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.     

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.     

Physical transport of young pollock larvae (Theragra chalcogramma) near Shelikof Strait as inferred from a hydrodynamic model

An advective model was used to simulate the drift of larval walleye pollock ( Theragra chalcogramma ) over a 40-day period (late April through early June) near Shelikof Strait, Alaska. This model was used: (i) to assess how much of the observed change in larval positions during that period can be explained by transport at fixed depth; (ii) to demonstrate that observed change can be related to mean large-scale meteorological forcing; and (iii) to investigate accumulation of larvae in specific areas near the coast. Based on availability of larval and circulation data, three years were studied: 1988, 1989 and 1991. Velocity fields generated from a hydrodynamic model driven by winds and runoff were used to advect particles seeded in accordance with observed larval distributions in late April of each year. The modelled larvae were tracked at 40 m depth, corresponding to the mean depth of sampled larvae and the depth of neutrally buoyant drifters employed in field studies. Specific features observed in late May larval surveys were reproduced by the model, such as the accumulation of larvae in a shoal area downstream of the strait. Differences among the modelled years include extensive flushing of larvae to the south-west in 1988 and 1991, vs. limited flushing in 1989. These differences appear related to the mean large-scale atmospheric pressure patterns for April-May of those years.     

Simulation of physically mediated variability in prey resources of a larval fish: a three-dimensional NPZ model

A three-dimensional biophysical nutrient–phytoplankton–zooplankton model was used to investigate the spatial and temporal dynamics of food resources for young walleye pollock in the western Gulf of Alaska, to further understanding of recruitment processes for pollock. We modeled nitrogen, phytoplankton, a large herbivorous grazer parameterized as Neocalanus spp. (the biomass dominant copepod in the Gulf), and the 13 stages (egg, naupliar and copepodite) of Pseudocalanus spp. (a major constituent of the diet of pollock) so that the appropriate size class of food for each size of larval pollock was represented. Model results identified an area between the Semidi and Shumagin Islands that may not be suitable as a nursery area early in the year due to low prey abundance. Modeled mesoscale eddies, previously hypothesized to be important for larval pollock retention in Shelikof Strait, contained higher prey concentrations than the surrounding waters when they were cyclonic. This work also help to understand the consistency of pollock spawning in time and space in Shelikof Strait, by examining the timing and location of prey availability which, along with transport, narrows the window for optimal spawning.     

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.     

Observed patches of walleye pollock eggs and larvae in Shelikof Strait, Alaska: their characteristics, formation and persistence

Using observations from 38 ichthyoplankton surveys conducted near Shelikof Strait, Alaska between 1979 and 1992, we characterized the horizontal distribution and spatial patchiness of the early life stages of walleye pollock ( Theragra chalcogtamma ). Lloyd's index of patchiness ranged from 3.9-6.1 for eggs and 3.9–16.2 for larvae. This index was size (age) dependent: low for eggs, high for newly hatched larvae, then decreasing through late larval stage. By the early juvenile stage, patchiness increased as pollock began to school. The percentage of larvae in a patch (defined as the percentage of larvae present at stations where larval counts exceeded the mean by one standard deviation during the given survey) varied greatly (26–92%). Larval distributions were used to deduce physical mechanisms responsible for patches. Three categories of patches were identified: those created by interaction of larvae with time-dependent currents, those in the vicinity of Sutwik Island, and those associated with eddies. Simulation experiments were utilized to examine processes influencing patch formation and the role of larval swimming. Between 5 and 6 weeks after hatching, larvae have swimming abilities that enable them to maintain a patch already created by physical mechanisms.     

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.     

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.     

Acoustic detection of mesoscale biophysical features in the Shelikof sea valley, and their relevance to pollock larvae in the Gulf of Alaska

Mesoscale features such as fronts and eddies can act to retain larval walleye pollock ( Theragra chalcogramma ) within the continental shelf zone in the western Gulf of Alaska. During two May cruises, we observed unusual patterns of backscattering with a 38 kHz acoustic system. Characteristics of this signal were a strong scattering layer at the surface and in midwater, with the water column between nearly void of sound scattering organisms. This signal appeared in several transects where satellite remote sensing indicated the presence of an eddy. Analysis of concomitant water properties and ADCP (153 kHz) data confirmed the existence of an anomalous physical feature at this location. Biological properties (chlorophyll and zooplankton) showed marked changes across the edge of the feature. Larval pollock densities were estimated to be an order of magnitude higher within this feature compared to those outside. Acoustic backscatter signals can be used to identify and characterize mesoscale biophysical features in the ocean, thereby enabling real-time studies of these features.     

