENVIRONMENTAL ASSOCIATIONS - The capelin is a marine fish of cold, deep water, found in the Atlantic Ocean on the offshore banks and in coastal areas, occasionally spending the winter and early spring months in deep bays off the east coast of Newfoundland. It is a plentiful species off the north shore of the Gulf of St. Lawrence but the largest concentrations in Canadian waters are found off Newfoundland and the Labrador coast. The presence of a spawning population on the Southeast Shoal of the Grand Bank was noted. This population may remain in deep water in this region throughout life, moving close to the bank to spawn *258*.   An intensive migration inshore by coastal population takes place prior to spawning activities on beaches. This inshore movement was noted in a study of the Atlantic cod off Newfoundland, which reported that tagged Atlantic cod migrated inshore following their preferred prey, the capelin *258*.   Water Depth: Pacific capelin have been collected at depths of 65-75 m in the fall and that Atlantic capelin inhabit depths to 200 m. Studies in eastern Canada and Norway have shown that capelin move from deeper water during the day to shallower water at night. Capelin in Canada disperse in the upper 20 m of water during the night and school mostly above 50 m but sometimes as deep as 100 m during the day. In Norway, capelin moved from 150 m depth during the day day to between 50 and 100 m at night *232*.   COMMENTS ON ADULT ENVIRONMENTAL ASSOC_ - Adults are pelagic fish *232*   Adult capelin have been trawled at depths of 70 m at a temperature of -1.7 to 1.3 oC in the fall *232*.   COMMENTS ON BREEDING ADULT ENVIRONMENTAL ASSOC_ - On the south and east coasts of Newfoundland and Labrador spawning begins around the first part of June and may continue through July depending on tides, winds, and water temperatures. Beach-spawning capelin spawn on coarse sand or fine gravel, where the eggs are buried by wave action and are presumably safe from flushing by tidal exchange and from predation while development takes place. Substrate characteristics of capelin-spawning beaches are quite specific. A study noted that the preferred pebble diameter at Holyrood Beach was 5-15 mm. At Bryant's Cove, NFLD, a study observed spawning occurred on beaches where pebble size ranged from 1 to 4 mm and during ebbing tide at or near the period of maximum tidal oscillation. Surface water temperature at Holyrood, Chapel's cove, and Fogo on the east coast of Newfoundland ranged from 5.6 to 7.2 oC,   Environment Associations - 3 (DRAFT) - Environment Associations Species capelin Species Id M010068 Date 27 AUG 96   averaging 6.2 oC. Capelin on the south coast at Grand Beach spawned at higher temperatures of 6.2-8.4 oC, averaging 7.3 oC. Six spawning periods were observed in eastern and southern localities. Water temperature was 6.5-10 oC in 1978 and 2.5-10.8 oC in 1979. Not all capelin spawn on beaches. A study docemented significant egg deposition on the bottom at depths of 2-3 m at Bryant's Cove, in 1978, 1979, and 1980. Offshore spawning also occurred on the Southeast Shoal of the Grand Bank in depths of 46-49 m. Bottom water temperatures in the sampling areas were 2.8-4.2 oC during 1950 and 1951. The sandy substrate, on which eggs were deposited, consisted of grains 0.5-2.2 mm in diameter, smaller than the selected pebble size on spawning beaches. Capelin in spawning condition occur offshore on various banks in depths up to 80 m *258*.   A study reported that mating on beaches was most intensive during periods of intermediate tide, when gravid females and ripe males began the spawning act. Rolling and swimming inshore near the crest of the waves brought the spawning fish onto the beaches where the eggs were deposited and fertilized on the substrate. Spent fish, sometimes stranded, appeared immobile for a short period before re-entering the water with the following waves *258*.   