Modelling the transport of larval sandeels on the north-west European shelf

This study considers the interannual variability in the advection of sandeel ( Ammodytes marinus ) larvae on the north-west European shelf. We use recently collated data on spawning location, age of larvae and the time of hatching (between 1955 and 1993) as input to a particle-tracking model. Utilizing hourly currents from a 39 year run of a two-dimensional sea circulation model, cohorts of sandeel larvae were represented as a distribution of particles in space and time, and particle drift was calculated for the larval life stage. The final distributions of particles were compared with actual distributions of 0-group sandeels in the North Sea, as determined from surveys conducted at around the end of the larval phase. Preliminary examination of the data between 1973 and 1993 indicates that differences in predicted drift could explain some of the large interannual variability in 0-group distributions. This approach may help to improve our understanding of the frequent colonization events that are typical of this species.     

Wind‐induced variability in coastal larval retention areas: a case study on Western Baltic spring‐spawning herring

The investigation of larval dispersal and retention, their variability and dependence on wind conditions, has become a major topic in fisheries research owing to potential effects on stock recruitment and stock structuring. The present study quantifies the wind‐induced variability of larval retention of herring in a highly productive coastal lagoon of the Western Baltic Sea. This lagoon, the Greifswalder Bodden, represents the main spawning area of Western Baltic Spring‐Spawning Herring, a stock that has recently undergone a continuous decline in recruitment. The study tests whether this decline was related to changes in larval retention, more precisely to changes in wind conditions, the main forcing of the lagoon's circulation. To answer this, a model approach was applied. Larvae were tracked as Lagrangian drifters under constant and variable wind conditions, examining the main drift patterns and reconstructing the incidents during the period of recruitment decline. For the latter, weekly cohorts of virtual larvae were released in the lagoon over the entire spawning period (April–June; >16 weeks). The fraction of retained larvae per cohort was related to observed larval abundances. On this basis, a new retention index was defined to evaluate the annual larval retention. The results presented cannot explain the observed recruitment decline but characterize the lagoon as an important larval retention area by virtue of unsteady wind conditions that prevent a steady outflow of larvae.     

The recruitment process of the Barents Sea capelin (Mallotus villosus) stock, 2001–2003

Oceanographic and predation processes are important modulators of fish larvae survival and mortality. This study addresses the hypothesis that immature Norwegian spring‐spawning herring (Clupea harengus), when abundant in the Barents Sea, determine the capelin reproduction success through consumption of Barents Sea capelin (Mallotus villosus) larvae. Combining a hydrodynamic model and particle‐tracking individual‐based model, a realistic spatio‐temporal overlap between capelin larvae and predatory immature herring was modelled for the summer seasons of 2001–2003. Capelin larvae originating from western spawning grounds became widely dispersed during the summer season, whereas those originating from eastern spawning grounds experienced a rapid drift into the southeastern Barents Sea. Herring caused a 3% mortality of the capelin larvae population in 2001 and a 16% mortality in 2003, but the effect of predation from herring on capelin larvae was negligible in 2002. Despite a strong capelin larvae cohort and a virtual absence of predatory herring, the recruitment from the capelin 2002 year class was relatively poor from a long‐term perspective. We show that the choice of capelin spawning grounds has a major impact on the subsequent capelin larvae drift patterns, constituting an important modulator of the capelin larvae survival. Variation in drift patterns during the summer season is likely to expose the capelin larvae to a wide range of hazards, including predation from young cod, sandeel and other predators. Such alternative predators might thus have contributed to the poor capelin recruitment during 2001–2003, leading to the collapse of the capelin stock in the subsequent years.     

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.     

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.     

Recent decline in cod stocks in the North Sea–Skagerrak–Kattegat shifts the sources of larval supply

