Cd van der Lingen

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.

Abrupt environmental shift associated with changes in the distribution of Cape anchovy Engraulis encrasicolus spawners in the southern Benguela

Cape anchovy Engraulis encrasicolus spawners in the southern Benguela showed an eastward shift in their distribution on the Agulhas Bank that occurred abruptly in 1996 and has since persisted. We assessed whether this shift was environmentally mediated by examining sea surface temperature data from different regions of the Agulhas Bank, which showed that in 1996 the inner shelf of the Agulhas Bank to the east of Cape Agulhas abruptly became 0.5°C colder than in previous years and has since remained that way. In addition, signals, coherent with the 1996 shift recorded in sea surface temperatures, were also found in atmospheric surface pressure and zonal wind data for that region; interannual coastal SST variability is also shown to be correlated with zonal wind-stress forcing. As a result, increased wind-induced coastal upwelling east of Cape Agulhas is proposed as the main driver of the observed cooling in the coastal region. The synchrony between the environmental and biological signals suggests that the eastward shift in anchovy spawner distribution was environmentally mediated and arose from a change in environmental forcing that altered the relative favourability for spawning between regions to the west and east of Cape Agulhas. The results highlight how a relatively minor change in environmental conditions can lead to a drastic spatial reorganisation of the life history of one species in an ecosystem.

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...

Multiple factors affecting South African anchovy recruitment in the spawning, transport and nursery areas

Despite high primary productivity, the yield of pelagic fish in the southern Benguela is relatively low compared to that in the Humboldt system. Part of the constraint may be the ability of pelagic fish to reproduce successfully in a strongly pulsed upwelling environment, where enrichment, retention and concentration mechanisms are less compatible than in Peru-Chile. Anchovy Engraulis capensis spawn upstream of the main upwelling centres on the food-poor, thermally stratified western Agulhas Bank, over a protracted summer season (October–February) when high wind speeds of 7–8 m˙s−1 are prevalent. Eggs spawned farther east, on the central or eastern Agulhas Bank, may be subject to increased cannibalism and advective losses, whereas those spawned farther west could be susceptible to heavy advective losses offshore during periods of strong southerly winds. Copepod concentrations are negatively correlated with spawner biomass on the western Bank and are inversely linked to high rates of gonad atresia in ancho...

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.

Shelf currents, lee-trapped and transient eddies on the inshore boundary of the Agulhas Current, South Africa: their relevance to the KwaZulu-Natal sardine run

The existence and strength of the annual KwaZulu-Natal (KZN) sardine run has long been a conundrum to fishers and scientists alike ― particularly that the sardine Sardinops sagax migrate along the narrow Transkei shelf against the powerful, warm Agulhas Current. However, examination of ship-borne acoustic Doppler current profiler (S–ADCP) data collected during two research surveys in 2005 indicated that northward-flowing coastal countercurrents exist at times between the Agulhas Bank and the KZN Bight, near Port Alfred, East London, Port St Johns and Durban. The countercurrent near Port Alfred extended as far east as the Keiskamma River, within an upwelling zone known to exist there. An ADCP mooring at a depth of 32 m off Port Alfred indicated that the countercurrent typically lasted a few days, but at times remained in the same direction for as long as 10 days. Velocities ranged between 20 and 60 cm s−1 with maximum values of ∼80 cm s−1. The S–ADCP data also highlighted the existence of cyclonic flow in the Port St Johns–Waterfall Bluff coastal inset, with a northward coastal current similarly ranging in velocity between 20 and 60 cm s−1. CTD data indicated that this was associated with shelf-edge upwelling, with surface temperatures 2–4 °C cooler than the adjacent core temperature (24–26 °C) of the Agulhas Current. Vertical profiles of the S–ADCP data showed that the countercurrent, about 7 km wide, extends down the slope to at least 600 m, where it appeared to link with the deep Agulhas Undercurrent at 800 m. S–ADCP and sea surface temperature (SST) satellite data confirmed the existence of the semi-permanent, lee-trapped, cyclonic eddy off Durban, associated with a well-defined northward coastal current between Park Rynie and Balito Bay. Analysis of three months (May–July 2005) of satellite SST and ocean colour data showed the shoreward core-boundary of the Agulhas Current (24 °C isotherm) to commonly be close to the coast along the KZN south coast, as well as between the Kei and Mbhashe rivers on the Transkei shelf. The Port St Johns–Waterfall Bluff cyclonic eddy was also frequently visible in these satellite data. Transient cyclonic eddies, which spanned 150–200 km of shelf, appeared to move downstream in the shoreward boundary of the Agulhas Current at a frequency of about once a month. These seemed to be break-away Durban eddies. Data collected by ADCP moorings deployed off Port Edward in 2005 showed that these break-away eddies and the well-known Natal Pulse are associated with temporary northward countercurrents on the shelf, which can last up to six days. It is proposed that these countercurrents off Port Alfred, East London and Port St Johns assist sardine to swim northwards along the Transkei shelf against the Agulhas Current, but that their progress north of Waterfall Bluff is dependent on the arrival of a transient, southward-moving, break-away Durban cyclonic eddy, which apparently sheds every 4–6 weeks, or on the generation of a Natal Pulse. This passage control mechanism has been coined the ‘Waterfall Bluff gateway’ hypothesis. The sardine run survey in June–July 2005 was undertaken in the absence of a cyclonic eddy on the KZN south coast, i.e. when the ‘gate’ was closed.