Capelin in the Canadian arctic have been reported spawning in warm, brackish water. Onshore winds initiate capelin spawning in Newfoundland, because the winds cause the nearshore waters to be warmer and less saline. Offshore spawning has been reported to depths of 280 m but usually occurs in the upper 75 m of the water column. Beach-spawning capelin found along the Newfoundland and Labrador coasts in June and July generally spawned in water temperatures of 5.5 to 8.5 oC, although spawning has been reported in temperatures as high as 10 oC. In the Canadian arctic near Baffin Island, the surface water temperatures during the spawning period varied from about 13 to 15 oC, while temperatures in southern British Columbia are about 10 to 12.5 oC. Water temperatures at the time of spawning of Bering Sea capelin are between 5.3 and 10.5 oC. Temperatures for offshore spawning range from 2 to 4.2 oC. Capelins spawn either offshore in deep water or on sand and gravel beaches *232*.   COMMENTS ON LARVAE ENVIRONMENTAL ASSOC_ - Hatching took place in the beach sediments averaging 9-11 d in high tide zone and 22-24 d in low-tide zone, the hatching inversely related to incubation temperature. The specific relationship between time to median hatch (Y) and average incubation temperature (X) is lnY= 4.27 - 0.63 ln(X+1) , r2 = 0.85, p > 0.001 *258*.   The emergence of capelin larvae from the beach gravel, and the onset of larval drift, is episodic and closely correlated with the occurrence of onshore winds. Onshore winds drive warmer, food-rich, predator-poor surface waters into the nearshore waters. Larvae emerge actively in response to the sharp temperature increases caused by this water mass exchange and, thus, become associated with a favorable predator/prey field. Onshore water mass exchange occurs synchronously over large areas of Newfoundland's east coast in response to largescale atmospheic systems. Larval emergence is, therefore, synchronous over large geographical areas *258*.   While in the gravel larvae exist on yolk-sac reserves. Time to complete Environment Associations - 4 (DRAFT) -   Environment Associations Species capelin Species Id M010068 Date 27 AUG 96   yolk-sac use varies from 3 to 8 d depending on temperatures in the gravel. When beach-residence times exceed time to yolk-sac absorption, larval condition declines rapidly and survival is poor. Approximately 38 percent of the annual variability in year-class strength of the eastern Newfoundland capelin stocks is attributable to wind-regulated variance in beach-residence time. Wind conditions during beach-residence together with sea temperatures during larval drift account for approximately 60 percent of year-to-year variance in year-class strength *258*.   Dispersal of the larvae is initially passive, but is later moderated by vertical migrations which bring the larvae in contact with different current regimes. Larval dispersal, followed by the wanderings of juveniles in search of food, forms the migratory pattern of this species in early life, bringing them inshore and near the surface in early summer and offshore into deeper waters in autumn *258*.   Capelin larvae have been found over a wide range of salinities from 4.8 to 32.6 part per thousand, generally moving to cooler, less saline water below the thermocline in the fall. Capelin larvae are mainly concentrated in the upper 50 m during the summer in eastern Canada and move to slightly deeper water in September as the surface waters cool. Yolk-sac larvae were most abundant in the upper 25 m of the water column. No evidence of vertical migration of the capelin larvae was seen during the summer months. Capelin larvae in northeastern Canada were collected at temperatures ranging from 0.2 to 14.4 oC. Larval capelin are pelagic fish *232*.   COMMENTS ON EGG ENVIRONMENTAL ASSOC - Beach-spawning capelin spawn on coarse sand or fine gravel, where the eggs are buried by wave action and are presumably safe from flushing by tidal exchange and from predation while development takes place. Substrate characteristics of capelin-spawning beaches are quite specific. A study noted that the preferred pebble diameter at Holyrood Beach was 5-15 mm. At Bryant's Cove, NFLD, a study observed spawning occurred on beaches where pebble size ranged from 1 to 4 mm and during ebbing tide at or near the period of maximum tidal oscillation. The sandy substrate, on which eggs were deposited, consisted of grains 0.5-2.2 mm in diameter, smaller than the selected pebble size on spawning beaches *258*.   Rolling and swimming inshore near the crest of the waves brought the spawning fish onto the beaches where the eggs were deposited and fertilized on the substrate. Eggs, which are spherical, demersal, and adhesive, can be buried 15 cm or more beneath the surface of the beach. They become attached to beach gravel or to bottom substrate, where they develop and hatch *258*.   