Residency of adult Atlantic cod (Gadus morhua) in the western Gulf of Maine

A mark and recapture study of cod in the western Gulf of Maine was conducted to study the seasonal movements of fish, particularly as they related to areas closed to commercial fishing. A total of 27,772 cod were tagged, and 1334 (4.8%) were recaptured with sufficiently detailed recapture location and date to be included in the study. Results indicated that the group is resident to the area and sedentary. Although there were a small percentage of fish (2.5%) that traveled long distances (>100 km), most were recaptured near their release location. There was no linear relationship between fish length and linear distances traveled, and linear distance traveled was only weakly related to days-at-large. For all groups of cod tagged and released in particular areas and months, mean distances traveled were small (<65 km), rates of travel were slow (<0.2 km/day), and rates of group dispersion were <50 km²/day. There were no recognizable spatial or temporal patterns in the mean angles of travel for groups released in various month/area combinations, and the angular deviations associated with the mean angles were quite large. Although there seems to be little pattern in the movement of cod in our study area, temporal and spatial changes in abundance indicate that movements are occurring.

The general pattern was a concentration of large cod into one small, inshore area (Area 133) in both the spring and winter, and dispersion from this area in the ensuing months. Monthly percentages of ripe cod in Area 133 provide evidence for two spawning groups; a winter group that spawns from November through January, and a spring group that spawns from April through July. Thus it is likely that the observed spring and winter concentrations of fish in Area 133 were associated with spawning. Fish in the spring group were the largest encountered in the study. The study also provided some evidence of natal homing, i.e. a return to the same spawning grounds year after year, for the spring spawning group. Although there is some exchange of fish between most of the closed areas, the timing of the closures appears to protect the largest aggregations of cod.     

Movements and spatiotemporal distribution of escaped farmed and local wild Atlantic cod (Gadus morhua L.)

Commercial farming of Atlantic cod (Gadus morhua L.) is now being developed in several countries. The ecological consequences of cod culture are poorly understood, but recent research suggests that Atlantic cod are more prone to escape from net pens than Atlantic salmon. Here, we describe the movements and the spatiotemporal distribution of farmed cod after escape relative to wild cod, both during and outside the natural spawning season. The experimental design included simulating escape incidents of farmed cod tagged with acoustic transmitters and using an array of automatic listening stations to monitor their dispersal and distribution. For comparison, local wild cod were monitored using the same array of receivers. The farmed cod dispersed rapidly after a simulated escape, they randomly distributed over large areas and their distribution overlapped with local wild cod. Moreover, escaped farmed fish were found at local cod spawning areas during the spawning season. The study also indicated that the recapture rate of escaped farmed cod was high compared with that of escaped farmed salmon. Thus, while our results showed that there is a considerable potential for ecosystem effects caused by escaped farmed cod, mitigating actions such as an efficient recapture fishery for escapees may be possible.     

Exploring connectivity between spawning and nursery areas of Mullus barbatus (L., 1758) in the Mediterranean through a dispersal model

Connectivity between spawning and nursery areas plays a major role in determining the spatial structure of fish populations and the boundaries of stock units. Here, the potential effects of surface current on a red mullet population in the Central Mediterranean were simulated using a physical oceanographic model. Red mullet larvae were represented as Lagrangian drifters released in known spawning areas of the Strait of Sicily (SoS), which represents one of the most productive demersal fishing‐grounds of the Mediterranean. To consider the effect of inter‐annual variability of oceanographic patterns, numerical simulations were performed for the spawning seasons from 1999 to 2012. The main goal was to explore connectivity between population subunits, in terms of spawning and nursery areas, inhabiting the northern (Sicilian‐Maltese) and southern (African) continental shelves of the SoS. The numerical simulations revealed a certain degree of connectivity between the Sicilian–Maltese and the African sides of the SoS. Connectivity is present in both directions, but it is stronger from the Sicilian–Maltese spawning areas to the African nurseries owing to the marine circulation features of the region. However, because the majority of the larvae are transported to areas unsuitable for settlement or outside the SoS, the dispersal process is characterized by a strong loss of potential settlers born in the spawning areas. These results are in agreement with the low genetic heterogeneity reported for this species in the Mediterranean Sea and support the existence of a metapopulation structure of red mullet in the SoS and the adjacent areas.     

