Reconstructing migrations of individual cod (Gadus morhua L.) in the Baltic Sea by using electronic data storage tags

A new methodology is presented to reconstruct migration pathways of individual fish inhabiting ecosystems with moderate‐to‐strong gradients in temperature or salinity. The method uses measurements of ambient pressure, temperature and salinity obtained from electronic data storage tags attached to individual fish and is particularly applicable in areas with negligible tides. We demonstrate the method with Baltic cod. Hydrographic fields obtained from hydrodynamic modelling were used as a geolocation database to identify daily positions of Baltic cod by comparison with the environmental data collected by the tags. Using randomly distributed individual parameter perturbations in the range of the instrument precision of the tag we simulated a cod migrating through the Baltic Sea. The distance between the prescribed and geolocated positions of this artificial cod was on average 2.9 ± 4.7 (SD) km. Subsequently, the method was used to reconstruct migration routes of 10 real cod tagged in the Bornholm Basin of the Baltic Sea in early spring 2003. Differences were compared between the tag data and the geolocation database. The uncertainty in geolocation at recapture day was on average 75 ± 23 (SD) km, as shown by comparison between geolocated position and recapture position.     

Processes controlling retention of spring‐spawned Atlantic cod (Gadus morhua) in the western Gulf of Maine and their relationship to an index of recruitment success

Atlantic cod, Gadus morhua, harvested in US waters are currently managed as a Gulf of Maine stock and as a stock comprising Georges Bank and southern New England populations. Over the past two and a half decades, success of age‐1 recruitment to the Gulf of Maine stock has varied by more than an order of magnitude. To investigate the hypothesis that this variation is related to variation in the transport of larval cod to nursery areas, we carried out model simulations of the movement of planktonic eggs and larvae spawned within the western Gulf of Maine during spring spawning events of 1995–2005. Results indicate that the retention of spring‐spawned cod, and their transport to areas suitable for early stage juvenile development, is strongly dependent on local wind conditions. Larval cod retention is favored during times of downwelling‐favorable winds and is least likely during times of upwelling‐favorable winds, during which buoyant eggs and early stage larvae tend to be advected offshore to the Western Maine Coastal Current and subsequently carried out of the Gulf of Maine. Model results also indicate that diel vertical migration of later stage larvae enhances the likelihood of retention within the western Gulf of Maine. Consistent with model results is a strong correlation between age‐1 recruitment success to the Gulf of Maine cod stock and the mean northward wind velocity measured in Massachusetts Bay during May. Based on these findings, we propose a wind index for strong recruitment success of age‐1 cod to the Gulf of Maine stock.     

Testing a model to track fish migrations in polar regions using pop‐up satellite archival tags

The use of pop‐up archival satellite tags (PSATs) to geolocate marine fishes in polar regions is challenging due to the brevity of periods during which there is a defined sunrise and sunset. Models using other environmental parameters are thus required to supplement geolocation data in the estimation of marine migratory routes. The objective of this work was to create a simple method that would estimate the migratory pathways of Atlantic salmon (Salmo salar) in polar regions using temperature and depth recordings. Validated geolocations from PSATs were used to test and constrict the model. The model’s predicted migratory routes were within 100 km of the light‐based geolocations calculated by the tags. By constraining the trajectories through the geolocations, bias was reduced. Sensitivity analyses demonstrated that slight alterations of the location and timing of the start and end points did not affect the mean migratory route estimates. This method is a management tool that can determine the primary habitat areas for any surface‐ or bottom‐dwelling marine species – especially in polar regions, where other methods may be impossible.     

Cod and haddock spawning on Georges Bank in relation to water residence times

The seasonal and geographical patterns of spawning for Georges Bank cod ( Gadus morhua ) and haddock ( Melanogrammus aeglefinus ) are estimated using composite distributions of stage I eggs derived from the Marine Resources Monitoring, Assessment and Prediction (MARMAP) data set (1977–1987). The inferred mean spawning locations are compared with patterns in particle residence times derived from a three-dimensional prognostic circulation model that estimates the tidal and seasonal mean circulation. The comparisons indicate that cod and haddock spawning occurs at times and locations characterized by model residence times in excess of 35 d. The results are discussed in the context of fish population regulation theory, particularly the member/vagrant hypothesis and the concept of abundance-dependent vagrancy.     