How “The Blob” affected groundfish distributions in the Gulf of Alaska

We investigated the distributional shifts of groundfish in response to anomalous ocean conditions, particularly the recent anomalously warm period (2014–2016; “The Blob”), based on data from ten Gulf of Alaska bottom trawl surveys conducted by the Alaska Fisheries Science Center during 1996–2015. Six groundfish species were considered: Pacific cod (Gadus macrocephalus), arrowtooth flounder (Atheresthes stomias), walleye pollock (Gadus chalcogrammus), Pacific ocean perch (Sebastes alutus), northern rock sole (Lepidopsetta polyxystra), and southern rock sole (Lepidopsetta bilineata). Ontogenetic differences were examined by dividing data for each fish species into size classes. Our study demonstrated that after accounting for size‐specific depth preferences, the spatial responses of groundfish to anomalous ocean conditions differed by species and foraging guild in the central Gulf of Alaska. Pacific cod and arrowtooth flounder showed similar responses to ocean warming, but different responses to cooling. In general, Pacific cod moved to deeper depths in warmer years and moved to shallower depths in colder years. Arrowtooth flounder also moved deeper in warmer years. However, in colder years, large arrowtooth flounder (>40 cm) shifted toward shallower depths while smaller‐sized fish shifted toward deeper depths. In warmer years, large pollock (>30 cm) moved to deeper waters while smaller pollock (10–20 cm) moved to shallower waters. Pacific ocean perch exhibited an opposite response to thermal changes in habitat compared with Pacific cod and arrowtooth flounder. They moved deeper in colder years, but there was no clear change in depth as a function of size in response to warmer habitat.     

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.     

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.     

Daily and sub-daily otolith increments of larval and juvenile walleye pollock, Theragra chalcogramma (Pallas), as validated by alizarin complexone experiments

Walleye pollock (Theragra chalcogramma) were reared from eggs to the juvenile life stage to study daily increment formation in the sagittae otoliths, which are routinely used for age and growth analyses. The apparent deposition of sub-daily growth increments becomes problematic for determining fish age from the late larval stage throughout the juvenile (young-of-the-year) development stage. Otolith marking experiments were conducted to determine interpretation criteria to differentiate between daily and sub-daily increments. Immersion of larval and transforming walleye pollock in 25 mg/l of alizarin complexone (ALC) for 6 h once a week produced a fluorescent mark on the day of staining. Evidence of six well defined and equally spaced increments counted between the weekly ALC marks validated the deposition of daily increments. The daily increments gradually increased in width as the fish/otolith grew. The criteria for determining the presence of sub-daily increments between the daily increments were (1) weak optical definition and (2) a sudden change in incremental distance that lasted for one or two increments and were approximately <0.5 μm in width. Growth problems that occurred during the experiments were identified on otoliths as reductions in daily incremental widths and optical definition, which continued for several days. Otoliths from field-collected fish have also shown similar changes in daily increment properties during the juvenile stage, which may be an indicator of an environmental influence. The criteria for defining different increment types help to resolve our current age determination issues for late larval and early juvenile stage walleye pollock from the Gulf of Alaska.     

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.     

Distribution, density and abundance of the western Baltic herring (Clupea harengus) in the Sound (ICES Subdivision 23) in relation to hydrographical features

