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.     

Revisiting Peruvian anchovy (Engraulis ringens) trophodynamics provides a new vision of the Humboldt Current system

The Peruvian anchovy or anchoveta (Engraulis ringens) forages on plankton and is a main prey for marine mammals, seabirds, fish, and fishers, and is therefore a key element of the food web in the Humboldt Current system (HCS). Here, we present results from the analysis of 21,203 anchoveta stomach contents sampled during 23 acoustic surveys over the period 1996–2003. Prey items were identified to the genus level, and the relative dietary importance of different prey was assessed by determination of their carbon content. Variability in stomach fullness was examined relative to the diel cycle, the distance from the coast, sea surface temperature, and latitude, using generalized additive models (GAMs). Whereas phytoplankton largely dominated anchoveta diets in terms of numerical abundance and comprised >99% of ingested prey items, the carbon content of prey items indicated that zooplankton was by far the most important dietary component, with euphausiids contributing 67.5% of dietary carbon followed by copepods (26.3%). Stomach fullness data showed that anchoveta feed mainly during daytime between 07h00 and 18h00, although night-time feeding also made a substantial contribution to total food consumption. Stomach fullness also varied with latitude, distance from the coast, and temperature, but with substantial variability indicating a high degree of plasticity in anchoveta feeding behaviour. The results suggest an ecological role for anchoveta that challenges current understanding of its position in the foodweb, the functioning of the HCS, and trophic models of the HCS.     

Ecological niches and areas of overlap of the squat lobster ‘munida’ (Pleuroncodes monodon) and anchoveta (Engraulis ringens) off Peru

The world’s largest mono-specific fishery, the Peruvian anchovy or anchoveta (Engraulis ringens) fishery, has been the subject of many studies since the 1960s. Details of its relationship with other species have mainly focused on alternations with sardine, Sardinops sagax, and little effort has so far been paid to interactions with other species sharing the same ecosystem. This is the case for Pleuroncodes monodon, the crustacean squat lobster or ’munida’, which has become highly abundant along the Peruvian coast since the mid-1990s. Munida is now an important prey for seabirds, mammals and coastal predatory fish. Knowledge of patterns of distribution and ecological niche of munida is scarce however off Peru. Here we describe and compare spatial patterns of distribution of anchoveta and munida and their ecological niches based on data from 26 acoustic surveys performed along the Peruvian coast between 1998 and 2006. The results indicate that munida and anchoveta share ecological niches but that munida is restricted to the coldest part of the productive cold coastal waters whereas anchoveta do not present any temperature preference over a large range (14–23 °C). The recent increase in munida abundance off Peru is concomitant with colder conditions; with their onset munida extended its range from central Chile northwards. Off Peru the very shallow oxycline keeps munida from its usual bottom habitat and has forced it to adopt pelagic behaviour.     

Climatic regimes and the recruitment rate of anchoveta, Engraulis ringens, off Peru

The recruitment rate of Peruvian anchoveta, Engraulis ringens, was studied to test the hypothesis that long-term environmental variation (regime shifts) had a significant impact on density-dependent processes governing the anchovy recruitment during the period 1963–2004. On the basis of previous defined regimes and turning points for the Humboldt Current System, we identified two groups of years for increased recruitment of anchoveta (1963–1971 and 1986–2004), and one unfavorable period (1972–1985). A common intercept and significantly different slopes were found when the recruitment rate was plotted as a function of the spawning stock biomass during those groups of years, suggesting that density-dependent effects on recruitment were affected during different climate regimes. The favorable (unfavorable) regime was characterized by higher (lower) zooplankton volumes, and with a higher frequency of colder (warmer) waters. Dome-shaped relationships between recruitment rate, spawning stock biomass and SST, were detected with a Generalized Additive Model for the favorable regime. Thus, recruitment could be explained by non-linear effects of environmental variables. Ultimately, climatic regimes are affecting the density-dependent effects on recruitment of anchoveta and the mechanisms involved may be associated with changes in the carrying capacity of the spawning habitat of anchoveta off Peru, which in turn are related with the effects of cold and warm regimes.     