Does vertical migratory behaviour retain fish larvae onshore in upwelling ecosystems? A modelling study of anchovy in the southern Benguela

A spatially explicit individual-based model (IBM) forced by 3D temperature and current fields simulated by a hydrodynamic model of the southern Benguela upwelling region was used to test two hypotheses concerning the role of diel vertical migration (DVM) by Cape anchovy Engraulis encrasicolus larvae and pre-recruits. These hypotheses were that: (1) DVM enhances alongshore transport of anchovy eggs and larvae from the spawning grounds to the nursery area while avoiding the lethal effect of low water temperatures in the upwelling system, and/or (2) DVM enhances the transport of larvae and pre-recruits from the offshore to the onshore domain of the nursery area, and then counteracts offshore advection by favouring retention. We tracked the trajectories of virtual particles in the model and calculated a pre-recruitment index as a proxy for transport success to the nursery area (onshore and offshore) and found that the index increased from 10% to 20% after the incorporation of larval vertical migration into the IBM, with virtual individuals held at depths of around 60 m showing maximal pre-recruitment index values. Hence, DVM does appear to enhance transport to the nursery area (offshore) for early and late larvae. Model outputs showed coarse-scale horizontal distribution patterns of larvae by age/size class that are similar to field observations for early, small larvae but not for large larvae and pre-recruits. Observations show that early/small larvae are located offshore whereas older/larger larvae and pre-recruits are found closer to the continental shelf and the inner nursery grounds. This disparity between model results and field observations does not support the hypothesis that DVM is one of the mechanisms involved in the onshore movement of early life-history stages, especially for large larvae.

A comparison of condition factor and gonadosomatic index of sardine Sardinops sagax stocks in the northern and southern Benguela upwelling ecosystems, 1984–1999

Time-series of condition factor (CF) and gonadosomatic index (GSI) were generated using general linear models (GLM) for sardine Sardinops sagax stocks in the northern and southern Benguela ecosystems over the period 1984–1999. During this period the biomass of sardine in the northern Benguela remained at relatively low levels of <500 000 tons, whereas that of southern Benguela sardine increased 40-fold to 1.3 million tons. The GLMs explained 27 and 45% of the observed variation in CF, and 32 and 28% of the observed variation in GSI, for sardine in the northern and southern Benguela subsystems respectively. Whereas the sardine CF in the northern Benguela remained stable over time, that for the southern Benguela stock declined steadily during the study period. Sardine CF showed a seasonal cycle in the southern but not in the northern Benguela. Time-series of GSI showed high interannual variability but no trends in either subsystem, and the seasonal pattern was similar for both stocks. The lack of coherence between the CF time-series for sardine in the two subsystems further suggests that sardine stocks in the northern and southern Benguela subsystems are independent.

Overview of the KwaZulu-Natal sardine run

This paper provides an introduction to, and overview of, the natural phenomenon known as the KwaZulu-Natal sardine run. Previous literature on this topic and hypotheses about the reasons why, and the mechanisms how, the run occurs are briefly synthesised and described. Papers contributing to this suite that detail more recent work on a variety of aspects of the sardine run, ranging from physical oceanography through sardine biology and ecology to socio-economic and ecological consequences, are outlined. Such studies will lead to improved understanding of the factors that regulate the timing and intensity of the run, which may permit predictions of whether it will occur, when fish will arrive on the KZN beaches, and how long it will persist. Such predictions would have substantial benefits for this ecologically and economically important event.

A knowledge base for management of the capital-intensive fishery for small pelagic fish off South Africa

As a contribution to South Africas move towards an ecosystem approach to fisheries management, this study explores the existence of common perceptions about South Africas pelagic fishery between resource users and scientists. It represents a collaborative research effort of social and natural scientists. A brief overview is given of the southern Benguela upwelling ecosystem and small pelagic fish resources, the fishery, and management of the fishery. Stakeholder knowledge and views were determined by conducting open-ended qualitative local knowledge interviews. Candidate indicators to address five major issues raised in the interviews were selected: length-at-50% maturity, total mortality, exploitation rate, proportion of bycatch, mean length of catch, and centre of gravity of catches. The indicator approach is shown to be a useful tool to manage the South African small pelagic fishery, and can be made compatible with existing management approaches. The foundation of a good adaptive fisheries management system is a data collection system that enables multi-disciplinary analysis and provides a basis on which decisions can be made.