Biomass and duration of the over-wintering period of the Rügen spring spawning herring stock (RHS) in the Sound (ICES Subdivison 23) were investigated as well as possible hydrographical factors affecting relative distribution and triggering southwards migration towards the spawning grounds. Monitoring was performed during 27 surveys over a 6-year period (1993–1998). Abundance of 45–165 000 t in August–February, 5–60 000 t in March–May, and <2000 t in June–July was found. This indicates a longer over-wintering period than hitherto assumed. The year classes 1988, 1991 and 1994 were relatively strong occurring in the Sound from age 2 (winter-rings) and abundance in autumn 1993 and 1996–1997 was relatively high. Relative year class strengths are in accordance with variations in larval indices at the spawning grounds. Decreasing abundance in late spring in the Sound was concordant with observed peaks in commercial landings and (subsequent) peak larval indices at the spawning grounds. Larger size groups seem to migrate southwards in spring before smaller herring size groups. Peak herring densities occur in the northern Sound in autumn and they do not concentrate near the southern Drogden threshold in spring before southwards spawning migration. Highest densities were found from 8 to 22 m below sea surface, and just below or in the halocline in areas with stratified water column. Indications of inflows to the Baltic triggering southwards migration can be observed at some occasions, however, these results are not conclusive because of relatively few biomass estimates in 1996–1998. The identified patterns in herring occurrence agree with some conclusions on migration routes for RHS from previously reported tagging studies, but the present investigations indicate prolonged over-wintering in the Sound. This report quantifies the abundance and distribution of herring in the Sound over the year on basis of repeated investigations, and investigates possible influence of hydrographical factors on distribution and migration.     

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.     

Spatial and temporal distribution and abundance of European hake, Merluccius merluccius, eggs and larvae in the Catalan coast (NW Mediterranean)

Plankton hauls were conducted on five surveys from November 1998 to November 1999 to study the spatial distribution of eggs and larvae of European hake off Catalonia (NW Mediterranean Sea). Also CTD casts to record hydrographic parameters were carried out on a closely spaced station grid. Merluccius merluccius eggs and larvae appeared mainly in late spring, summer and autumn surveys and were very scarce in winter. Strong differences in terms of egg and larval densities were observed between the two November surveys, which could be attributed to the anomalous hydrographic situation during November 1998. M. merluccius egg and larvae were mainly distributed over the continental shelf, with peak abundances between the 100 m isobath and the edge of the shelf. On the evidence of larval size frequency distributions in the different sampling sectors and the closely overlapping distribution patterns for the eggs and the adult spawning stock, drifting of hake eggs and larvae was not a major factor. The larval distribution extended only slightly further offshore than the egg distribution. Using the hydrographic information and the larval distribution data, an attempt was made to relate the different seasonal productivity levels over the spawning period and the distribution of the larvae.     

Ichthyophonus‐infected walleye pollock Theragra chalcogramma (Pallas) in the eastern Bering Sea: a potential reservoir of infections in the North Pacific

In 2003, the Alaska walleye pollock industry reported product quality issues attributed to an unspecified parasite in fish muscle. Using molecular and histological methods, we identified the parasite in Bering Sea pollock as Ichthyophonus. Infected pollock were identified throughout the study area, and prevalence was greater in adults than in juveniles. This study not only provides the first documented report of Ichthyophonus in any fish species captured in the Bering Sea, but also reveals that the parasite has been present in this region for nearly 20 years and is not a recent introduction. Sequence analysis of 18S rDNA from Ichthyophonus in pollock revealed that consensus sequences were identical to published parasite sequences from Pacific herring and Yukon River Chinook salmon. Results from this study suggest potential for Ichthyophonus exposures from infected pollock via two trophic pathways; feeding on whole fish as prey and scavenging on industry‐discharged offal. Considering the notable Ichthyophonus levels in pollock, the low host specificity of the parasite and the role of this host as a central prey item in the Bering Sea, pollock likely serve as a key Ichthyophonus reservoir for other susceptible hosts in the North Pacific.     

Seasonal abundance of two important forage species in the North Pacific Ocean,Pacific herring and walleye pollock

Pacific herring (Clupea pallasii) and walleye pollock (Theragra chalcogramma) are important forage species in the North Pacific Ocean, but their seasonal abundance patterns are poorly known. During three consecutive years of monthly acoustic surveys in Lynn Canal, southeastern Alaska, large schools of herring dominated during winter and were present in a 60-km long submarine gully; this gully appears to provide critical winter habitat for herring when their prey are less available and energy expenditure must be reduced. The salient change in pollock distribution is a shift from shallow waters during summer to deeper waters during winter, such that shallow (<40 m) waters are nearly devoid of pollock during winter. The shift presumably occurs in response to the build-up of secondary productivity during summer and predator avoidance during winter. The seasonal changes in herring abundance drove seasonal changes in predator abundance, as Steller sea lions (Eumetopias jubatus) and humpback whales (Megaptera novaeangliae) preyed upon the winter aggregation of herring. Such seasonal links likely serve an important role in structuring trophic relationships in the North Pacific Ocean ecosystem.