Environmental factors and recruitment of mackerel, Scomber scombrus L. 1758, along the north-east Atlantic coasts of Europe

Increasingly, scientific publications refer to some of the environmental factors affecting the recruitment in fish species. However, presently, there is little information available concerning the influence of the environment on the recruitment of Scomber scombrus, the North-east Atlantic mackerel (NEAM). In this contribution, North Atlantic Oscillation (NAO), upwelling and turbulent mixing (or `turbulence') series were compared with recruitment estimates, at age-0. The results show that the southern prespawning migration pattern of the Atlantic mackerel is directed towards areas with low turbulent mixing at spawning time, providing a `stable environment' for egg and larval survival. In the southern areas, where the spawning starts, the turbulence conditions of prespawning and spawning periods have the largest influence on the success of recruitment; this is related, possibly, to the more `stable' weather in the subsequent months and for the remainder of the year. In contrast, in the northern areas, the role of turbulence over the entire year becomes increasingly more relevant; this is related, possibly, to the high levels of turbulence during autumn and winter, which may become limiting to the survival of juveniles. About 50% of the variability in the Atlantic mackerel recruitment may be explained by means of environmental variables, such as turbulence. Other variables, such as upwelling and NAO, are only slightly, or not, statistically significant.     

Tracking shifts in Atlantic mackerel (Scomber scombrus) larval habitat suitability on the Northeast U.S. Continental Shelf

Climate change has altered the oceanographic environment and subsequently the habitats of marine species. Fish and invertebrate populations’ responses to habitat include movement with latitude and depth to remain within their fundamental niches. The northwest Atlantic mackerel (Scomber scombrus) population has fluctuated over the last century due in part to changes in the environment. We used species distribution models to understand the influence of the physical (temperature) and biological (zooplankton) environment on mackerel larval abundance, and how such relations have determined larval habitat suitability in the Northeast U.S. Shelf. Atlantic mackerel larval presence and abundance correlated with sea temperature and copepod abundances, suggesting that larval survival may be sensitive to specific temperatures and zooplankton prey. Predicted abundances were spatially interpolated to estimate Atlantic mackerel larval suitable habitat. Metrics for habitat quality indicate that the Mid‐Atlantic Bight has become less suitable over time. Since the 1970s, the proportion of Northeast U.S. Shelf suitable habitat located in the Mid‐Atlantic Bight has decreased, as southern New England and the western Gulf of Maine regions have become more suitable. Habitat suitability within the Northeast U.S. Shelf has shifted northeast: from the Mid‐Atlantic Bight‐southern New England border towards the northeast portion of southern New England. While total Northeast U.S. Shelf habitat suitability has decreased since the 1970s, the decline in the time series trend was not statistically significant. Thus, while select ecoregions have decreased in habitat suitability, larval habitat does not appear to be the only contributor to decreases in the U.S. Atlantic mackerel contingent.     

Stock size fluctuations in chub mackerel (Scomber japonicus) in the East China Sea and the Japan/East Sea

The stock size of chub mackerel (Scomber japonicus) in the East China Sea and the Japan/East Sea has shown a continuous decrease, with fluctuations, from 1975 to 1990. After a remarkable increase in 1992–96, the stock size fell to its lowest level in 1997–99. The spawning success (number of recruits/spawning biomass) fluctuates periodically with a period of several years, although there are no clear relationships between the spawning biomass and the number of recruits. Spawning success was inversely correlated with sea surface temperature (SST). On the contrary, the catch amount and the catch per unit effort (CPUE) of spotted mackerel (Scomber australasicus) has increased since 1997.     