The specificity of beach spawning sites used by capelin results in dense concentrations of eggs (>800/cm2) in the beach gravel. Studies concluded that mortalities vary annually with changes in biological, meteorological, and hydrological conditions. Lack of required oxygen resulting from egg density, water and air temperature effects, substrate characteristics, amount of rainfall, and accumulation of excretory products are hypothesized to affect egg development and egg mortality *258*.   Hatching took place in the beach sediments averaging 9-11 d in high tide   Environment Associations - 5 (DRAFT) - Environment Associations Species capelin Species Id M010068 Date 27 AUG 96   zone and 22-24 d in low-tide zone, the hatching inversely related to incubation temperature. The specific relationship between time to median hatch (Y) and average incubation temperature (X) is lnY= 4.27 - 0.63 ln(X+1) , r2 = 0.85, p > 0.001. Incubation temperature during spawning periods in June and July in 1978 and 1979 at Bryant's Cove was in the range 1-19 oC with an average temperature of approximately 10 oC *258*.   The eggs are covered with sand or gravel by tidal action. The eggs are approximately 1 mm indiameter and stick to the substrate. Hatching occurs in 55 days at 0 oC, 30 days at 5 oC, and 15 days at 10 oC *232*.   Environment Associations - 6 (DRAFT) - Life History Species capelin Species Id M010068 Date 27 AUG 96 LIFE HISTORY Feeding is seasonal, intensifying in later winter and early spring in the prespawning period, declining as the spawning season approaches, and virtually ceasing during spawning. Several weeks after the spawning period, surviving capelin commence feeding and continue until cessation in early winter *258*. Reproduction: Spawning begins in the 3rd year of life, mass spawning taking place when fish are 3-4 yr old. On the south and east coasts of Newfoundland and Labrador spawning begins around the first part of June and may continue through July depending on tides, winds, and water temperatures. Late spawning is not unusual; in some years observed as late as 30 August. A study found a significant relationship between latitude and date of first spawning. Capelin may spawn more than once. Off the coast of the Gaspe Peninsula, Que, on the north shore of the Gulf of St. Lawrence and northeast shore of Cape Breton Island, spawning on beaches was observed June and July. A study cites evidence of repeated spawning by females *258*. Beach-spawning capelin spawn on coarse sand or fine gravel, where the eggs are buried by wave action and are presumably safe from flushing by tidal exchange and from predation while development takes place. Substrate characteristics of capelin-spawning beaches are quite specific. A study noted that the preferred pebble diameter at Holyrood Beach was 5-15 mm. At Bryant's Cove, NFLD, a study observed spawning occurred on beaches where pebble size ranged from 1 to 4 mm and during ebbing tide at or near the period of maximum tidal oscillation. Surface water temperature at Holyrood, Chapel's cove, and Fogo on the east coast of Newfoundland ranged from 5.6 to 7.2 oC, averaging 6.2 oC. Capelin on the south coast at Grand Beach spawned at higher temperatures of 6.2-8.4 oC, averaging 7.3 oC. Six spawning periods were observed in eastern and southern localities. A study noted that spawning occurred at two distinct time intervals during the study period at Bryant's Cove in 1978 and 1979. Fisrt spawning in the 1978 period took place 14-20 June, the second 5-9 July, 15 d later. In 1979 first spawning was observed 4-8 June and second 12 d later, 21 June. Water temperature was 6.5-10 oC in 1978 and 2.5-10.8 oC in 1979. Not all capelin spawn on beaches. A study docemented significant egg deposition on the bottom at depths of 2-3 m at Bryant's Cove, in 1978, 1979, and 1980. Offshore spawning also occurred on the Southeast Shoal of the Grand Bank in depths of 46-49 m. Bottom water temperatures in the sampling areas were 2.8-4.2 oC during 1950 and 1951. The sandy substrate, on which eggs were deposited, consisted of grains 0.5-2.2 mm in diameter, smaller than the selected pebble size on spawning beaches. Capelin in spawning condition occur offshore on various banks in depths up to 80 m *258*. A study reported that mating on beaches was most intensive during periods of