Ecology of Juvenile Walleye Pollock, Theragra chalcogramma: Papers from the workshop "The Importance of Prerecruit Walleye Pollock to the Bering Sea and North Pacific Ecosystems" Seattle, Washington, 28-30 October 1993

The Alaska Fisheries Science Center (AFSC), National Marine Fisheries Service (NMFS), hosted an international workshop, 'The Importance of Prerecruit Walleye Pollock to the Bering Sea and North Pacific Ecosystems," from 28 to 30 October 1993. This workshop was held in conjunction with the annual International North Pacific Marine Science Organization (PICES) meeting held in Seattle. Nearly 100 representatives from government agencies, universities, and the fishing industry in Canada, Japan, the People's Republic of China, Russia, and the United States took part in the workshop to review and discuss current knowledge on juvenile pollock from the postlarval period to the time they recruit to the fisheries. In addition to its importance to humans as a major commercial species, pollock also serves as a major forage species for many marine fishes, birds, and mammals in the North Pacific region. (PDF file contains 236 pages.)

Regional variation in the annual feeding cycle of juvenile walleye pollock (Theragra chalcogramma) in the western Gulf of Alaska

Juvenile fish in temperate coastal oceans exhibit an annual cycle of feeding, and within this cycle, poor wintertime feeding can reduce body growth, condition, and perhaps survival, especially in food-poor areas. We examined the stomach contents of juvenile walleye pollock (Theragra chalcogramma) to explain previously observed seasonal and regional variation in juvenile body condition. Juvenile walleye pollock (1732 fish, 37–250 mm standard length) of the 2000 year class were collected from three regions in the Gulf of Alaska (Kodiak, Semidi, and Shumagin) representing an area of the continental shelf of ca. 100,000 km2 during four seasons (August 2000 to September 2001). Mean stomach content weight (SCW, 0.72% somatic body weight) decreased with fish body length except from winter to summer 2001. Euphausiids composed 61% of SCW and were the main determinant of seasonal change in the diets of fish in the Kodiak and Semidi regions. Before and during winter, SCW and the euphausiid dietary component were highest in the Kodiak region. Bioenergetics modeling indicated a relatively high growth rate for Kodiak juveniles during winter (0.33 mm standard length/d). After winter, Shumagin juveniles had relatively high SCW and, unlike the Kodiak and Semidi juveniles, exhibited no reduction in the euphausiid dietary component. These patterns explain previous seasonal and regional differences in body condition. We hypothesize that high-quality feeding locations (and perhaps nursery areas) shift seasonally in response to the availability of euphausiid

An allometric smoothing function to describe the relation between otolith and somatic growth over the lifespan of walleye pollock (Theragra chalcogramma)

We propose a new equation to describe the relation between otolith length (OL) and somatic length (fork length [FL]) of fish for the entire lifespan of the fish. The equation was developed by applying a mathematical smoothing method based on an allometric equation with a constant term for walleye pollock (Theragra chalcogramma) —a species that shows an extended longevity (>20 years). The most appropriate equation for defining the relation between OL and FL was a four-phase allometric smoothing function with three inflection points. The inflection points correspond to the timing of settlement of walleye pollock, changes in sexual maturity, and direction of otolith growth. Allometric smoothing functions describing the relation between short otolith radius and FL, long otolith radius and FL, and FL and body weight were also developed. The proposed allometric smoothing functions cover the entire lifespan of walleye pollock. We term these equations “allometric smoothing functions for otolith and somatic growth over the lifespan of walleye pollock.”

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.