Simulation of mackerel (Scomber scombrus) recruitment with an individual-based model and comparison with field data

An individual‐based model (IBM) for the simulation of year‐to‐year survival during the early life‐history stages of the north‐east Atlantic stock of mackerel (Scomber scombrus) was developed within the EU funded Shelf‐Edge Advection, Mortality and Recruitment (SEAMAR) programme. The IBM included transport, growth and survival and was used to track the passive movement of mackerel eggs, larvae and post‐larvae and determine their distribution and abundance after approximately 2 months of drift. One of the main outputs from the IBM, namely distributions and numbers of surviving post‐larvae, are compared with field data as recruit (age‐0/age‐1 juveniles) distribution and abundance for the years 1998, 1999 and 2000. The juvenile distributions show more inter‐annual and spatial variability than the modelled distributions of survivors; this may be due to the restriction of using the same initial egg distribution for all 3 yr of simulation. The IBM simulations indicate two main recruitment areas for the north‐east Atlantic stock of mackerel, these being Porcupine Bank and the south‐eastern Bay of Biscay. These areas correspond to areas of high juvenile catches, although the juveniles generally have a more widespread distribution than the model simulations. The best agreement between modelled data and field data for distribution (juveniles and model survivors) is for the year 1998. The juvenile catches in different representative nursery areas are totalled to give a field abundance index (FAI). This index is compared with a model survivor index (MSI) which is calculated from the total of survivors for the whole spawning season. The MSI compares favourably with the FAI for 1998 and 1999 but not for 2000; in this year, juvenile catches dropped sharply compared with the previous years but there was no equivalent drop in modelled survivors.     

Effect of environmental variability on body condition and recruitment success of Atlantic Mackerel (Scomber scombrus L.) in the Gulf of St. Lawrence

The objective of this study was to assess the effect of environmental variability on the dynamics of the Atlantic mackerel (Scomber scombrus L.) stock in the Gulf of St. Lawrence (GSL). We first described the dominant modes of physical and biological (zooplankton) variability using Principal Components Analyses of 40 variables. Two principal modes of variability were identified, a long‐term mode (15–20 yr) associated with a warming of the GSL and a second mode describing alternating cold and warm periods at a higher frequency (5–10 yr). A strong link between physical forcing and the dynamics of zooplankton species known to be important for mackerel was shown. Second, a set of Generalized Additive Models (GAM) was developed to explore how these environmental variations could influence mackerel condition (Fulton's K) and recruitment success (R_s). Optimal GAMs including variations in abundance, species composition and phenology of key copepods improved model performance by 40–50% relative to those considering only physical environmental conditions. The results are consistent with the match–mismatch hypothesis and illustrate the key role of zooplankton dynamics in modulating variations in mackerel K and R_s. Finally, this study showed that large variations in R_s could be caused by varying environmental conditions independently of the influence of stock biomass. Our results strongly indicate that the effect of environmental variability should be considered in the implementation of an ecosystem‐based approach to Atlantic mackerel stock management.     

Occurrence of postmortem myoliquefactive kudoosis in Atlantic mackerel, Scomber scombrus L., from the North Sea

Members of the myxosporean genus Kudoa occur in various marine teleosts worldwide. Several species are of concern to the fishery and aquaculture industries as they may produce unsightly cysts in the fish host's musculature or are associated with postmortem myoliquefaction of the fish muscle, commonly referred to as 'soft flesh'. This study describes the occurrence and effects on a host of a Kudoa species in Atlantic mackerel, Scomber scombrus , from the northern North Sea. Generalized postmortem myoliquefaction associated with Kudoa sp. occurred in 0.8% of the examined fish, i.e. 11 of 1339 mackerel developed 'soft flesh'. There was a significant difference in the prevalence of myoliquefaction between medium sized (400–600 g) and large mackerel (>600 g). The prevalence reached 8.9% in the latter host size group. No subclinical infections of Kudoa sp. were detected when examining fresh muscle ( n  = 103) and blood ( n  = 165) samples for spores using light microscopy. Affected mackerel developed generalized myoliquefaction after 38–56 h post-catch. No inflammatory host response was associated with the presence of plasmodia within single body muscle fibres of 'soft flesh' affected fish. Based on comparison of myxospore dimensions and analysis of the nuclear small subunit (SSU) ribosomal DNA, the present Kudoa species is assigned to Kudoa thyrsites . However, due to the species' apparently very wide geographical distribution and host range, its varying effect on different fish host species, together with the still unknown life cycle of Kudoa spp., the taxonomic status of K. thyrsites appears not to be fully resolved.     

