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.     

Winter monsoon promotes the transport of Japanese temperate bass Lateolabrax japonicus eggs and larvae toward the innermost part of Tango Bay,the Sea of Japan

Northwesterly cold winds characteristic of the East Asian Winter Monsoon (EAWM) dictate winter climatic conditions over the Japanese Archipelago. Japanese temperate bass Lateolabrax japonicus is a commercially important coastal fish that spawns offshore in winter and uses shallow waters as nursery habitats. To investigate the effects of EAWM on the planktonic period of L. japonicus, eggs, larvae, and juveniles were quantitatively collected in Tango Bay on the Sea of Japan side in winter and spring from 2007 to 2017. Although eggs occurred close to the mouth of the bay, planktonic larvae occurred further inside as they developed. The horizontal distribution of planktonic larvae, combined with water velocity data obtained from mooring observations, indicated that planktonic larvae are transported south‐ to westward through Ekman current and an anticyclonic circulation, which are driven by northwesterly winds. To evaluate survival during the planktonic period in each year class, the abundance of benthic larvae/juveniles was divided by winter total landings of Lateolabrax spp. (proxy of the spawning stock size). This survival index exhibited a positive correlation with the northwesterly component of winter winds, and a negative correlation with winter air temperature (average from December to February, Spearman's correlation, p < .05). There was, however, no significant correlation with winter water temperature or winter freshwater discharge in the bay. We conclude that northwesterly cold winds of EAWM play a critical role in transporting L. japonicus eggs and larvae toward nursery habitats, specifically beaches and estuaries fringing the innermost part of Tango Bay.     

Early life history connectivity of Antarctic silverfish (Pleuragramma antarctica) in the Ross Sea

A recent population hypothesis for Antarctic silverfish (Pleuragramma antarctica), a critical forage species, argued that interactions between life history and circulation associated with glacial trough systems drive circumpolar distributions over the continental shelf. In the Ross Sea, aggregations of eggs and larvae occur under fast ice in Terra Nova Bay, and the hypothesis predicted that dispersing larvae encounter outflow along the western side of Drygalski Trough. The outflow advects larvae towards the shelf‐break, and mixing with trough inflow facilitates return toward the inner shelf. To examine the hypothesis, we compared samples of P. antarctica collected near Coulman Island in the outflow, along Crary Bank in the inflow, and a third set taken over the rest of the Ross Sea. We ruled out misidentification using an innovative genetic validation. Silverfish larvae comprised 99.5% of the catch, and the highest population densities were found in Drygalski Trough. The results provided no evidence to reject the population hypothesis. Abundance indices, back‐calculated hatching dates, length distributions and growth were congruent with a unified early life history in the western Ross Sea, constrained by cryopelagic early stages in Terra Nova Bay. By contrast, a sample in the Bay of Whales revealed much smaller larvae, suggesting either a geographically separate population in the eastern Ross Sea, or westward connectivity with larvae spawned nearby by fish sourced from troughs upstream in the Amundsen Sea. These results illustrate how hypotheses that integrate population structure with life history can provide precise spatial predictions for subsequent testing.     

Observation on the spawning of Sardina and Sardinella off the south Moroccan Atlantic coast (21–26°N)

This paper provides information on the early life stages of the Moroccan Atlantic sardine, Sardina pilchardus, and two species of Sardinella: Sardinella aurita and S. maderensis, between Cape Blanc (21°N) and Cape Boujdor (26°N), dealing with the spawning grounds, the nursery areas and the optimal spawning temperature and salinity range.

The spawning areas for Sardina are located North of Dakhla (24°30′–25°30′N) and near Cintra Bay (23°N). The larval development area is described together with seasonal and annual characteristics. The nurseries of Sardina are near Dakhla and to the south between 21 and 22°N but precise locations vary with season. The maximum occurrence of eggs was during winter, and was at temperatures from 16 to 18 °C in winter and between 18 and 18.5 °C in summer.

The main spawning area of Sardinella species is between Cape Blanc and Cintra Bay (21–23°N) with a maximum occurrence of eggs and larvae in July. For Sardinella off the southern region of Morocco, the optimal temperature interval for spawning is between 18 and 21.14 °C. No conclusions can be drawn on the relationship between spawning and water salinity.

The area north of Cintra Bay has a broad, but shallow continental shelf. This topography leads to decreasing dispersion effects and the zone constitutes a favorable area for larval retention and development.     

