Decadal changes in fish assemblages in waters near the Ieodo ocean research station (East China Sea) in relation to climate change from 1984 to 2010

We compiled and analyzed past time-series data to evaluate changes in oceanographic conditions and marine ecosystems near the Ieodo ocean research station (IORS) in the East China Sea (N 31°15??C33°45??, E 124°15??C127°45??) in relation to longterm changes in climate and global warming. The environment data we used was a depth-specific time-series of temperature and salinity for the water columns at 175 fixed stations along 22 oceanographic lines in Korean waters, based on bimonthly measurements since 1961 taken by the National Fisheries Research & Development Institute. As an indicator for the ecosystem status of the waters off Ieodo, we analyzed species composition in biomass of fishes caught by Korean fishing vessels in the waters near the IORS (1984?C2010) and summarized the data in relation to the environmental changes using canonical correspondence analysis (CCA). To detect step changes in the time-series of environmental factors, we applied a sequential t-test analysis of regime shift. Correspondence analysis detected a major shift in fish assemblage structure between 1990 and 1993: the dominant species was filefish during 1981?C1992, but chub mackerel during 1992?C2007. This shift in fish assemblage structure seemed to be related to the well-established 1989 regime shift in the North Pacific, which was confirmed again with respect to temperature in the Yellow Sea and the Korea Strait (but not in the waters off the IORS). In overall from 1984 to 2010, salinity was more important than water temperature in CCA, implying that the fluctuation of the Tsushima warm current is a most important force driving the long-term changes in fish assemblage structure in the waters off the IORS. Further multidisciplinary researches are required to identify oceanographic and biological processes that link climate-driven physical changes to fish recruitment and habitat range fluctuations.     

Variations in the abundance of fisheries resources and ecosystem structure in the Japan/East Sea

Evidence supports the hypothesis that two climatic regime shifts in the North Pacific and the Japan/East Sea, have affected the dynamics of the marine ecosystem and fisheries resources from 1960 to 2000. Changes in both mixed layer depth (MLD) and primary production were detected in the Japan/East Sea after 1976. The 1976 regime shift appears to have caused the biomass replacement with changes in catch production of major exploited fisheries resources, including Pacific saury, Pacific sardine and filefish. Both fisheries yield and fish distribution are reflected in these decadal fluctuations. In the 1960s and 1990s, common squid dominated the catches whereas in the 1970s and 1980s, it was replaced by walleye pollock. In the post-1988 regime shift, the distribution of horse mackerel shifted westward and southward and its distributional overlap with common mackerel decreased. The habitat of Pacific sardine also shifted away from mackerel habitats during this period. To evaluate changes in the organization and structure of the ecosystem in the Japan/East Sea, a mass-balanced model, Ecopath, was employed. Based on two mass-balanced models, representing before (1970–75) and after (1978–84) the 1976 regime shift, the weighted mean trophic level of catch increased from 3.09 before to 3.28 after. Total biomass of species groups in the Japan/East Sea ecosystem increased by 15% and total catch production increased by 48% due to the 1976 regime shift. The largest changes occurred at mid-trophic levels, occupied by fishes and cephalopods. The dominant predatory species shifted from cephalopods to walleye pollock due to the 1976 regime shift. It is concluded that the climatic regime shifts caused changes in the structure of the ecosystem and the roles of major species, as well as, large variations in biomass and production of fisheries resources.     