Low-fidelity homing behaviour of Biwa salmon <Emphasis Type="Italic">Oncorhynchus</Emphasis> sp. landlocked in Lake Biwa as inferred from otolith elemental and Sr isotopic compositions

Biwa salmon Oncorhynchus sp. is endemic to Lake Biwa, Japan, where it is an important commercial and recreational fisheries species. However, no information is currently available on its population structure and migration ecology. Therefore, here we evaluated whether otolith Sr/Ca, Ba/Ca and ⁸⁷Sr/⁸⁶Sr ratios can be used as natural signatures in Biwa salmon and then used these to determine the natal origins of lake-migration-phase individuals and spawning adults, and the homing ability of spawning adults in the Lake Biwa water system. Quadratic discriminant function analysis demonstrated that the lake-migration school comprised individuals with multiple origins, including rivers to the east, west and north of Lake Biwa, and that the homing rate of spawning adults was low (18 out of 80 individuals), with ca. 78% of fish straying into non-natal rivers. However, this straying behaviour was not spatially random, with fish tending to migrate upstream in rivers neighbouring their natal rivers. The high rate of straying in spawning adults is considered important for establishing and maintaining this species, which is highly adapted to life in the Lake Biwa water system where environmental disturbances often occur.     

The spatial distribution of cod (Gadus morhua L.) spawning grounds in the Kattegat, eastern North Sea

Similar to many other commercial marine fish species, Atlantic cod (Gadus morhua L.) migrate towards specific sites at spawning. These temporal aggregations are generally the most targeted by the commercial fishery. The Kattegat cod has undergone a substantial reduction over the past 25 years and both stock size and spawning stock biomass have remained at very low levels since the end of the 1990s. There is, therefore, an urgent need to map and document spawning grounds still in use. In the present study, spawning sites of Atlantic cod were identified in the Kattegat using a combination of commercial and fishery independent data from 1996 to 2004. Moreover, putative spawning and non-spawning areas were also sampled before and during the reproductive season between 2002 and 2006, and the proportion of mature females together with the individual physiological status were used to validate and strengthen the spatial analyses.

The spatial analyses identified several spawning areas in the region and two areas in the southeastern part of the Kattegat which appeared to be the most important. The results showed the presence of cod spawning aggregations, although reduced in size, in areas that have been utilized for more than 25 years according to historical information. Some local spawning grounds may have also disappeared. The proportion of mature females was higher in putative spawning than in non-spawning areas (p < 0.001) and females from spawning areas had higher gonadosomatic (p < 0.05) and hepatosomatic (p < 0.001) indices than those from non-spawning areas.

Knowledge of stock spatial and temporal distribution is essential in designing recovery strategies for depleted fish populations. The unambiguous stability of the locations of spawning aggregations over time, as shown in this study, represents a useful aid in order to efficiently implement a recovery plan for the collapsing cod stock in this area.     

Spatiotemporal distribution of Atlantic cod (Gadus morhua L.) with intact and blocked olfactory sense during the spawning season in a Norwegian fjord with intensive salmon farming

In several areas along the Norwegian coast local fishermen claim that wild migrating Atlantic cod (Gadus morhua L.) have changed their spawning migratory behaviour following establishment of salmon farms, and no longer enter their natural spawning grounds in the fjords. This has created a heated debate in Norway and, in some areas, passed down a moratorium on establishing new salmon farms. Research has so far not been able to show a causal connection, but it has been suggested that water-soluble odorants from high density salmon stocks in commercial fish farms might be responsible for the avoidance of areas with fish farming and abandonment of nearby traditional spawning grounds. The aim of this study was to test this hypothesis by studying the spatiotemporal distribution of wild migrating cod tagged with acoustic transmitters using an array of automatic listening stations in Øksfjord, a Norwegian fjord with intensive fish farming and traditional spawning grounds for cod. The tagged cod were released either in the outer parts of Øksfjord, i.e. in the area they were caught, or in the inner part of the fjord close to their traditional spawning areas. The olfactory sense was physically blocked for 50% of the fish. Only one cod released in the outer part of the fjord visited the traditional spawning grounds. The majority of the cod that were released in the inner part of the fjord remained in this area for approximately three weeks before they left the fjord. There were no differences in spatiotemporal distribution between cod with intact and cod with blocked olfactory sense. Thus, the results provided no evidence that migrating wild cod avoid areas with fish farming as a response to odorants, but the results do not per se contradict the fishermen's observation that coastal cod have changed their migratory pattern in this fjord.     