Cod stocks in the North Sea, including the Kattegat and the Skagerrak, have declined dramatically since the 1970s. Occasionally there is a high recruitment of juveniles in Kattegat/Skagerrak, without leading to the rebuilding of adult cod stocks despite reduced fishing mortality. In a biophysical model of egg and larval drift, we examined the potential importance of extant and historical spawning grounds for recruitment of cod in the Kattegat/Skagerrak seas using data of spawning stock biomass from the 1970s and from today's reduced stocks. The results suggest that Kattegat in the 1970s relied on largely locally retained (83%) larvae with little annual variation in recruitment. Kattegat also provided a substantial proportion of larvae recruiting in Swedish Skagerrak (72%). This is in contrast to present conditions where the Kattegat spawning stock has been reduced by 94%, and Kattegat only provides 34% of locally retained larvae and 30% to Swedish Skagerrak. Instead, the protected area in the Öresund and the Belt Sea are expected today to provide most larvae recruiting in Kattegat. Also, the inflow of larvae from the North Sea to Skagerrak and Kattegat can be significant although highly variable between years, with a positive correlation to the North‐Atlantic Oscillation index (NAO). The rebuilding of healthy spawning areas in the Kattegat may be key for restoring local cod stocks in both Kattegat and along the Skagerrak coast. This poses a management challenge if cod with local ‘Kattegat’ adaptations, e.g., in terms of egg density and migration patterns, are lost or reduced to non‐resilient densities.     

Productivity and recovery of forage fish under climate change and fishing: North Sea sandeel as a case study

Forage fish occupy a central position in marine food‐webs worldwide by mediating the transfer of energy and organic matter from lower to higher trophic levels. The lesser sandeel (Ammodytes marinus) is one of the ecologically and economically most important forage fish species in the North‐east Atlantic, acting as a key prey for predatory fish and sea birds, as well as supporting a large commercial fishery. In this case study, we investigate the underlying factors affecting recruitment and how these in turn affect productivity of the North Sea sandeel using long‐term data and modelling. Our results demonstrate how sandeel productivity in the central North Sea (Dogger Bank) depends on a combination of external and internal regulatory factors, including fishing and climate effects, as well as density dependence and food availability of the preferred zooplankton prey (Calanus finmarchicus and Temora longicornis). Furthermore, our model scenarios suggest that while fishing largely contributed to the abrupt stock decline during the late 1990s and the following period of low biomass, a complete recovery of the stock to the highly productive levels of the early 1980s would only be possible through changes in the surrounding ecosystem, involving lower temperatures and improved feeding conditions. To that end, we stress the need for ecosystem‐based management accounting for multiple internal and external factors occurring within the broader context of the ecosystem in which forage fish species, such as sandeel, play an important and integral part.     

Variations in the drift of larval cod (Gadus morhua L.) in the Baltic Sea: combining field observations and modelling

Coupled three-dimensional (3-D) physical oceanographic modelling and field sampling programmes were carried out in May 1988 and August 1991 to investigate the potential drift of larval cod ( Gadus morhua L.) in the Bornholm Basin of the Baltic Sea. The goals were to predict the transport of cod larvae, thus aiding the identification of physical processes influencing larval retention/dispersal. Numerical simulations were performed using a 3-D eddy-resolving baroclinic model based on the Bryan–Cox–Semtner code adapted for the Baltic Sea. Within the Bornholm Basin, the model was initialized with ground truth data of physical parameters obtained on the research cruises, and all simulations were forced with actual wind data. Outside the basin, generalized hydrographic features of the Baltic Sea were utilized by incorporation of simulated hydrographic fields from previous model runs typical for the time periods considered. Larval drift was simulated either by incorporation of passive drifters, or as the initial horizontal distribution of larvae implemented into the model. Drift model simulations of larval transport agreed relatively well with field observations. The influence of variations in the vertical distribution on a smaller scale, i.e. vertical deviations of ± 6 m from the observed mean centre of mass, on the drift was examined, revealing no significant differences in the drift of larvae depending on their vertical distribution. The different wind forcing during the investigated time periods was linked to a retention situation in May 1988 and to a dispersal situation in August 1991. Finally, observed spatial distribution patterns of 1-group cod based on Baltic Young Fish Surveys (BYFS) were compared with their predicted transport in the larval phase and examined with respect to recruitment.     

The influence of life history dynamics and environment on the determination of year class strength in North Sea herring (Clupea harengus L.)

The inter-annual variability in year class strength (1976–2000) of North Sea herring (Clupea harengus) was investigated using Paulik diagrams based on survey data and Virtual Population Analysis. The herring life cycle was split into five stages: spawning stock biomass (SSB), egg production, larvae, fish with 0 winter rings on the otolith (0-wr), 1-wr and 2-wr. Surveys were used as indices and Paulik analysis revealed relationships between stages. In 80% of the years, year class strength reflected SSB. Poorer than expected year classes were determined during the larva to 0-wr phase, whilst stronger than expected year classes were apparently determined during the 0-wr to 1-wr stage. There was no clear relationship between survival of young stages of herring and the abundance of Calanus finmarchicus but the year class strength of 0-wr and 1-wr had a negative relationship to bottom water temperature. Lower sea water temperatures in the North Sea are associated with higher Calanus abundance. The analysis shows that the strength of aberrant year classes of North Sea herring is determined between the pelagic larval and the juvenile stages.     

