Early life history studies of Yellowfin tuna, Thunnus albacares. I. Food selection of Yellowfin tuna, Thunnus albacares, larvae reared in the laboratory. II. Age validation and growth of Yellowfin tuna, Thunnus albacares, larvae reared in the laboratory

English: Food selection of first-feeding yellowfin tuna larvae was studied in the laboratory during October 1992. The larvae were hatched from eggs obtained by natural spawning of yellowfin adults held in sea pens adjacent to Ishigaki Island, Okinawa Prefecture, Japan. The larvae were fed mixed-prey assemblages consisting of size-graded wild zooplankton and cultured rotifers. Yellowfin larvae were found to be selective feeders during the first four days of feeding. Copepod nauplii dominated the diet numerically, by frequency of occurrence and by weight. The relative importance of juvenile and adult copepods (mostly cyclopoids) in the diet increased over the 4-day period. Rotifers, although they comprised 31 to 40 percent of the available forage, comprised less than 2.1 percent of the diet numerically. Prey selection indices were calculated taking into account the relative abundances of prey, the swimming speeds of yellowfin larvae and their prey, and the microscale influence of turbulence on encounter rates. Yellowfin selected for copepod nauplii and against rotifers, and consumed juvenile and adult copepods in proportion to their abundances. Yellowfin larvae may select copepod nauplii and cyclopoid juveniles and adults based on the size and discontinuous swimming motion of these prey. Rotifers may not have been selected because they were larger or because they exhibit a smooth swimming pattern. The best initial diet for the culture of yellowfin larvae may be copepod nauplii and cyclopoid juveniles and adults, due to the size, swimming motion, and nutritional content of these prey. If rotifers alone are fed to yellowfin larvae, the rotifers should be enriched with a nutritional supplement that is high in unsaturated fatty acids. Mouth size of yellowfin larvae increases rapidly within the first few days of feeding, which minimizes limitations on feeding due to prey size. Although yellowfin larvae initiate feeding on relatively small prey, they rapidly acquire the ability to add relatively large, rare prey items to the diet. This mode of feeding may be adaptive for the development of yellowfin larvae, which have high metabolic rates and live in warm mixed-layer habitats of the tropical and subtropical Pacific. Our analysis also indicates a strong potential for the influence of microscale turbulence on the feeding success of yellowfin larvae. --- Experiments designed to validate the periodicity of otolith increments and to examine growth rates of yellowfin tuna larvae were conducted at the Japan Sea-Farming Association’s (JASFA) Yaeyama Experimental Station, Ishigaki Island, Japan, in September 1992. Larvae were reared from eggs spawned by captive yellowfin enclosed in a sea pen in the bay adjacent to Yaeyama Station. Results indicate that the first increment is deposited within 12 hours of hatching in the otoliths of yellowfin larvae, and subsequent growth increments are formed dailyollowing the first 24 hours after hatching r larvae up to 16 days of age. Somatic and otolith gwth ras were examined and compared for yolksac a first-feeding larvae reared at constant water tempatures of 26�and 29°C. Despite the more rapid develo of larvae reared at 29°C, growth rates were nnificaifferent between the two treatments. Howeve to poor survival after the first four days, it was ssible to examine growth rates beyond the onset of first feeding, when growth differences may become more apparent. Somatic and otolith growth were also examined for larvae reared at ambient bay water temperatures during the first 24 days after hatching. timates of laboratory growth rates were come to previously reported values for laboratory-reared yelllarvae of a similar age range, but were lower than growth rates reported for field-collected larvae. The discrepancy between laboratory and field growth rates may be associated with suboptimal growth conditions in the laboratory. Spanish: Durante octubre de 1992 se estudió en el laboratorio la seleccalimento por larvaún aleta amarillmera alimentación. Las larvas provinieron de huevos obtenidosel desove natural de aletas amarillas adultos mantenidos en corrales marinos adyacentes a la Isla Ishigaki, Prefectura