Changes in the distribution of sardine eggs and larvae off Portugal, 1985–2000

Generalized additive models (GAMs) were fitted to sardine (Sardina pilchardus) egg distribution data from three daily egg production method surveys. The results showed that the area of egg cover off Portugal decreased significantly from 11 800 km² in 1988 to 7000 km² in 1997 and 7400 km² in 1999. This is because of a significant reduction in sardine egg presence off northern Portugal, GAM estimated areas being similar or higher in the late 1990s for southwestern and southern Portugal. The distributional area covered by larvae was not estimated for 1988 (larval distribution extended beyond the survey area), although it was probably higher than the 9600 km² for 1997 and 5500 km² for 1999. In 1997 and 1999, the Gulf of Cadiz was also sampled, indicating extensive areas with sardine eggs and larvae (more than 50% of the total area of distribution off Portugal). Standardized data from 15 ichthyoplankton surveys between 1985 and 2000 show a decline in the mean probability of egg presence within the Portuguese continental shelf from the mid‐1980s to the late‐1990s, because of a marked reduction in egg presence off northern Portugal. Sardine larval data from the same surveys suggest that the reduction in mean probability of presence in the north is less marked than for eggs (although this comparison ignores the presence of sardine larvae beyond the continental shelf in the 1980s). Similar changes off northern Portugal and western Galicia are observed in commercial sardine catches and the acoustically estimated area of fish distribution. It is possible that the observed decline in spawning area off northwestern Iberia during the 1990s is indirectly reflecting the prevalence of environmental conditions detrimental to sardine recruitment (northerly winds during winter that favour coastal upwelling and offshore transport), which have reduced the spawning contribution of young fish in that area.     

The effect of vertical and horizontal distribution on retention of sardine (Sardinops sagax) larvae in the Northern Benguela – observations and modelling

In the present study, a modelling experiment is conducted to simulate the transport of sardine (Sardinops sagax) eggs and larvae in the Northern Benguela. Based on historical and newly obtained data, different scenarios of vertical and horizontal distribution are applied and the effects on retention are discussed. The simulations showed that vertical and horizontal distribution were important for retention of sardine larvae in the Northern Benguela. By using age‐dependent data on vertical distribution, it was shown that retention of particles in the simulation was substantially enhanced compared with a scenario where particles were distributed in the offshore moving Ekman layer. Retention was lowest during October–December (when upwelling intensity is high) and highest during February–April (when upwelling intensity is somewhat lower). When different spawning areas were considered, highest retention was observed in an area near Walvis Bay. It is concluded that the behaviour of sardine larvae is adapted to the circulation system in the Northern Benguela in a way that promotes retention of the larvae in inshore nursery areas.     

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.     

Spatial variations in feeding and condition of juvenile pink and chum salmon off Vancouver Island, British Columbia

Spatial variations in feeding and condition of juvenile pink (Oncorhynchus gorbuscha) and chum (Oncorhynchus keta) salmon, and their implications for growth, were examined on the Vancouver Island continental shelf in early summer 1992. Juvenile pink salmon off northern Vancouver Island had more material in their stomachs, were in better condition, and had higher potential growth rates (from a bioenergetics model) than pink salmon off southern Vancouver Island. These variations were consistent with spatial differences in zoo-plankton biomass, there being more plankton in the northern region. There was a significant positive relationship between condition of pinks and the amount of material in their stomachs, suggesting a positive feedback on feeding success. Juvenile chum in the north also had more material in their stomachs than chum to the south. However, condition factor was not significantly different between southern and northern regions nor was there a significant relationship between condition factor and the weight of stomach contents for chum on the southern shelf. A bioenergetics model suggests that chum in the south were food limited. Stable carbon isotope data also indicated different feeding histories for some chum in the southern region, which may have been recent migrants onto the continental shelf from near-shore areas, or possibly a nearby hatchery. Estimation of the energy required by juvenile salmon to migrate north in a continental shelf area with low zooplankton biomass and a weak northerly current (inner shelf), compared with an area with higher zooplankton biomass but a strong southerly current (outer shelf), indicated sufficient surplus energy only in the inner shelf, consistent with observations of northward migrations predominantly through this area. Spatial variations in current velocity and zooplankton biomass can affect feeding, condition, and potential growth of juvenile pink and chum salmon off Vancouver Island.     

Connectivity of the bay scallop (Argopecten irradians) in Buzzards Bay,Massachusetts, U.S.A.