Regime shifts in the Humboldt Current ecosystem

Of the four major eastern boundary currents, the Humboldt Current (HC) stands out because it is extremely productive, dominated by anchovy dynamics and subject to frequent direct environmental perturbations of the El Niño Southern Oscillation (ENSO). The long-term dynamics of the HC ecosystem are controlled by shifts between alternating anchovy and sardine regimes that restructure the entire ecosystem from phytoplankton to the top predators. These regime shifts are caused by lasting periods of warm or cold temperature anomalies related to the approach or retreat of warm subtropical oceanic waters to the coast of Peru and Chile. Phases with mainly negative temperature anomalies parallel anchovy regimes (1950–1970; 1985 to the present) and the rather warm period from 1970 to 1985 was characterized by sardine dominance. The transition periods (turning points) from one regime to the other were 1968–1970 and 1984–1986. Like an El Nino, the warm periods drastically change trophic relationships in the entire HC ecosystem, exposing the Peruvian anchovy to a multitude of adverse conditions. Positive temperature anomalies off Peru drive the anchovy population close to the coast as the coastal upwelling cells usually offer the coolest environment, thereby substantially decreasing the extent of the areas of anchovy distribution and spawning. This enhances the effects of negative density-dependent processes such as egg and larval cannibalism and dramatically increases its catchability. Increased spatial overlap between anchovies and the warmer water preferring sardines intensifies anchovy egg mortality further as sardines feed heavily on anchovy eggs.Food sources for juvenile and adult anchovies which prey on a mixed diet of phyto- and zooplankton are drastically reduced because of decreased plankton production due to restricted upwelling in warm years, as demonstrated by lower zooplankton and phytoplankton volumes and the diminution of the fraction of large copepods, their main food source.Horse mackerel and mackerel, the main predators of anchovy, increase predation pressure on juvenile and adult anchovies due to extended invasion into the anchovy habitat in warmer years. In contrast to these periods of warm and cold temperature anomalies on the decadal scale, ENSO events do not play an important role for long-term anchovy dynamics, as the anchovy can recover even from strong ENSO events within 1–2 years. Consequently, the strong 1972–1973 ENSO event (in combination with overfishing) was not the cause of the famous crash of the Peruvian anchovy fishery in the 1970s.     

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.     

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.     

Schooling behaviour and environmental forcing in relation to anchoveta distribution: An analysis across multiple spatial scales

The Peruvian anchovy or anchoveta (Engraulis ringens) supports the highest worldwide fishery landings and varies in space and time over many scales. Here we present the first comprehensive sub-mesocale study of anchoveta distribution in relation to the environment. During November 2004, we conducted a behavioural ecology survey off central Peru and used a series of observational and sampling tools including SST and CO₂ sensors, Niskin bottles, CTD probes, zooplankton sampling, stomach content analysis, echo-sounder, multibeam sonar, and bird observations. The sub-mesoscale survey areas were chosen from mesoscale acoustic surveys. A routine coast-wide (2000 km) acoustic survey performed just after the sub-mesoscale surveys, provided information at an even larger population scale. The availability of nearly concurrent sub-mesoscale, mesoscale and coast-wide information on anchoveta distribution allowed for a unique multi-scale synthesis. At the sub-mesoscale (100s m to km) physical processes (internal waves and frontogenesis) concentrated plankton into patches and determined anchoveta spatial distribution. At the mesoscale (10s km) location relative to the zone of active upwelling (and age of the upwelled water) and the depth of the oxycline had strong impacts on the anchoveta. Finally, over 100s km the size of the productive area, as defined by the upwelled cold coastal waters, was the determining factor. We propose a conceptual view of the relative importance of social behaviour and environmental (biotic and abiotic) processes on the spatial distribution of anchoveta. Our ecological space has two y-axis; one based on self-organization (social behaviour), and the other based on the environmental processes. At scales from the individual (10s cm), to the nucleus (m), social behaviour (e.g. the need to school) drives spatial organization. At scales larger than the school, environmental forces are the main driver of fish distribution. The conceptual ecosystem models presented in this paper may provide the final links needed to develop accurate forecasts of the spatial distribution of anchoveta over multiple scales.     

