Spawning site selection by European anchovy (Engraulis encrasicolus) in relation to oceanographic conditions in the Strait of Sicily

European anchovy egg occurrence and density data from summer surveys (1998–2007) and oceanographic data were examined to study the mechanisms that control the spatial distribution of anchovy spawning habitat in the Strait of Sicily. Quotient analysis indicated habitat preference for temperature (18–19°C), bottom depth (50–100 m), water column stability (13–14 cycle h^?1), fluorescence (0.10–0.15 μg m^?3 Chl a), salinity (37.5–37.6 PSU), current speed (0.20–0.25 m s^?1) and density (26.7–26.8 kg m^?3, σ_t). Canonical discriminant analysis identified temperature, column stability and fluorescence as major drivers of anchovy spawning habitat. Three of the 4 years which had lower egg abundance were warmer years, with low values of primary productivity. A geostrophic current flowing through the Strait (the Atlantic Ionic Stream, AIS) was confirmed as the main source of environmental variability in structuring the anchovy spawning ground by its influence on both the oceanography and distribution of anchovy eggs. This 10‐yr data series demonstrates recurrent but also variable patterns of oceanographic flows and egg distribution. A lack of freshwater flow in this area appears to depress productivity in the region, but certain and variable combinations of environmental conditions can elevate production in some sub‐areas in most years or other sub‐areas in fewer years. These temporal and spatial patterns are consistent with an ocean triad theory postulating that processes of oceanographic enrichment, concentration, and retention may help predict fishery yields.     

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.     

Assessing the variability of hydrographic processes influencing the life cycle of the Sicilian Channel anchovy, Engraulis encrasicolus, by satellite imagery

Three oceanographic surveys carried out in the Sicilian Channel during the spawning season (June to July) of anchovy (Engraulis encrasicolus) showed a close relationship between anchovy reproductive strategy and important hydrographic structures. A time series of satellite‐derived sea surface temperature images of the Sicilian Channel were analysed by means of empirical orthogonal functions and the dominant empirical modes were studied in detail. The first empirical mode captured much of the original variance and reproduced the trajectory of the Atlantic Ionian Stream (AIS), the principal hydrodynamic feature of the area. The time coefficients of modes 1 and 2 had seasonal signals which, when combined, accounted for the enhancement of the thermal front, clearly visible off Cape Passero (southernmost coast of Sicily) during summer. As the area constituted the principal nursery ground of the Sicilian Channel anchovy, the combination of the time coefficients of these modes was considered a potential indicator of the food particle concentration usually associated with oceanic fronts, which provided the energy requirements for larval growth. Mode 3 described the north/south displacements of the mean AIS trajectory, which modified the surface temperature regime of the anchovy spawning habitat. Therefore, the time coefficients of this mode were used as a potential indicator of anchovy spawning habitat variability. The capability of time coefficients of modes 2 and 3 to modify the main pattern depicted by mode 1 were tested successfully against in situ oceanographic observations.     

Variation in the condition factors of California pelagic fishes and associated environmental factors

Time series of condition factors for mackerel, Scomber japonicus , jack mackerel, Truchurus symmetricus , and northern anchovy, Engraulis mordax , stocks in the Southern California region were compared with time series of oceanographic indices to develop hypotheses concerning physical environmental forcing of the population dynamics and energetics of small pelagic fishes. Mackerel and jack mackerel condition factor time series showed decade-scale variation, whereas those of anchovy showed coherent fluctuations for 1 to 2 years. Mackerel, and to a lesser extent jack mackerel, condition factors were correlated with proxy indices of alongshore advection (sea level), offshore advection (Ekman transport), ambient temperature (shore station temperature), and ambient salinity (shore station salinity). The condition factor of anchovy was much less correlated with environmental variables. Multiple regression analyses which included sea level, upwelling and salinity proxies explained 80% (33%) of the variance in the annual (monthly) condition factor of mackerel. The first-order variation in condition factors of mackerel and jack mackerel suggests that they are responding to very large-scale perturbations of the California Current system which are at least partially described by variations in sea level. The population size of mackerel is apparently also responding to these large-scale perturbations, making it difficult to isolate environmental dependence of condition factors from density dependence. The second-order variation is more regional in nature and unexpectedly it appears to be associated with upwelling in the Baja California region.     

