Anomalous transport of walleye pollock larvae linked to ocean and atmospheric patterns in May 1996

Larvae from a large aggregation of walleye pollock spawning in early spring in Shelikof Strait, Gulf of Alaska, are normally transported to the south-west in the vigorous Alaska Coastal Current. In the spring of 1996, anomalous winds resulted in unusually weak transport in the Shelikof Strait sea valley. The main aggregation of larval pollock in the Shelikof region was surveyed four times in 1996 over a period of about 40 days, including finer-scale sampling of the leading south-western edge of the larval distribution. The south-western edge of the larval distribution showed weak transport up the sea valley for a period of about 10 days, corresponding to the observations of currents, after which many larvae were transported over the shelf region to the west. These observations are unique in over 15 years of monitoring larval transport patterns and demonstrate how anomalous weather, and hence current patterns, influence variability in larval transport.     

Spatial and temporal patterns of walleye pollock (Theragra chalcogramma) spawning in the eastern Bering Sea inferred from egg and larval distributions

Walleye pollock Theragra chalcogramma (pollock hereafter) is a key ecological and economic species in the eastern Bering Sea, yet detailed synthesis of the spatial and temporal patterns of pollock ichthyoplankton in this important region is lacking. This knowledge gap is particularly severe considering that egg and larval distribution are essential to reconstructing spawning locations and early life stages drift pathways. We used 19 yr of ichthyoplankton collections to determine the spatial and temporal patterns of egg and larval distribution. Generalized additive models (GAMs) identified two primary temporal pulses of pollock eggs, the first occurring from 20 February to 31 March and the second from 20 April to 20 May; larvae showed similar, but slightly lagged, pulses. Based on generalized cross‐validation and information theory, a GAM model that allowed for different seasonal patterns in egg density within three unique areas outperformed a GAM that assumed a single fixed seasonal pattern across the entire eastern Bering Sea. This ‘area‐dependent’ GAM predicted the highest densities of eggs (i.e., potential spawning locations) in three major areas of the eastern Bering Sea: near Bogoslof Island (February–April), north of Unimak Island and the Alaska Peninsula (March–April), and around the Pribilof Islands (April–August). Unique temporal patterns of egg density were observed for each area, suggesting that pollock spawning may be more spatially and temporally complex than previously assumed. Moreover, this work provides a valuable baseline of pollock spawning to which future changes, such as those resulting from climate variability, may be compared.     

Walleye pollock recruitment in Shelikof Strait: applied fisheries oceanography

Fisheries-Oceanography Coordinated Investigations (FOCI) is a National Oceanic and Atmospheric Administration (NOAA) research programme seeking to understand recruitment processes of commercially exploited Alaskan fishes. The FOCI is mainly comprised of scientists at the Pacific Marine Environmental Laboratory and the Alaska Fisheries Science Center who study both the biotic and abiotic environment, including processes within larval patches through integrated field, laboratory, and modelling studies. The initial focus of studies was walleye pollock ( Theragra chakogramma ) spawning in Shelikof Strait, Gulf of Alaska. The choice of this population for our research was based on development of a large fishery and the substantial variation in recruitment that was observed in the late 1970s and early 1980s. Also, the early life history of this population is quite predictable and restricted both temporally and spatially. Walleye pollock spawn consistently in a small part of Shelikof Strait in early spring from which a large patch of eggs and later larvae is produced. In most years this concentration of larvae drifts to the south-west through the strait during April and May. Large numbers of larvae are often found in eddies which frequent the area and we have observed improved feeding conditions for larvae, in as opposed to out of eddies. We have found that first-feeding larvae have higher survival rates during calm periods, rather than in storms, and that in many years recruitment is largely set by the end of the larval period, although in some years age-0 juvenile mortality is also important. FOCI now generates information that is being used for management of this resource.     

A review: Walleye pollock in the North Pacific–how much difference do they really make?