Geographical patterns of straying of fall chinook salmon,Oncorhynchus tshawytscha (Walbaum), from Columbia River (USA) hatcheries

The large-scale spatial patterns of straying of marked chinook salmon, Oncorhynchus tshawytscha (Walbaum), produced at hatcheries in the Columbia River (north-west USA) are described. Straying rates were extremely variable among hatcheries (from 1% to 95%). Fish produced near the mouth of the river strayed more commonly than those produced farther upriver. While the large-scale straying patterns were well explained by the distance between the hatchery of origin and alternative destinations, substantial deviations from this pattern were observed at smaller scales. Three distinct areas of origin were identified and most straying occurred between hatcheries and natural spawning grounds within each area. The location of the stream where the hatchery of origin is located affects the patterns of straying. Fish reared at hatcheries in the main stem of the river did not stray into tributaries as often as predicted by a model based exclusively on linear distance between alternative locations. Likewise, fish released from large tributaries strayed to main stem hatcheries less often than predicted by the model. Fish produced at hatcheries on north bank tributaries appeared to stray less into southern streams than fish released from southern streams. These findings indicate that salmon are more likely to stray to spawning areas similar to their natal site than to dissimilar sites, in addition to the tendency to stray to nearly sites. In addition to the general patterns, examination of data from two types of transplants indicated that there is a genetic component to homing/straying behaviour within the watershed. However, salmon transplanted into the Columbia River watershed from elsewhere showed homing precision comparable to that of local fish, although they retained their ancestral marine distribution.     

Behavioural responses in wild cod (Gadus morhua L.) exposed to fish holding water

Local fishermen in several areas of Norway assert that salmon farms have caused the wild migrating cod to change their migratory behaviour, so that they no longer enter their natural spawning grounds in the fjords. If the asserted changes in behaviour of wild cod populations can be linked to establishment of salmon farms, water-soluble odorants are then possible candidates to explain such a connection. Chemical stimulants are important to the individual fish's conception of the surrounding environment. High density stocks of fish in a farm are expected to release large amounts of waterborne information. The present laboratory experiments were conducted to test behavioural responses in Atlantic cod exposed to water containing metabolites and waste from farmed salmon. The trials were conducted on single fish in a multiple chamber preference system. The results show that migrating wild Atlantic cod chose to spend more time in chambers without addition of water from the salmon tank, regardless of their maturation status, and even at very low concentrations (0.2%). The avoidance is probably due to presence of chemical compounds with olfactory properties from salmon, since anosmic cod did not elicit such response. Farmed cod, on the other hand, does not avoid water from the salmon tank, and stationary wild cod caught nearby a fish farm had a less pronounced response as compared to wild migrating cod. The response seen is not species specific, as wild migrating cod responded similar to water from a tank holding farmed cod as to water from salmon. These results do not preclude fishermen's observations that cod change their behaviour in areas with fish farming activity. Such a change in behaviour could be a response to water-soluble odorants, but needs to be validated and detailed in further laboratory experiments as well as in nature before any conclusions can be made.     

Historical spatial baselines in conservation and management of marine resources

Increased knowledge on the spatial distribution of marine resources is crucial for the implementation of a true ecosystem approach to management and the conservation of marine organisms. For exploited fish species characterized by aggregation behaviour during spawning time, the identification and tracking of spawning areas is essential for a correct assessment of their productivity and population abundance. To elucidate this concept, we reconstructed the spatio‐temporal distribution of adult plaice (Pleuronectes platessa, Pleuronectidae) during spawning time along the 20^th century. Historical data reveal that not only the abundance but also the former population richness was much higher than previously estimated and has declined because of protracted over‐exploitation during the last 30 years. We conclude that forecast of stock recovery to former levels of abundance neglecting spatial reorganizations might be over‐optimistic and shaded by a lost memory of the past population richness. These results reinforce the importance of managing exploited marine resources at a greater spatial resolution than has been carried out in the history of fishery management.     