Temporal Variation in Egg and Larval Productivity of Eyestalk Ablated Spawners of the Prawn Penaeus monodon from Cook Bay, Australia

This paper reports on the variability in spawner performance and viability of the eggs and larvae of Penueus monodon from a single geographical region (Cook Bay, Australia). Data is presented from seven separate experiments spawning eyestalk ablated P. monodon conducted between Sep tember 1992 and July 1994. These experiments used 200 spawners from which 290 spawns were assayed. Total egg and larval (Nauplius, and ha,) production values per prawn were calculated and were significantly different ( P < 0.05) between experiments. Spawner weight and number of spawns per prawn also varied significantly between experiments and, although these factors afiected egg and larval productivity, they accounted for only a small percentage of the observed variability. The cause of this variation is unknown but it may be due to differences in the age of the spawners and to seasonal environmental effects. The variation in reproductive success of wildcaught spawners probably reflects differences in overall condition of the spawners and/or their nutritional status at different times. Understanding the causes of variation in productivity is important if hatcheries are to improve the reliability of supply of larvae for the prawn farming industry.     

Egg and larval distributions of seven fish species in north-east Atlantic waters

The distribution of egg and larvae of mackerel, horse mackerel, sardine, hake, megrim, blue whiting and anchovy along the European Atlantic waters (south Portugal to Scotland) during 1998 is described. Time of the year, sea surface temperature and bottom depth are used to define the spawning habitat of the different species. Mackerel, horse mackerel, and sardine eggs and larvae presented the widest distribution, whereas megrim and anchovy showed a limited distribution, restricted to the Celtic Sea and the Bay of Biscay respectively. Correspondingly mackerel, horse mackerel and sardine showed the highest aggregation indices. Blue whiting larvae were found at the lowest temperatures, whereas anchovy eggs and larvae were found in the warmest waters. The analysis is a basis for evaluation of ongoing changes in the pelagic ecosystem of the north‐east Atlantic.     

Fish larvae distribution off Mejillones Peninsula (northern Chile) during a coastal upwelling event in Spring 1999: interactions with the cold upwelling plume

We examined the interaction between vertical and horizontal distribution of fish larvae off Mejillones Peninsula (23°S), northern Chile, under conditions of active coastal upwelling. An oceanographic survey covered spatial variability in temperature, chlorophyll- a (chl a ), dissolved oxygen, salinity and water density. Fish larvae were sampled during daytime and nighttime periods through two consecutive days in four stations: two inside and two outside of a well-developed upwelling plume, and at three depth strata: 0–20, 20–80 and 80–200 m. Eighteen taxa were analysed, of which the Myctophidae Diogenychthys atlanticus , Diogenichthys laternatus , and the anchovy Engraulis ringens , were most abundant. Our data showed little evidence for diel vertical migration and larvae were more abundant at depth (>80 m) under low temperature (∼12°C) and low chl a (∼2 mg m^–3), below the highly advective upper layer. The exploratory K -means analysis allowed the separation of data into two distinct habitats: upwelling and nonupwelling types. Most taxa were allocated in nonupwelling waters, i.e. outside the cold plume. However, short-term variations (<24 h) in the position of the upwelling plume influenced both horizontal and vertical occurrence as well as abundance of taxa, and caused variability in temperature, oxygen and chl a . These changes in oceanographic conditions, caused by upwelling circulation and the dynamics of the cold plume, may sharply modify the habitat of fish larvae and have an important role in survivorship and recruitment success.     