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.     

Comparative trophodynamics of anchovy Engraulis encrasicolus and sardine Sardinops sagax in the southern Benguela: are species alternations between small pelagic fish trophodynamically mediated?

The results of detailed morphological, experimental, field and modelling studies on various aspects of the trophic ecology of sardine Sardinops sagax and anchovy Engraulis encrasicolus in the Benguela ecosystem are synthesised, and differences in the trophodynamics of these two species are highlighted. Anchovy possess a relatively coarse branchial apparatus; feed predominantly by particulate-feeding and maximise their net energetic gain by using this feeding mode; show higher weight-standardised clearance rates than do sardine for prey >580μm; are most efficient at assimilating nitrogen from zooplankton and excrete <50% of ingested nitrogen; feed inefficiently on phytoplankton and derive the bulk of their dietary input from larger zooplankton; and maximise their scope for growth on mesozooplankton. In contrast, sardine possess a relatively fine branchial apparatus; feed predominantly by filter-feeding and maximise their net energetic gain by using this feeding mode; show higher weightstandardised clearance rates than do anchovy for prey <580μm; are most efficient at assimilating nitrogen from zooplankton but excrete >50% of ingested nitrogen; are able to feed on phytoplankton but derive the bulk of their dietary input from smaller zooplankton; and maximise their scope for growth on microzooplankton. These differences provide compelling evidence that anchovy and sardine are trophically distinct, and indicate that the two species show resource partitioning based on zooplankton size. The implications of these trophic differences for ecosystem functioning are discussed, and support the hypothesis that species alternations between anchovy and sardine, both in the southern Benguela and in other upwelling ecosystems, are likely to be trophodynamically mediated.     

Climate variability and pelagic fisheries in northern Chile

A time series analysis of long-term climate variability in northern Chile (18°21′–24°00′S) shows anomalies associated with the El Niño events and the longer warm period observed since 1976, followed by a cooling trend since mid 1980s. The succession of pelagic fisheries, anchovy (Engraulis ringens) and sardine (Sardinops sagax), occurring in this fishing zone was analyzed taking into account the landings, the CPUE abundance index, the fishing effort, and the environmental variables. The anchovy production model is a negative linear function of fishing effort and turbulence. For sardine, the production model is a negative linear function of fishing effort and a quadratic function of the sea surface temperature.An analysis of the relationship between recruitment, adult biomass and the environment shows that the annual recruitment of anchovy increases with turbulence intensity until wind speed reaches a value of 5.46 m s⁻¹, decreasing for higher values. For sardine, the recruitment increases with turbulence intensity until 5.63 m s⁻¹, stabilizing thereafter.It is deduced that the climatic variations associated to the El Niño events affect the abundance of coastal pelagic fishes, without forgetting the most likely effects upon its distribution and the fishing effort. However, it is the long-term variability that mainly affects the fishing activity.     

Interdecadal variability of anchoveta abundance and overcapacity of the fishery in Peru

Paleontological and historical stock abundance estimates indicate that pelagic fish populations inhabiting upwelling ecosystems undergo large interdecadal variations in abundance with amplitudes equal to, if not larger than, the interannual variability. The interdecadal variability is characterized by periods of high and low abundance, termed “pseudo-cycles”, because of their irregular periodicity. Fisheries targeting small pelagic fish suffer from overall overcapitalization, like many other fisheries, but also from an additional overcapitalization problem: a phase displacement between rapid fish population decreases and a slower disinvestment which follows. This lag produces economic hardship.Here we document the fish:fishery relationship for the Peruvian anchoveta. Anchoveta pseudo-cycles of 20 to >100 years are observed, with the present stock abundance most likely located near upper part of the cycle. Fleet overcapacity expressed as the proportion of unused present capacity is estimated at 72% and processing overcapacity at 89%. A simple bio-economic model demonstrates the risks associated with the pseudo-periodicity in fish stock abundance in conjunction with fishery investment, open access, and overcapacity: a timing bomb for the fishing sector. The lag between disinvestment and decrease in fish abundance is quantified. A reduction of the fishing and processing capacity and measures to decrease the investment lag are recommended to limit the social, economical and political tensions that will result from the expected decrease in stock abundance. Finally, some management options to reduce these risks are discussed.     