Hatching date, nursery grounds, and early growth of juvenile walleye pollock (Theragra chalcogramma) off northern Japan*

Walleye pollock (Theragra chalcogramma) is widely distributed in the North Pacific Ocean and plays an important role in coastal subarctic ecosystems. The Japanese Pacific population of this species is one of the most important demersal fishes for commercial fisheries in northern Japan. The population is distributed along the Pacific coast of Hokkaido and the Tohoku area (Fig. 1), which is the southern limit of distribution of the species in the western North Pacific. In Funka Bay, the main spawning ground for this population, pollock spawn from December to March (Kendall and Nakatani, 1992). Planktonic eggs and larvae are transported into the bay, where juveniles usually remain until late July when they reach 60−85 mm in total length (Hayashi et al., 1968; Nakatani and Maeda, 1987). These juvenile pollock then migrate from Funka Bay eastward to the Doto area off southeastern Hokkaido (Honda et al., 2004). Many studies on eggs, larvae, and juveniles of the species have been conducted in or near Funka Bay, but little information is available on the ecology of the early life stages in the Tohoku area. Hashimoto and Ishito (1991) suggested that eggs are transported from Funka Bay southward to the Tohoku area by the coastal branch of the Oyashio Current, but there has been no study to verify this hypothesis.

Variation in the distribution of walleye pollock (Theragra chalcogramma) with temperature and implications for seasonal migration

Aspects of the feeding migration of walleye pollock (Theragra chalcogramma) in the eastern Bering Sea (EBS) were investigated by examining the relationship between temperatures and densities of fish encountered during acoustic and bottom trawl surveys conducted in spring and summer between 1982 and 2001. Bottom temperature was used as an indicator of spring and summer warming of the EBS. Clusters of survey stations were identified where the density of walleye pollock generally increased or decreased with increasing water temperature. Inferences about the direction and magnitude of the spring and summer feeding migration were made for five length categories of walleye pollock. Generally, feeding migrations appeared to be northward and shoreward, and the magnitude of this migration appeared to increase with walleye pollock size up to 50 cm. Pollock larger then 50 cm showed limited migratory behavior. Pollock may benefit from northward feeding migrations because of the changes in temperature, zooplankton production, and light conditions. Ongoing climate changes may affect pollock distribution and create new challenges for pollock management in the EBS.

Geographic variation among age-0 walleye pollock (Theragra chalcogramma): evidence of mesoscale variation in nursery quality?

Nurseries play an important part in the production of marine f ishes. Determining the relative importance of different nurseries in maintaining the parental population, however, can be difficult. In the western Gulf of Alaska, the Kodiak Island vicinity may be particularly well suited as a pollock nursery because of a prey-rich nearshore environment. Our objectives were 1) to examine age-0 pollock body condition, growth, and diet for evidence of a nearshore-shelf effect, and 2) to determine if variation in the potential prey field of zooplankton was associated with this effect. This was a pilot study that occurred in three bays and over the adjacent shelf off east Kodiak Island during 5−18 September 1993. Sampling occurred only during night at locations where echo sign indicated the presence of age-0 pollock. Echo sign was targeted to increase the chance of collecting fish given the limited vessel time. Fish condition was indicated by length-specific body weight. Growth rate indices were estimated for three different periods by using fish lengthage data and daily otolith increment widths: 1) from hatching date to capture, 2) 1−5 d before capture, and 3) 6−10 d before capture. Fish diet was determined from gut content analysis. Considerable variation among areas was evident in zooplankton composition, and fish condition, growth, and diet. However, relatively high prey densities, as well as fish condition and growth rates indicated that Chiniak Bay was particularly well suited as a pollock nursery. Hatching-date distributions indicated that most of the age-0 walleye pollock from bays were spawned earlier than were those from the shelf. The benefit of being reared in nearshore areas is therefore realized more by individuals that were spawned early than by individuals spawned relatively late.

Survey Assessment of Semi-pelagic Gadoids: The Example of Walleye Pollock, Theragra chalcogramma, in the Eastern Bering Sea