de Okinawa (Japón). Se alimentó a las larvas con presas mixtas de zooplancton silvestre clasificado por tamaño y rotíferos cultivados. Se descubrió que las larvas de aleta amarilla se alimentan de forma selectiva durante los cuatro primeros días de alimentación. Los nauplios de copépodo predominaron en la dieta en número, por frecuencia de ocurrencia y por peso. La importancia relativa de copépodos juveniles y adultos (principalmente ciclopoides) en la dieta aumentó en el transcurso del período de 4 días. Los rotíferos, pese a que formaban del 31 al 40% del alimento disponible, respondieron de menos del 2,1% de la dieta en número. Se calcularon índices de selección de presas tomando en cuenta la abundancia relativa de las presas, la velocidad de natación de las larvas de aleta amarilla y de sus presas, y la influencia a microescala de la turbulencia sobre las tasas de encuentro. Los aletas amarillas seleccionaron a favor de nauplios de copépodo y en contra de los rotíferos, y consumieron copépodos juveniles y adultos en proporción a su abundancia. Es posible que las larvas de aleta amarilla seleccionen nauplios de copépodo y ciclopoides juveniles y adultos con base en el tamaño y movimiento de natación discontinuo de estas presas. Es posible que no se hayan seleccionado los rotíferos a raíz de su mayor tamaño o su patrón continuo de natación. Es posible que la mejor dieta inicial para el cultivo de larvas de aleta amarilla sea nauplios de copépodo y ciclopoides juveniles y adultos, debido al tamaño, movimiento de natación, y contenido nutritivo de estas presas. Si se alimenta a las larvas de aleta amarilla con rotíferos solamente, se debería enriquecerlos con un suplemento nutritivo rico en ácidos grasos no saturados. El tamaño de la boca de las larvas de aleta amarilla aumenta rápidamente en los primeros pocos días de alimentación, reduciendo la limitación de la alimentación debida al tamaño de la presa. Pese a que las larvas de aleta amarilla inician su alimentación con presas relativamente pequeñas, se hacen rápidamente capaces de añadir presas relativamente grandes y poco comunes a la dieta. Este modo de alimentación podría ser adaptivo para el desarrollo de larvas de aleta amarilla, que tienen tasa metabólicas altas y viven en hábitats cálidos en la capa de mezcla en el Pacífico tropical y subtropical. Nuestro análisis indica también que la influencia de turbulencia a microescala es potencialmente importante para el éxito de la alimentación de las larvas de aleta amarilla. --- En septiembre de 1992 se realizaron en la Estación Experimental Yaeyama de la Japan Sea- Farming Association (JASFA) en la Isla Ishigaki (Japón) experimentos diseñados para validar la periodicidad de los incrementos en los otolitos y para examinar las tasas de crecimiento de las larvas de atún aleta amarilla. Se criaron las larvas de huevos puestos por aletas amarillas cautivos en un corral marino en la bahía adyacente a la Estación Yaeyama. Los resultados indican que el primer incremento es depositado menos de 12 horas después de la eclosión en los otolitos de las larvas de aleta amarilla, y que los incrementos de crecimiento subsiguientes son formados a diario a partir de las primeras 24 horas después de la eclosión en larvas de hasta 16 días de edad. Se examinaron y compararon las tasas de crecimiento somático y de los otolitos en larvas en las etapas de saco vitelino y de primera alimentación criadas en aguas de temperatura constante entre 26°C y 29°C. A pesar del desarrollo más rápido de las larvas criadas a 29°C, las tasas de crecimiento no fueron significativamente diferentes entre los dos tratamientos. Debido a la mala supervivencia a partir de los cuatro primeros días, no fue posibación, uando las diferencias en el crecimiento podrían hacerse más aparentes. Se examinó también el crecimiento somático y de los otolitos para larvas criadas en temperaturas de agua ambiental en la bahía durante los 24 días inmediatamente después de la eclosión. Nuestras estimaciones de las tasas de crecimiento en el laboratorio fueron comparables a valores reportados previamente para larvas de aleta amarilla de edades similares criadas en el laboratorio, pero más bajas que las tasas de crecimiento reportadas para larvas capturadas en el mar. La discrepancia entre las tasas de crecimiento en el laboratorio y el mar podría estar asociada con condiciones subóptimas de crecimiento en el lab