Populations of several species of marine birds and mammals in the Bering Sea and Gulf of Alaska have been declining since the mid-1970s, with numbers of one, the Steller sea lion (Eumetopias jubatus) , so depressed it was listed as threatened under the Endangered Species Act in spring 1990. All of the declining populations depend to an important extent on walleye pollock (Theragra chakogramma) for food, although they eat numerous other species as well. In contrast, certain animals that compete with pollock for common prey have been increasing in abundance. All of these changes could be related through food web connections mediated by pollock. Pollock is also important to people–it presently supports the largest single-species commercial fishery in the world, in large part because of its great biomass, which has averaged about 15 × 10⁶t in the Bering Sea over the past 15 years. Pollock consume an inordinate proportion of the pelagic production in the Bering Sea, which further supports the conclusion that it is a key species in the ecosystem. However, there are conflicting hypotheses about the importance of the roles played by pollock as predator and prey, and about the effect that changes in pollock abundance might have on biomass yield at higher trophic levels.     

Variations in larval walleye pollock feeding and condition: a synthesis

There was a strong association among concentrations of microzooplankton prey sampled from the walleye pollock, Theragra chalcogramma , larval habitat, gut contents of larvae, and their nutritional condition. Subsequently, hypothesized survival potentials linked to food availability were validated by independently determined mortality rates. We present evidence that a significant number of walleye pollock larvae were starving in 1991 but that fewer were starving in 1992. At some stations where prey levels were anomalously low in 1991, up to 40% of the larvae were in poor condition. There appears to be a 2-week period after first feeding when walleye pollock are vulnerable to starvation.     

Observation and analysis of fishery processes: larval pollock at Shelikof Strait, Alaska

Conditions affecting distributions of larval walleye pollock ( Theragra chalcogramma ) were examined at Shelikof Strait, Alaska, during springtime, 1986 and 1987. Abundance and distribution of larval pollock southwest of the Strait's southern entrance was determined with oblique plankton tows taken each year in May. Infrared images of sea surface temperature patterns were derived from AVHRR scenes obtained by NOAA satellites during each April and May. Pattern displacements between 24-hour-interval images were used to estimate surface motion. Each spring, measurements were taken by remote weather stations and ships, and a nearsurface current meter record was obtained during 1987. Treated as quasi-synoptic, spatial relations between sets of surface temperature, surface flow, and larval pollock distributions show coincidences between submesoscale physical and biological features. The highest larval abundances occurred as patches within a cold plume (1986) and an eddy (1987). These confirm that physical features can retain larval pollock on the continental shelf. Observations are examined for evidence of physical and biological events that jointly can cause such coincidences and foster alternatives for survival during transport to nursery grounds. Explanations for presence of cohorts observed within the 1987 eddy are given in terms of spatial and temporal relationships evident between spawning and hatching areas, hatch date distributions, meanders, eddy generation and movement, background flow, and advection times. The observations, analyses, and results are consistent with the concept of a coupled, fluctuating biophysical process that can emulate variations in larval abundance and provide a multiplicity of system pathways for early-life stages representations.     

Use of juvenile salmon growth and temperature change indices to predict groundfish post age‐0 yr class strengths in the Gulf of Alaska and eastern Bering Sea

Juvenile marine growth (SW1) of salmon and a new temperature change (TC) index were evaluated as ecosystem indicators and predictors for the post age‐0 year class strength (YCS) of groundfish in the Gulf of Alaska (GOA) and eastern Bering Sea (EBS). Our hypothesis was that SW1, as measured on the scales of adult Pacific salmon (Oncorhynchus spp.), is a proxy for ocean productivity on the continental shelf, a rearing area for young salmon and groundfish. Less negative TC index values are the result of a cool late summer followed by a warm spring, conditions favorable for groundfish YCS. In the GOA, SW1 was a positive predictor of age‐1 pollock (Theragra chalcogramma), but not age‐2 sablefish (Anoplopoma fimbria) YCS, indicating that the growth of the Karluk River sockeye salmon that enter Shelikof Strait is a proxy for ocean conditions experienced by age‐0 pollock. Contrary to our hypotheses, the TC index was a negative predictor of GOA pollock YCS; and the SW1 a negative predictor of EBS pollock and cod YCS since the 1980s. Recent fisheries oceanography survey results provide insight into possible mechanisms to support the inverse SW1 and YCS relationship. For the EBS, the TC index was a significant positive predictor for pollock and cod YCS, supporting the hypothesis that a cool late summer followed by a warm spring maximizes the over‐wintering survival of pollock and cod (Gadus macrocephalus), especially since the 1980s. The TC and SW1 index showed value for the assessment of pollock and cod, but not sablefish.     