Influences of mean advection and simple behavior on the distribution of cod and haddock early life stages on Georges Bank

Results of a modeling study designed to explore the influences of physical advection and certain biological mechanisms on the distribution of cod (Gadus morhua) and haddock (Melarwgrammus aeglefinus) early life stages on Georges Bank are described. Using a late-winter/early-spring 3-D circulation field driven by the M₂ tidal current, mean wind stress and Scotian Shelf inflow, we examine the distribution of cod and haddock larvae spawned on the Northeast Peak of the Bank. The sensitivity to a March-April baroclinic field is also explored. Results indicate that larvae remaining in the surface Ekman layer are generally advected off-bank. However, downwelling associated with Ekman layer convergence near the shelf break provides a mechanism for larvae to exit from the off-bank surface drift. Larvae below the surface layer are transported south-westward along the southern flank of Georges Bank and are retained on the Bank if their position immediately upstream of the Great South Channel is shoalward of (roughly) the 70 m isobath. Within the Great South Channel region and between the 50 and 70 m isobaths, retention can depend on the phase of the tide. Spawning shoalward of the 50 m isobath on the Northeast Peak greatly increases the chances of retention. These results apply to passive larvae and to those with specified vertical distributions and migration based on observations. Directional on-bank swimming at rates of 0.5 to 1 body length per second would substantially enhance shoalward displacement, resulting in larval distributions during the first 2 months that are consistent with field observations.     

Optimum temperature and food-limited growth of larval Atlantic cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) on Georges Bank

We estimated recent growth of Atlantic cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) larvae collected on the southern flank of Georges Bank in May 1992–94 from the ratio of RNA to DNA (R/D) and water temperature. Growth of both species increased with water temperature to about 7°C and then decreased. The highest growth rates were observed in May 1993 at water temperatures around 7°C. These data confirm an earlier observation of comparable temperature optima for growth of Atlantic cod and haddock larvae in the north‐west Atlantic. Comparisons of field growth rates and temperature optima with data for larvae cultured at high temperatures and prey densities in the laboratory suggest that growth may have been food‐limited at higher temperatures on Georges Bank. Given that 7°C is the long‐term mean water temperature on the southern flank in May and that climate models predict a possible 2–4°C rise in water temperatures for the western North Atlantic, our findings point to a possible adverse effect of global warming on Atlantic cod and haddock.     

Winter distribution and movements of northern Atlantic cod (Gadus morhua) along the Newfoundland—Labrador continental shelf edge derived from observations on commercial trawlers

Catch-effort data recorded by observers aboard commercial trawlers fishing the Newfoundland-Labrador continental shelf during the winters of 1980–1991 were used to define the distribution and movements of pre-spawning and spanning Atlantic cod, Gadus morhua. Although cod were widely distributed over the outer continental shelf in the months of January to April, commercial concentrations were consistently located in three areas along the shelf edge. These three areas, constituting only 16% of the 190000 km² fishing ground for northern cod, were centred north-east of Belle Isle Bank, north-east of Funk Island Bank, and between Funk Island Bank and the northern Grand Bank Tow positions with catch rates 5000 kg h^-1 were plotted on a weekly basis to define movements of cod schools during the winters of 1986–1988. It was assumed that change in the collective position of trawlers taking large quantities of cod reflected shoal movements. Analyses indicate that cod moved southeast along the shelf edge in January and February of each year, crossing from NAFO Division 2J into 3K Each March, cod in Division 3K apparently moved northwest against prevailing ocean currents into the Hamilton Bank region of 2J. These movements do not appear to be a direct response to ocean temperature or pack ice conditions, although cod moving along the shelf break were generally in bottom waters of 24°C. We suggest these patterns reflect the migratory behaviour of pre-spawning and spawning 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.     