Effects of temperature, wind and spawning stock biomass on recruitment of Arcto-Norwegian cod

Sea temperature has earlier been shown to have a large influence on the recruitment of Arcto-Norwegian cod, Gadus morhua. We here hypothesize that this linkage is partly due to the direct effect of temperature on larval and juvenile growth. Secondly, temperature acts as a proxy for both biotic and abiotic factors influencing recruitment. Indices of abundance of early juvenile cod (2–3 months old), 0-group cod (4–5 months old) and 3-year-old cod are analysed in more detail against the environmental temperature, wind stress components, wind-induced turbulent energy and the spawning stock biomass. To deal with autocorrelation, non-stationar-time and nun-normality, which complicate a statistical time series analysis, randomization and Box-Jenkins methods are applied. In addition to the important effect of high sea temperature during the early life stage in forming strong year classes, the results show that the spawning stock biomass is nearly as important. Also, alongshore southerly wind stress anomalies during the period of pelagic drift (from April through summer) and offshore wind stress anomalies during egg and early larval stages (in April) act favourably on recruitment. The beneficial effect of southerly wind anomalies could he linked partly to high temperature, but the flux of zooplankton-rich water from the Norwegian Sea into the feeding areas of the Barents Sea may also be increased. The favourable influence of offshore winds in April is less predominant and causal links are also less clear; possible explanations for this might be increased offshore spreading of eggs and early larvae, resulting in reduced risk of predation, and increased compensation inflow of intermediate Norwegian Sea water which, in this restricted period of time, has a high concentration of spawning copepods suitable as prey for the developing cod larvae.     

Bio-physical modelling of the early life stages of haddock, Melanogrammus aeglefinus, in the North Sea

An individual-based modelling approach was developed to investigate the spatial and temporal patterns in the recruitment processes of North Sea haddock, Melanogrammus aeglefinus . The approach was based on the realization that the survivors to recruitment of an annual cohort are most probably not drawn at random from the initial population of eggs, but represent the fastest-growing individuals. Individual growth rates reflect the unique exposure of each larva to the environment along its drift trajectory. In this context, the environment refers to a wide range of factors affecting growth such as food, turbulence and temperature. A combination of a model of egg production by the adult stock, a particle-tracking scheme, and a model of larval growth and mortality rate was used to simulate the dispersal trajectories, and the survival of haddock larvae spawned at different times and locations on the continental shelf. The particle tracking was driven by flowfields from a climatological implementation of the Hamburg Shelf–Ocean Model (HAMSOM) for the North Sea and NE Atlantic. The system was able to resolve spatial and temporal patterns in the recruitment process and indicated that the surviving population of larvae was drawn from a restricted part of the spawning distribution. The results have the potential to guide the development of future conservation measures in fisheries management.     

Modelling recruitment dynamics of hake, Merluccius merluccius, in the central Mediterranean in relation to key environmental variables

Hake recruitment has been examined in relation to environmental variables in two of the main reproductive areas of the central Mediterranean, the northern and central Tyrrhenian Sea. Seventeen years time series data from trawl surveys revealed high fluctuations in recruit abundance that could not be just explained by spawning biomass estimations. Generalized additive models were developed to investigate hake recruitment dynamics in the Tyrrhenian Sea in relation to spawner abundance and selected key oceanographic variables. Environmental data were explored in attempt to explain survival processes that could affect early life history stages of hake and that accounted for high fluctuations in its recruitment.Thermal anomalies in summer, characterised by high peaks in water temperature, revealed a negative effect on the abundance of recruits in autumn, probably due to a reduction in hake egg and larval survival rates. In the northern Tyrrhenian, recruitment was reduced when elevated sea-surface temperatures were coupled with lower levels of water circulation. Enhanced spring primary production, related to late winter low temperatures could affect water mass productivity in the following months, thus influencing spring recruitment. In the central Tyrrhenian a dome-shaped relationship between wind mixing in early spring and recruitment could be interpreted as an “optimal environmental window” in which intermediate water mixing level played a positive role in phytoplankton displacement, larval feeding rate and appropriate larval drift. Results are discussed in relation to the decline in hake stock biomass and within the present climate change and global warming context.     