The harvest of bay scallops (Argopecten irradians) from Buzzards Bay, Massachusetts, U.S.A. undergoes large interannual fluctuations, varying by more than an order of magnitude in successive years. To investigate the extent to which these fluctuations may be due to yearly variations in the transport of scallop larvae from spawning areas to suitable juvenile habitat (settlement zones), a high‐resolution hydrodynamic model was used to drive an individual‐based model of scallop larval transport. Model results revealed that scallop spawning in Buzzards Bay occurs during a time when nearshore bay currents were principally directed up‐bay in response to a persistent southwesterly sea breeze. This nearshore flow results in the substantial transport of larvae from lower‐bay spawning areas to settlement zones further up‐bay. Averaged over the entire bay, the spawning‐to‐settlement zone connectivity exhibits little interannual variation. However, connectivities between individual spawning and settlement zones vary by up to an order of magnitude. The model results identified spawning areas that have the greatest probability of transporting larvae to juvenile habitat. Because managers may aim to increase scallop populations either locally or broadly, the high‐connectivity spawning areas were divided into: (i) high larval retention and relatively little larval transport to adjoining settlement areas, (ii) both significant larval retention and transport to more distant settlement areas, and (iii) little larval retention but significant transport to distant settlement areas.     

Association between the planktonic larval and benthic stages of Manila clam <Emphasis Type="Italic">Ruditapes philippinarum</Emphasis> in eastern Hokkaido,Japan

The association between planktonic larval and benthic stages of Manila clam Ruditapes philippinarum and factors determining successful recruitment were studied in Akkeshi-ko estuary and Akkeshi Bay, Hokkaido, northern Japan. The field study showed that planktonic larvae were localized mainly in the bay after dispersal from the estuary and recruitment to the estuarine fishing grounds was not always successful. A numerical analysis suggested that estuarine circulation transported tracers, used to simulate planktonic larva, from the estuary to the bay but the number of returning tracers varied depending on the river discharge and wind direction. The planktonic larvae were affected by unsuitable conditions for survival in the estuary, such as high turbidity. We conclude that most larvae develop within the bay, where conditions are more saline and less turbid, and that although return to the estuary is essential for recruitment, it is not necessarily related to the abundance of larvae.     

Springtime ichthyoplankton of the slope region off the north-eastern United States of America: larval assemblages, relation to hydrography and implications for larval transport

Larval transport in the slope region off north‐eastern North America influences recruitment to juvenile habitats for a variety of fishes that inhabit the continental shelf. In this study, collections of larval fishes were made during springtime over the continental slope to provide insights into larval distributions and transport. Ichthyoplankton composition and distribution mirrored the physical complexity of the region. Three larval fish assemblages were defined, each with different water mass distributions. A Gulf Stream assemblage was found predominantly in the Gulf Stream and associated with filaments of discharged Gulf Stream water in the Slope Sea. Larvae of this assemblage originated from oceanic and shelf regions south of Cape Hatteras. Several members of this assemblage utilize habitats in the Middle Atlantic Bight (MAB) as juveniles (Pomatomus saltatrix, Peprilus triacanthus) and other members of the assemblage may share this life cycle (Mugil curema, Sphyraena borealis, Urophycis regia). A Slope Sea assemblage was found in all water masses, and was composed of epi‐ and mesopelagic fish larvae, as well as larvae of benthic shelf/slope residents. Larvae of one member of this assemblage (U. tenuis) are spawned in the Slope Sea but cross the shelf‐slope front and use nearshore habitats for juvenile nurseries. A MAB shelf assemblage was found in MAB shelf water and was composed of larvae that were spawned on the shelf. Some of these species may cross into the Slope Sea before returning to MAB shelf habitats (e.g. Enchelyopus cimbrius, Glyptocephalus cynoglossus). Previous studies have examined the effect of warm‐core rings on larval distributions, but this study identifies the importance of smaller‐scale features of the MAB shelf/slope front and of filaments associated with Gulf Stream meanders. In combination with these advective processes, the dynamic nature of larval distributions in the Slope Sea appears to be influenced, to varying degrees, by both vertical and horizontal behaviour of larvae and pelagic juveniles themselves.     

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

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

Hydrographic phenomena influencing early life stages of the Sicilian Channel anchovy