Abundance estimation of Northeast Atlantic mackerel based on tag recapture data—A useful tool for stock assessment?

In the present study we utilize tag recapture data to estimate year class abundance and spawning stock biomass of mackerel (Scomber scombrus L.) in the Northeast Atlantic for the period 1986–2008. On average 20,000 jigged mackerel have been tagged annually with internal steel tags in the spawning area west of Ireland and the British Isles, and the tags have been recaptured in commercial catches screened through metal detectors. The spawning stock biomass estimates derived from two different tag-based models were highly variable but were on average 2 and 2.3 times higher than the ICES official estimate. The official estimate is considered uncertain and most likely an underestimate of the actual biomass, due to unregistered mortality in the fisheries and lack of fishery-independent, age-disaggregated data. Hence, tag-based estimates could potentially improve the current assessment if included in the ICES stock assessment on a regular basis. These estimates also involve some uncertainty that needs consideration, especially related to variable tagging mortality, detector efficiency and migrations of the stock.     

Conserving a subpopulation of the northern Atlantic cod metapopulation with a marine protected area

• 1. Marine reserves can play an important role in the conservation of subpopulations of marine fish metapopulations. The population spatial structure of northern Atlantic cod of Newfoundland and Labrador has characteristics of a metapopulation. Subpopulations of northern Atlantic cod on the continental shelf were decimated by decades of overfishing, and have not recovered. The remaining northern cod are concentrated in coastal areas.
• 2. A Marine Protected Area (MPA) was established in Gilbert Bay, Labrador by the Government of Canada in 2005 to protect the bay's resident subpopulation of northern Atlantic cod. Conservation of Gilbert Bay cod will help protect the genetic diversity of the northern cod metapopulation.
• 3. Unlike some other MPAs, Gilbert Bay is not a harvest refugium or ‘no‐take’ reserve. Aboriginal subsistence fisheries for salmonids with a bycatch of cod are allowed in designated areas of the MPA. A recreational fishery for Atlantic cod by angling open to all people is under consideration. Management of the MPA must ensure that fishing activities do not endanger the local cod population.
• 4. The population dynamics of Gilbert Bay cod were simulated using an age‐structured Leslie matrix model to estimate the total mortality under various recreational fishing scenarios. The level of sustainable harvest by a recreational fishery depends on the natural mortality of the Gilbert Bay cod population, which is unknown. Therefore, there is risk in permitting a recreational fishery in the MPA.
• 5. There may be benefits to the northern cod metapopulation, if the Gilbert Bay subpopulation is allowed to rebuild to the carrying capacity of the bay. If the abundance of Gilbert Bay cod exceeds the level which the local marine ecosystem can support, some cod may emigrate from the bay and recolonize adjacent coastal areas. The potential for Gilbert Bay cod to recolonize continental shelf areas is less certain.

Copyright © 2008 John Wiley & Sons, Ltd.     

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.     

Larval lobster (Homarus americanus) distribution and drift in the vicinity of the Gulf of Maine offshore banks and their probable origins