Seasonal changes in pelagic fish biomass around the Chiswell Island Steller sea lion rookery in 2003

Fisheries acoustics surveys were conducted around the Chiswell Island rookery in the northern Gulf of Alaska at night in April and August 2003 to assess seasonal changes in prey available to Steller sea lions (Eumetopias jubatus) foraging around the rookery. Adult walleye pollock (Theragra chalcogramma) ≥28 cm fork length was the dominant biomass in the upper 50 m of the water column in both months, increasing from 122.8 kg/nmi² in April to 457.9 kg/nmi² in August. A similar pattern was observed for Pacific herring (Clupea pallasii), which averaged 2.8 and 65.6 kg/nmi² in April and August, respectively. Incidental trawl catch suggested the appearance of age-0 pollock and juvenile salmonids (Oncorhynchus spp.) around the rookery in August as well. The increased biomass of these key prey species is linked to increased foraging trip durations by lactating sea lions from Chiswell Island, and supports the general view that sea lions in the northern Gulf of Alaska are not food limited during summer months.     

A review of the distribution, ecology and population dynamics of age-0 walleye pollock in the Gulf of Alaska

This review paper synthesizes published research and unpublished data on the abundance and distribution patterns, ecology and population dynamics of walleye pollock ( Theragra chalcogramma ) during their first year of life (age-0) in the Gulf of Alaska. Distribution patterns have been described using mainly trawl catches, but recently, acoustic methodology has been employed, especially in examining vertical distributions. Although age-0 pollock are found throughout the Gulf, the highest catches occurred west of Kodiak Island. Pollock are pelagic for at least their first 6 months of life but show an ontogenetic increase in depth distribution superimposed on a pronounced diel vertical migration at a larger size. Daily growth rates are variable depending on year, season and area, and growth generally ceases during the winter. The diet of age-0 pollock shifts from mainly copepods in early juveniles to euphausiids by fall, with epibenthic organisms becoming important during the winter months. Feeding occurs mainly at night in surface waters. Age-0 pollock are most frequently associated with gelatinous zooplankton (medusae) and older pollock. Many predators on age-0 pollock have been identified; the most important are arrowtooth flounder ( Atheresthes stomias ), adult pollock, puffins ( Fratercula spp.), murres ( Una spp.), harbor seals ( Phoca vitulina richardsi ) and Steller sea lions ( Eumetopiasjubatus ). Modelling provides some insight into the population dynamics of these juveniles and environmental conditions which interannually affect their survival. These results are discussed relative to the importance of age-0 pollock in the recruitment of this species and to their role in the pelagic ecosystem.     

Cohort survival patterns of walleye pollock, Theragra chalcogramma, in Shelikof Strait, Alaska: a critical factor analysis

A series of age-specific life tables for walleye pollock ( Theragra chalcogramma ) in the western Gulf of Alaska was compiled for the 1980-91 year classes. The life tables were utilized to perform an exploratory key factor analysis to examine the timing of critical periods in the recruitment process, evidence of density-dependence at different stages and trends in mortality rates. Early larval mortality was significantly correlated with generational mortality (In recruits/spawning bio-mass), but patterns in juvenile mortality also were similar to generational mortality and in some years were clearly dominant in determining the fate of a cohort. Density-dependent mortality, based on the correlation between mortality and initial abundance, was indicated only for the late larval to early juvenile stage. Time trends were marginally significant for juvenile mortality. It is speculated that the observed increase in juvenile mortality is associated with increasing abundance of arrowtooth flounder. Weaknesses in the data base are discussed; these along with the short time series involved make our conclusions tentative and subject to further study. We hypothesize that pollock recruitment levels can be established at any life stage depending on sufficient supply from prior stages, a type of dynamics which can be termed supply dependent multiple life stage control.     