The 2005 KwaZulu-Natal sardine run survey sheds new light on the ecology of small pelagic fish off the east coast of South Africa

Despite much public awareness surrounding the annual migration of sardine Sardinops sagax northward along the east coast of South Africa in winter each year, relatively little research effort has been expended to improve understanding of the ‘sardine run’. For this reason, a dedicated multidisciplinary survey, timed to coincide with the annual sardine run, was conducted off the East Coast in June and July of 2005. The major objective of the survey was to estimate the biomass of sardine off the East Coast during the run, and to compare this with biomass estimates collected during previous surveys conducted in this area during the late 1980s when the South African sardine population was at a considerably smaller size. We also collected data on the distribution of sardine and other small pelagic fish species and their eggs, the biological characteristics of sardine during the run, and data on the hydrography (temperature and currents) and lower trophic levels (phytoplankton and zooplankton) of the region. Results suggest that the biomass of sardine off the East Coast in winter remains relatively small and consistent, regardless of overall sardine population size. The narrow continental shelf to the east of Port Alfred, which is dominated offshore by the fast-flowing warm Agulhas Current, constrains the amount of suitable habitat for sardine and other clupeoids such as anchovy Engraulis encrasicolus, West Coast round herring Etrumeus whiteheadi and East Coast round herring Etrumeus teres, and hence precludes these species from attaining a high biomass in this region. Additionally, primary and secondary productivity levels are much lower than elsewhere on the western and eastern Agulhas Bank off the south coast of South Africa, suggesting that the sardine run is not a feeding migration. A previous hypothesis that the run is mainly a result of an expansion of the distributional range of these fish as conditions become favourable in winter due to sporadic cooling off the East Coast is also not entirely supported by results from the survey. It is suggested that a migration for the purposes of spawning off this coast when conditions become favourable is a more likely incentive for sardine to undertake this arduous journey, despite increased predation and poor feeding conditions.     

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.     

Comparison of assemblages and some life-history traits of seabirds in the Humboldt and Benguela systems

There are 21 and 15 species of seabirds that breed in the Humboldt and Benguela upwelling systems respectively. Only two species of gull are common to both systems, one as an endemic subspecies to the Benguela system. Eleven species and two subspecies are endemic (or nearly so) to the Humboldt system; seven species and one subspecies to the Benguela system. Each system has an endemic penguin, sulid, cormorant and tern that feed mainly on anchovy Engraulis spp., sardine Sardinops sagax or both these fish. The Peruvian pelican Pelecanus thagus also feeds primarily on these prey items. A plentiful availability of food has resulted in many of these seabirds attaining high levels of abundance. For the four pairs of species that feed on anchovy and sardine, those in the Humboldt system all have a biology that enables them to increase more rapidly than their Benguela counter-parts. This reflects the higher frequency of environmental perturbations that depress seabird populations in the Humboldt system. In addition, both systems have a small endemic cormorant that feeds near the coast and a small endemic tern that breeds in the adjacent mainland desert and feeds at the sea surface. Several seabirds endemic to a system have no obvious ecological equivalent in the other system: the pelican, a diving-petrel, four storm-petrels and a gull in the Humboldt system; a cormorant and a gull in the Benguela system. Some species with tropical or subantarctic affinities breed at the boundaries of the systems. Others breed also in freshwater systems. The grey gull Larus modestus, which feeds in the Humboldt system, breeds in montane deserts.     