Comparative feeding ecology of two elasmobranch species, Squalus blainville and Scyliorhinus canicula, off the coast of Portugal

The small-spotted catshark (Scyliorhinus canicula) (Linnaeus, 1758) and the longnose spurdog (Squalus blainville) (Risso, 1826) are two species occurring in the European and western African continental shelves with a wide geographical distribution. In this study, the diet of S. blainville and S. canicula off the Portuguese western Atlantic coast was investigated in 2006 by collecting monthly samples of these two species from local fishing vessels. In the stomachs of both species, crustaceans and teleosts were the dominant prey items, and molluscs, polychaetes, echinoderms, and sipunculids were found in lower abundance. In S. canicula, urochordate and chondrichthyan species were also observed in stomachs and were classified as accidental prey items. Scyliorhinus canicula consumed a broader group of prey items than did S. blainville. A significant diet overlap was observed, despite both species occupying different depth ranges over the continental shelf. Scyliorhinus canicula exhibited a consistency in diet composition among seasons, sexes, and maturity stages. Nonetheless, for both adults and juveniles, an increase in relative abundance of teleosts in the diet was observed in the spring and summer. This study provides evidence of the importance of S. canicula and S. blainville as benthic and pelagic predators along the western Atlantic coast.

Ontogenetic and temporal variability in the fat content and fatty acid composition of Atlantic herring (Clupea harengus) from the Bay of Fundy, Canada

Atlantic herring (Clupea harengus) is an ecologically and economically valuable species in many food webs, yet surprisingly little is known about the variation in the nutritional quality of these fish. Atlantic herring collected from 2005 through 2008 from the Bay of Fundy, Canada, were examined for variability in their nutritional quality by using total lipid content (n=889) and fatty acid composition (n=551) as proxies for nutritional value. A significant positive relationship was found between fish length and total lipid content. Atlantic herring also had significantly different fatty acid signatures by age. Fish from 2005 had significantly lower total lipid content than fish from 2006 through 2008, and all years had significantly different fatty acid signatures. Summer fish were significantly fatter than winter fish and had significantly different fatty acid signatures. For all comparisons (ontogenetic, annual, and seasonal) percent concentrations of omega-3, -6, and long-chain monounsaturated fatty acids were the most important for distinguishing between the fatty acid signatures of fish. This study underscores the importance of quantifying variation in prey quality synoptically with prey quantity in food webs over ontogenetic and temporal scales when evaluating the effect of prey nutritional quality on predators and on modeling trophic dynamics.

Spatial and seasonal relationships between Pacific harbor seals (Phoca vitulina richardii) and their prey, at multiple scales

Knowing where pinnipeds forage is vital to managing and protecting their populations, and for assessing potential interactions with fisheries. We assessed the spatial relationship between the seasonal distribution of Pacific harbor seals (Phoca vitulina richardii) outfitted with satellite transmitters and the seasonal distributions of potential harbor seal prey species in San Francisco Bay, California. Pearson’s correlation coefficients were calculated between the number of harbor seal locations in an area of the San Francisco Bay and the abundance of specific prey species in the same area. The influence of scale on the analyses was assessed by varying the scale of analysis from 1 to 10 km. There was consistency in the prey species targeted by harbor seals year-round, although there were seasonal differences between the most important prey species. The highest correlations between harbor seals and their prey were found for seasonally abundant benthic species, located within about 10 km of the primary haul-out site. Probable foraging habitat for harbor seals was identified, based on areas with high abundances of prey species that were strongly correlated with harbor seal distribution. With comparable local data inputs, this approach has potential application to pinniped management in other areas, and to decisions about the location of marine reserves designed to protect these species.

Reconstruction of original body size and estimation of allometric relationships for the longfin inshore squid (Loligo pealeii) and northern shortfin squid (Illex illecebrosus)

Quantification of predator-prey body size relationships is essential to understanding trophic dynamics in marine ecosystems. Prey lengths recovered from predator stomachs help determine the sizes of prey most influential in supporting predator growth and to ascertain size-specific effects of natural mortality on prey populations (Bax, 1998; Claessen et al., 2002). Estimating prey size from stomach content analyses is often hindered because of the degradation of tissue and bone by digestion. Furthermore, reconstruction of original prey size from digested remains requires species-specific reference materials and techniques. A number of diagnostic guides for freshwater (Hansel et al., 1988) and marine (Watt et al., 1997; Granadeiro and Silva, 2000) prey species exist; however they are limited to specific geographic regions (Smale et al., 1995; Gosztonyi et al., 2007). Predictive equations for reconstructing original prey size from diagnostic bones in marine fishes have been developed in several studies of piscivorous fishes of the Northwest Atlantic Ocean (Scharf et al., 1998; Wood, 2005). Conversely, morphometric relationships for cephalopods in this region are scarce despite their importance to a wide range of predators, such as finfish (Bowman et al., 2000 ; Staudinger, 2006), elasmobranchs (Kohler, 1987), and marine mammals (Gannon et al., 1997; Williams, 1999).