Comparing trophic flows in the southern Benguela to those in other upwelling ecosystems

A balanced trophic flow model of the southern Benguela ecosystem is presented, averaging the period 1980–1989 and emphasizing upper trophic levels. The model is based largely on studies conducted within the framework of the Benguela Ecology Programme and updates the results of an expert workshop held in Cape Town in September 1989. Small pelagic fish other than anchovy Engraulis capensis and sardine Sardinops sagax, mainly round herring Etrumeus whiteheadi and mesopelagic fish, were important components of the food web in the southern Benguela. Severe balancing difficulties were encountered with respect to the semi-pelagic resources (hake Merluccius spp.) and demersal top predators (sharks), indicating the need for further research on the interaction of these groups with their ecosystem. The model is compared to other existing trophic flow models of ecosystems in major upwelling areas, i.e. the northern Humboldt Current (4–14°S), the California Current (28–42°N) and the southern Canary Current (l2–25°N), and to two independently constructed models of the northern Benguela ecosystem. These models are compared using network analysis routines of the ECOPATH software, focusing on the interactions between the five dominant fish species (anchovy, sardine, horse mackerel Trachurus trachurus capensis, chub mackerel Scomber japonicus and hake) that support important fisheries in all systems. The upwelling systems rank by size rather than species dominance. The ratio of catches and primary production differs between systems, partly because of differences in fishing regimes. Predation on the five dominant fish groups by other fish in the system was the most important cause of fish mortality in all models. Fishery catches are generally a larger cause of mortality for these groups than predation by mammals. The ecological cost of fishing appears to be comparatively low in the southern Benguela, because catches are low compared with the primary production, but also because the fishery is relatively low in the foodweb. However, in view of the very tight foodweb demonstrated in the model. it is likely that an increase in fishing pressure would cause severe trade-offs with respect to other components of the southern Benguela ecosystem.     

Interannual variations of the occurrence of epipelagic fish in the diets of the seabirds breeding on Teuri Island, northern Hokkaido, Japan

The diets of breeding seabirds can be a good monitor of marine environmental changes. From 1984 to 2001 we monitored the diets of black-tailed gulls (Larus crassirostris) (“surface foragers”), rhinoceros auklets (Cerorhinca monocerata) (“epipelagic divers”), and Japanese cormorants (Phalacrocorax filamentotus) (“bottom divers”) that breed on Teuri Island at the northern boundary of the Tsushima Warm current in the Sea of Japan/East Sea. Between 1984 and 1987, both the gulls and the auklets foraged on the sardine (Sardinops melanostictus), but after 1992, they switched to the anchovy (Engraulis japonica). This change might reflect the collapse of the sardine stock in the late 1980s. In the 1990s, the year-to-year variations of the percentage of anchovy in the diets of the three seabird species showed similar trends: High in 1994 and 1998–2001; and low in 1992–1993 and 1995–1997. The estimated stock size of the anchovy population in the Tsushima Current area was positively correlated with the percentage of mass of anchovy in the seabirds’ diets. Thus, the short-term annual changes of the total anchovy availability, which might reflect SST or the volume transport of Tsushima Current, possibly affected the seabirds diets on this island.     

The relationship of anchovy and sardine to water masses in the Peruvian Humboldt Current System from 1983 to 2005

The Humboldt Current System (HCS) is dominated by two pelagic species; Peruvian anchovy or anchoveta (Engraulis ringens) and sardine (Sardinops sagax). Using data from 43 acoustic surveys conducted from 1983 through 2005 by the Peruvian Marine Institute (IMARPE), we examined the distribution of these two species relative to water masses. We tested the hypothesis that anchovy was found more frequently in upwelled cold coastal water (CCW) and mixed waters (MCW) than in other water types and that sardine was more associated with more offshore oceanic surface subtropical water (SSW). Surface temperature, salinity, latitude, season and distance to the coast data were used to define water masses. Results using generalized additive models (GAM), modelling sardine and anchovy presence–absence as a function of year, water body, bottom depth and latitude, showed that anchovy were primarily found in CCW and MCS, while sardine were more ubiquitous relative to water masses with some predilection for SSW. These results were supported by various indexes of anchovy and sardine distribution versus water mass as well as temporal and location variables.     