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

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

Temperature conditions in the Argentine chub mackerel (Scomber japonicus) fishing ground: implications for fishery management

We describe the seasonal migrations of Argentine chub mackerel ( Scomber japonicus ) into the fishing ground in relation to sea surface temperature conditions and analyse the temperature evolution during the fishing season on the basis of 1955–97 time-series. The upper temperature limit for the presence of schools was around 19°C. At higher values, most schools leave coastal waters (< 50 m) for the shelf. An inverse relationship was observed between catches per unit of effort (CPUE) of the inshore fishery and sea surface temperature. The highest standardized CPUE values corresponded to temperatures lower than 19°C. Above this temperature, the probability of obtaining a standardized CPUE higher than 1 was extremely low, and the standard deviations were minimal. The economic benefit from catches was also analysed through the different months of the fishing season. It was found that the mean benefit corresponding to temperatures lower than 19°C exceeded approximately 15 times that obtained at higher temperatures. A simple fishing strategy was proposed to the commercial fleet in order to improve the economic benefit of the catches. The migratory pattern followed by mackerel during the spawning period was also analysed in relationship to other environmental factors like food availability. Zooplankton composition, size structure and abundance in coastal and shelf waters are described, and their implications for feeding ecology of larvae and adults of the species are discussed.     

Dual effects of reversed winter–spring temperatures on year‐to‐year variation in the recruitment of chub mackerel (Scomber japonicus)

To clarify the effects of temperature on the recruitment of chub mackerel (Scomber japonicus) in the North Pacific, we investigated the influence of winter surface temperature (WST) on spawners at the time of maturity around the spawning grounds and the influence of ambient spring temperature on larvae using estimated temperature (ET) obtained from particle tracking experiments. We found a significant positive correlation between ET approximately 10 days following hatching and the recruitment per spawning stock biomass (RPS) after 2000. The closer (more meandering) the Kuroshio Current (KC) axis was in relation to the spawning ground, the higher (lower) the spring surface temperature and the higher (lower) RPS was in the spawning ground. In contrast, WST inside KC near the maturity/spawning ground was significantly negatively correlated with RPS. A significant negative correlation between the temperatures in winter and spring was detected in the area after 2000, when the conditions of the Pacific decadal oscillation index and the stability of the Kuroshio Extension were synchronous, indicating that KC shifted northward during this time. The reversed temperature pattern was consistent with the winter–spring movement of KC axis in the offshore direction and was correlated with the winter–spring difference in the intensity of the Aleutian low. These results suggest that the annual variation in chub mackerel recruitment after 2000 was strongly affected by the combined effects of ambient temperature because of the reversal of conditions that occurred between winter and spring around the maturity/spawning ground, which was related to the KC path.     

Covariation in climate, zooplankton biomass and mackerel recruitment in the southern Gulf of St Lawrence