Surveys for lobster larvae in offshore waters of the north‐eastern Gulf of Maine in 1983, 1987 and 1989 confirm that local hatching occurs mainly at depths <100 m over the banks, including Georges and Browns Banks. Detailed studies in the vicinity of Georges Bank in late July of both 1987 and 1989 indicate that the first and second moult stages were located primarily over the bank whereas stages III and IV lobster were collected both over and off the bank. At times stage IV lobster were more abundant off the bank than over it. The condition of stage III and IV lobster, as measured by a lipid index, was better off than over Georges Bank in 1988 and 1989 indicating a possible physiological advantage to being off the bank. In addition, the higher surface temperatures off Georges Bank would shorten larval development time to settlement. To determine the probable hatch sites of stage IV lobster collected off of Browns Bank in 1983 and off of Georges in 1987 and 1989, a 3‐D circulation model of the Gulf of Maine was used to simulate larval lobster drift backwards in time. In all cases, areas off Cape Cod, MA, and off Penobscot Bay, ME were suggested as the source of the larvae, although most of the larval trajectories never reached these near‐shore waters that are well‐known, larval hatching areas. The model‐projected larval release times match most closely the observed inshore hatch off Massachusetts but model uncertainties mean that coastal Maine cannot be ruled out as a source. Georges Bank is also a potential source because the present model does not take into account short‐term wind events, off‐bank eddy transport or the possibility of directed off‐bank larval swimming. Examination of weather records prior to and during our 1988 and 1989 sampling periods indicates that winds were not of sufficient intensity and duration to induce larval transport off Georges Bank. The shedding of eddies from the northern flank of Georges Bank into the Gulf of Maine are a relatively common phenomenon during summer but not enough is known about them to evaluate their contribution to possible cross‐bank transport of lobster larvae. Directed larval swimming is another possible source for the stage IV lobster found near Georges Bank. Plankton distributions across the northern frontal zone of Georges Bank in 1988 were used as proxies for the scarce larval lobsters. The more surface distribution of the microplankton, in particular, supports the possibility that wind and eddy events may be important in the transport of stage III and IV lobsters off of Georges Bank. Further studies are needed to evaluate these possible additional sources of advanced stage lobster larvae found off of the offshore banks.     

Horizontal movements of bigeye tuna (Thunnus obesus) near Hawaii determined by Kalman filter analysis of archival tagging data

Geolocation data were recovered from archival tags applied to bigeye tuna near Hawaii. A state‐space Kalman filter statistical model was used to estimate geolocation errors, movement parameters, and most probable tracks from the recovered data. Standard deviation estimates ranged from 0.5° to 4.4° latitude and from 0.2° to 1.6° longitude. Bias estimates ranged from ?1.9° to 4.1° latitude and from ?0.5° to 3.0° longitude. Estimates of directed movement were close to zero for most fish reaching a maximum magnitude of 5.3 nm day^?1 for the one fish that moved away from its release site. Diffusivity estimates were also low, ranging from near zero to 1000 nm² day^?1. Low values of the estimated movement parameters are consistent with the restricted scale of the observed movement and the apparent fidelity of bigeye to geographical points of attraction. Inclusion of a time‐dependent model of the variance in geolocation estimates reduced the variability of latitude estimates. The state‐space Kalman filter model appears to provide realistic estimates of in situ geolocation errors and movement parameters, provides a means to avoid indeterminate latitude estimates during equinoxes, and is a potential bridge between analyses of individual and population movements.     

Movement of adult male snow crab, Chionoecetes opilio, in the southern Gulf of St. Lawrence and eastern Nova Scotia, Canada

A total of 12,755 adult male snow crabs (Chionoecetes opilio) were tagged with spaghetti–Carlin tags in 33 sites throughout the southern Gulf of St. Lawrence (sGSL) and eastern Nova Scotia (eNS) between 1993 and 2003. Of the 1971 tag returns obtained between 1994 and 2004, 1703 had accurate geographical recapture positions. The average distance travelled was 16.7 km for sGSL (maximum 165 km) and 61.5 km for eNS (maximum 368 km). The release site and the number of days at liberty before recapture were significant factors affecting the distance travelled by the crabs. Size had no significant effect on the distance travelled or the direction of movement. The present observations support the sedentary nature previously reported for the sGSL snow crab stock, but not the eNS stock, where the movement of snow crab was more dynamic. In both regions, there were movements between management areas, which indicates that managing the fisheries in discrete units is not warranted.     

Juvenile cod (Gadus morhua) mortality and the importance of bottom sediment type to recruitment on Georges Bank