The influence of mesoscale physical processes on the larval fish community in the Canaries CTZ, in summer

We have studied the relation between the hydrography, the composition and horizontal structure of the larval fish community, and the horizontal distribution patterns of larval fish abundances in an area characterised by strong mesoscale oceanographic activity, located between the Canary Islands and the African coast (the Canaries Coastal Transition Zone), during August 1999. Upwelling, upwelling filaments, cyclonic and anticyclonic eddies and island wakes are typical mesoscale features of the northwest African coast in summer. A single upwelling filament off Cabo Juby was joined in mid-August by a second that originated about 100 km to the north. The two filaments flowed together and merged 100 km offshore. The merged filament was partially entrained around a cyclonic eddy, trapped between the Canary Islands and the African coast, and interacted with cyclonic and anticyclonic eddies shed from Gran Canaria. Mesoscale oceanographic features strongly influenced the horizontal distributions of fish larvae. Eddies acted as a mechanism of concentration, while upwelling filaments were dispersive, transporting larvae from the African neritic zone into oceanic areas and towards the Canary archipelago. This transport was the major cause of the predominance of neritic larvae in the composition of the larval fish community of the area. The results also suggest: (1) that anchovy larvae are good indicators of the offshore displacement of upwelled water; (2) that the alternation between anchovy and sardine as species dominant in the larval fish community of the area during summer depends upon the water temperature in the African upwelling region, anchovy dominating at higher temperature; (3) that a coupling of anchovy and sardine spawning with the mesoscale oceanographic structure formed by the upwelling filaments and trapped eddy overcomes the negative effect that Ekman transport has on their populations.     

Characterising and comparing the spawning habitats of anchovy Engraulis encrasicolus and sardine Sardinops sagax in the southern Benguela upwelling ecosystem

The spawning habitats of anchovy Engraulis encrasicolus and sardine Sardinops sagax in the southern Benguela upwelling ecosystem were characterised by comparing their egg abundances with environmental variables measured concomitantly during two different survey programmes: the South African Sardine and Anchovy Recruitment Programme (SARP), which comprised monthly surveys conducted during the austral summers of 1993/94 and 1994/95; and annual pelagic spawner biomass surveys conducted in early summer (November/December) from 1984 to 1999. Eggs were collected using a CalVET net. Physical variables measured included sea surface temperature (SST), surface salinity, water depth, mixed-layer depth, and current and wind speeds; biological variables measured included phytoplankton biomass, and zooplankton biomass and production. Spawning habitat was identified by construction of quotient curves derived from egg abundance data and individual environmental variables, and relationships between these variables were determined using multivariate co-inertia analysis. SARP data showed that anchovy spawning was associated with cool water and moderate wind and current speeds, whereas sardine spawning was related to warmer water and more turbulent and unstable conditions (i.e. high wind speeds and strong currents) than for anchovy. SARP data also showed significant differences in selection of spawning habitat of the two species for all environmental variables. The relationship between anchovy egg abundance and salinity was strongly positive, but strongly negative with water depth, phytoplankton biomass and zooplankton production. Sardine egg abundance was strongly positively related to current speed. The spawner biomass survey data demonstrated that the spawning habitat of anchovy was characterised by warm water and high salinity, whereas sardine spawning was associated with cool water and low salinity. The survey data showed significant differences in spawning habitat selection by anchovy and sardine for SST, salinity and zooplankton biomass, but not for the other environmental variables. There was a positive relationship between anchovy egg abundance and SST, salinity and mixed-layer depth, and a negative relationship with water depth, phytoplankton biomass and zooplankton production. For sardine there was a strong positive relationship between egg abundance and current speed and wind speed. Differences in the results between the two survey programmes could be attributable to differences in their spatio-temporal coverage. Spawning habitats of anchovy and sardine appear to be substantially different, with anchovy being more specific than sardine in their preference of various environmental conditions.     