Between 1982 and 1991, an annual survey of stage I egg production of Atlantic mackerel ( Scomber scombrus ) was conducted in June/early July in the southern Gulf of St Lawrence. We investigated the relationship between interannual variability in biomass of zooplankton, determined from the archived survey plankton samples, and mackerel recruitment, estimated from the proportion of three-year-olds in the catch of the commercial fishery. Zooplankton biomass varied by a factor of 2.5, primarily owing to fluctuations in the >1000 μm size fraction. The index of mackerel recruitment fluctuated by a factor of ≈20 and was positively related (linear regression: P  < 0.05; n  = 10) to variations in the zooplankton biomass. Both mackerel recruitment and zooplankton biomass were negatively related (linear regression: P  < 0.05) to RIVSUM, a measure of freshwater discharge from the St Lawrence River system and an index of variability in the region's climate. Three hypotheses are put forward to explain these observations: (1) there is a strong link between interannual variation in abundance of copepod females, which produce prey for mackerel larvae, and larval survival; the exceptional recruitment and subsequent year class resulted from an exceptional production of Calanus finmarchicus nauplii; (2) years of high zooplankton biomass provide better feeding conditions and consequently higher survival of mackerel juveniles; and (3) mackerel recruitment and zooplankton biomass are independently under the control of an underlying physical process, without strong trophic linkage. The first hypothesis is supported by a study of copepod species composition and female abundance conducted for four of the survey years. At the present time, none of these hypotheses can be ruled out.     

Climate,oceanography, and recruitment: the case of the Bay of Biscay anchovy (Engraulis encrasicolus)

The Bay of Biscay anchovy has experienced, since 2001, a succession of low recruitments, resulting in the collapse of the stock in 2005; this has led to successive closures of the fishery. This study investigates the possible impact of different controlling factors [North Atlantic Oscillation (NAO), East Atlantic (EA) pattern, turbulence, upwelling, and river flow] upon anchovy recruitment and fishery catches. Fifty‐five percent of the recruitment variability of this fishery can be explained by upwelling over the spawning area; this is related, in turn, to the EA pattern. The conceptual understanding of the system proposed for the Bay of Biscay anchovy suggests that negative EA periods are associated with northeasterly wind circulation, which produces weak upwelling over the continental shelf. This pattern results in hydrodynamic stability over the area, leading, probably, to adequate food availability. A positive EA (which extends onwards, from 1998) is associated with southwesterly winds and downwelling over the continental shelf; this leads, probably, to the dispersion of anchovy food and larvae, together with increasing mortality.     

Biophysical modelling of larval drift, growth and survival for the prediction of anchovy (Engraulis encrasicolus) recruitment in the Bay of Biscay (NE Atlantic)

Fish recruitment is the result of the integration of small‐scale processes affecting larval survival over a season and large oceanic areas. A hydrodynamic model was used to explore and model these physical–biological interaction mechanisms and then to perform the integration from individual to population scales in order to provide recruitment predictions for fisheries management. This method was applied to the case of anchovy (Engraulis encrasicolus) in the Bay of Biscay (NE Atlantic). The main data available to investigate survival mechanisms were past growth (otolith) records of larvae and juveniles sampled at sea. The drift history of these individuals was reconstructed by a backtracking procedure using hydrodynamic simulations. The relationships between (real) growth variation and variations in physical parameters (estimated by hydrodynamic simulations) were explored along the individual trajectories obtained. These relationships were then used to build and adjust individual‐based growth and survival models. Thousands of virtual buoys were released in the hydrodynamic model in order to reproduce the space–time spawning dynamics. Along the buoy trajectories (representative of sub‐cohorts), the biophysical model was run to simulate growth and survival as a function of the environment encountered. The survival rate after 3 months of drift was estimated for each sub‐cohort. The sum of all these survival rates over the season constituted an annual recruitment index. This index was validated over a series of recruitment estimations. The modelling choices, model results and the potential use of the recruitment index for fisheries management are discussed.     

The influence of environment and spawning distribution on the survival of anchovy (Engraulis encrasicolus) larvae in the Bay of Biscay (NE Atlantic) investigated by biophysical simulations