Does larval mortality influence population dynamics? An analysis of North Sea herring (Clupea harengus) time series

The daily mortality rates of North Sea herring early‐stage larvae are found to vary over decades. Larval abundance data were used with a spatio‐temporal oceanographic model to reconstruct temperature histories of the observed larvae. The histories were used in conjunction with a temperature‐based growth model to estimate larval age. Mean daily mortality rates were then estimated for the four spawning components (Downs, Banks, Buchan and Orkney/Shetland) using the vertical life table approach, which considers instantaneous abundances across all ages rather than following distinct cohorts. All spawning components, but especially Downs (in the south), exhibited a steady rise in mortality associated with increasing population size. In addition, the three northern components shared a distinct trend in mortality that was significantly correlated with ambient water temperatures experienced by the larvae during the respective time periods after hatching. This trend was also significantly negatively correlated with the residuals of the whole stock‐recruitment relationship. These findings were generally robust to assumptions about growth and hatch length of larvae. The compensatory increase in productivity in the late 1980s and poor recruitment since 2000 coincide with changes in the mortality of larvae younger than 30 days post hatch and covary with larval density and temperature. Thus we suggest that the mortality of early‐stage larvae does impact on the population dynamics in North Sea herring in its current productivity regime, implying a critical period in the determination of year class strength.     

From egg to juvenile in the Bay of Biscay: spatial patterns of anchovy (Engraulis encrasicolus) recruitment in a non‐upwelling region

In this study the spatial distribution of eggs, larvae and juveniles of European anchovy (Engraulis encrasicolus) was followed in 2004 and 2005 during three consecutive cruises each year in spring–summer to test what the limits of retention are in a non‐upwelling area. Eggs, small larvae and large juveniles were mainly distributed over the shelf, whereas large larvae and small juveniles were found mainly off the shelf. Although overall distributions were similar, the 2 yr differed in that there was more of a coastal distribution of individuals in 2004, whereas in 2005 more individuals were found off the shelf. There were no significant differences in the length–weight relationships for individuals found on and off the shelf or between years. The correspondence in circulation patterns and the lack of difference in the length–weight relationships suggest that a single population is present, larvae drifting off the shelf due to currents and returning as mobile juveniles. Quantile regression analysis of the long‐term recruitment index suggests that transport off the shelf may favour good recruitments. This would suggest that in non‐upwelling regions the retention area resulting in good recruitment may not be restricted to the shelf.     

Modelling the dispersal of herring and hake larvae in the Strait of Georgia for the period 2007–2009

The Strait of Georgia (SoG), between Vancouver Island and mainland British Columbia, is a larval rearing ground for both hake and herring stocks, which are commercially important. Year‐to‐year variability in larval retention within the strait is examined by simulating drift tracks of larvae for these species using an ocean circulation model and a particle‐tracking model. Larvae with different vertical swimming behaviors were tracked in the springs of 2007, 2008, and 2009. Since herring larvae mostly stay near the surface, their distribution is heavily influenced by the wind. Strong winds to the north soon after the hatching period tend to wash herring larvae out of SoG and winds to the south help retain herring larvae inside the Strait. In 2007, the model indicates a massive wind‐driven export of herring larvae which may have led to the observed failure of herring production. In contrast, hake larvae reside deeper in the water column (50–200 m). Their distribution is less sensitive to surface forcing but is shaped by a deep gyre with cross‐strait currents. This study also suggests that the northern and southern SoG are weakly connected for herring larvae dispersal, which makes both regions potentially important to recruitment.     

Larval transport and retention of the spiny lobster, Panulirus argus, in the coastal zone of the Florida Keys, USA

The spiny lobster Panulirus argus is of ecological and commercial importance in the South Florida coast of the continental USA and throughout the Intra‐Americas Sea. Essential spiny lobster habitat in South Florida is primarily located in the Florida Keys coastal zone (including the Dry Tortugas), where the dynamic regional circulation coupled with the long planktonic larval duration (6–12 months) of P. argus raises questions of larval retention and recruitment. Locally spawned phyllosomata entrained in the Florida Current are likely to be expatriated out of the Straits of Florida, which implies that the local spiny lobster population is sustained by the transport of larval recruits from upstream locations. We examined the physical processes that may influence recruitment. Transport processes in the Keys coastal zone are spatially variable. Observed and modelled data suggest that the upper Keys is a point of onshore larval transport via the inshore meandering of the Florida Current, and the lower Keys to Dry Tortugas region apoint of retention through wind‐driven onshore/countercurrents and eddy recirculation. Eddies that propagate between the Dry Tortugas and the lower Keys facilitate the exchange of larvae between the Florida Current and the coastal zone. Northerly wind events associated with cold fronts can enhance recirculation of larvae in the upper Keys. The association of older larvae with the Florida Current front supports the hypothesis that spiny lobster larval recruits come from upstream sources in the Caribbean.     