An overview of the Oyashio ecosystem

The Oyashio shelf region and the seasonally ice-covered areas north of Hokkaido are highly productive, supporting a wide range of species including marine mammals, seabirds and commercially important species in the western subarctic Pacific. The fishes include gadids, such as walleye pollock and Pacific cod, and subarctic migratory pelagic fishes such as chum salmon and pink salmon. It is also an important summer feeding ground for subtropical migrants such as the Japanese sardine, Japanese anchovy, Pacific saury, mackerels, Japanese common squid, whales and seabirds. In recent decades, some components of the Oyashio ecosystem (i.e., phytoplankton, mesozooplankton, gadid fish, and subtropical migrants) have shown changes in species abundance or distribution that are correlated with environmental changes such as the 1976/1977 and 1988/1989 regime shifts. The First Oyashio Intrusion moved northward from the mid-1960s until the late 1970s, when it moved southward until the 1980s, after which it returned to the north again after the mid-1990s. The sea-surface temperature in spring decreased after the late 1970s, increased after the late 1980s, and remained high during the 1990s. The extent of ice cover in the Sea of Okhostk also decreased during the latest warming in the 1980–1990s but has increased again since the late 1990s. This and other variabilities affect the Oyashio ecosystem and the surrounding region.     

Long-term changes in the fish community structure from the Tsushima warm current region of the Japan/East Sea with an emphasis on the impacts of fishing and climate regime shift over the last four decades

Japanese fisheries production in the Japan/East Sea between 1958 and 2003 increased to their peak (1.76 million tons) in the late 1980s and decreased abruptly with the collapse of Japanese sardine. Catch results for 58 fisheries and various environmental time-series data sets and community indices, including mean trophic level (MTL) and Simpson’s diversity index (DI), were used to investigate the impacts of fishing and climate changes on the structure of the fish community in the Tsushima warm current (TWC) region of the Japan/East Sea. The long-term trend in fisheries production was largely dependent on the Japanese sardine that, as a single species, contributed up to 60% of the total production in the Japanese waters of the Japan/East Sea during the late 1980s. Excluding Japanese sardine, production of the small pelagic species was higher during 1960s and 1990s but lower during 1970s and 1980s. This variation pattern generally corresponds with the trend in water temperature, warmer before early 1960s and after 1990s but colder during 1970s and 1980s. The warm-water, large predatory fishes and cold water demersal species show opposite responses to the water temperature in the TWC region, indicating the significant impact of oceanic conditions on fisheries production of the Japan/East Sea. Declines in demersal fishes and invertebrates during 1970s and 1980s suggested some impact of fishing. MTL and DI show a similar variation pattern: higher during 1960s and 1990s but lower during 1970s and 1980s. In particular, the sharp decline during the 1980s resulted from the abundant sardine catches, suggesting that dominant species have a large effect on the structure of the fish community in the Japan/East Sea. Principal component analysis for 58 time-series data sets of fisheries catches suggested that the fish community varied on inter-annual to inter-decadal scales; the abrupt changes that occurred in the mid-1970s and late 1980s seemed to correspond closely with the climatic regime shifts in the North Pacific. These results strongly suggest that the structure of the fish community in the Japan/East Sea was largely affected by climatic and oceanic regime shifts rather than by fishing. There is no evidence showing “fishing down food webs” in the Japan/East Sea. However, in addition to the impacts of abrupt shifts that occurred in the late 1980s, the large predatory and demersal fishes seem to be facing stronger fishing pressure with the collapse of the Japanese sardine.     

250 years of sardine and anchovy scale deposition record in Mejillones Bay, northern Chile