Predicting early life connectivity of Antarctic silverfish,an important forage species along the Antarctic Peninsula

The early life stages of the Antarctic silverfish (Pleuragramma antarctica), an important prey species for higher predators in the Southern Ocean ecosystem, dominate the larval fish assemblages of the Bransfield Strait, one of the most important areas for larval retention off the Antarctic Peninsula. Nevertheless, the spatial location of areas where they were spawned and the timing of larval hatching remain unknown. By linking Lagrangian particle tracking simulations with age data obtained using otolith microincrements from fish caught north of Joinville Island in a pelagic survey, we estimated the distribution of hatch dates and subsequent growth rates of silverfish reaching the Bransfield Strait, and predicted the areas where they were spawned. Larval hatching peaked during the last week of December, and the inner shelf and shelf break, east of the Larsen Ice shelf, were the dominant areas predicted to contribute to larval assemblages in the Bransfield Strait. Over simulated periods of 600–630 days, 35–40% of particles remained within the Bransfield Strait, suggesting an important source of supply to higher predators feeding off the northern Antarctic Peninsula. The daily growth rate at the mean size of 22.3 mm was 0.18 mm, corresponding to a daily change in size of approximately 0.82% standard length (SL), and large variability in growth rate suggested a wide range of environmental conditions experienced during the period of advection from the spawning areas. These results provide spatial predictions that can be tested empirically in future studies, using the simulated trajectories to inform sampling design and spatial coverage.     

Geographical variations in infection by larval Anisakis simplex and Contracaecum osculatum (Nematoda, Anisakidae) in walleye pollock Theragra chalcogramma stocks off Hokkaido, Japan

ABSTRACT: Stocks of walleye pollock Theragra chalcogramma collected from: (i) the Sea of Japan (off Rebun Island and Kumaishi); (ii) the Pacific coast (off Shikabe and eastern Hokkaido); and (iii) Nemuro Strait off Hokkaido, northern Japan, were examined for anisakid nematodes during December 1999 to February 2000, and the prevalence and abundance of Anisakis simplex and Contracaecum osculatum larvae were compared among the various sampling sites for fish of the same size and age. Anisakis simplex was generally more abundant than C. osculatum . Infection by A. simplex varied between the aforementioned stocks of walleye pollock as well as within stocks, whereby fish from off Rebun Island and Nemuro Strait were infected the most, followed by those from off the Pacific coast and Kumaishi. Infection by C. osculatum differed between the host stocks, and C. osculatum was the most abundant among the fish from Nemuro Strait. The infection variations seemed to be due to differences in host growth rate, host feeding habit, and the distribution of marine mammal final hosts. The results indicate that these two larval nematodes are useful biological indicators for the population study of walleye pollock in Japanese waters.     

Interannual variability in hatching period and early growth of juvenile walleye pollock, Theragra chalcogramma, in the Pacific coastal area of Hokkaido

Juvenile walleye pollock of the Japanese Pacific population were collected from the Funka Bay [spawning ground; 16–64 mm fork length (FL)] in spring and the Doto area (nursery ground; 70–146 mm FL) in summer. Hatch dates were estimated by subtracting the number of otolith daily increments from sampling dates, and their early growth was back‐calculated using otolith radius–somatic length relationships. Interannual change of the hatching period was observed during 2000–02, and the peaks ranged from mid‐February in 2000 to early‐April in 2002. In 2000, when a strong year class occurred, early life history of the surviving juveniles could be characterized by early hatching and slower growth in the larval stage (<22 mm length). Higher growth rate in 2001 and 2002 did not always lead to good survival and recruitment success. Even though their growth was slow in 2000, the larvae hatched early in the season had larger body size on a given date than faster‐growing larvae hatched in later season in 2001 and 2002. Bigger individuals at a certain moment may have advantage for survival. The delay of hatching period may result in higher size‐selective mortality, and as a necessary consequence, back‐calculated growth in 2001 and 2002 could shift towards higher growth rate, although abundance of such a year class would be at the lower level. Variability in spawning period, early growth and their interaction might have a strong relation to larval survival through cumulative predation pressure or ontogenetic changes in food availability.     

