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.     

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.     

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.     

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.     

Characterizing the potential habitat of European anchovy Engraulis encrasicolus in the Mediterranean Sea,at different life stages

Identification of the potential habitat of European anchovy (Engraulis encrasicolus) at different life stages in relation to environmental conditions is an interesting subject from both ecological and management points of view. For this purpose, acoustic data from different seasons and different parts of the Mediterranean Sea along with satellite environmental and bathymetry data were modelled using generalized additive models. Similarly, egg distribution data from summer ichthyoplankton surveys were used to model potential spawning habitat. Selected models were used to produce maps presenting the probability of anchovy presence (adults, juveniles and eggs) in the entire Mediterranean basin, as a measure of habitat adequacy. Bottom depth and sea surface chlorophyll concentration were the variables found important in all models. Potential anchovy habitats were located over the continental shelf for all life stages examined. An expansion of the potential habitat from the peak spawning (early summer) to the late spawning season (early autumn) was observed. However, the most suitable areas for the presence of anchovy spawners seem to maintain the same size between seasons. Potential juvenile habitats were associated with highly productive inshore waters, being less extended and closer to coast during winter than late autumn. Potential spawning habitat in June and July based on ichthyoplankton surveys overlapped but were wider in extent compared with adult potential habitat from acoustics in the same season. Similarities and dissimilarities between the anchovy habitats as well as comparisons with sardine habitats in the oligotrophic Mediterranean Sea and other ecosystems with higher productivity are discussed.     

Simulation of the impact of climate change on the egg and larval transport of Japanese anchovy (Engraulis japonicus) off Kyushu Island,the western coast of Japan

In order to investigate the impact of climate change on egg and larval transport of Japanese anchovy (Engraulis japonicus) off Kyushu Island western Japan, we conducted particle‐tracking simulations on transport success/failure to fishing grounds from 1960 to 2007. The modeled transport success since the mid‐1990s increased and decreased in the offshore and coastal zones, respectively, compared with the 1960s and 1970s. The estimated northward shift of the spawning ground and weakened Tsushima Warm Current contributed to increase in modeled transport success to the offshore zone. Conversely, the weakening trend of the modeled onshore current in the Goto‐Nada Sea combined with the northward shift of the spawning ground resulted in unsuccessful larval transport. These results suggest that fluctuations in juvenile and subadult anchovy catches in this area may be attributable to changes in the physical environment. The present study showed that changes in transport success induced by oceanographic fluctuations related to climate change, have the potential to affect anchovy recruitment off the western coast of Japan.     

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.     

Historical fluctuations in spawning location of anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) in the Bay of Biscay during 1967–73 and 2000–2004

The spatial extent of small pelagic fish spawning habitat is influenced by environmental factors and by the state of the adult population. In return, the configuration of spawning habitat affects recruitment and therefore the future structure of the adult population. Interannual changes in spatial patterns of spawning reflect variations in adult population structures and their environment. The present study describes the historical changes in the spatial distribution of spawning of anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) in the Bay of Biscay during two periods: 1967–72 and 2000–2004. Using data from egg surveys conducted in spring, the spatial distributions of anchovy and sardine eggs are characterized by means of geostatistics. For each survey, a map of probability of egg presence is constructed. The maps are then compared to define (1) recurrent spawning areas, (2) occasional spawning areas and (3) unfavourable spawning areas during each period. Sardine spawning habitat is generally fragmented and appears spatially limited by the presence of cold bottom water. It is confined to coastal or shelf break refuge areas in years of restricted spawning extent. For anchovy, recurrent spawning sites are found in Gironde and Adour estuaries whilst spawning can extend further offshore in years of more intense spawning. For both species, the mean pattern of spawning has changed between 1967–72 and 2000–2004. Noticeably, the spatial distribution of anchovy eggs in spring has expanded northward. This trend possibly results from changes in environmental conditions during the last four decades.     