Marine oxygen-deficient environments with high sedimentation rates and high primary productivity can provide relevant information regarding variations of ocean–climatic conditions in the past. In the Humboldt current ecosystem, which now hosts huge populations of pelagic fishes (mainly anchovy and sardine), fish scale abundance in the sedimentary record may be useful indicators of environmental change. Here we assess such a proxy record in a 42 cm-long sedimentary core collected from 80 m in Mejillones Bay (23°S, northern Chile). We also analyse fish remains in surface sediment sampled along a bathymetric transect (from 10 to 110 m water depth) in the same bay. In the core-top record, the fluctuations of sardine and anchovy scale deposition rates (SDR) agreed with those of industrial catches for these two species in northern Chile, tending to validate the SDR as a proxy of local fish biomass when bottom anoxic conditions prevail. However, apparent SDR for records prior to 1820 have probably been influenced by dissolution processes linked to the oxygenation of the bottom environment of Mejillones Bay, as suggested by other proxy records. After 1820, the fluctuations in the relative abundance of sardine and anchovy scales point to alternating warm and cold conditions during about 30 years and then a progressively cooler period. Since ca. 1870, marked fluctuations of SDR of both species are observed, probably as a consequence of the onset of a different oceanographic regime characterized by intensified upwelling, stronger subsurface oxygen deficiency, higher primary productivity, and enhanced “ENSO-like” interdecadal variability. While anchovy SDR fluctuated in periods of 25–40 years, only two peak periods of sardine SDR occurred (late 19th century and late 20th century), suggesting that sardine abundance depends on other ocean–climatic factors.     

Abundance and distribution of rhynchoteuthion larvae of Illex argentlnus (Cephalopoda: Ommastrephidae) in the south-western Atlantic

The results of ichthyoplankton surveys carried out by the Instituto Nacional de Pesca (INAPE) in the Argentine–Uruguayan Common Fishing Zone (AUCFZ) in summer 1980 and the winters of 1980–1982 are presented. In addition, the results of joint research with INIDEP, the Argentine and Japan Fisheries Agency, on larvae, juveniles and adults of the short-finned squid Illex argentinus carried out in winter 1989 are given, as are data obtained during the joint sardine and anchovy recruitment project carried out by R.V. Meteor in November/December 1989. The short-finned squid Illex argentinus is the most important cephalopod resource of the area, and an extended fishery is carried out on the Patagonian shelf and in the proximity of the Falkland Islands. The fishing season in the AUCFZ, for pre-spawning and spawning animals, extends from early autumn to winter (March–August). Abundance of rhynchoteuthion larvae (417–485 larvae·10 m^?2) was greatest in winter in surface temperatures of 12,1–16,9°C and surface salinities of 34,8 – 35,2 × 10^?3. The distribution of the larvae is related to the western boundary of the Brazil Current and the convergence front.     

Seasonal comparison of the diets of juvenile European anchovy Engraulis encrasicolus and sardine Sardina pilchardus in the Gulf of Lions

Anchovy and sardine in the Mediterranean are known to share the same habitat and, consequently, to interact with one another. These two sympatric pelagic species are planktivorous and consume a wide range of planktonic prey items during all their developmental stages, potentially overlapping their ecological niches, although the feeding interactions between these species have been poorly investigated. Here we compare the dietary habits of the juvenile phases of anchovy and sardine during different seasons in the northwestern Mediterranean Sea, through analysis of their stomach contents and of their feeding-related anatomical characteristics. In this study we show that juveniles of anchovy and sardine do not compete for food, and we describe significant dietary differences between anchovy and sardine due to their different alimentary tract morphology.     

Influence of oceanographic fronts and low oxygen on the distribution of ichthyoplankton in the Benguela and southern Angola currents

The study focuses on ichthyoplankton populations in the southern Angola Current, the Angola-Benguela Front and the coastal upwelling area of the northern and central Benguela Current. The horizontal and vertical distributions of eggs and larvae of sardine Sardinops sagax, anchovy Engraulis encrasicolus and horse mackerel Trachurus trachurus capensis were examined in relation to distribution patterns of temperature, salinity and dissolved oxygen. Samples were collected during February–March 2002 using a Bongo net and a multiple opening/closing net in five depth layers (0–30m, 30–60m, 60–100m, 100–140m and 140–220m). Eggs and larvae were restricted mainly to the shelf areas in the Angola-Benguela Font and southern Angola Current waters. Their vertical distributions in the Angola-Benguela Front and northern Benguela upwelling region showed a sharp decline in abundance at about 60m depth. This decline appeared to be correlated to the distribution of dissolved oxygen in the water column It is hypothesised that low oxygen concentrations have a strong impact on the development and survival of the early life-cycle stages of fish, and that recruitment of sardine and other pelagic species relies more on the upward extension of the oxygen minimum layer than was previously thought.     

