Pelagic ecology of a northern boundary current system: effects of upwelling on the production and distribution of sardine (Sardinops sagax), anchovy (Engraulis australis) and southern bluefin tuna (Thunnus maccoyii) in the Great Australian Bight

Shelf waters of southern Australia support the world's only northern boundary current ecosystem. Although there are some indications of intense nitrate enrichment in the eastern Great Australian Bight (GAB) arising from upwelling of the Flinders Current, the biological consequences of these processes are poorly understood. We show that productivity in the eastern GAB is low during winter, but that coastal upwelling at several locations during the austral summer–autumn results in localized increases in surface chlorophyll a concentrations and downstream enhancement of zooplankton biomass. Sardine (Sardinops sagax) and anchovy (Engraulis australis) eggs and larvae are abundant and widely distributed in shelf waters of the eastern and central GAB during summer–autumn, with high densities of sardine eggs and larvae occurring in areas with high zooplankton biomass. Egg densities and distributions support previous evidence suggesting that the spawning biomass of sardine in the waters off South Australia is an order of magnitude higher than elsewhere in southern Australia. Sardine comprised >50% of the identified prey species of juvenile southern bluefin tuna (SBT, Thunnus maccoyii) collected during this study. Other studies have shown that the lipid content of sardine from the GAB is relatively high during summer and autumn. We suggest that juvenile SBT migrate into the eastern and central GAB during each summer–autumn to access the high densities of lipid‐rich sardines that are available in the region during the upwelling period. Levels of primary, secondary and fish production in the eastern GAB during summer–autumn are higher than those recorded in other parts of Australia, and within the lower portion of ranges observed during upwelling events in the productive eastern boundary current systems off California, Peru and southern Africa.     

Entrainment and advection of larval sardine,Sardinops sagax,by the East Australian Current and retention in the western Tasman Front

The poleward flowing East Australian Current (EAC) drives sporadic upwelling, entrains coastal water and forms the western Tasman Front (wTF), creating a mosaic of water types and larval transport routes along south eastern Australia. The spatial distribution, otolith chemistry and growth rates of larval sardine (Sardinops sagax) were examined to infer spawning location and larval transport. A gradient of increasing larval size from north to south along the shelf was not detected but was evident between the shelf and offshore in the wTF. Here larvae were larger and older. Based on the occurrence of newly hatched larvae, spawning by S. sagax between southern Queensland and mid New South Wales (NSW) was more extensive than previously reported. The otolith chemistry from two wTF larval size classes differed, implying different origins. The otolith chemistry of wTF post‐flexion larvae was similar to larvae from northern NSW, whereas wTF flexion larvae were similar to larvae observed nearby from mid‐NSW. Two possible larval transport routes, direct and indirect, are inferred from otolith chemistry, current velocities and a previously published particle tracking study. Either larvae from northern NSW were advected south and entrained with younger larvae directly into the wTF, or larvae from a range of shelf regions were advected around the southern edge of an anticyclonic eddy, to join younger larvae directly entrained into the wTF. Based on the co‐occurrence of larval ages and sizes in the wTF and their advection routes, the wTF appears to be an important larval retention zone.     

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.     

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.     

Advection of anchovy (Engraulis encrasicolus) larvae along the Catalan continental slope (NW Mediterranean)

The Gulf of Lions is one of the main anchovy (Engraulis encrasicolus) spawning areas in the NW Mediterranean. During the spring, low‐salinity surface water from the outflow of the Rhône is advected by the shelf‐slope current along the continental slope off the Catalan coast. In June 2000, a Lagrangian experiment tracking these low‐salinity surface waters was conducted to assess the importance of this transport mechanism for anchovy larvae and to determine the suitability of the tracked surface waters for survival of anchovy larvae. The experiment consisted of sampling the tracked water parcel for 10 days with three drifters launched at the core of the shelf‐slope current where low‐salinity surface waters were detected. The survey was completed by sampling the surrounding waters. Anchovy larvae from the spawning area in the Gulf of Lions were advected towards the south in the low‐salinity waters. The size increase of anchovy larvae throughout the Lagrangian tracking closely followed the general growth rate calculated by otolith analysis (0.65 mm day⁻¹). However, advection by the current was not the only mechanism of anchovy larval transport. A series of anticyclonic eddies, originated in the Gulf of Lions and advected southwards, seemed to play a complementary role in the transport of larvae from the spawning ground towards the nursery areas. These eddies not only contributed to larval transport but also prevented their dispersion. These transport and aggregation mechanisms may be important for anchovy populations along the Catalan coast and require further study.     