Assessment of walleye pollock, Theragra chalcogramma, in the eastern Bering Sea is complicated because the species is semi-pelagic in habit. Annual bottom trawl surveys provide estimates of demersal abundance on the eastern Bering Sea shelf. Every third year (starting in 1979), an extended area of the shelf and slope is surveyed and an echo integration-midwater trawl survey provides estimates of pollock abundance in midwater. Overall age-specific population and biomass estimates are obtained by summing the demersal and midwater results, assuming that the bottom trawl samples only pollock inhabiting the lower 3 m of the water column. Total population estimates have ranged from 134 x 109 fish in 1979 to 27 x 109 fish in 1988. The very high abundance observed in 1979 reflects the appearance of the unusually large 1978 year class. Changes in age-specific abundance estimates have documented the passage of strong (1978, 1982, and 1984) and weak year classes through the fishery. In general, older fish are more demersally oriented and younger fish are more abundant in midwater, but this trend was not always evident in the patterns of abundance of 1- and 2-year-old fish. As the average age of the population has increased, so has the relative proportion of pollock estimated by the demersal surveys. Consequently, it is unlikely that either technique can be used independently to monitor changes in abundance and age composition. Midwater assessment depends on pelagic trawl samples for size and age composition estimates, so both surveys are subject to biases resulting from gear performance and interactions between fish and gear. In this review, we discuss survey methodology and evaluate assumptions regarding catchability and availability as they relate to demersal, midwater, and overall assessment.

Using measurements of muscle cell nuclear RNA with flow cytometry to improve assessment of larval condition of Walleye Pollock (Gadus chalcogrammus)

Nuclear RNA and DNA in muscle cell nuclei of laboratory-reared larvae of Walleye Pollock (Gadus chalcogrammus) were simultaneously measured through the use of flow cytometry for cell-cycle analysis during 2009–11. The addition of nuclear RNA as a covariate increased by 4% the classification accuracy of a discriminant analysis model that used cell-cycle, temperature, and standard length to measure larval condition, compared with a model without it. The greatest improvement, a 7% increase in accuracy, was observed for small larvae (<6.00 mm). Nuclear RNA content varied with rearing temperature, increasing as temperature decreased. There was a loss of DNA when larvae were frozen and thawed because the percentage of cells in the DNA synthesis cell-cycle phase decreased, but DNA content was stable during storage of frozen tissue.

A method to improve size estimates of walleye pollock (Theragra chalcogramma) Atka mackerel (Pleurogrammus monopterygius) consumed by pinnipeds: digestion correction factors applied to bones and otoliths recovered in scats

The lengths of otoliths and other skeletal structures recovered from the scats of pinnipeds, such as Steller sea lions (Eumetopias jubatus), correlate with body size and can be used to estimate the length of prey consumed. Unfortunately, otoliths are often found in too few scats or are too digested to usefully estimate prey size. Alternative diagnostic bones are frequently recovered, but few bone-size to prey-size correlations exist and bones are also reduced in size by various degrees owing to digestion. To prevent underestimates in prey sizes consumed techniques are required to account for the degree of digestion of alternative bones prior to estimating prey size. We developed a method (using defined criteria and photo-reference material) to assign the degree of digestion for key cranial structures of two prey species: walleye pollock (Theragra chalcogramma) and Atka mackerel (Pleurogrammus monopterygius). The method grades each structure into one of three condition categories; good, fair or poor. We also conducted feeding trials with captive Steller sea lions, feeding both fish species to determine the extent of erosion of each structure and to derive condition-specific digestion correction factors to reconstruct the original sizes of the structures consumed. In general, larger structures were relatively more digested than smaller ones. Mean size reduction varied between different types of structures (3.3−26.3%), but was not influenced by the size of the prey consumed. Results from the observations and experiments were combined to be able to reconstruct the size of prey consumed by sea lions and other pinnipeds. The proposed method has four steps: 1) measure the recovered structures and grade the extent of digestion by using defined criteria and photo-reference collection; 2) exclude structures graded in poor condition; 3) multiply measurements of structures in good and fair condition by their appropriate digestion correction factors to derive their original size; and 4) calculate the size of prey from allometric regressions relating corrected structure measurements to body lengths. This technique can be readily applied to piscivore dietary studies that use hard remains of fish.