intermediate tide, when gravid females and ripe males began the spawning act. Spawning took place at night or during times of heavy overcast and ceased during hours of sunlight. Rolling and swimming inshore near the crest of the waves brought the spawning fish onto the beaches where the eggs were deposited and fertilized on the substrate. A male would be observed to press against the side of the female, and on many occasions a second male would take position on the opposite side of the female. Eggs, which are Life History - 1 (DRAFT) - Life History Species capelin Species Id M010068 Date 27 AUG 96   spherical, demersal, and adhesive, can be buried 15 cm or more beneath the surface of the beach. They become attached to beach gravel or to bottom substrate, where they develop and hatch. Average diameter of 100 capelin eggs was 0.965 mm. Size of eggs in the ovary of a large female 25.2 cm TL and weighing 59 g, captured at Trinity Bay, NFLD, in March 1967, was 0.35-0.37 mm in diameter. The number of eggs increases with size of the female, a large female producing up to 50,000 eggs *258*.   Spent fish, sometimes stranded, appeared immobile for a short period before re-entering the water with the following waves. Mass mortalities occur, leading to the mistaken belief that capelin may spawn only once *258*.   The specificity of beach spawning sites used by capelin results in dense concentrations of eggs (>800/cm2) in the beach gravel. Studies concluded that mortalities vary annually with changes in biological, meteorological, and hydrological conditions. Lack of required oxygen resulting from egg density, water and air temperature effects, substrate characteristics, amount of rainfall, and accumulation of excretory products are hypothesized to affect egg development and egg mortality *258*.   Hatching took place in the beach sediments averaging 9-11 d in high tide zone and 22-24 d in low-tide zone, the hatching inversely related to incubation temperature. The specific relationship between time to median hatch (Y) and average incubation temperature (X) is lnY= 4.27 - 0.63 ln(X+1) , r2 = 0.85, p > 0.001. Incubation temperature during spawning periods in June and July in 1978 and 1979 at Bryant's Cove was in the range 1-19 oC with an average temperature of approximately 10 oC *258*.   The emergence of capelin larvae from the beach gravel, and the onset of larval drift, is episodic and closely correlated with the occurrence of onshore winds. Onshore winds drive warmer, food-rich, predator-poor surface waters into the nearshore waters. Larvae emerge actively in response to the sharp temperature increases caused by this water mass exchange and, thus, become associated with a favorable predator/prey field. Onshore water mass exchange occurs synchronously over large areas of Newfoundland's east coast in response to largescale atmospheric systems. Larval emergence is, therefore, synchronous over large geographical areas *258*.   While in the gravel larvae exist on yolk-sac reserves. Time to complete yolk-sac use varies from 3 to 8 d depending on temperatures in the gravel. When beach-residence times exceed time to yolk-sac absorption, larval condition declines rapidly and survival is poor. Approximately 38 percent of the annual variability in year-class strength of the eastern Newfoundland capelin stocks is attributable to wind-regulated variance in beach-residence time. Wind conditions during beach-residence together with sea temperatures during larval drift account for approximately 60 percent of year-to-year variance in year-class strength *258*.   Dispersal of the larvae is initially passive, but is later moderated by vertical migrations which bring the larvae in contact with different current regimes. Larval dispersal, followed by the wanderings of juveniles in search of food, forms the migratory pattern of this species in early life, bringing them inshore and near the surface in early summer and offshore into deeper waters in autumn *258*.   Life History - 2 (DRAFT) - Life History Species capelin Species Id M010068 Date 27 AUG 96   Growth: At age 1+, length of immature capelin taken from cod stomachs averages 86.6 mm. The estimates of capelin age, and hence growth, may be biased as a result of mistaken interpretation of a metamorphic check as a true annulus. Larval growth rates average 25 mm per day *258*.   