A synthesis of biological and physical processes affecting the feeding environment of larval walleye pollock (Theragra chalcogramma) in the eastern Bering Sea

Biological and physical phenomena that affect conditions for larval survival and eventual recruitment differ in the oceanic and shelf regions. In the oceanic region, eddies are a common feature. While their genesis is not well known, eddies have unique biophysical characteristics and occur with such regularity that they likely affect larval survival. High concentrations of larval pollock often are associated with eddies. Some eddies are transported onto the shelf, thereby providing larvae to the Outer Shelf Domain. Advection, rather than local production, dominated the observed springtime increase in chlorophyll (often a correlate of larval food) in the oceanic region. Over two-thirds of the south-eastern shelf, eddies are absent and other phenomena are important. Sea ice is a feature of the shelf region: its interannual variability (time of arrival, persistence, and areal extent) affects developmental rate of larvae, timing of the phytoplankton bloom (and potentially the match/mismatch of larvae and prey), and abundance and distribution of juvenile pollock. In the oceanic region, interannual variation in food for first-feeding pollock larvae is determined by advection; in the shelf region, it is the coupled dynamics of the atmosphere–ice–ocean system.     

First‐year survival of North East Atlantic mackerel (Scomber scombrus) from 1998 to 2012 appears to be driven by availability of Calanus,a preferred copepod prey

Mackerel (Scomber scombrus) is one of the ecologically and economically most important fish species in the Atlantic. Its recruitment has, for unknown reasons, been exceptional from 1998 to 2012. The majority (75%) of the survivors in the first winter were found north of an oceanographic division at approximately 52°N, despite the fact that mackerel spawns over a wide range of latitudes. Multivariate time series modelling of survivor abundance in the north revealed a significant correlation with the abundance of copepodites (stage I–IV) of Calanus sp. in the spawning season (April to June). The copepodites were a mix of C. helgolandicus (dominating) and C. finmarchicus. The growth of mackerel larvae is known to be positively related to the availability of nauplii and copopodites of preferred prey species, namely, large calanoid copepod species such as Calanus. The statistical relationship between mackerel survivors and abundance of Calanus, therefore, most likely, reflected a causal relationship: high availability of Calanus probably reduced starvation, stage‐specific predation and cannibalism (owing to prey switching). The effects of other abundant, but less preferred zooplankton taxa, (Acartia sp., Branchiopoda spp. and Echinodermata spp. larvae), as well as stock size, temperature and wind‐induced turbulence were not found to be significant. However, stock size was retained in the final model because of a significant interaction with Calanus in oceanic areas west of the North European continental shelf. This was suggested to be a consequence of a density driven expansion of the spawning area that increased the overlap between early life stages of mackerel and food (Calanus) in new areas.     

Annual variation in otolith increment widths of walleye pollock (Gadus chalcogrammus) larvae in Funka Bay,Hokkaido, Japan

To clarify relationships between year‐class strength and larval growth of walleye pollock (Gadus chalcogrammus), and oceanographic conditions in the Pacific stock off Hokkaido and Tohoku, Japan, we undertook conductivity/temperature/depth (CTD) observations and investigated larval densities, larval otolith increment widths and larval prey densities (of copepod nauplii) of the 2008, 2009, 2010 and 2011 yr classes in Funka Bay. Oyashio Coastal Water (OCW) flowed into the bay in late February in 2008, 2010 and 2011, and the mean water temperatures decreased to 1.9–3.1 °C in March. OCW was not observed in 2009, and it was warm in late February (≥3.4 °C). Increment widths of lapillar otoliths during the yolk‐sac stage were wide in 2009 and 2011, medium in 2010 and narrow in 2008. Increment widths during the first‐feeding stage tended to become wider as the hatch month progressed, and the annual variation during the first‐feeding stage was larger than that of the yolk‐sac stage. The densities of the primary food for the larvae were high in 2008 when larval increment widths were narrowest, so the effect of prey abundance on larval growth appeared to be small. The ranking of the larval abundance in March was nearly coincident with that of the increment width during the larval stage. We, therefore, suggest that the larval growth rate is associated with the mortality rate and that the growth–mortality hypothesis may be applicable to walleye pollock in Funka Bay. Feeding success under warm water conditions may be an important factor that contributes towards high growth rates.     