Sizes of walleye pollock (Theragra chalcogramma) and Atka mackerel (Pleurogrammus monopterygius) consumed by the western stock of Steller sea lions (Eumetopias jubatus) in Alaska from 1999 to 2000

Prey-size selectivity by Steller sea lions (Eumetopias jubatus) is relevant for understanding the foraging behavior of this declining predator, but studies have been problematic because of the absence and erosion of otoliths usually used to estimate fish length. Therefore, we developed regression formulae to estimate fish length from seven diagnostic cranial structures of walleye pollock (Theragra chalcogramma) and Atka mackerel (Pleurogrammus monopterygius). For both species, all structure measurements were related with fork length of prey (r2 range: 0.78−0.99). Fork length (FL) of walleye pollock and Atka mackerel consumed by Steller sea lions was estimated by applying these regression models to cranial structures recovered from scats (feces) collected between 1998 and 2000 across the range of the Alaskan western stock of Steller sea lions. Experimentally derived digestion correction factors were applied to take into account loss of size due to digestion. Fork lengths of walleye pollock consumed by Steller sea lions ranged from 3.7 to 70.8 cm (mean=39.3 cm, SD=14.3 cm, n=666) and Atka mackerel ranged from 15.3 to 49.6 cm (mean=32.3 cm, SD=5.9 cm, n=1685). Although sample sizes were limited, a greater proportion of juvenile (≤20 cm) walleye pollock were found in samples collected during the summer (June−September) on haul-out sites (64% juveniles, n=11 scats) than on summer rookeries (9% juveniles, n=132 scats) or winter (February−March) haul-out sites (3% juveniles, n=69 scats). Annual changes in the size of Atka mackerel consumed by Steller sea lions corresponded to changes in the length distribution of Atka mackerel resulting from exceptionally strong year classes. Considerable overlap (>51%) in the size of walleye pollock and Atka mackerel taken by Steller sea lions and the sizes of these species caught by the commercial trawl fishery were demonstrated.

Sizes of walleye pollock (Theragra chalcogramma) consumed by the eastern stock of Steller sea lions (Eumetopias jubatus) in Southeast Alaska from 1994 to 1999

Lengths of walleye pollock (Theragra chalcogramma) consumed by Steller sea lions (Eumetopias jubatus) were estimated by using allometric regressions applied to seven diagnostic cranial structures recovered from 531 scats collected in Southeast Alaska between 1994 and 1999. Only elements in good and fair condition were selected. Selected structural measurements were corrected for loss of size due to erosion by using experimentally derived condition-specific digestion correction factors. Correcting for digestion increased the estimated length of fish consumed by 23%, and the average mass of fish consumed by 88%. Mean corrected fork length (FL) of pollock consumed was 42.4 ±11.6 cm (range=10.0−78.1 cm, n=909). Adult pollock (FL>45.0 cm) occurred more frequently in scats collected from rookeries along the open ocean coastline of Southeast Alaska during June and July (74% adults, mean FL=48.4 cm) than they did in scats from haul-outs located in inside waters between October and May (51% adults, mean FL=38.4 cm). Overall, the contribution of juvenile pollock (≤20 cm) to the sea lion diet was insignificant; whereas adults contributed 44% to the diet by number and 74% by mass. On average, larger pollock were eaten in summer at rookeries throughout Southeast Alaska than at rookeries in the Gulf of Alaska and the Bering Sea. Overall it appears that Steller sea lions are capable of consuming a wide size range of pollock, and the bulk of fish fall between 20 and 60 cm. The use of cranial hard parts other than otoliths and the application of digestion correction factors are fundamental to correctly estimating the sizes of prey consumed by sea lions and determining the extent that these sizes overlap with the sizes of pollock caught by commercial fisheries.