Discrete spatial dynamics in a marine broadcast spawner: Re-evaluating scales of connectivity and habitat associations in Atlantic cod (Gadus morhua L.) in coastal Newfoundland

We examined the spatial dynamics of Atlantic cod, Gadus morhua, through its life history on the northeast and southeast coasts of Newfoundland in order to delineate dispersal and critical habitat requirements as potential determinants of appropriate spatial scales for management. Hydroacoustic surveys across the northeast Newfoundland shelf in 1990, 1995, and 2000 showed the distribution of the remaining large aggregations of cod to be concentrated in inshore embayments, which is contrary to historic patterns. Inshore surveys on the south coast identified local spawning aggregations at specific geographic locations, to which adults homed in subsequent years. Dispersal of eggs and larvae to the offshore occurred during spring, but was localized (within bay) in the warm summer months. Abundance of age-0 cod increased with proximity to spawning location inshore, suggesting self-recruitment within local embayments. Age-2–3 juveniles actively selected structurally complex habitat at scales of less than tens of kilometers, with home ranges typically less than 10 ha. Juveniles were associated with specific habitat types, especially at age 0, in both lab and field experiments. Our results suggest that self-recruitment and critical life-history events (e.g., spawning, settlement, and maturity) may occur within local embayments, which currently assume much greater importance to overall cod stock dynamics than in the past. Given evidence for limited dispersal, and the inshore presence of critical habitat for early life-history stages, spatial management tools that consider local dynamics may provide a greater degree of protection for coastal Newfoundland cod than previously thought.     

Elemental fingerprinting in otoliths reveals natal homing of anadromous Baltic Sea pike (Esox lucius L.)

We examined the element pattern in the otoliths of a migratory fish species that inhabit the coastal areas in the brackish of the Baltic Sea. The northern pike (Esox lucius) show migratory behaviour, spawning in streams and rivers and foraging in the sea. We examined spawning migration in four nearby streams in the south‐west part of the Baltic. Otolith analysis by microPIXE revealed unique elemental patterns (Sr, Zn, Br, Co and Mn) for the juveniles in each of the different streams. The strontium signal in the otolith of the juveniles was used as an indicator of freshwater origin and the time spent in the stream. Adult pike in their migrating spawning phase were caught in each of the streams. The elemental composition in otoliths in their freshwater phase (using juvenile pike in the streams as references) was determined. A principal component analysis showed that the elemental fingerprint during the freshwater phase several years back in time was similar for the adult fish and for juveniles inhabiting the stream today. The results indicated natal homing of the adults to a specific stream, a conclusion that was strengthened by the fact that marked fish returned to spawn over consecutive years. Anadromous pike in the Baltic Sea may thus be divided in subpopulations. The results of the study may have implications for fishery management, as pike in the Baltic Sea cannot be seen as homogenous population.     

Oceanographic influences on the distribution of Baltic cod, Gadus morhua, during spawning in the Bornholm Basin of the Baltic Sea

The Baltic Sea is a stratified, semienclosed sea typified by a low-salinity surface layer and a deep saline layer of varying volume, salinity, temperature and oxygen concentration. The relationships between these oceanographic factors and the distribution of Baltic cod are presented, utilizing results from a survey carried out during the 1995 spawning period in the Bornholm Basin, at present the main spawning area of this stock. Cod distribution, abundance and population structure were estimated from hydroacoustic and trawl data and related to hydrographic parameters as well as to bottom depth. In the central basin, cod were aggregated in an intermediate layer about 15 m thick. This area of peak abundance was defined at its upper limit by the halocline and at the lower limit by oxygen content. The majority of individuals caught in the basin centre were in spawning or pre-spawning condition with a high proportion of males to females. On the basin slopes, aggregations of cod were found near the bottom. These individuals were mainly immature and maturing stages with an increasing proportion of females to males with size. Salinity and oxygen conditions were found to be the major factors influencing the vertical and horizontal distribution of adult cod. Abundance of immature cod was also positively related to decreasing bottom depths. The effect of temperature was minor. The observed size- and sex-dependent spawning aggregation patterns, in association with habitat volume and stock size, may influence cod catchability and thereby the assessment and exploitation patterns of this stock.     

Influence of larval transport and temperature on recruitment dynamics of North Sea cod (Gadus morhua) across spatial scales of observation

The survival of fish eggs and larvae, and therefore recruitment success, can be critically affected by transport in ocean currents. Combining a model of early‐life stage dispersal with statistical stock–recruitment models, we investigated the role of larval transport for recruitment variability across spatial scales for the population complex of North Sea cod (Gadus morhua). By using a coupled physical–biological model, we estimated the egg and larval transport over a 44‐year period. The oceanographic component of the model, capable of capturing the interannual variability of temperature and ocean current patterns, was coupled to the biological component, an individual‐based model (IBM) that simulated the cod eggs and larvae development and mortality. This study proposes a novel method to account for larval transport and success in stock–recruitment models: weighting the spawning stock biomass by retention rate and, in the case of multiple populations, their connectivity. Our method provides an estimate of the stock biomass contributing to recruitment and the effect of larval transport on recruitment variability. Our results indicate an effect, albeit small, in some populations at the local level. Including transport anomaly as an environmental covariate in traditional stock–recruitment models in turn captures recruitment variability at larger scales. Our study aims to quantify the role of larval transport for recruitment across spatial scales, and disentangle the roles of temperature and larval transport on effective connectivity between populations, thus informing about the potential impacts of climate change on the cod population structure in the North Sea.     