Capelin exhibit sexual dimorphism in size. Males grow faster than females until they reach maturity, after which time rate of growth is approximately the same. There is a general north-south cline. Those from the Grand Bank and southern Newfoundland areas grow more quickly than those from the Labrador area, until similar maximum size is attained. Labrador capelin tend to mature 1 yr later than those from the Grand Bank *258*.   The majority of capelin do not live longer than 5 yr, but in Greenland, where growth rate is slower, 7-yr-old fish are known *258*.   Size of mature specimens is generally 13-20 cm in length. The largest capelin recorded was 5-yr-old, female, 25.2 cm TL, weighing 59 g, captured in Trinity Bay, NFLD *258*.   Description: Body slender, elongate, laterally compressed; males appear angular during breeding season because of enlarged lateral line scales. Head pointed; snout elongate; mouth terminal, oblique, lower jaw protruding, minute teeth on jaws, tongue, vomer, palatine, maxillary, and premaxillary; eye large. Gill rakers, 8-10 + 24-27, long, slender. Branchiostegals 8. Fins: soft rayed; dorsal 12-14, dorsal adipose present, low, long; caudal deeply forked; anal 19-21, low, base longer than dorsal, base elevated on a hump on males; pelvics 8, abdominal, no axillary process; pectorals 18-20, inserted just behind gill opening, elongated in males. Scales 175-205 in lateral line, cycloid, very small, deciduous; mature males with 4 rows of scales along lateral line forming prominent ridge at spawning time. Lateral line complete, straight. Pyloric caeca 5-8. Vertebrae 62-69. Size generally to 13-20 cm long. Color translucent, olive to bottle green above, sides silvery below lateral line, scale margins dotted with dusky specks, belly white *258*.   Predation and Competition: The role of capelin as food for other fishes, birds, and marine mammals is most significant and has been estimated to be in the order of 5 million t annually at virgin population levels. A study listed 10 fish species, 5 species of marine mammals, and 9 marine bird species known to forage extensively on capelin *258*.   Capelin are preyed upon heavily by the Atlantic cod, Gadus morhua, probably their chief predator. They are followed inshore during the spawning migration. During the period from June to early August 98 percent of the food of the Atlantic cod was capelin. Offshore Atlantic cod also feed heavily on this species. Capelin and their eggs were eaten extensively by haddock, Melanogrammus aeglefinus, on the Southeast Shoal of the Grand Bank during times when haddock were plentiful in the area. They are also consumed in large quantities by haddock on the spawning grounds of Norway and are the chief food of the Atlantic salmon, Salmo salar, in the northwest Atlantic *258*.   Life History - 3 (DRAFT) - Life History Species capelin Species Id M010068 Date 27 AUG 96   Juveniles and larvae often form part of the food of smaller fishes such as Atlantic herring, Clupea harengus harengus. Many other fishes, such as flounders, dogfish, sculpins, and eelpout, prey on them *258*.   Several species of whales, notably minke whales (Balaenoptera acutorostrata), feed intensively on capelin and they are frequently food of fin whales (B. physalus). Their role as food for other species extends to seals, especially the harp seal (Phoca groenlandica) but including ringed seal, and seabirds. A study reported the occurrence of capelin in the stomachs of thick-billed murres captured at Digges Sound, Hudson Bay. Winter flounder feed heavily and systematically on capelin eggs, supporting up to 25 percent of their annual somatic growth from this food source *258*.   Parasites and Disease: The capelin in Atlantic waters appears to be only lighly parasitized. A study listed one species of cestode, Eubothrium parvum; two species of nematodes, Anisakis sp. larva and Contracaecum sp.; and the protozoan Glugea sp., for which the capelin was described as a new host *258*.   A study redescribed and published new locality records of monogenean trematodes taken from the gills of capelin in the northwest Atlantic and Icelandic waters *258*.   Offshore Spawning: Offshore spawning has been reported to depths of 280 m but usually occurs in water less than 75 m deep. When waters became too warm on the beach, spawning continues in deeper coastal waters *232*.   