The demersal settlement of pelagic juvenile fish has been considered a critical period in which the final adjustment is made to the size of a year class. Distribution patterns of pelagic and recently settled juvenile cod (Gadus morhua) were examined from nine surveys on Georges Bank during the summer over 5 years, 1984–1989, to relate juvenile survival to the sedimentary environment. Pelagic juveniles were widespread across Georges Bank in June, and by mid‐July they occurred on all bottom types from sand to gravel on eastern Georges Bank. However, by late July‐early August they were mostly abundant on the northeastern edge gravel deposit, which with its complex relief, provides abundant prey and refuge from predators. A bank‐wide estimate of pelagic juvenile abundance in 1986 and 1987 was used to assess mortality of the recently settled juveniles and to evaluate the relative importance of survivors from the northeastern edge gravel area to recruitment of the Georges Bank population. Settlement mortality rates over 1–2 months on the northeastern gravel area ranged from 3 to 8% day^?1, which compared reasonably with other studies. The seasonal abundance of the pelagic juveniles was almost an order of magnitude higher in 1987 than 1986; however, recruitment at age 1 was similar, indicating that a high mortality of the demersal juveniles occurred in 1987. The limited northeastern gravel area on Georges Bank may represent a survival bottleneck depending on the variability in the distribution and abundance of juvenile cod settlement in relation to that of their predators.     

Sensitivity analysis of sea scallop (Placopecten magellanicus) larvae trajectories to hydrodynamic model configuration on Georges Bank and adjacent coastal regions

The previous larval-trajectory modeling studies on Georges Bank were assessed through process-oriented Lagrangian-tracking comparison experiments using the high-resolution Gulf of Maine/Georges Bank Finite-Volume Coastal Ocean Model (GOM-FVCOM). The results indicate that in a strong nonlinear system such as Georges Bank, the passive tracer movement is driven by a fully three-dimensional Lagrangian flow field that varies in space and time due to large tidal excursion and steep bottom topography. The particle-tracking methods developed based on the assumption of weak nonlinearity of the flow field are not applicable to Georges Bank. The results of previous larval transport studies driven by circulation fields constructed under the weak-nonlinearity assumption need to be interpreted with caution. In the present work, the influence of model physical setups on sea scallop larval dispersal and settlement on Georges Bank and adjacent shelf regions is examined. Distinct differences in the spatial distribution of the passive larvae predicted by the model under various physical conditions suggest that a fully nonlinear model driven by realistic spatially and temporally varying forcing should be employed for Lagrangian-based studies of fishery population dynamics on Georges Bank.     

Characterization of seabird bycatch in eastern Canadian waters, 1998–2011, assessed from onboard fisheries observer data

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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.     

Recruitment variability and oceanographic stability

We examine the hypothesis that recruitment is more variable in populations on isolated offshore banks than nearby shelf populations. Recruitment of cod (Gadus morhua) and American plaice (Hippoghssoides plates-soides) on Flemish Cap is more variable than in any comparable population. Recruitment of haddock (Melanogrammus aeglefinus) on Rockall Bank is also more variable than in surrounding populations. These results are confirmed both by estimates obtained by virtual population analysis and by research surveys. Recruitment of haddock and herring (Clupea harengus) on Georges Bank is also more variable than in surrounding populations; however, the results for two other groundfish populations, cod and yellowtake flounder (Limanda ferruginae), on Georges Bank are ambiguous. We conclude that marine fish populations on isolated banks are more variable than those on nearby shelf regions.     

Vertical habitat use of sailfish (Istiophorus platypterus) in the Atlantic and eastern Pacific,derived from pop‐up satellite archival tag data

Vertical habitat use of sailfish (Istiophorus platypterus) was evaluated using pop‐up satellite archival tag data from the eastern tropical Atlantic, western North Atlantic, and eastern tropical Pacific. Data included Argos transmitted depth, temperature, and light level frequency histograms binned at 1–8‐h intervals, and four recovered pop‐up satellite archival tags that provided high resolution archival data recorded at 30‐s intervals. We tabulated the proportions of time spent within each degree of water temperature relative to the surface temperature (Delta T) and proportions of time at temperature, as these are major input variables for habitat standardization models used in stock assessment procedures. Frequency distributions were calculated for daylight, darkness, and twilight for each of the three regions and for all regions combined. Vertical habitat envelopes indicated greater use of deeper strata in the western North Atlantic, compared to the hypoxia‐based habitat compressed regions of the eastern Atlantic and Pacific. However, there were no significant differences in Delta T distributions when comparing the three regions, affirming this metric for its application in habitat standardization models.