`Ocean triads' in the Mediterranean Sea: physical mechanisms potentially structuring reproductive habitat suitability (with example application to European anchovy, Engraulis encrasicolus)

Summaries of maritime weather reports and mean seasonal satellite-sensed ocean colour distributions for the Mediterranean Sea are used to identify characteristic configurations of physical mechanisms promoting (i) nutrient enrichment, (ii) concentration of larval food distributions, and (iii) local retention of eggs and larvae. Five subbasin scale `ocean triads', hypothesized to be particularly favourable groupings, are identified in the Aegean Sea, the Gulf of Lions and nearby Catalan Coast, the Alboran Sea, the Straits of Sicily/Tunisian Coast, and the Adriatic Sea. These are examined in relation to available knowledge of anchovy spawning grounds. All areas are characterized by patterns of linked wind-driven Ekman upwelling and downwelling. All areas except the Straits of Sicily are found to have inputs of less saline surface waters offering raised nutrient concentrations, enhanced upper layer stability, and frontal density contrasts, and to have areas where the characteristic rate of turbulent mixing energy input by the wind fall below a reference intensity level. All areas, except the Sicilian Channel/Tunisian Coast, also contain abundant locally reproducing anchovy populations.     

Changes in the spawning habitat of two small pelagic fish in the Northwestern Mediterranean

We investigated the effects of three sea surface oceanographic variables (temperature, salinity, and chlorophyll a) on the abundance of eggs and larvae of two summer‐spawning species in the NW Mediterranean Sea, the anchovy Engraulis encrasicolus and the round sardinella Sardinella aurita, based on data from ichthyoplankton surveys carried out in the 1980s, 2000s, and 2010s. The environmental data showed an increase in seawater temperature and salinity along time, coupled with a decrease in chlorophyll a (proxy for primary production). These long‐term directional changes in environmental conditions helped explain the important reduction observed in the abundance of eggs and larvae of anchovy, as well as shrinking of spawning habitat in this species. At the same time, the probability of occurrence of round sardinella has increased from practically zero in the 1980s to probabilities near 1 along the coastal area of the study region in the two decades of the 21st century. Given that the trends observed in the environmental variables along the three decades of study are expected to continue during the 21st century, as a consequence of climate change, the spawning habitat of anchovy is expected to continue decreasing, while round sardinella habitat can expand. Considering that anchovy is of high commercial importance in NW Mediterranean fisheries, while round sardinella has very low commercial interest, our results show that the viability of small pelagic fisheries in the area may be compromised.     

The relationship between the skipjack tuna (Katsuwonus pelamis) fishery and seasonal temperature variability in the south-western Atlantic

The spatio‐temporal distribution of tuna fishing effort has been related to oceanographic circulation and features in several seas of the world. Understanding the relationship between environmental variables and fishery resource dynamics is important for management decisions and to improve fishery yields. The relationship between sea temperature variability and the pole‐and‐line skipjack tuna (Katsuwonus pelamis) fishery in the south‐western Atlantic Ocean was investigated in this work. Data from logbooks, satellite images (sea surface temperature), and oceanographic surveys were used in the analyses. Skipjack are caught in warm tropical waters of the Brazil Current (BC). The north–south displacement of fishing effort was strongly associated to seasonal variation of the surface temperature, which was coupled to the tropical BC flow. Oceanographic fronts from autumn to spring and a shallow thermocline in summer probably induces the aggregation of skipjack schools over the shelfbreak, favouring fishing operations. Hypotheses are proposed to explain the relationship between peaks of fishing events and the presence of topographic peculiarities of the shelfbreak.     

Larval growth rates differ in response to seasonal temperature variations among clupeoid species inhabiting the Pacific coastal waters of Japan

Growth rates of round herring Etrumeus teres, Japanese anchovy Engraulis japonicus, and Japanese sardine Sardinops melanostictus larvae respond differently to seasonal temperature variations in Sagami Bay on the Pacific coast of central Japan. Larvae of these three clupeoid species were collected from small seine net catches between February 2008 and April 2009. Correlations between otolith growth increment widths and ambient temperatures were examined at five body sizes and indexed to otolith radii of 50–175 μm, which corresponded to notochord lengths of 15–24 mm. Round herring larvae showed positive correlations between growth rate and habitat temperature at all five body sizes. In contrast, Japanese sardine larvae did not show significant correlations between growth rate and ambient temperature at any of the five body sizes. Japanese anchovy larvae showed significant but weaker correlations between growth rate and temperature than round herring. Although these three species share habitats with some differences in seasonal occurrence in the bay, they showed different species-specific responses to seasonal habitat temperature variations. These differences in response to temperature variations may constitute an ecological basis for different migration patterns among the three species.     