Beach Spawning: Spawning occurs on sand, gravel, or pebble beaches. Spawning occurs at high tide and generally during periods of onshore winds for Atlantic capelin. However, capelin have been reported to spawn in offshore winds in the Kodiak area. Spawning usually occurs at night or during dull, cloudy weather; however, capelin spawn during daylight in the Canadian arctic when there are 24 hours of light *232*.   Reproductive Seasonality: Capelin spawn in May and June in the Kodiak area and in late spring in Bristol Bay. In the Bering Sea, they spawn in summer, slightly later near Pt. Barrow, and in the Beaufort Sea in August and September *232*.   Reproductive Behavior: At the spawning grounds, capelin segregate into schools by sex. Males school near the beaches, while the females remain in slightly deeper water. As the females ripen, individuals proceed to the beaches to spawn, followed by small groups of males. Generally, two males squeeze the female between them on the beach, extruding milt over eggs deposited in a slight hollow. The eggs are covered with sand or gravel by tidal action. Spawning occured for over four weeks in Norton Sound *232*.   Age at Sexual Maturity: During the first year, males and females are the same size, but during the second year, males grow at a faster rate. A few capelin mature and spawn at age 2, but most maturation occurs at age 3. Spawning capelin in Norton Sound and Togiak consist mainly of ages 2 and 3 fish; on Kodiak most were age 2 *232*.   Life History - 4 (DRAFT) - Life History Species capelin Species Id M010068 Date 27 AUG 96   Frequency of Breeding: During and following the spawning season, large numbers of dead capelin can be observed floating on the surface or stranded on the beach. The mortality rate of spawning capelin is high, and most investigators believe that postspawning survival is negligible *232*.   Fecundity: Fecundity in capelin varies widely with geographic location. Fecundity in Pacific capelin ranges from 15,300 to 39,300 eggs. Capelin in southern British Columbia, however, produce 4,600 to 6,700 eggs, and in the Sea of Japan fecundity ranges from 15,000 to 57,000 eggs. The eggs are approximately 1 mm in diameter and stick to the substrate *232*.   Incubation Period/Emergence: The incubation period in capelin is influenced by the average temperature, which varied between years and between positions on the beach. Incubation temperatures were determined by water temperature, air temperature, and hours of sunlight. Hatching occurs in 55 days at 0 oC, 30 days at 5 oC, and 15 days at 10 oC. Emergence of larvae from the beach is a clearly defined, abrupt transition to a pelagic existence. Atlantic capelin larvae accumulate on the beach as they hatch and are released synchronously by onshore winds. Because no food is available to the larvae while in the sediments, rapid physical deterioration occurs if larvae are unable to leave the beach before yolk sac reserves are depleted. Environmental conditions during the early larval period can influence growth efficiency, size at first feeding, and the prey-capture ability of larvae, independently of the yolk reserves at hatching or the food available *232*.   Movements Associated With Life Functions: Capelin in Newfoundland overwinter in large, inactive schools in cold water near the bottom at depths from 140 to 200 m. In the spring, these fish move into warming surface waters and disperse into smaller feeding schools. About a month before spawning, capelin migrate to shallow water and segregate into schools by sex, with the males closest to shore. During the summer and early fall, juvenile and adult Atlantic capelin were widely distributed in small schools. In November, the capelin concentrated into dense schools and moved to deeper water *232*.   