The information collected from a European Union funded project on the ‘Distribution Biology and Biomass Estimates of the Sicilian Channel Anchovy (Engraulis encrasicolus)’ was used to analyse the linkage between the general circulation pattern of the Atlantic Ionian Stream (AIS) and the reproductive strategy of the Sicilian Channel anchovy. The main spawning ground is located in the NW region of the southern Sicilian coast. This region is a stable area of low current produced by the impingement towards the coast of the AIS and its bifurcation into two branches. The main branch heads towards the SE end of the Sicilian coast (Cape Passero) acting as a transport mechanism for the anchovy eggs and larvae. Along the AIS trajectory, there is a density front to the left of the current, facing downstream. This front is a consequence of the shoreward sloping of isopycnals that maintains the geostrophic flow, facilitating the mixing of deeper waters with surface layers and fertilization of coastal waters. The front enhances primary production assuring food availability for anchovy larvae during their advection by the AIS. The highest concentrations of larval anchovy were found off the SE Sicilian coast, in the area off Cape Passero. The greater average sizes of larvae found in this region, and their estimated age, support the evidence of advection by the AIS. The hydrographic features observed in this area, such as the existence of a well‐defined cyclonic vortex, implies the existence of upwelling in its centre, providing a suitable environment for sustained enhanced rates of primary production and allowing the larval population to maintain their relative position. This retention area is conceived as favourable for providing the necessary feeding conditions. The data acquired from a survey carried out to evaluate the anchovy recruitment strength confirm that larvae reach the juvenile stage in the south‐eastern coast of Sicily, since most of the young‐of‐the‐year anchovy were located in the Cape Passero region.     

Do Sardinella aurita spawning seasons match local retention patterns in the Senegalese–Mauritanian upwelling region?

Sardinella aurita is the most abundant small pelagic fish in the Senegalese–Mauritanian region. The success of its reproduction crucially depends on the local circulation as this determines whether larvae reach coastal nursery areas favorable to their survival or are dispersed into the open ocean. As a first step towards evaluating sardinella vulnerability to climate‐driven hydrodynamical changes, this study aims at underpinning how transport pathways drive optimal spatial and seasonal patterns for sardinella reproduction. We have used two estimates of the Senegalese–Mauritanian coastal seasonal circulation simulated by two hydrodynamical model configurations that differ in their forcing and topography. Nursery areas are determined by evaluating coastal retention with a Lagrangian individual‐based model that accounts for processes such as diel vertical migration and mortality as a result of lethal temperature exposure. Our results suggest that the shelf zones located at the Arguin Bank (19.5°N–21°N) and south of Senegal (12°N–14.75°N) are highly retentive. We find maximum retention rates in July–August and November–December over the Arguin Bank; from February–July and November–December over the southern Senegalese shelf; and lower retention rates over the central region (14.75°N–19.5°N) that are locally maximum in June–July when the upwelling weakens. These retention areas and their seasonality are in agreement with previously reported spawning patterns, suggesting that the Sardinella aurita spawning strategy may result from a trade‐off between retention patterns associated with the seasonal circulation and food availability. Exposure to lethal temperatures, although not well studied, could be a further limiting factor for spawning. The Lagrangian analysis reveals important connectivity between sub‐regions within and south of the system and hence underlines the importance for joint management of the Sardinella aurita stock.     

Tests of larval retention in a tidally energetic environment reveal the complexity of the spatial structure in herring populations

Dispersion during the larval phase is of central importance in the dynamics of marine fish and invertebrate populations. Rapid transport or dispersion of larvae may contribute to connectivity and mixing, whereas spatial persistence (retention) is hypothesized to favour stock complexity and local subpopulations. Larval retention, while rarely quantified, may be defined in species with protracted spawning by the spatial co‐occurrence of larvae of different sizes or ages. The spatial distributions of larval Atlantic herring (Clupea harengus) were examined from 22 annual autumn surveys (1975–1998) and 9 spring surveys (1975–1984) from the Bay of Fundy, a region with large tides and residual flow. Larvae of all sizes (3–27 mm in length, from hatch to nearly 4 months post‐hatch) were observed each year in two major aggregations; one off southwestern Nova Scotia, and the other in the mid‐inner Bay of Fundy off the northwestern shore of Nova Scotia. Two similar aggregations were evident over 5 months later from 9 spring surveys (1975–1984), despite the residual flow that would have swept the larvae from the region within 1 month. Larval retention was apparent from overlapping centres of mass of different size (=age) classes of larvae, and tested using a size diversity index, based on the co‐occurrence of 1‐mm‐size categories, derived from protracted spawning of several weeks. Geospatial ‘hot spots’ (G_i* statistic) of four size (age) classes were evident at specific stations in the 50–100 m bathymetric zone and not elsewhere. These metrics provide quantitative measures of retention that may be applied to many ichthyoplankton data sets. One of the three main spawning areas collapsed during the study period after a period of intense fishing and failed to rebuild, but there was no substantial change in the location of larval hotspots in subsequent years. While larval retention does not directly relate to each spawning location, larval retention in the Bay of Fundy contributes to the complex ‘metapopulation’ structure of herring stocks in the western Atlantic.     