Fluctuation in abundance of larval anchovy and environmental conditions in coastal waters off south-western Taiwan as associated with the El Niño–Southern Oscillation

Monthly abundance ( CPUE ) of larval anchovy in the coastal waters off south-western Taiwan from 1980 to 1992 (156 months) fluctuated at intervals corresponding to the 4.3- and 2.2-year cycles of the southern oscillation index ( SOI ). Also, CPUE was significantly correlated with sea surface temperature with a time lag of 3 months and nearly significantly to river flow with a time lag of 4 months, which in turn correlated with SOI at lags of 13–14 months (cross-correlation and transfer function analyses). The results suggested the presence of linkage between recruitment of the larvae and ENSO episodes, perhaps through oceanographic and meteorological conditions that affect coastal upwelling and river discharge. The Kuroshio Current, which is the western extension of the North Equatorial Current, may be one of the important mechanisms of ENSO's teleconnections affecting local climate and fisheries in the western Pacific region.     

Larval population structure of Engraulis encrasicolus in the Strait of Sicily as revealed by morphometric and genetic analysis

In order to investigate the larval population structure, specimens of Engraulis encrasicolus larvae from five different locations in the Strait of Sicily were analyzed by means of otolith readings, morphometric and genetic techniques. The distribution of age in day, identified by means of the otolith readings and associated with the oceanographic parameters, was useful to identify possible spawning areas and transport dynamics. The presence of more than one spawning area suggested the possibility that two or more sub‐populations may co‐exist in the study area. The morphometric characteristics were more adequate than genetic parameters to discriminate the different larval groups. The most relevant variables for the separation were the mouth length (ML) and the body diameter (BD). The population structure by means of genetic data reported the presence of two phylogroups co‐occurring among samples in each sampling locations specimens. The pattern of genetic divergence among anchovy larvae in the Strait of Sicily was congruent with previous studies conducted on adult populations present in other Mediterranean areas with different molecular markers. The habitat‐specific nature of the morphological variation and the lack of corresponding genetic variation among larvae from the different locations suggested that the observed differences in morphology could be linked to environmental parameters. The body form differences among different larvae samples could reflect the nutritional status of larvae. In fact, these differences were found among anchovy larvae collected in areas with different oxygen and fluorescence, which is an index of primary productivity and is linked to the availability of food for anchovy larvae.     

Temporal variation of seasonality of egg production and the spawning biomass of Pacific anchovy, Engraulis japonicus, in the southern waters of Korea in 1983–1994

Embryonic mortality, egg production and the spawning stock biomass of Pacific anchovy, Engraulis japonicus , off Southern Korea during 1983–1994, and their biological response to oceanographic features in spring and summer, were analysed. The instantaneous mortality rate (IMR) of embryonic stages decreased in spring and increased in summer, with a range of 0.33–1.23 day^–1 in spring and 0.78–1.69 day^–1 in summer. Egg production in summer was three times that during spring and production was low in the late 1980s. Mean lengths of yolk-sac larvae and adult females were greater in spring than in summer, whereas spawning fraction and spawning stock ratio (spawning biomass:adult biomass) were lower in spring than summer. Estimated mean spawning stock biomass ranged from 141 × 10³ to 380 × 10³ MT in spring and from 221 × 10³ to 557 × 10³ MT in summer. Statistically, the seasonal and long-term trends of embryonic mortality, egg production and spawning stock biomass of Pacific anchovy can be explained largely by spring warming, summer cooling and by less abundant zooplankton in the late 1980s.     