Remotely sensed spawning habitat of Pacific sardine (Sardinops sagax) and Northern anchovy (Engraulis mordax) within the California Current

We use trivariate kernel density estimation to define spawning habitat of northern anchovy ( Engraulis mordax ) and Pacific sardine ( Sardinops sagax ) in the California Current using satellite data and in situ egg samples from the Continuous Underway Fish Egg Sampler (CUFES) deployed during surveys in April by the California Cooperative Oceanic Fisheries Investigations (CalCOFI). Observed egg distributions were compared with monthly composite satellite sea surface temperature (SST) and surface chlorophyll a (chl a ) data. Based on the preferred spawning habitat, as defined in SST and chl a space, the satellite data were used to predict potential spawning habitat along two areas of the west coast of North America. Data from the southern area (21.5 to 39°N) were compared to observations from the CUFES data for the period 1998–2005. Northern anchovy and Pacific sardine exhibited distinctly different spawning habitat distributions. A significant relationship was found between satellite-based spawning area and that measured during surveys for sardine. CUFES area estimated for sardine was similar in magnitude to that estimated from satellite data (∼60 000 km²). In contrast, spawning habitat of anchovy averaged between 1000 and 200 000 km² for the period 1998–2005, for CUFES and satellite estimates, respectively. Interannual variability in the area (km²) and duration (months) of estimates of suitable habitat varied between species and between the northern (39 to 50.5°N) and southern portions of the California Current. Long-term monitoring of habitat variability using remote sensing data is possible in the southern portion of the California Current, and could be improved upon in the northern area with the addition of surveys better timed to describe relationships between observed and estimated spawning habitats.     

Environmental processes driving anchovy and sardine distribution in a highly variable environment: the role of the coastal structure and riverine input

Engraulis encrasicolus and Sardina pilchardus stocks are highly variable in terms of recruitment, biomass and spatial distribution. Changes in habitat conditions may influence both the survival of the early life stages and the adult stages. Detailed studies on the spatial distribution and habitat selection of such species have been performed in different areas of the world, highlighting the importance of environmental processes. The present study analyzes the spatial distribution of anchovy and sardine in the Tyrrhenian Sea in relation to environmental heterogeneity. Four acoustic surveys were carried out in this area in the period 2009–2014. Analysis of the environmental dataset permitted identification, in two specific areas, of a pattern of variables driving enrichment processes and impacting on the habitat suitability of the two species. In the northern and central parts of the study area, both anchovy and sardine showed a marked preference for shallower areas characterized by lower salinity. In these areas, PCA results on an environmental dataset highlighted a strong link between primary production, particulate organic carbon, distance from the mouth of the river, salinity and depth. A less clear picture was obtained for the southern part of the Tyrrhenian sea, characterized by a narrow continental shelf, moderately complex coastline morphology and the presence of very small rivers. Most of the anchovy biomass was found to be located in enclosed areas (gulfs) under the influence of relatively small rivers. This finding, taking into account that the surveys were carried out during the anchovy spawning period, highlights for such species a positive effect of the interaction between coastal morphology and riverine input, probably favoring food supply and retention of spawning products.     

Distribution and spawning dynamics of capelin (Mallotus villosus) in Glacier Bay, Alaska: a cold water refugium

Pacific capelin (Mallotus villosus) populations declined dramatically in the Northeastern Pacific following ocean warming after the regime shift of 1977, but little is known about the cause of the decline or the functional relationships between capelin and their environment. We assessed the distribution and abundance of spawning, non‐spawning adult and larval capelin in Glacier Bay, an estuarine fjord system in southeastern Alaska. We used principal components analysis to analyze midwater trawl and beach seine data collected between 1999 and 2004 with respect to oceanographic data and other measures of physical habitat including proximity to tidewater glaciers and potential spawning habitat. Both spawning and non‐spawning adult Pacific capelin were more likely to occur in areas closest to tidewater glaciers, and those areas were distinguished by lower temperature, higher turbidity, higher dissolved oxygen and lower chlorophyll a levels when compared with other areas of the bay. The distribution of larval Pacific capelin was not sensitive to glacial influence. Pre‐spawning females collected farther from tidewater glaciers were at a lower maturity state than those sampled closer to tidewater glaciers, and the geographic variation in the onset of spawning is likely the result of differences in the marine habitat among sub‐areas of Glacier Bay. Proximity to cold water in Glacier Bay may have provided a refuge for capelin during the recent warm years in the Gulf of Alaska.     