Zooplankton research off Peru: A review

A review of zooplankton studies conducted in Peruvian marine waters is given. After a short history of the development of zooplankton research off Peru, we review zooplankton methodology, taxonomy, biodiversity, spatial distribution, seasonal and interannual variability, trophodynamics, secondary production, and modelling. We review studies on several micro-, meso-, macro-, and meroplankton groups, and give a species list from both published and unpublished reports. Three regional zooplankton groups have been identified: (1) a continental shelf group dominated by Acartia tonsa and Centropages brachiatus; (2) a continental slope group characterized by siphonophores, bivalves, foraminifera and radiolaria; (3) and a species-rich oceanic group. The highest zooplankton abundances and biomasses were often found between 4–6°S and 14–16°S, where continental shelves are narrow. Species composition changes with distance from the shore. Species composition and biomass also vary strongly on short time scales due to advection, peaks of larval production, trophic interactions, and community succession. The relation of zooplankton to climatic variability (ENSO and multi-decadal) and fish stocks is discussed in the context of ecological regime shifts. An intermediate upwelling hypothesis is proposed, based on the negative effects of low upwelling intensity in summer or extremely strong and enduring winter upwelling on zooplankton abundance off Peru. According to this hypothesis, intermediate upwelling creates an optimal environmental window for zooplankton communities. Finally, we highlight important knowledge gaps that warrant attention in future.     

Diet of the Cape fur seal Arctocephalus pusillus pusillus at the Robberg Peninsula,Plettenberg Bay,and implications for local fisheries

Cape fur seals Arctocephalus pusillus pusillus were harvested to extirpation on the Robberg Peninsula, Plettenberg Bay, on the south-east coast of South Africa, between the 17th and early 20th centuries. Seals returned to Robberg in small numbers during the early 1990s and their numbers subsequently increased. We studied the diet of this increasing population using faecal (scat) sampling to determine: the species composition and size of prey in the diet of Cape fur seals at Robberg; to explore temporal variation in the diet; and to investigate the potential for competition between seals and the fisheries around Plettenberg Bay. Of the 445 scats collected, 90% contained hard prey remains and 15 teleost prey species were represented in the 3 127 otoliths that could be identified. The seals’ most important prey species in terms of numerical abundance, frequency of occurrence and mass in the diet, were anchovy Engraulis encrasicolus, sardine Sardinops sagax, horse mackerel Trachurus capensis, sand tongue-fish Cynoglossus capensis and shallow-water hake Merluccius capensis (in decreasing order of importance for numerical abundance). The proportion of anchovy in the diet increased during the study period (2003–2008), whereas the proportion of sardine decreased. The estimated average annual consumption of sardine by seals was higher than the average annual catch made by purse-seine fisheries in this area, suggesting resource competition between seals and purse-seiners, especially in the light of continuing growth of seal numbers in the area. However, direct competition between seals and linefisheries appeared to be minimal. Scat sampling of Cape fur seals holds potential to serve a useful and cost effective indicator of temporal changes in sardine abundance.     

Revised estimates of abundance of South African sardine and anchovy from acoustic surveys adjusting for echosounder saturation in earlier surveys and attenuation effects for sardine

Hydro-acoustic surveys have been used to provide annual estimates of May recruitment and November spawner biomass of the South African sardine Sardinops sagax and anchovy Engraulis encrasicolus resources since 1984. These time-series of abundance estimates form the backbone of the assessment of these resources, and consequently the management of the South African sardine and anchovy is critically dependent on them. Upgrades to survey equipment over time have resulted in recent surveys providing more accurate estimates of abundance, yet in order to maintain comparability across the full time-series, estimates of biomass mimicking the old equipment were used for a number of years. In this paper we develop a method to revise the earlier part of the time-series to correct for receiver saturation in the older generation SIMRAD EK400 and EKS-38 echo sounders and to account for attenuation in dense sardine schools. This is applied to provide a revised time-series of biomass estimates for the South African sardine and anchovy resources with associated variance–covariance matrices. Furthermore, the time-series presented here are based on updated acoustic target strength estimates, making this the most reliable time-series currently available for both resources.