A growth and survival model of the early life stages was run along virtual drift trajectories tracked in a hydrodynamic model to simulate the annual recruitment process of anchovy (Engraulis encrasicolus) in the Bay of Biscay (NE Atlantic). These biophysical simulations concerning three different years were analysed in order to investigate the influence of environment and spawning dynamics on the survival of larvae and juveniles. The location of space–time survival windows suggested major environmental mechanisms involved in simulated recruitment variability at the different scales – retention of larvae and juveniles in favourable habitats over the shelf margins and turbulence effects. These small‐scale and meso‐scale mechanisms were related to the variations in wind direction and intensity during spring and summer. Survival was also variable according to the origin of the drift trajectories, that is spawning distribution in space and time. The observed spawning distribution (according to field surveys) was compared with the spawning distribution that would maximize survival (according to the biophysical model) on a seasonal scale, which revealed factors not considered in the biophysical model (e.g. spawning behaviour of the different age classes). The variation of simulated survival according to spawning distribution was examined on a multi‐annual scale and showed a coherent pattern with past and present stock structures. The interaction processes between the population (influence on spawning) and its environment (influence on survival) and its implications on recruitment and stock dynamics are discussed.     

Relationships between anchovy (Engraulis encrasicolus) recruitment and environment in the Bay of Biscay (1967–1996)

Various hypotheses have been put forward to explain the mechanisms leading to recruitment variability in fish populations. These have been based on either physical (wind stress, upwelling) or biological (food and predation) processes. In the Bay of Biscay, the hypothesis of a physical influence on anchovy recruitment has been confirmed. Oceanographic conditions in the Bay of Biscay in the spring and summer, influenced by north-easterly winds of medium and low intensity, seem to induce good levels of recruitment to the anchovy ( Engraulis encrasicolus ) population. An index of upwelling was significantly correlated with annual recruitment of anchovy ( P  < 0.0001) for the period 1967–1996. This physical parameter explains about 59% of the variability in the recruitment of the Bay of Biscay anchovy. Two factors affecting productivity in the Bay of Biscay may be linked to north-easterly winds, namely weak upwelling and an extension of the area influenced by river outflows. Both of these factors, together with low turbulence and stability, may act to enhance survival of the early life-history stages of anchovy by increasing food availability. The potential use of this upwelling index to forecast the recruitment of the Bay of Biscay anchovy offers possibilities for improving the management of this population.     

Distribution of jack mackerel (Trachurus japonicus) larvae and juveniles in the East China Sea, with special reference to the larval transport by the Kuroshio Current

Horizontal distribution patterns of jack mackerel (Trachurus japonicus) larvae and juveniles were investigated in the East China Sea between 4 February and 30 April 2001. A total of 1549 larvae and juveniles were collected by bongo and neuston nets at 357 stations. The larvae were concentrated in the frontal area between the Kuroshio Current and shelf waters in the upstream region of the Kuroshio. The abundance of small larvae (<3 mm notochord length) was highest in the southern East China Sea (SECS) south of 28°N, suggesting that the principal spawning ground is formed in the SECS from late winter to spring. Jack mackerel also spawned in the northern and central East China Sea (NECS and CECS, respectively), as some small larvae were also collected in these areas. In the SECS, the abundance of small larvae was highest in February and gradually decreased from March to April. The habitat temperature of small larvae in the SECS and CECS (20–26°C) was higher than that in the NECS (15–21°C), suggesting higher growth rates in the SECS and CECS than in the NECS. The juveniles (10‐ to 30‐mm standard length) became abundant in the NECS off the west coast of Kyushu Island and CECS in April and were collected in association with scyphozoans typical of the Kuroshio waters. However, juveniles were rarely collected in the SECS, where the small larvae were concentrated. Considering the current systems in the study area, a large number of the eggs and larvae spawned and hatched in the SECS would be transported northeastward by the Kuroshio and its branches into the jack mackerels’ nursery grounds, such as the shallow waters off the west coast of Kyushu and the Pacific coast of southern Japan.     