Natural Factors Influencing Populations: Physical and biological factors such as the availabilty of food, water temperatures, and winds can contribute to the survival of emergent and larval capelin. On shore winds replace cold, food-poor, predator-laden upwelling waters, which occupy the nearshore areas during offshore winds, with warmer, food-rich, predator-deficient surface waters. Haddock and winter flounder consume large quantities of Atlantic capelin eggs during the spawning period. Egg mortality from predation may be significantly greater at deep-water dermersal spawning sites and may have a significant impact on recruitment of demersal spawners. Many fish are predators of adult and juvenile capelin, including cod and salmon in the Atlantic and arctic char, saffron cod, and starry flounder in the Canadian arctic. Marine mammals also consume capelin as a primary part of their diet. In the Gulf of Alaska, harbor seals and Steller sea lions are predators, especially in nearshore waters in the spring and summer. Other capelin predators include ribbon seals in the Pribilof Islands area, spotted seals in the southeastern Bering Sea in the spring, and belukha whales in the coastal northern Bering and Chukchi seas in summer. Competition between capelin and other zooplankton-feeding species may occur. In the Newfoundland area, the diets of capelin, arctic cod, and squid overlap. In years when capelin have reduced their zooplankton consumption, the abundance of arctic cod and squid has increased *232*.   Life History - 5 (DRAFT) - Life History Species capelin Species Id M010068 Date 27 AUG 96   Human-related Factors Influencing Populations: Capelin have adhesive, demersal eggs, and any contamination of the beach or offshore substrate when caplein spawn could result in heavy mortality. Juvenile and adult capelin are pelagic, relying on planktonic food sources, and thus are vulnerable to contamination of surface waters. A summary of possible impacts from human-related activities includes: changes in biological oxygen demand or nutrient loading; changes in the chemical composition of water; changes in dissolved oxygen, temperature, pH, or salinity; changes in sedimentation rate, turbidity, or suspended solids; changes in substrate composition and location; increased susceptibility to harvest or predation; and shock waves, blasting *232*.   LIFE HISTORY CODES - Breeding/Spawning Season: June Breeding/Spawning Season: July Breeding/Spawning Season: August Spawning Site: Sand Spawning Site: Gravel Periodicity: Active at night Average Number of Offspring/Reproductive Effort: Grea Gestation/Incubation Period: 1-2 weeks Gestation/Incubation Period: 3-4 weeks Periodicity: Most active in winter Periodicity: Most active in early spring Dispersion: Clumped Foraging Strategy: Filtering Breeding/Spawning Season: May Breeding/Spawning Season: June Average Number of Offspring/Reproductive Effort: 1,00 Gestation/Incubation Period: 1-2 months Breeding/Spawning Season: September   REFERENCES FOR LIFE HISTORY- 258 and 232   Life History - 6 DRAFT) - Management Practices   Species capelin Species Id M010068 Date 27 AUG 96 MANAGEMENT PRACTICES RESULT MANAGEMENT PRACTICE Adverse Salinity alteration Beneficial Developing/maintaining suitable salinity Beneficial Developing/maintaining suitable pH Beneficial Controlling sedimentation Beneficial Controlling pollution [thermal, chemical, physical] Adverse Underwater explosions Beneficial Regulating harvest of species being described Beneficial Other management practices [specified in comments] Adverse Other management practices [specified in comments]   REFERENCES FOR BENEFICIAL MANAGEMENT PRACTICES - 232   REFERENCES FOR ADVERSE MANAGEMENT PRACTICES - 232   COMMENTS ON MANAGEMENT PRACTICES - Capelin have adhesive, demersal eggs, and any contamination of the beach or offshore substrate when caplein spawn could result in heavy mortality. Juvenile and adult capelin are pelagic, relying on planktonic food sources, and thus are vulnerable to contamination of surface waters. A summary of possible impacts from human-related activities includes: changes in biological oxygen demand or nutrient loading; changes in the chemical composition of water; changes in dissolved oxygen, temperature, pH, or salinity; changes in sedimentation rate, turbidity, or suspended solids; changes in substrate composition and location; increased susceptibility to harvest or predation; and shock waves, blasting *232*.   Management Practices - 1 (DRAFT) - References   Species capelin Species Id M010068 Date 27 AUG 96 References 258 * Scott, W.B., M.G. Scott. 1988. Atlantic Fishes of Canada. Canadian Bulletin of Fisheries and Aquatic Sciences (219) (ed.). University of Toronto Press Toronto, Canada:731. 232 * State of Alaska Department of Fish, and Game. 1986. Alaska Habitat Management Guide: Life Histories and Habitat Requirements of Fish and Wildlife. (ed.). Alaska Department of Fish and Game Juneau, Alaska:763. References - 1

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