Elemental signatures in otoliths of larval walleye pollock (Theragra chalcogramma) from the northeast Pacific Ocean

The population structure of walleye pollock (Theragra chalcogramma) in the northeastern Pacific Ocean remains unknown. We examined elemental signatures in the otoliths of larval and juvenile pollock from locations in the Bering Sea and Gulf of Alaska to determine if there were significant geographic variations in otolith composition that may be used as natural tags of population affinities. Otoliths were assayed by using both electron probe microanalysis (EPMA) and laser ablation inductively coupled plasma mass spectrometry (ICP-MS). Elements measured at the nucleus of otoliths by EPMA and laser ablation ICP-MS differed significantly among locations. However, geographic groupings identified by a multivariate statistical approach from EPMA and ICP-MS were dissimilar, indicating that the elements assayed by each technique were controlled by separate depositional processes within the endolymph. Elemental profiles across the pollock otoliths were generally consistent at distances up to 100 μm from the nucleus. At distances beyond 100 μm, profiles varied significantly but were remarkably consistent among individuals collected at each location. These data may indicate that larvae from various spawning locations are encountering water masses with differing physicochemical properties through their larval lives, and at approximately the same time. Although our results are promising, we require a better understanding of the mechanisms controlling otolith chemistry before it will be possible to reconstruct dispersal pathways of larval pollock based on probe-based analyses of otolith geochemistry. Elemental signatures in otoliths of pollock may allow for the delineation of fine-scale population structure in pollock that has yet to be consistently revealed by using population genetic approaches.

Changes over time in the spatial distribution of walleye pollock (Theragra chalcogramma) in the Gulf of Alaska, 1984-1996

Triennial bottom trawl survey data from 1984 to 1996 were used to evaluate changes in the summer distribution of walleye pollock in the western and central Gulf of Alaska. Differences between several age groups of pollock were evaluated. Distribution was examined in relation to several physical characteristics, including bottom depth and distance from land. Interspecies associations were also analyzed with the Bray-Curtis clustering technique to better understand community structure. Our results indicated that although the population numbers decreased, high concentrations of pollock remained in the same areas during 1984–96. However, there was an increase in the number of stations where low-density pollock concentrations of all ages were observed, which resulted in a decrease in mean population density of pollock within the GOA region. Patterns emerging from our data suggested an alternative to Mac-Call’s “basin hypothesis” which states that as population numbers decrease, there should be a contraction of the population range to optimal habitats. During 1984–96 there was a concurrent precipitous decline in Steller sea lions in the Gulf of Alaska. The results of our study suggest that decreases in the mean density of adult pollock, the main food in the Steller sea lion diet, combined with slight changes in the distribution of pollock (age-1 pollock in particular) in the mid-1980s, may have contributed to decreased foraging efficiency in Steller sea lions. Our results support the prevailing conceptual model for pollock ontogeny, although there is evidence that substantial spawning may also occur outside of Shelikof Strait.

An examination of spatial and temporal genetic variation in walleye pollock (Theragra chalcogramma) using allozyme, mitochondrial DNA, and microsatellite data

We used allozyme, microsatellite, and mitochondrial DNA (mtDNA) data to test for spatial and interannual genetic diversity in wall-eye pollock (Theragra chalcogramma) from six spawning aggregations representing three geographic regions: Gulf of Alaska, eastern Bering Sea, and eastern Kamchatka. Interpopulation genetic diversity was evident primarily from the mtDNA and two allozyme loci (SOD-2*, MPI*). Permutation tests ˆindicated that FST values for most allozyme and microsatellite loci were not significantly greater than zero. The microsatellite results suggested that high locus polymorphism may not be a reliable indicator of power for detecting population differentiation in walleye pollock. The fact that mtDNA revealed population structure and most nuclear loci did not suggests that the effective size of most walleye pollock populations is large (genetic drift is weak) and migration is a relatively strong homogenizing force. The allozymes and mtDNA provided mostly concordant estimates of patterns of spatial genetic variation. These data showed significant genetic variation between North American and Asian populations. In addition, two spawning aggregations in the Gulf of Alaska, in Prince William Sound, and off Middleton Island, appeared genetically distinct from walleye pollock spawning in the Shelikof Strait and may merit management as a distinct stock. Finally, we found evidence of interannual genetic variation in two of three North American spawning aggregations, similar in magnitude to the spatial variation among North American walleye pol-lock. We suggest that interannual genetic variation in walleye pollock may be indicative of one or more of the following factors: highly variable reproductive success, adult philopatry, source-sink metapopulation structure, and intraannual variation (days) in spawning timing among genetically distinct but spatially identical spawning aggregates.