Evaluation of a Fast Method Based on the Presence of Two Restriction Sites in the Mitochondrial ND5 (mt ND5) Gene for the Identification of Scomber Species

The purpose of this work was to evaluate the suitability of a method based on the presence of two restriction sites (for Hae III and Hindf I) in the mitochondrial NADH dehydrogenase subunit 5 (mt ND5) gene to identify Scomber species. The evaluation was performed on 144 reference and market samples by sequencing of the entire 505-bp fragment of the mt ND5 gene and of a 464-bp fragment of the Kocher fragment of the cytochrome b gene (mt Cytb). Sequence analysis of any of the two fragments allows the identification of each of the four Scomber species, but S. japonicus and S. colias had the same restriction sites at the ND5 amplicon and would not have been differentiated by this analysis. Similarly, loss of the Hae III site in some S. scombrus individuals would have misidentified them as not being Scomber. All the market products were correctly labeled except one acquired in Spain labeled as originating in the Atlantic and containing S. japonicus.     

Ichthyoplankton-based spawning dynamics of blue mackerel (Scomber australasicus) in south-eastern Australia: links to the East Australian Current

We describe findings of three ichthyoplankton surveys undertaken along south‐eastern Australia during spring (October 2002, 2003) and winter (July 2004) to examine spawning habitat and dynamics of blue mackerel (Scomber australasicus). Surveys covered ～860 nautical miles between southern Queensland (Qld; 24.6°S) and southern New South Wales (NSW; 41.7°S), and were mainly centred on the outer shelf including the shelf break. Egg identifications were verified applying mtDNA barcoding techniques. Eggs (n = 2971) and larvae (n = 727; 94% preflexion) occurred both in spring and winter, and were confined to 25.0–34.6°S. Greatest abundances (numbers per 10 m²) of eggs (1214–7390) and larvae (437–1172) occurred within 10 nm shoreward from the break in northern NSW. Quotient analyses on egg abundances revealed that spawning is closely linked to a combination of bathymetric and hydrographic factors, with the outer shelf as preferred spawning area, in waters 100–125 m deep with mean temperatures of 19–20°C. Eggs and larvae in spring occurred in waters of the East Australian Current (EAC; 20.6–22.3°C) and mixed (MIX; 18.5–19.8°C) waters, with none occurring further south in the Tasman Sea (TAS; 16.0–17.0°C). Results indicate that at least some of the south‐eastern Australian blue mackerel stock spawns during winter‐spring between southern Qld and northern NSW, and that no spawning takes place south of 34.6°S due to low temperatures (<17°C). Spawning is linked to the EAC intrusion, which also facilitates the southward transport of eggs and larvae. Since spring peak egg abundances came from where the EAC deflects offshore, eggs and larvae are possibly being advected eastwards along this deflection front. This proposition is discussed based on recent data on blue mackerel larvae found apparently entrained along the Tasman Front.     

Effect of low salinity on the expression profile of Na+/K+‐ATPase and the growth of juvenile chub mackerel (Scomber japonicus Houttuyn)

Chub mackerel (Scomber japonicus Houttuyn) is one of the most commercially important scombroid fish used as a food resource. Recently, there has been a demand for efficient rearing methods of this fish for a full‐life cycle aquaculture. In the present study, we evaluated the physiological responses in the juvenile S. japonicus to different ambient salinities. A significantly higher gain of the body mass was observed in the juveniles reared in 24 g/L and 13 g/L seawater than in those reared in natural seawater (34 g/L) within 40 days of the experimental period without affecting mortality. A principal enzyme for osmoregulation, Na⁺/K⁺‐ATPase, was expressed in the ionocytes located in the gill filaments of the juveniles. The number and the cell size of ionocytes and the enzymatic activity of Na⁺/K⁺‐ATPase in the gills decreased within 10 days after the low‐salinity challenge, which implies the reduction of the energy‐consuming active ion secretion under the low‐salinity environment. The physiological capacity for adaptation to low‐salinity seawater in chub mackerel could be basic knowledge to carry out culturing of these fish in coastal sea pens where ambient salinity fluctuates. The improvement of the growth performance by rearing in low‐salinity seawater will contribute to the efficient production of the seed juveniles for aquaculture.     

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.