Entrainment of larval fish assemblages from the inner shelf into the East Australian Current and into the western Tasman Front

Entrainment and transport of larval fish assemblages by the East Australian Current (EAC) were examined from the coastal waters of northern New South Wales (NSW) to the western Tasman Front, via the separation of the EAC from the coast, during the austral spring of 2004. Shore‐normal transects from the coast to the EAC off northern NSW revealed an inner shelf assemblage of near‐shore families (Clupeidae, Engraulidae, Platycephalidae and Triglidae), an EAC assemblage dominated by Myctophidae and Gonostomatidae, and a broadly distributed assemblage over the continental shelf dominated by Scombridae and Carangidae. Further south and after the EAC had separated from the coast, we observed a western Tasman Front assemblage of inner shelf and shelf families (Clupeidae, Engraulidae, Serranidae, Scombridae, Carangidae, Bothidae and Macroramphosidae). The abundance of these families declined with distance from the coast. Surprisingly, there was no distinctive or abundant larval fish assemblage in the chlorophyll‐ and zooplankton‐enriched waters of the Tasman Sea. Water type properties (temperature‐salinity, T‐S), the larval fish assemblages and family‐specific T‐S signatures revealed the western Tasman Front to be an entrained mix of EAC and coastal water types. We found an abundance of commercially important species including larval sardine (Sardinops sagax, Clupeidae), blue mackerel (Scomber australasicus, Scombridae) and anchovy (Engraulis australis, Engraulidae). The entrainment and transport of larval fish from the northern inner shelf to the western Tasman Front by the EAC reflects similar processes with the Gulf Stream Front and the Kuroshio Extension.     

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.     

Importance of the Shatsky Rise Area in the Kuroshio Extension as an offshore nursery ground for Japanese anchovy (Engraulis japonicus) and sardine (Sardinops melanostictus)

Recent findings suggest that recruitment of Japanese anchovy (Engraulis japonicus) and sardine (Sardinops melanostictus) depends on survival during not only the first feeding larval stage in the Japanese coastal waters and the Kuroshio front but also during the post‐larval and juvenile stages in the Kuroshio Extension. Spatial distributions of juvenile anchovy and sardine around the Shatsky Rise area in the Kuroshio Extension region and the Kuroshio–Oyashio transition region are described, based on a field survey in the late spring using a newly developed mid‐water trawl for sampling juveniles. All stages of anchovy from post‐larvae to juveniles were obtained in the northern Shatsky Rise area. The Kuroshio Extension bifurcates west of the Shatsky Rise area and eddies are generated, leading to higher chlorophyll concentrations than in the surrounding regions in April and May. When Japanese anchovy and sardine spawn near the Kuroshio front or the coastal waters south‐east of Japan, their larvae are transported by the Kuroshio Extension and are retained in the Shatsky Rise area, which forms an important offshore nursery ground, especially during periods of high stock abundance.     

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.     

Does upwelling intensity determine larval fish habitats in upwelling ecosystems? The case of Senegal and Mauritania

European sardine (Sardina pilchardus) and round sardinella (Sardinella aurita) comprise two‐thirds of total landings of small pelagic fishes in the Canary Current Eastern Boundary Ecosystem (CCEBE). Their spawning habitat is the continental shelf where upwelling is responsible for high productivity. While upwelling intensity is predicted to change through ocean warming, the effects of upwelling intensity on larval fish habitat expansion is not well understood. Larval habitat characteristics of both species were investigated during different upwelling intensity regimes. Three surveys were carried out to sample fish larvae during cold (permanent upwelling) and warm (low upwelling) seasons along the southern coastal upwelling area of the CCEBE (13°–22.5°N). Sardina pilchardus larvae were observed in areas of strong upwelling during both seasons. Larval habitat expansion was restricted from 22.5°N to 17.5°N during cold seasons and to 22.5°N during the warm season. Sardinella aurita larvae were observed from 13°N to 15°N during cold seasons and 16–21°N in the warm season under low upwelling conditions. Generalized additive models predicted upwelling intensity driven larval fish abundance patterns. Observations and modeling revealed species‐specific spawning times and locations, that resulted in a niche partitioning allowing species' co‐existence. Alterations in upwelling intensity may have drastic effects on the spawning behavior, larval survival, and probably recruitment success of a species. The results enable insights into the spawning behavior of major small pelagic fish species in the CCEBE. Understanding biological responses to physical variability are essential in managing marine resources under changing climate conditions.     