Effects of seasonal variations in meteorology and oceanography on the Brazilian sardine fishery

Seasonal variations in meteorological and oceanographic processes in the South Brazil Bight were studied to assess their influence on the Brazilian sardine Sardinella brasiliensis fishery. According to its environmental characteristics, the South Brazil Bight was divided into three sectors. The northern and southern sectors were similar during the spring and summer, showing areas of coastal upwelling and low rainfall rates. The middle sector was characterized by the presence of low-salinity coastal waters in the inner shelf and by high rainfall. During autumn and winter, the middle and southern sectors were similar, both being under the influence of sub-Antarctic waters. Multiple regression analyses showed significant correlation between environmental data and catches, although at a lower level in the northern sector ( r ² = 0.57). In the middle and southern sectors, environmental data were highly correlated with the catches ( r ² = 0.90 and 0.84). The sardine landings in the middle sector are principally affected by meteorological parameters, and in the south by oceanographic ones. In the middle sector, an inverse correlation between landings and rainfall is shown, while in the south, landings are positively correlated with salinity and sea temperature. The conclusion is that the seasonal landings of the Brazilian sardine within the South Brazil Bight are influenced by the distribution of less saline waters, the coastal waters and sub-Antarctic waters.     

Remotely sensed mesoscale oceanography and the distribution of Illex argentinus in the South Atlantic

In this study, we consider the influence of mesoscale oceanographic processes around the Falkland Islands (Islas Malvinas) in the South-west Atlantic, during the period in which the commercial squid fishery for Illex argentinus operates. Spatially referenced fishery data and satellite-derived advanced very high resolution radiometry (AVHRR) sea surface temperature (SST) data were examined using geographic information system (GIS) techniques. The distribution and relative abundance of I. argentinus in the Falkland Islands fishery was examined for the period 1989–96. Three consistent areas of high abundance were observed to the north-east (shelf-break region) and north-west (shelf region) of the islands, and close to the northern coast of East Falkland. Areas of high sea surface temperature gradients (thermal gradients) extracted from remotely sensed satellite images were used as an indicator of mesoscale oceanographic activity and compared with the location of the fishery. I. argentinus were found to be associated with areas of thermal gradients, commonly seen at the interface of Falkland Current and Patagonian shelf waters. The techniques used in this analysis allow the overlay and analysis of physical oceanographic and fishery data with potential applications in fisheries management and operational fisheries oceanography.     

Alternating dominance of postlarval sardine and anchovy caught by coastal fishery in relation to the Kuroshio meander in the Enshu-nada Sea

In the mid 1970s, the fishery catch of postlarval Japanese anchovy (Engraulis japonica) in a shelf region of the Enshu‐nada Sea, off the central Pacific coast of Japan, started to decline corresponding to a rapid increase of postlarval sardine (Sardinops melanostictus). In late 1980s, sardine started to decline, and it was replaced by anchovy in the 1990s. This alternating dominance of postlarval sardine and anchovy corresponded to the alternation in egg abundance of these two species in the spawning habitat of this sea. It was also noteworthy that during the period of sardine decline, sardine spawning occurred in April–May, a delay of two months compared with spawning in the late 1970s. The implication of oceanographic changes in the spawning habitat for the alternating dominance of sardine and anchovy eggs was explored using time‐series data obtained in 1975–1998, focusing on the effect of the Kuroshio meander. Large meanders of the Kuroshio may have enhanced the onshore intrusion of the warm water into the shelf region and contributed to an increase in temperature in the spawning habitat. This might favour sardine, because its egg abundance in the shelf region was more dependent on the temperature in early spring than was that of anchovy. In addition, enhanced onshore intrusion could contribute to transport of sardine larvae from upstream spawning grounds of the Kuroshio region. On the other hand, anchovy egg abundance was more closely related to lower transparency at the shelf edge, which may indicate the prevalence and prolonged residence of the coastal water, and therefore higher food availability, frequently accompanying non‐meandering Kuroshio. The expansion/shrinkage of the spawning habitat of sardine and anchovy in the shelf region, apparently responding to the change in the Kuroshio, possibly makes the alternation in dominance of postlarval sardine and anchovy most prominent in the Enshu‐nada Sea, in combination with changes in the abundance of spawning adults, which occurred almost simultaneously in the overall Kuroshio region. The implication of this rather regional feature for the alternating dominance of sardine and anchovy populations on a larger spatial scale is also discussed.     