Kittiwake breeding success in the southern North Sea correlates with prior sandeel fishing mortality

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The importance of individual behaviour for successful settlement of juvenile plaice (Pleuronectes platessa L.): a modelling and field study in the eastern Irish Sea

To study the transport of plaice (Pleuronectes platessa L.) eggs and larvae in the eastern Irish Sea, we constructed a 3D‐baroclinic physical model and coupled it to a particle‐tracking scheme that allowed aspects of larval behaviour to be simulated. Starting positions for eggs were based upon data from a series of ichthyoplankton surveys and final positions were compared with results of settled plaice distributions from two beam trawl surveys conducted on beaches around the eastern Irish Sea. If simulated larval behaviour was limited to passive drift or horizontal swimming, the particles diffused away from the spawning areas but failed to reach nursery grounds in significant numbers (85–90% remaining offshore). In contrast, switching on circatidal vertical swimming significantly increased the numbers of larvae reaching the coast (only 23–30% remained offshore). Particles tended to accumulate in bays and estuaries and this pattern compared well with the distribution of settled plaice from the field surveys. Studies in the southern North Sea (where spawning and nursery grounds are widely separated) have also demonstrated the importance of selective tidal stream transport for successful recruitment of settling plaice to nursery grounds. Although our understanding of the ontogeny of this behaviour is still poor, the model results presented suggest that this aspect of behaviour is a key factor influencing plaice settlement success.     

Migrations and hydrography determine the abundance fluctuations of blue whiting (Micromesistius poutassou) in the Barents Sea

The Barents Sea is the north‐eastern fringe of the distribution of blue whiting (Micromesistius poutassou). Fluctuations in distribution and abundance of blue whiting in the area have been marked. Two hypotheses are put forward to explain these fluctuations. First, rich year classes in the main Atlantic stock of blue whiting may contribute to increased abundance in the Barents Sea. Second, variations in hydrography, such as influx of warm Atlantic water, may be particularly important in this fringe area. We investigated these hypotheses using data from bottom trawl surveys conducted during the period 1981–2006. Variations in abundance (measured either as incidence or density) and distribution were correlated with recruitment in the Atlantic stock of blue whiting as well as hydrographic conditions. Regression analyses indicated that the abundance fluctuations are primarily determined by variations in recruitment of Atlantic blue whiting, a strong year class leading to high abundance in the Barents Sea the year after spawning. However, salinity anomaly in the Fugløya–Bear Island transect during the previous year, an indicator of high inflow of Atlantic water, had also a significant, positive effect. Thus, the data suggested a climatic modulation of dynamics that were primarily determined by recruitment of blue whiting in the main Atlantic stock. Analyses of size structure as well as earlier studies on population genetics supported this conclusion.     

Modelling the advection and diffusion of eggs and larvae of Greenland halibut (Reinhardtius hippoglossoides) in the north-east Arctic

Since the late 1980s there has been considerable uncertainty in recruitment levels of the north‐east Arctic stock of Greenland halibut (Reinhardtius hippoglossoides). The abundance of several year classes, originally considered very low at 0–3 yr age, appeared higher than expected at the age of 6+. This may be due to poor targeting of recruitment surveys of the younger year classes. The present work considers the transport and dispersion of eggs and larvae of Greenland halibut by numerical modelling in order to predict the locations of the initial recruitment grounds. Current fields from a 3D baroclinic hydrodynamic model are fed into a Lagrangian particle‐tracking model developed for the Barents Sea area. The particles are released into the current at the spawning field along the shelf slope from Lofoten to Bear Island (69–75°N). Vertically, the particles can follow a predefined depth‐by‐age curve or be kept at a fixed depth. This model system is used for different years to examine changes in the drift pattern. The results indicate that spawning location, transport depth and inflowing activity to the Barents Sea are important factors influencing the distribution of juveniles.