The trophic dynamics of summer flounder (Paralichthys dentatus) in Chesapeake Bay

Data on the trophic dynamics of fishes are needed for management of ecosystems such as Chesapeake Bay. Summer flounder (Paralichthys dentatus) are an abundant seasonal resident of the bay and have the potential to impact foodweb dynamics. Analyses of diet data for late juvenile and adult summer flounder collected from 2002−2006 in Chesapeake Bay were conducted to characterize the role of this flatfish in this estuary and to contribute to our understanding of summer flounder trophic dynamics throughout its range. Despite the diversity of prey, nearly half of the diet comprised mysid shrimp (Neomysis spp.) and bay anchovy (Anchoa mitchilli). Ontogenetic differences in diet and an increase in diet diversity with increasing fish size were documented. Temporal (inter- and intra-annual) changes were also detected, as well as trends in diet reflecting peaks in abundance and diversity of prey. The preponderance of fishes in the diet of summer flounder indicates that this species is an important piscivorous predator in Chesapeake Bay.

Diets of Steller sea lions (Eumetopias jubatus) in Southeast Alaska, 1993−1999

The diet of Steller sea lions (Eumetopias jubatus) was determined from 1494 scats (feces) collected at breeding (rookeries) and nonbreeding (haulout) sites in Southeast Alaska from 1993 to 1999. The most common prey of 61 species identified were walleye pollock (Theragra chalcogramma), Pacific herring (Clupea pallasii), Pacific sand lance (Ammodytes hexapterus), Pacific salmon (Salmonidae), arrowtooth flounder (Atheresthes stomias), rockfish (Sebastes spp.), skates (Rajidae), and cephalopods (squid and octopus). Steller sea lion diets at the three Southeast Alaska rookeries differed significantly from one another. The sea lions consumed the most diverse range of prey categories during summer, and the least diverse during fall. Diet was more diverse in Southeast Alaska during the 1990s than in any other region of Alaska (Gulf of Alaska and Aleutian Islands). Dietary differences between increasing and declining populations of Steller sea lions in Alaska correlate with rates of population change, and add credence to the view that diet may have played a role in the decline of sea lions in the Gulf of Alaska and Aleutian Islands.

Factors influencing cannibalism and abundance of walleye pollock (Theragra chalcogramma) on the eastern Bering Sea shelf, 1982–20

Cannibalism is thought to be an inf luential top-down process affecting walleye pollock (Theragra chalcogramma) recruitment in the eastern Bering Sea (EBS). In summer, many age-1 pollock occupy the same depths as those of adult walleye pollock, making them vulnerable to cannibalism. We examine factors that inf luence the occurrence and amount of cannibalism, as well as the abundance and co-occurrence of predator and prey walleye pollock. Large walleye pollock were generally found in deeper waters and avoided cold temperatures; whereas, age-1 walleye pollock were found in broader bottom depth and temperature ranges. The occurrence of cannibalism was highest in the area where predator and prey walleye pollock co-occurred and the amount of cannibalism was highest on the middle and outer EBS shelf. Both the occurrence and amount of cannibalism were influenced by location, bottom temperature and bottom depth, and the abundance of prey walleye pollock. The abundance of both large and small walleye pollock decreased during the 1982–2006 survey period in the EBS and, hence, the occurrence and amount of cannibalism also decreased. The occurrence and amount of cannibalism observed in the diet samples from the summer survey were good indicators of year class strength, as estimated by the stock assessment model. There was more cannibalism of age-1 walleye pollock when predicted recruit abundance was highest, indicating that summer cannibalism on age-1 walleye pollock, a top-down process, does not control walleye pollock recruitment in the EBS.