Instantaneous areal population density of entire Atlantic cod and herring spawning groups and group size distribution relative to total spawning population

The wide‐area group behaviour of spawning Atlantic cod and herring is investigated. By a combination of Ocean Acoustic Waveguide Remote Sensing (OAWRS) and conventional sensing methods, first‐look images of the instantaneous population density are obtained of entire Atlantic cod spawning groups, stretching for tens of kilometres in the Nordic Seas. This structural information made it possible to quantify the spawning group size distribution of cod over a roughly 30‐year period from conventional line‐transect data acquired annually by vertical echo sounding in the Nordic Seas. The size distribution is found to be consistent with the log‐normal probability density often found in growth processes that depend on many independent parameters. Nordic Seas cod populations are found to distribute into many vast behavioural groups during spawning with relatively stable mean size despite larger variations in total annual population. When sustained at pre‐industrial levels, the total spawning population is found to greatly exceed the mean spawning group size. As an apparent consequence of this large differential, when the total population, or overall scale, declined to within a standard deviation of this mean cod spawning group quantum, or inner‐group‐behavioural scale, return to pre‐industrial levels required decades. Findings for Atlantic herring are similar, where summing the spawning group populations measured in a single instantaneous OAWRS image per day over the 8‐day peak spawning period enabled accurate enumeration of the entire Georges Bank herring spawning population to within 7% of the independent NOAA estimate for 2006. These results may be relevant to other oceanic fish.     

Spawning of North Sea fishes linked to hydrographic features

Spawning of fishes takes place across a wide area of the North Sea. However, more intense spawning is seen in restricted areas, indicating that such areas present favorable conditions. To update information on fish spawning in the North Sea and analyze potential linkages to hydrographic characteristics, an internationally coordinated survey was conducted in the winter/spring of 2004. Oblique hauls for fish eggs and larvae and vertical profiles of temperature and salinity were carried out at 393 stations across the entire North Sea. The hydrography was strongly influenced by the interfacing of water masses of different salinity, and frontal zones were seen along all coastal areas and off the Dogger and Fisher Banks. Total abundances of eggs and larvae, including fish species such as cod, haddock, plaice, long rough dab and sandeel, peaked in the vicinity of the frontal areas. Hence our findings indicate that the main spawning locations of fish are linked to recurrent hydrographic features such as salinity fronts. Such a linkage may provide survival advantages, as the fronts present favorable feeding conditions, and the related physical processes may confine egg and larval dispersal and transport them towards suitable nursery habitats.     

Anadromy and homing: two life‐history traits with adaptive synergies in salmonid fishes?

Anadromy and homing are two traits found widely amongst teleost fishes. They co-occur in salmonid fishes, and probably also in other fish families. Anadromy provides fish with the opportunity for more rapid growth, larger size, and higher fecundity through the exploitation of rich food resources and favourable growing temperatures in the sea, but may result in higher predation mortalities and has the tendency to disperse stocks very widely. Homing fosters adaptation of stocks to favourable local spawning conditions, but the dispersive effects of anadromy, in the absence of homing, may tend to break down such local adaptation. A low percentage of straying in species that home may have long-term evolutionary advantages. This paper explores the question of whether homing may be a preadaptation for successful evolution of anadromy, or whether the two attributes have coevolved.     

Migratory behaviour and range in Atlantic cod: inference from a century of tagging

A century of tagging experiments on 174 Atlantic cod (Gadus morhua) groups is reviewed and the frequency and spatial distribution of four migratory behaviours documented. Of all cod groups, 41% were categorized as sedentary, 18% as accurate homers, 20% as inaccurate homers and 20% as dispersers (ranging over large areas without recognizable return migrations). All behaviours occurred over the full spatial range of cod. Coastal groups did not differ significantly from offshore (shelf) groups in the relative frequency of migratory behaviours. However, the north‐east Atlantic had more sedentary and accurate homing groups than did the north‐west Atlantic, which had more dispersing groups. Overall, sedentary cod groups had lower maximum historical biomass than did other groups, confirming that migration/dispersal begets abundance. Maximum historical biomass was strongly related to the area occupied [log (biomass, tonnes)] = 1.58 log (range, km²) + 1.529; r² = 0.9), irrespective of migratory type (for 23 major groups a total of 22 million tonnes over 3.3 million km² with an average density of 7 tonnes km^?2). Historical densities were not related to area occupied, although all large groups exhibited high densities (10–12 tonnes km^?2 ; smaller groups had a wide range of density). The four migratory strategies in cod enable entrainment by diverse physical, oceanographic and biological ecosystem properties, and is key to cod success in the North Atlantic.     