Investigating the ‘northern Humboldt paradox’ from model comparisons of small pelagic fish reproductive strategies in eastern boundary upwelling ecosystems

Eastern boundary upwelling ecosystems are highly productive and sustain the world’s largest fisheries, usually dominated by sardine and anchovy species. Stock size is highly variable from year to year due to the impact of the unstable physical environment on fish early stages. Biophysical models of early life‐stage dispersal of marine organisms have been built by coupling (i) hydrodynamic models and (ii) life history models (i.e. egg and larva stages), and are therefore useful tools to investigate physical–biological interactions. Here, we review biophysical models of anchovy and sardine ichthyoplankton dispersals developed in the Benguela, Humboldt and Canary Current upwelling ecosystems. We also include a similar study conducted in the California Current upwelling on zooplankton. We then integrate this information into a comparative analysis of sardine and anchovy reproductive strategies in the different systems. We found that the main spawning periods match the season of (i) maximal simulated ichthyoplankton retention over the continental shelf in the northern Benguela, southern Humboldt and Canary (for sardine); (ii) maximal food concentration in the southern Benguela, California and Canary (for anchovy); and (iii) maximal shelf retention of ichthyoplankton and food concentration in the northern Humboldt (for both anchovy and sardine). This specificity of the northern Humboldt ecosystem could explain why it sustains the largest small pelagic fish stock. Finally, the possible effects of climate change on these patterns are discussed.     

Impact of freshwater input and wind on landings of anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) in shelf waters surrounding the Ebre (Ebro) River delta (north-western Mediterranean)

Time series analyses (Box–Jenkins models) were used to study the influence of river runoff and wind mixing index on the productivity of the two most abundant species of small pelagic fish exploited in waters surrounding the Ebre (Ebro) River continental shelf (north‐western Mediterranean): anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus). River flow and wind were selected because they are known to enhance fertilization and local planktonic production, thus being crucial for the survival of fish larvae. Time series of the two environmental variables and landings of the two species were analysed to extract the trend and seasonality. All series displayed important seasonal and interannual fluctuations. In the long term, landings of anchovy declined while those of sardine increased. At the seasonal scale, landings of anchovy peaked during spring/summer while those of sardine peaked during spring and autumn. Seasonality in landings of anchovy was stronger than in sardine. Concerning the environmental series, monthly average Ebre runoff showed a progressive decline from 1960 until the late 1980s, and the wind mixing index was highest during 1994–96. Within the annual cycle, the minimum river flow occurs from July to October and the wind mixing peaks in winter (December–April, excluding January). The results of the analyses showed a significant correlation between monthly landings of anchovy and freshwater input of the Ebre River during the spawning season of this species (April–August), with a time lag of 12 months. In contrast, monthly landings of sardine were significantly positively correlated with the wind mixing index during the spawning season of this species (November–March), with a lag of 18 months. The results provide evidence of the influence of riverine inputs and wind mixing on the productivity of small pelagic fish in the north‐western Mediterranean. The time lags obtained in the relationships stress the importance of river runoff and wind mixing for the early stages of anchovy and sardine, respectively, and their impact on recruitment.     

Turbulent mixing and mesoscale distributions of late-stage fish larvae on the NW Shelf of Western Australia

Light traps were deployed to describe vertical and cross‐shelf distributions of late‐stage larval fishes during five cruises in each of the 1997/98 and 1998/99 summers in the region of the Gulf of Exmouth on the southern North West Shelf of Western Australia. At each light trap station on a cross‐shelf transect we measured water temperature, salinity and chlorophyll a and used vertical plankton tows to estimate zooplankton biomass and copepod abundance. Current meters were deployed on moorings near the transect and the data used to model flows and mixing on the NW Shelf and in the Gulf. The majority of reef, pelagic and baitfish larvae (81, 83 and 66% respectively) were collected at only two stations that marked the boundary between stratified waters offshore and well‐mixed water within the Gulf. Most baitfishes (primarily Spratelloides spp.) were captured by traps deployed near the seabed, while reef fishes (mostly pomacentrids, lethrinids and siganids) and pelagic species (mostly scombrids and carangids) were captured in traps deployed near surface. Catch composition varied between summers with 64% of baitfishes collected in the first summer, while the majority of reef and pelagic fishes (81 and 80% respectively) were captured in the second summer. Modelling of circulation showed that the velocity of tidal currents was enhanced by constriction of flow between NW Cape and South Muiron Island and by shallowing of the shelf. Flood‐tide intrusions of water allowed the thermocline to move up the continental shelf, upwelling cool nutrient‐rich water that was then mixed throughout the entire water column at stations in the mouth of the Gulf. This upwelling and mixing resulted in higher chlorophyll a concentrations and copepod abundances either as a result of local in situ growth or advection/aggregation processes, and may account for the great abundances of late‐stage fish larvae in the mouth of the Gulf.     