Environmental and biological monitoring for forecasting anchovy recruitment in the southern Benguela upwelling region

Environmental and biological sampling and monitoring have been carried out in the southern Benguela since 1988. The overall goal of this research is to investigate environmental factors affecting anchovy recruitment and to develop the ability to forecast anchovy recruitment from year-to-year using field data obtained during the spawning season (August to March). Sampling was conducted at three different temporal and spatial scales: during annual (November) broad-scale hydro-acoustic surveys to determine spawner biomass on the entire spawning ground and in the core transport and recruitment areas; during monthly surveys in the core spawning, transport and recruitment regions over two entire spawning seasons (1993/94 and 1994/95); and during weekly sampling (since 1995) along a single transect downstream from the spawning area. Annual surveys provide the best spatial coverage, but are inadequate for representing environmental conditions and anchovy spawning success over a prolonged season. Weekly sampling provides the best temporal coverage, but logistical constraints restrict information to a limited portion of the spawning area and a reduced number of variables. Monthly surveys provide intermediate coverage in time and space, but are expensive and labour-intensive. Forecasting anchovy recruitment has been based on two different approaches: the establishment of empirical relationships, and the development of rule-based expert systems. Forecasts from deterministic expert systems have compared well with final estimates of recruitment strength, and indicate that environmental and biological variables may be used in a structured way to forecast anchovy recruitment.     

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.     

Fine scale spatial pattern of Pacific sardine (Sardinops sagax) and northern anchovy (Engraulis mordax) eggs

Pacific sardine (Sardinops sagax) and northern anchovy (Engraulis mordax) eggs exhibited different spatial structure on the scale of 0.75–2.5 km in two egg patches sampled in the Southern California Bight in April 2000. Plankton samples were collected at 4‐min intervals with a Continuous Underway Fish Egg Sampler (CUFES) on 5 × 5 km grids centered on surface drifters. Variograms were calculated for sardine and anchovy eggs in Lagrangian coordinates, using abundances of individual developmental stages grouped into daily cohorts. Model variograms for sardine eggs have a low nugget effect, about 10% of the total variance, indicating high autocorrelation between adjacent samples. In contrast, model variograms for anchovy eggs have a high nugget effect of 50–100%, indicating that most of the variance at the scales sampled is spatially unstructured. The difference between observed spatial patterns of sardine and anchovy eggs on this scale may reflect the behavior of the spawning adults: larger, faster, more abundant fish may organize into larger schools with greater structure and mobility that create smoother egg distributions. Size and mobility vary with population size in clupeoids. The current high abundance of sardines and low abundance of anchovy off California agree with the greater autocorrelation of sardine egg samples and the observed tendency for locations of anchovy spawning to be more persistent on the temporal scale of days to weeks. Thus the spatial pattern of eggs and the persistence of spawning areas are suggested to depend on species, population size and age structure, spawning intensity and characteristic physical scales of the spawning habitat.     

`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.     

Exploring connectivity between spawning and nursery areas of Mullus barbatus (L., 1758) in the Mediterranean through a dispersal model

Connectivity between spawning and nursery areas plays a major role in determining the spatial structure of fish populations and the boundaries of stock units. Here, the potential effects of surface current on a red mullet population in the Central Mediterranean were simulated using a physical oceanographic model. Red mullet larvae were represented as Lagrangian drifters released in known spawning areas of the Strait of Sicily (SoS), which represents one of the most productive demersal fishing‐grounds of the Mediterranean. To consider the effect of inter‐annual variability of oceanographic patterns, numerical simulations were performed for the spawning seasons from 1999 to 2012. The main goal was to explore connectivity between population subunits, in terms of spawning and nursery areas, inhabiting the northern (Sicilian‐Maltese) and southern (African) continental shelves of the SoS. The numerical simulations revealed a certain degree of connectivity between the Sicilian–Maltese and the African sides of the SoS. Connectivity is present in both directions, but it is stronger from the Sicilian–Maltese spawning areas to the African nurseries owing to the marine circulation features of the region. However, because the majority of the larvae are transported to areas unsuitable for settlement or outside the SoS, the dispersal process is characterized by a strong loss of potential settlers born in the spawning areas. These results are in agreement with the low genetic heterogeneity reported for this species in the Mediterranean Sea and support the existence of a metapopulation structure of red mullet in the SoS and the adjacent areas.     