Effects of ontogeny, temperature, and light on vertical movements of larval Pacific cod (Gadus macrocephalus)

The role of behavior, especially vertical migration, is recognized as a critical component of realistic models of larval fish dispersion. Unfortunately, our understanding of these behaviors lags well behind our ability to construct three-dimensional flow-field models. Previous field studies of vertical behavior of larval Pacific cod ( Gadus macrocephalus ) were limited to small, preflexion stages (≤11 mm SL) in a narrow range of thermal conditions. To develop a more complete picture of larval behavior, we examined the effects of ontogeny, temperature, and light on vertical responses of larval Pacific cod in experimental columns. While eggs were strictly demersal, yolk-sac larvae displayed a strong surface orientation as early as 1 day post hatch (∼ 5 mm SL). Consistent with field observations, small preflexion larvae (<10 mm SL) showed no response to varying light levels. However, there was a direct effect of temperature on larval behavior: Pacific cod larvae exhibited a stronger surface orientation at 4°C than at 8°C. The behavior of larger, postflexion larvae (>15 mm SL) in experimental columns was consistent with a diel vertical migration and independent of water temperature: fish were more widely distributed in the column, and median positions were consistently deeper at higher light levels. These laboratory observations are combined with observations from discrete-depth (MOCNESS) sampling in the Gulf of Alaska to characterize the vertical distribution of larval Pacific cod and contrast ontogenetic patterns with walleye pollock ( Theragra chalcogramma ). The vertical movements of larval Pacific cod described here will be applied in the development of dispersal projections from Gulf of Alaska spawning grounds.     

Integrating seabird dietary and groundfish stock assessment data: Can puffins predict pollock spawning stock biomass in the North Pacific?

Information on the annual variability in abundance and growth of juvenile groundfish can be useful for predicting fisheries stocks, but is often poorly known owing to difficulties in sampling fish in their first year of life. In the Western Gulf of Alaska (WGoA) and Eastern Bering Sea (EBS) ecosystems, three species of puffin (tufted and horned puffin, Fratercula cirrhata, Fratercula corniculata, and rhinoceros auklet, Cerorhinca monocerata, Alcidae), regularly prey upon (i.e., “sample”) age-0 groundfish, including walleye pollock (Gadus chalcogramma, Gadidae) and Pacific cod (Gadus microcephalus, Gadidae). Here, we test the hypothesis that integrating puffin dietary data with walleye pollock stock assessment data provides information useful for fisheries management, including indices of interannual variation in age-0 abundance and growth. To test this hypothesis, we conducted cross-correlation and regression analyses of puffin-based indices and spawning stock biomass (SSB) for the WGoA and EBS walleye pollock stocks. For the WGoA, SSB leads the abundance of age-0 fish in the puffin diet, indicating that puffins sample the downstream production of the WGoA spawning stock. By contrast, the abundance and growth of age-0 fish sampled by puffins lead SSB for the EBS stock by 1–3 years, indicating that the puffin diet proxies incoming year class strength for this stock. Our study indicates connectivity between the WGoA and EBS walleye pollock stocks. Integration of non-traditional data sources, such as seabird diet data, with stock assessment data appears useful to inform information gaps important for managing US fisheries in the North Pacific.     

Interannual variation in the coastal distribution of a juvenile gadid in the northeast Pacific Ocean: The relevance of wind and effect on recruitment

Drift of propagules occurs within many populations inhabiting flow fields. This affects the number of propagules that rejoin their source population (recruitment) and plays a role in adaptive spatial redistribution. We focus on the cause and consequence of interannual variation in geographic distribution of population density among five cohorts of young‐of‐the‐year (age‐0) juvenile walleye pollock Gadus chalcogrammus in the western Gulf of Alaska (GOA). The coastal GOA is a wind‐driven advective system. Walleye pollock spawn during spring and their eggs and larvae drift southwestward; by late summer, age‐0 juveniles are variously distributed over the shelf. We found that high population densities of age‐0 juveniles (ca. 6 months old) near the southwestward exit of the Alaska Coastal Current from the GOA corresponded with high abundance of larvae from the major spawning area upstream, but did not translate into high abundance at older ages. Further, offshore and upwelling‐favorable winds were associated with the high downstream abundance and presumed export. In contrast, downwelling‐favorable (northeasterly) wind during and shortly after spawning (April–May) was associated with high recruitment at age 1. Finally, we found that recruitment also increased with apparent retention of age‐0 juveniles in favorable habitat upstream near the main spawning area. We hypothesize that wind‐related retention in superior upstream habitat favors recruitment. Our results argue for including wind‐driven transport in future walleye pollock recruitment models. We encourage more work on the juvenile stage of marine fishes aimed at understanding how transport and species‐specific habitat suitability interact to affect population response to large‐scale forcing.