Estimating consumption rates of juvenile sandbar sharks (Carcharhinus plumbeus) in Chesapeake Bay, Virginia, using a bioenergetics model*

Using a bioenergetics model, we estimated daily ration and seasonal prey consumption rates for six age classes of juvenile sandbar sharks (Carcharhinus plumbeus) in the lower Chesapeake Bay summer nursery area. The model, incorporating habitat and species-specific data on growth rates, metabolic rate, diet composition, water temperature (range 16.8−27.9°C), and population structure, predicted mean daily rations between 2.17 ±0.03 (age-0) and 1.30 ±0.02 (age-5) % body mass/day. These daily rations are higher than earlier predictions for sandbar sharks but are comparable to those for ecologically similar shark species. The total nursery population of sandbar sharks was predicted to consume ~124,000 kg of prey during their 4.5 month stay in the Chesapeake Bay nursery. The predicted consumption rates support the conclusion that juvenile sandbar sharks exert a lesser top-down effect on the Chesapeake Bay ecosystem than do teleost piscivores and hu

Application of two methods for determining diet of northern fur seals (Callorhinus ursinus)

Examination of hard parts recovered from scats (feces) is currently the most common method for determining the diet of pinnipeds. However, large or sharp prey remains may be spewed (regurgitated) biasing prey composition and size estimations in diet studies based on scats. Percent frequency of occurrence (FO%) and age or size of selected prey remains recovered from northern fur seal (Callorhinus ursinus) scat (n=3444) and spew samples (n=267) collected from rookeries on St. George Island and St. Paul Island, Alaska, between 1990 and 2000 were compared to determine if a bias in prey composition and age or size estimations existed between scats and spews. Overall prey composition was similar between sample type and location, but the relative FO% of primary prey (≥5%) varied by sample type and location. Age or size estimates of walleye pollock (Theragra chalcogramma) and of two species of gonatid squids (Gonatopsis borealis and Berryteuthis magister) were significantly larger in spews than in scats. Observed differences in FO% and estimated age or size of prey species whose remains were found in scats and spews likely result from size-selective digestion of prey remains. Scats were biased toward smaller prey remains, whereas spews were biased toward larger prey remains and cephalopod beaks. The percent overlap between age classes of walleye pollock caught by the commercial trawl fishery and age classes of walleye pollock consumed by northern fur seals varied noticeably between sample types for both islands (scats: St. George=15. 5%; St. Paul=4.1%; spews: St. George=94.6%; St. Paul=89.6%). These results demonstrate that the inclusion of multiple sampling methods allows for a more accurate assessment of northern fur seal prey occurrence and prey age and size.

Estimating the encounter rate of Atlantic capelin (Mallotus villosus) with fish eggs, based on stomach content analysis

The number of pelagic fish eggs (cod and cunner) found in stomachs of capelin (Mallotus villosus) sampled in coastal Newfoundland was used to estimate the encounter rates between capelin and prey, and thus the effective volume swept by capelin. Fish eggs were found in 4−8% of capelin stomachs, represented an average of 1% of prey by numbers, and their abundance increased as relative stomach fullness decreased. The average number of eggs per stomach doubled for each 5-cm increase in length of capelin. The effective volume swept for eggs by capelin ranged from 0.04 to 0.84 m3/h—a rate that implies either very slow capelin swimming speeds (<1 cm/s) or that fish eggs are not strongly selected as prey. The predation rate estimated from stomach contents was higher than that predicted from laboratory studies of feeding pelagic fish and lower than that predicted by a simple foraging model. It remains uncertain whether capelin play an important regulatory role in the dynamics of early life stages of other fish.

The effect of predation (current and historical) by humpback whales (Megaptera novaeangliae) on fish abundance near Kodiak Island, Alaska

Humpback whales (Megaptera novaeangliae) are significant marine consumers. To examine the potential effect of predation by humpback whales, consumption (kg of prey daily) and prey removal (kg of prey annually) were modeled for a current and historic feeding aggregation of humpback whales off northeastern Kodiak Island, Alaska. A current prey biomass removal rate was modeled by using an estimate of the 2002 humpback whale abundance. A historic rate of removal was modeled from a prewhaling abundance estimate (population size prior to 1926). Two provisional humpback whale diets were simulated in order to model consumption rate. One diet was based on the stomach contents of whales that were commercially harvested from Port Hobron whaling station in Kodiak, Alaska, between 1926 and 1937, and the second diet, based on local prey availability as determined by fish surveys conducted within the study area, was used to model consumption rate by the historic population. The latter diet was also used to model consumption by the current population and to project a consumption rate if the current population were to grow to reach the historic population size. Models of these simulated diets showed that the current population likely removes nearly 8.83

Seasonal marine growth of Bristol Bay sockeye salmon (Oncorhynchus nerka) in relation to competition with Asian pink salmon (O. gorbuscha) and the 1977 ocean regime shift