Dispersal and tributary immigration by juvenile coho salmon contribute to spatial expansion during colonisation

Anadromous fishes are frequently restricted by artificial barriers to movement such as dams and culverts, so measuring dispersal helps identify sites where improved connectivity could promote range expansion and population viability. We used a combination of DNA‐based parentage analysis and mark–recapture techniques to evaluate dispersal by juvenile coho salmon (Oncorhynchus kisutch) in a population in the initial stages of colonisation following installation of fish passage structures at a previously impassable dam on the Cedar River, WA, USA. The spatial distribution of individuals within maternal families revealed that dispersal was common. Among the offspring of radio‐tagged mothers, 28% were collected outside the spawning reach and dispersed up to 6.3 km (median = 1.5 km). Most juveniles captured in a tributary (Rock Creek, where few adults spawned) had immigrated from the Cedar River and represented many different families. Juvenile dispersal therefore provided a secondary phase of spatial expansion following initial colonisation by adults. Consistent with the condition‐dependent dispersal hypothesis, juveniles that dispersed farther upstream in the tributary were larger than fish collected near the tributary mouth. Overall, the results demonstrated widespread dispersal in a system with low coho salmon densities, and this might increase the rate of population growth if it reduces the effects of local density dependence. By implication, juveniles can take advantage of rearing habitats reconnected through barrier removal, even when such areas are located several kilometres from adult breeding grounds.     

Dynamic in-lake spawning migrations by female sockeye salmon

Abstract –  Precise homing by salmon to natal habitats is considered the primary mechanism in the evolution of population-specific traits, yet few studies have focused on this final phase of their spawning migration. We radio tagged 157 female sockeye salmon ( Oncorhynchus nerka ) as they entered Lake Clark, Alaska, and tracked them every 1–10 days to their spawning locations. Contrary to past research, no specific shoreline migration pattern was observed (e.g., clockwise) nor did fish enter a tributary unless they spawned in that tributary. Tributary spawning fish migrated faster (mean = 4.7 km·day⁻¹, SD = 2.7, vs. 1.6 km·day⁻¹, SD = 2.1) and more directly (mean linearity = 0.8, SD = 0.2, vs. 0.4, SD = 0.2) than Lake Clark beach spawning fish. Although radio-tagged salmon migrated to within 5 km of their final spawning location in an average of 21.2 days (SD = 13.2), some fish migrated five times the distance necessary and over 50 days to reach their spawning destination. These results demonstrate the dynamic nature of this final phase of migration and support studies indicating a higher degree of homing precision by tributary spawning fish.     

Larval group differentiation in Atlantic cod (Gadus morhua) inside and outside the Gullmar Fjord

The spatial and temporal occurrence of pelagic fish stages and their biological variability may affect their dispersal and survival, and ultimately fish recruitment. We collected Atlantic cod larvae at one station inside and at one station outside the Gullmar Fjord, eastern Skagerrak, in order to investigate small-scale larval group differentiation. Rectangular midwater trawl hauls were taken every 6 h (during 24 h) from three separate depth intervals between the surface and 70 m depth. About 80% and 20% of all larvae occurred above the halocline at the Fjord station and the Coastal station, respectively. Hatching (at both stations) occurred from the 3rd week in February to the 1st week in May, indicating that cod larvae were present for at least 5 months (from late February to early August). The length and hatch date frequency distributions of larvae from the surface layer were unimodal inside the fjord but bimodal outside the fjord. Analyses of seven microsatellite DNA loci indicated that larvae collected inside the fjord (where local spawning occurs) were genetically distinct from larvae sampled on the outside (F_ST = 0.0026). The two age cohorts outside the fjord were not, however, genetically different, nor were larvae collected at different depths. We conclude that small-scale variability of vertical concentration and larval life history variability should have consequences for interpreting models of larval dispersal and survival, and subpopulation structure analyses.