Recruitment of common sardine (Strangomera bentincki) and anchovy (Engraulis ringens) off central-south Chile in the 1990s and the impact of the 1997–1998 El Niño

The recruitment of Strangomera bentincki (common sardine) and Engraulis ringens (anchovy) and the relationships with oceanographic conditions in the upwelling ecosystem of central-south Chile were investigated from 1990 to 1998, with emphasis on the 1997–1998 El Niño. Time series of recruitment, biomass, local sea surface temperature, and a coastal upwelling index were used to explore relationships during the spawning (July–August) and pre-recruitment (August–December) periods. The 1997–1998 El Niño caused physical changes in the small pelagic fish habitat off central-south Chile. Anomalous sea surface temperatures (SST) and upwelling indexes began to be detected from May 1997 and persisted into 1998. Recruitment of common sardine showed significantly negative relationship with SST anomalies during the pre-recruitment period, as well as with the upwelling index during the peak of spawning. However, the recruitment of anchovy did not seem to be affected by the environmental changes observed in the 1990s. Instead, the recruitment rate of anchovy showed negative relationship with the recruitment rate of common sardine. We conclude that the conditions of the 1997–1998 El Niño off central-south Chile affected the survival of common sardine offspring, and that the recruitment success of anchovy could be determined by less-abundant cohorts of common sardine through a biological mechanism of interaction.     

Impacts of climate change on the complex life cycles of fish

To anticipate the response of fish populations to climate change, we developed a framework that integrates requirements in all life stages to assess impacts across the entire life cycle. The framework was applied on plaice (Pleuronectes platessa) and Atlantic herring (Clupea harengus) in the North Sea, Atlantic cod (Gadus morhua) in the Norwegian/Barents Seas and European anchovy (Engraulis encrasicolus) in the Bay of Biscay. In each case study, we reviewed habitats required by each life stage, habitat availability, and connectivity between habitats. We then explored how these could be altered by climate change. We documented environmental processes impacting habitat availability and connectivity, providing an integrated view at the population level and in a spatial context of potential climate impacts. A key result was that climate‐driven changes in larval dispersion seem to be the major unknown. Our summary suggested that species with specific habitat requirements for spawning (herring) or nursery grounds (plaice) display bottlenecks in their life cycle. Among the species examined, anchovy could cope best with environmental variability. Plaice was considered to be least resilient to climate‐driven changes due to its strict connectivity between spawning and nursery grounds. For plaice in the North Sea, habitat availability was expected to reduce with climate change. For North Sea herring, Norwegian cod and Biscay anchovy, climate‐driven changes were expected to have contrasting impacts depending on the life stage. Our review highlights the need to integrate physiological and behavioural processes across the life cycle to project the response of specific populations to climate change.     

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.     

Habitat and prey selection of common minke,sei, and Bryde’s whales in mesoscale during summer in the subarctic and transition regions of the western North Pacific

This study represents the first quantitative analysis of the characteristics of the distribution areas and stomach contents of common minke whale Balaenoptera acutorostrata, sei whale B. borealis, and Bryde’s whale B. edeni in relation to oceanographic and prey environments in mid summer in the western North Pacific. Common minke whales were distributed within subarctic regions and the northernmost region of the transitional domain, coinciding with the main habitat of their preferred prey, Pacific saury Cololabis saira. Sei whales were mainly found in the northernmost part of the transition zone and showed prey preference for Japanese anchovy Engraulis japonica, which was significantly more abundant in the main distribution area of the whale than in its adjacent areas. “Hot spots” of Bryde’s whales were found in several regions of the transition zone between the subarctic boundary and the Kuroshio front. This whale species preferred Japanese anchovy as prey, for which the distribution density was significantly higher in the main distribution area of the whale than in the adjacent areas. These results indicate that the summer distributions of Pacific saury and Japanese anchovy greatly influence the distributions of these whale species, suggesting that the whales’ habitat selection is closely related to their prey selection.