Abundance and Distribution Patterns of Nekton and Micronekton in the Northern California Current Transition Zone

The epipelagic and mesopelagic nekton communities of the northern California Current have been sampled somewhat continuously over the last four decades with bottom and pelagic trawls, small midwater trawls, and purse seines. We review the zoogeography and community and environmental associations of the dominant pelagic micronekton and nekton species in this region with a view to understanding their role in this dynamic marine ecosystem. As is typical of many upwelling eastern boundary current regions, the pelagic biomass is dominated by a few species that fluctuate dramatically through time. The abundance trends of pelagic nekton caught in this region demonstrated large-scale ecosystem changes about the time of the regime shifts of 1976/77 and 1989 and possibly another beginning in 1999. The rapidity of the changes in composition indicates that the response was due to a change in migration or distribution patterns as opposed to recruitment patterns. The 1989 regime shift led to a dramatic increase in sardine and a decrease in anchovy populations. The most pronounced interannual signals were attributed to strong El Niño/Southern Oscillation (ENSO) conditions in 1983 and 1998 that altered the latitudinal ranges and proximity to the coast of many pelagic species. Variations in abundance and cross-shelf distribution patterns were noted for both pelegic nekton and micronektonic from surveys off California, Oregon, and Washington.     

Taxonomic diversity of fish assemblages in the Changjiang Estuary and its adjacent waters

Taxonomic diversity of fish assemblages in the Changjiang Estuary(Yangtze River Estuary) and its adjacent waters was analyzed based on bottom trawl surveys carried out by R/V Beidou in June,August and October 2006.Four groups of fishes were identified for each survey by the twoway indicator species analysis(TWIA).Taxonomic distinctness,species richness and phylogenetic diversity were used to detect changes in taxonomic structure of fish assemblages.Most dominant fish species mainly belonged to Perciformes,Pleuronectiformes,Clupeiformes,Scorpaeniformes and Tetraodontiformes.Species richness in offshore waters of the survey area was higher than that in the other area.The average taxonomic distinctness value(AvTD,△+ ) of each fish assemblage was close to the average taxonomic distinctness of master list in the Changjiang Estuary(79.9),and had a positive correlation with species richness and Shannon diversity index(H ).A negative correlation was found between variations in taxonomic distinctness(VarTD,Λ + ) and traditional diversity indices,which were caused by fish species component that led to longer average path lengths among species.Taxonomic diversity index(△) had the similar results with species richness,H ,Simpson diversity index(D) and Pielou's evenness index(J ).VarTD also kept stable,which further certificated that fish assemblages and ecological environment were in equilibrium.Taxonomic distinctness index(△* ) was relatively stable in most of fish assemblages,and had higher values in some fish assemblages for a few absolutely dominant species.The present study showed that fish community formed new equilibrium stability in 2006 in the Changjiang Estuary when compared with those in the corresponding months of annual survey from 1985 to 1986.And AvTD in each sampling station was lower than AvTD of master list in the Changjiang Estuary,so some ecological niches were absent in each sampling stations from taxonomic or phylogenetic relationships,these would be helpful to stock natural resource and maintain ecological equilibruim of fish assemblages.     

Spatio‐temporal variations in demersal fish assemblages and diversity in the northern Gulf of St. Lawrence (Canada)