罗源湾夏季鱼卵和仔稚鱼种类组成与数量分布

分析研究了2009年7月福建罗源湾鱼卵、仔稚鱼的种类组成及数量分布。共鉴定19种鱼类的鱼卵和仔稚鱼,隶属于13科。其中,鱼卵10种、仔稚鱼14种。夏季,罗源湾水平拖网鱼卵和仔稚鱼平均密度分别为1.80 ind/m3和0.54 ind/m3;垂直拖网鱼卵和仔稚鱼平均密度分别为4.11 ind/m3和2.63 ind/m3。鱼卵、仔稚鱼以鳀科数量最多,优势种类为中颌棱鳀(Thrissa mystax)和康氏小公鱼(Stolephoruscommersonli)。与历史资料相比,不但种类数减少,且优势种由多科鱼类并存向鳀科鱼类为主转变。夏季鱼卵、仔稚鱼主要分布在罗源湾中北部海域及湾口附近。     

Estimating larval supply of Ezo abalone <Emphasis Type="Italic">Haliotis discus hannai</Emphasis> in a small bay using a coupled particle-tracking and hydrodynamic model: insights into the establishment of harvest refugia

Most juveniles of Haliotis discus hannai have been found to be descendants of wild individuals, although most adults were artificially produced (released) individuals as a result of restocking inside the refugium located near the head of Oshoro Bay, Hokkaido, Japan. To estimate the larval supply from released and wild individuals into the refugium and to compare the suitability of locations as larval sources, we simulated larval dispersal using a coupled hydrodynamic and particle-tracking model. The simulation results indicated that more larvae may be supplied from the wild adults inside the bay to the refugium than from the released adults. These results are consistent with the observed high abundance of wild juveniles in the refugium. Most larvae from the refugium were predicted to disperse out of the bay. We found that larval retention in the bay was at least one order of magnitude higher than that in the refugium. Thus, it may be more effective in terms of self-replenishment and reproduction if the refugium were to be expanded to the bay scale. There were only minor differences among the compared sites at the head of the bay in terms of their suitability as larval sources. Consequently, the establishment of new refugia in this area could be expected to provide an effectiveness equal to that of the current refugium.     

Circulation of the eastern North Atlantic and north-west European continental shelf – a hydrodynamic modelling study

As part of the interdisciplinary investigation of Calanus finmarchicus migrations between oceanic and shelf areas off north-west Europe (the ICOS project), a three-dimensional, prognostic baroclinic circulation model system, consisting of a coarse and a nested fine scale model, was constructed. The aim was to simulate the flow fields around Iceland, the Faroe Islands and the north-west European continental shelf and slope, including the northern North Sea. The coarse scale model covered the northern North Atlantic and provided the far field flow to the finer scale model. The latter was initialized and forced with climatological seasonal (winter and spring) means of temperature, salinity, river run-off, and other parameters necessary for the calculation of heat fluxes. Seasonal average simulations of flow and hydrographic conditions were performed, together with simulations of specific scenarios of uniform wind forcing from different directions. A feature common to all the results was a topographically guided northwards flow along the continental shelf edge, with a southwards counter-flow in the vicinity of the Faroe–Shetland Channel. The extent and strength of the shelf edge current, of its counter flow, and of the re-circulation cell in the northern North Sea, were dependent on the prevailing wind direction. The modelling results were consistent with the accepted circulation patterns in the area, and a comparison with field measurements in the Faroe–Shetland Channel showed that the main characteristics of the flow were well replicated, although the estimated strength of the slope current was somewhat less than in the field observations.     

Spawning habitat of the Atlantic menhaden in Onslow Bay, North Carolina

The Continuous, Underway Fish Egg Sampler (CUFES) was used to sample pelagic eggs of the Atlantic menhaden ( Brevoortia tyrannus ) from 3-m depth off North Carolina in winter 1993–94 and 1994–95. Simultaneous measurements were made of temperature, salinity, and the concentration of chlorophyll a . The maximal concentration of eggs was 346 eggs m⁻³. Eggs were highly aggregated in patches which occurred between the Gulf Stream and mid-shelf fronts (17–23°C, 36.0–36.4‰). Unexpectedly, eggs were found almost exclusively in water of 20–60 m (mode 20 m) bottom depth. Thus, spawning appears related to bathymetry as well as hydrography. Variograms for egg concentration indicated a mean (± SE) patch scale of 3.6 ± 1.7 km and a high degree of spatial variance explained by CUFES sampling. Lagrangian modelling of particles moving in response to tides, winds, and a prescribed flow from the north indicated that the region of observed, maximal occurrence of eggs is favourable for the retention of eggs and larvae on the shelf adjacent to inlets used to enter nursery areas.