Modelling the effect of buoyancy on the transport of anchovy (Engraulis capensis) eggs from spawning to nursery grounds in the southern Benguela: an IBM approach

An individual‐based model (IBM) was used to investigate the effects of physical and biological variables on the transport via a jet current of anchovy (Engraulis capensis) eggs from spawning to the nursery grounds in the southern Benguela ecosystem. As transport of eggs and early larvae is considered to be one of the major factors impacting on anchovy recruitment success, this approach may be useful to understand further the recruitment variability in this economically and ecologically important species. By coupling the IBM to a 3D hydrodynamic model of the region called Plume, and by varying parameters such as the spatial and temporal location of spawning, particle buoyancy, and the depth range over which particles were released, we could assess the influences of these parameters on transport success. A sensitivity analysis using a General Linear Model identified the primary determinants of transport success in the various experimental simulations, and model outputs were examined and compared with patterns observed in field studies. Model outputs compared well with observed patterns of vertical and horizontal egg distribution. Particle buoyancy and area of particle release were the major single determinants of transport success, with an egg density of 1.025 g cm^?3 maximizing average particle transport success and the western Agulhas Bank being the most successful spawning area. This IBM may be useful as a generic prototype for other upwelling ecosystems.     

The global distribution of wind-induced upwelling

The global distribution of wind-induced upwelling at the base of the surface Ekman layer has been computed from the COADS monthly wind stress from 1950 to 1988. In contrast to previous studies where the Ekman transport equations became invalid near the Equator, this paper computed upwelling right to the equator by incorporating friction in the surface transport equations. Comparing the seasonal changes in the equatorial upwelling between the Atlantic and the Pacific, we found that the Atlantic had the larger magnitude change while the Pacific had the larger latitudinal shift. The anomalous wind-induced upwelling/downwelling associated with El Niño were computed over several latitudinal bands in the Pacific by compositing the data from six El Nino events. By plotting the July zonally averaged upwelling anomalies in the Pacific in a latitude-time contour plot, we found northward phase propagation at about 3.5° latitude per decade in the low latitudes, a feature not found in the January upwelling. Long-term upwelling trends were computed and their potential influence on fisheries are discussed.     

Modelling potential spawning habitat of sardine (Sardina pilchardus) and anchovy (Engraulis encrasicolus) in the Bay of Biscay

Large amplitude variations in recruitment of small pelagic fish result from interactions between a fluctuating environment and population dynamics processes such as spawning. The spatial extent and location of spawning, which is critical to the fate of eggs and larvae, can vary strongly from year to year, as a result of changing population structure and environmental conditions. Spawning habitat can be divided into ‘potential spawning habitat’, defined as habitat where the hydrographic conditions are suitable for spawning, ‘realized spawning habitat’, defined as habitat where spawning actually occurs, and ‘successful spawning habitat’, defined as habitat from where successful recruitment has resulted. Using biological data collected during the period 2000–2004, as well as hydrographic data, we investigate the role of environmental parameters in controlling the potential spawning habitat of anchovy and sardine in the Bay of Biscay. Anchovy potential spawning habitat appears to be primarily related to bottom temperature followed by surface temperature and mixed‐layer depth, whilst surface and bottom salinity appear to play a lesser role. The possible influence of hydrographic factors on the spawning habitat of sardine seems less clear than for anchovy. Modelled relationships between anchovy and sardine spawning are used to predict potential spawning habitat from hydrodynamical simulations. The results show that the seasonal patterns in spawning are well reproduced by the model, indicating that hydrographic changes may explain a large fraction of spawning spatial dynamics. Such models may prove useful in the context of forecasting potential impacts of future environmental changes on sardine and anchovy reproductive strategy in the north‐east Atlantic.