Species diversity is generally considered one of the key factors of ecosystem resilience in response to anthropogenic pressures, including fishing. In this context, the spatial and temporal changes in demersal fish assemblages and species diversity were investigated in the northern Gulf of St. Lawrence (Canada), over a 20‐year period (1990–2010). Data were obtained from the summer research survey conducted by the Department of Fisheries and Oceans, and include commercial and non‐commercial species. The study covers the period of groundfish fishery collapse, the moratorium period, and the post‐moratorium period, and reflects various modifications in management. Multivariate statistical methods revealed two communities. A coastal community corresponds to strata located above 200 m depth and a deeper community located in the deep channels. Interannual differences in the composition of fish assemblages were observed and are mainly due to the changes in the relative biomass of some dominant species. Three diversity indices (Shannon–Wiener, Simpson's Index of Diversity and Motomura's constant) indicate a slight but significant increase of the diversity over time. This trend is due to the increase of the relative biomass of low‐rank species, which may have been favoured by the prohibition of groundfish trawling after 1997 in that region. The geographical distribution of the Shannon–Wiener index also shows temporal dynamics reflecting the biomass distribution of dominant demersal species.     

Regime shifts in upwelling ecosystems: observed changes and possible mechanisms in the northern and southern Benguela

A regime shift is considered to be a sudden shift in structure and functioning of a marine ecosystem, affecting several living components and resulting in an alternate state. According to this definition, regime shifts differ from species replacement or alternation of species at similar trophic levels, whereby the ecosystem is not necessarily significantly altered in terms of its structure and function; only its species composition changes. This paper provides an overview of regime shifts, species replacements and alternations that have been observed in the northern and southern Benguela ecosystems over the past few decades. Bottom-up control, initiating and sustaining regime shifts or species replacements via environmental forcing, is documented for both the southern and the northern Benguela ecosystems. Fishing (a case of top-down control) appears to have played an important role in regime shift processes in the Namibian ecosystem. Very low biomass levels of exploited fish stocks associated with less efficient energy transfer in the northern Benguela are indicative of a regime shift. Very high biomass levels have been reached in the southern Benguela in the 2000s. However the alternation between sardine and anchovy that has been observed in the southern Benguela over the last two decades appears not to have had major effects on the overall functioning of the ecosystem. The consequences of regime shifts for exploitation are highlighted, suggesting that fisheries managers should move towards a more effective ecosystem approach to fisheries.     

The State of the Adriatic Sea Centered on the Small Pelagic Fish Populations

Abstract. The paper provides an overview of the state of the small pelagic fish in the Adriatic Sea. The North Adriatic is reviewed for the 1976–1998 period and the South Adriatic for the 1987–1998 period. First the fluctuations in time and in space of the pelagic biomass, as a whole and per species, are presented. Then the dramatic collapse of anchovy stock and its apparent association with the decrease of surface temperature is discussed. Finally the changes of the anchovy and sardine catches are compared and analysed with respect to their abundances in the sea. The conclusion is that acoustic and satellite methods are able to provide a large amount of information on the variability in the pelagic populations; this is essential for an appropriate management of these resources. There is still a need, however, to refine the methods and to integrate this kind of information with further environmental data (including natural predators).     

Diet of sardine Sardinops sagax in the southern Benguela upwelling ecosystem

The diet of sardine Sardinops sagax in the southern Benguela was investigated by microscopic examination of stomach contents. The relative dietary importance of prey size and prey type was assessed by calculating the carbon content of prey items. Sardine is an omnivorous clupeoid, ingesting both phytoplankton and zooplankton, with the relative importance of these two food types varying both spatially and temporally. Stomach contents were numerically dominated by small prey items, principally dinoflagellates, followed by crustacean eggs, cyclopoid copepods, calanoid copepods and diatoms. Virtually all prey items ingested by sardine were <1.2 mm maximum dimension, the particle size below which sardine only filter-feed. Despite the numerical dominance by phytoplankton, zooplankton contributed the major portion to sardine dietary carbon, small calanoid and cyclopoid copepods, anchovy eggs and crustacean eggs being the primary prey types. These results indicate that, like anchovy Engraulis capensis, sardine in the southern Benguela are primarily zoophagous, and contrast with earlier dietary studies on sardine in the region. However, the two species appear to partition their prey on the basis of size; sardine consume small zooplankton, whereas anchovy consume large zooplankton. This difference has been observed in other upwelling ecosystems where the two genera co-exist and is likely to contribute to the regime shifts observed between sardine and anchovy.