Spawning patterns provide further evidence for multiple stocks of sardine (Sardinops sagax) off eastern Australia

Previous studies have suggested that sardine, Sardinops sagax, off eastern Australia spawns across its entire range when habitat conditions are suitable. However, recent studies have suggested that separate sub‐populations and spawning groups may occur in the region. Spawning patterns off eastern Australia were investigated using data collected during nine ichthyoplankton surveys conducted between 1997–2015, and adult reproductive data obtained from ad hoc commercial sampling off New South Wales (NSW). The egg surveys covered the known distribution of sardine off eastern Australia and included year‐round sampling in the northern and southern parts of this range. Egg distributions and analysis of gonadosomatic indices identified two spatio‐temporally separate spawning groups; one occurring off southern Queensland to northern NSW during late winter and early spring, and a smaller group off eastern Tasmania to southern NSW during summer. Most eggs were collected from waters 50–90 m deep, with sea surface temperatures of 18–23°C. Additive modelling indicated depth was the most significant factor driving selection of spawning habitat, followed by the interaction of month and latitude. Low egg densities were recorded in waters between 34–37°S, despite conditions within the ranges suitable for spawning. The presence of two spawning groups of sardine off eastern Australia supports recent findings that two sub‐populations occur in the region. Findings of this study will help to optimise the spatio‐temporal extent of future egg surveys and further confirm the need to coordinate future management of each sub‐population among relevant jurisdictions.     

Transport and environmental temperature variability of eggs and larvae of the Japanese anchovy (Engraulis japonicus) and Japanese sardine (Sardinops melanostictus) in the western North Pacific estimated via numerical particle-tracking experiments

Numerical particle-tracking experiments were performed to investigate the transport and variability in environmental temperature experienced by eggs and larvae of Pacific stocks of the Japanese anchovy ( Engraulis japonicus ) and Japanese sardine ( Sardinops melanostictus ) using high-resolution outputs of the Ocean General Circulation Model for the Earth Simulator (OFES) and the observed distributions of eggs collected from 1978 to 2004. The modeled anchovy individuals tend to be trapped in coastal waters or transported to the Kuroshio–Oyashio transition region. In contrast, a large proportion of the sardines are transported to the Kuroshio Extension. The egg density-weighted mean environmental temperature until day 30 of the experiment was 20–24°C for the anchovy and 17–20°C for the sardine, which can be explained by spawning areas and seasons, and interannual oceanic variability. Regression analyses revealed that the contribution of environmental temperature to the logarithm of recruitment per spawning (expected to have a negative relationship with the mean mortality coefficient) was significant for both the anchovy and sardine, especially until day 30, which can be regarded as the initial stages of their life cycles. The relationship was quadratic for the anchovy, with an optimal temperature of 21–22°C, and linear for the sardine, with a negative coefficient. Differences in habitat areas and temperature responses between the sardine and anchovy are suggested to be important factors in controlling the dramatic out-of-phase fluctuations of these species.     

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

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

Subtropical water influences temporal fluctuations of early life stages of Vinciguerria lucetia (Osteichthyes: Phosichthyidae) in the Humboldt Current System (1998–2004)

In the last decades, the fish Vinciguerria lucetia (Garman) has been of important interest to the fisheries sector; nonetheless, the spawning and nursery zones in the Humboldt Current System (HCS) have not yet been defined. By using a temporal series of 23 oceanographic surveys from austral spring of 1998 to autumn 2004 off northern Chile, the spatial and temporal distribution and abundance of eggs and larvae of V. lucetia were studied. The relationships with environmental conditions (sea surface temperature, water column stratification, salinity, dissolved oxygen) were modeled using generalized additive models (GAMs). Seasonal variations in eggs and larval abundances were recorded, and higher abundances were observed in spring and summer, respectively. The main spawning areas were located at approximately between 40 and 80 nautical miles offshore. The largest abundances of V. lucetia eggs were found during spring 2003; however, larval abundances reached the highest values following the strongest ENSO event 1997–98. GAMs predicted that offshore location, sea surface temperature, and the deepening of the oxygen minimum zone, characteristics of the subtropical waters (22–24°C, >34.9, 3–6 ml/L) drove eggs and larval distributions of V. lucetia in the HCS during 1998–2004, toward areas with scarce food availability for larvae. These results suggest that spawning and larval development of this oceanic species occur in oligotrophic waters as a loophole strategy, in order to reduce predation risk during early life stages.     

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.     

`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.     

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.