Recent research demonstrated significantly lower growth and survival of Bristol Bay sockeye salmon (Oncorhynchus nerka) during odd-numbered years of their second or third years at sea (1975, 1977, etc.), a trend that was opposite that of Asian pink salmon (O. gorbuscha) abundance. Here we evaluated seasonal growth trends of Kvichak and Egegik river sockeye salmon (Bristol Bay stocks) during even- and odd-numbered years at sea by measuring scale circuli increments within each g rowth zone of each major salmon age group between 1955 and 2000. First year scale growth was not significantly different between odd- and even-numbered years, but peak growth of age-2 smolts was significantly higher than age-1. smolts. Total second and third year scale growth of salmon was significantly lower during odd- than during even-numbered years. However, reduced scale growth in odd-numbered years began after peak growth in spring and continued through summer and fall even though most pink salmon had left the high seas by late July (10−18% growth reduction in odd vs. even years). The alternating odd and even year growth pattern was consistent before and after the 1977 ocean reg ime shift. During 1977−2000, when salmon abundance was relatively great, sockeye salmon growth was high during specific seasons compared with that during 1955−1976, that is to say, immediately after entry to Bristol Bay, after peak growth in the first year, during the middle of the second growing season, and during spring of the third season. Growth after the spring peak in the third year at sea was relatively low during 1977−2000. We hypothesize that high consumption rates of prey by pink salmon during spring through mid-July of odd-numbered years, coupled with declining zooplankton biomass during summer and potentially cyclic abundances of squid and other prey, contributed to reduced prey availability and therefore reduced growth of Bristol Bay sockeye salmon during late spring through fall of odd-numbered years.

Using bone measurements to estimate the original sizes of bluefish (Pomatomus saltatrix) from digested remains

The ability to estimate the original size of an ingested prey item is an important step in understanding the community and population structure of piscivorous predators (Scharf et al., 1998). More specifically, knowledge of original prey size is essential for deriving important biological information, such as predator consumption rates, biomass of the prey consumed, and selectivity of a predator towards a specific size class of prey (Hansel et al., 1988; Scharf et al., 1997; Radke et al., 2000). To accurately assess the overall “top-down” pressure a predator may exert on prey community structure, prey size is crucial. However, such information is often difficult to collect in the field (Trippel and Beamish, 1987). Stomach-content analyses are the most common methods for examining the diets of piscivorous fish, but the prey items found are often thoroughly digested and sometimes unidentifiable. As a result, obtaining a direct measurement of prey items is frequently impossible.

Biological Characteristics and Fishery Assessment of Alaska Plaice, Pleuronectes quadrituberculatus, in the Eastern Bering Sea

Alaska plaice, Pleuronectes quadrituberculatus, is one of the major flatfishes in the eastern Bering Sea ecosystem and is most highly concentrated in the shallow continental shelf of the eastern Bering Sea. Annual commercial catches have ranged from less than 1,000 metric tons (t) in 1963 to 62,000 t in 1988. Alaska plaice is a relatively large flatfish averaging about 32 cm in length and 390 g in weight in commercial catches. They are distributed from nearshore waters to a depth of about 100 m in the eastern Bering Sea during summer, but move to deeper continental shelf waters in winter to escape sea ice and cold water temperatures. Being a long-lived species (>30 years), they have a relatively low natural mortality rate estimated at 0.20. Maturing at about age 7, Alaska plaice spawn from April through June on hard sandy substrates of the shelf region, primarily around the 100 m isobath. Prey items primarily include polychaetes and other marine worms. In comparison with other flatfish, Alaska plaice and rock sole, Pleuronectes bilineatus, have similar diets but different habitat preferences with separate areas of peak population density which may minimize interspecific competition. Yellowfin sole, Pleuronectes asper, while sharing similar habitat, differs from these two species because of the variety of prey items in its diet. Competition for food resources among the three species appears to be low. The resource has experienced light exploitation since 1963 and is currently in good condition. Based on the results of demersal trawl surveys and age-structured analyses, the exploitable biomass increased from 1971 through the mid-1980’s before decreasing to the 1997 level of 500,000 t. The recommended 1998 harvest level, Allowable Biological Catch, was calculated from the Baranov catch equation based on the FMSY harvest level and the projected 1997 biomass, resulting in a commercial harvest of 69,000 t, or about 16% of the estimated exploitable biomass.