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.     

The role of acoustic surveys in the assessment of pelagic fish resources on the South African continental shelf

Acoustic data on the abundance and distribution of anchovy Engraulis capensis, pilchard Sardinops ocellatus and round herring Etrumeus whiteheadi on the South African continental shelf have been collected from 21 echo-integrator surveys between 1984 and 1991. Most effort has been concentrated on estimating adult biomass of anchovy and pilchard in November (spring) and anchovy recruitment in autumn. Distribution maps from all surveys are presented and the biomass estimates considered most reliable documented. A series of distribution maps tracing movements of three anchovy year-classes over a four-year period is presented to illustrate the usefulness of the surveys in migration studies. The major findings of the survey programme have been that anchovy are generally considerably more abundant and widespread than was thought to be the case prior to the surveys, that the pilchard resource has recovered substantially in recent years, and that the round herring resource, about which little was known prior to the surveys, is probably of the same order of magnitude as the anchovy resource and is probably underexploited. The anchovy and pilchard resources are currently managed through procedures based largely on the acoustic estimates of biomass and their estimated precision. The role of these estimates in the management procedures is discussed in some detail.     

The 2004 re-assessment of the South African sardine and anchovy populations to take account of revisions to earlier data and recent record abundances

Hydroacoustic surveys off the coast of South Africa over the early years of the 21st century indicated that both the sardine Sardinops sagax and anchovy Engraulis encrasicolus populations had simultaneously reached record abundances. The South African pelagic fishery is regulated using an Operational Management Procedure (OMP). The OMP in use at that time had been developed using data from the two populations prior to this rapid and substantial increase in abundances. This paper documents the revised assessments that were urgently required to provide a basis to update the OMP. These assessments resulted in a changed perception of the status and productivity of these populations. In particular, estimates of the stock-recruitment relationships and the extent of variation about them, which play a key role in evaluating risk when developing OMPs, altered substantially from estimates derived from earlier assessments.     

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.     

Top predators in the Benguela ecosystem — implications of their trophic position

Many top predators in the Benguela ecosystem feed on prey species targeted by commercial fisheries. Their roles as indicators of the state of exploited prey resources, as competitors with commercial fisheries for resources, and as susceptible to impact from commercial fishing on those resources are briefly considered. Trends in the occurrence of anchovy Engraulis capensis and pilchard Sardinops ocellatus in the diet of Cape gannets Morus capensis off South Africa's west coast are related significantly to survey estimates of the abundance of these fish species, and they provide useful confirmation of those estimates. In the 1980s, anchovy decreased in the diet of Cape gannets, but pilchard increased. In both the northern and southern portions of the Benguela system, groundfish were thought to eat most (66–73%) of the total quantity of cephalopods and vertebrates consumed by predators and man in the 1980s. South African fur seals Arctocephalus pusillus pusillus, predatory pelagic fish and man removed roughly equal amounts, with squids, seabirds and cetaceans having a smaller impact. In the 1980s, man and seals removed about two million tons live mass more than in 1930. Indices of the rate of natural mortality of anchovy and pilchard attributable to Cape gannets are not related to biomass of the prey species. That for anchovy was high in 1989 when a poor anchovy year-class was formed. Decreased abundance of anchovy led to poor breeding by Cape cormorants Phalacrocorax capensis in 1989 and 1990. A model linking the Cape cormorant population with anchovy is used to explore the impact of possible exploitation strategies for anchovy on Cape cormorants.     

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.     

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.     

Testing for the occurrence of pilchard herpesvirus (PHV) in South African sardine Sardinops sagax

Catches of South African sardine Sardinops sagax have declined in recent years from about 200 000 t harvested annually during the period 2002–2006 to less than 100 000 t. Consequently, some companies are now importing sardine from sources elsewhere in the world to meet local demand for canned sardine and bait. This importation has the potential for the introduction of sardine pathogens, in particular the pilchard herpesvirus (PHV), which could have a negative impact on the currently small South African sardine population. The aims of the current study were to determine whether PHV is present in the local sardine population and to assess the extent to which sardine is being imported into the country and whether imported fish are from countries where the virus is known to be endemic. Fish sampled from South Africa’s western (n = 150), southern (n = 182) and eastern (n = 96) putative stocks of S. sagax were analysed for the presence of PHV using real-time quantitative polymerase chain reaction (qPCR). The origin and amount of potentially infected material imported into South Africa during the period 2010–2014 was also assessed. None of the South African sardine collected during this study tested positive for PHV, suggesting that active PHV was not prevalent in the local population of S. sagax at the time of this study. Between 56 000 and 71 000 t of frozen sardine was imported annually into South Africa from countries where S. sagax occurs, including some from areas (Australia and New Zealand) where sardine infection by PHV is known to be endemic. Hence, it is plausible that the PHV pathogen, capable of perpetuating infections in local sardine populations, could be imported into South Africa along with the importation of frozen sardine. Should local sardine be naïve to the virus, as suggested by this study, then the population is at risk of infection and precautions against such must be taken.     

Variable spatial structure of schooling pelagic fish off Namibia: implications for acoustic surveys

Schools of horse mackerel Trachurus trachurus capensis, sardine Sardinops sagax and round herring Etrumeus whiteheadi were surveyed by conventional echo-integration along systematic transects. The transects were surveyed twice, in opposite directions. Clupeoid biomass was concentrated in a few dense aggregations, whereas horse mackerel were less dense and distributed over larger areas. The influence of aggregation patterns of the different species on the precision of the acoustic estimates was analysed with respect to spatial variability and diurnal effects. Isotropic variograms computed from values of acoustic back-scattering strength showed little or no structure for all three species. Indicator variograms showed relatively large structures (>15 miles) for all three species at the lowest cut-off level, confirming the influence of high values on the variograms. The autocorrelation range decreased with an increase in cut-off value for horse mackerel, whereas variograms of sardine and round herring showed no structure at larger cut-off values. There were large variances in estimated biomass for sardine (77%) and round herring (90%) between surveys, whereas biomass estimates of horse mackerel were relatively similar between surveys.     

Can anchovy age structure be estimated from length distribution data collected during surveys?

Historically, a time-series of proportions-at-age 1 from annual November hydro-acoustic surveys has been used to inform the assessment of, and in particular the choice of appropriate values for juvenile and adult natural mortality for, the South African anchovy Engraulis encrasicolus. However, information from direct ageing is limited and almost two decades old. A new method is developed to estimate the annual proportions of anchovy at age 1 directly from the population length distributions estimated from samples taken during the November surveys. This method involves modelling the annual length distributions of age 1 and age 2+ anchovy. The analysis provides a new time-series of proportions-at-age 1, together with associated standard errors, for input into assessments of the resource. The results also caution against the danger of scientists reading more information into data than is really there.     

Spatial variability in branchial basket meristics and morphology of southern African sardine Sardinops sagax

We examined spatial variability in meristic and morphological characteristics of the branchial basket of sardine Sardinops sagax collected from four geographical regions around the southern African coast, namely Namibia and the South African west, south and east coasts. Our analysis tested the hypothesis of three putative sardine stocks off South Africa, one in each of the three geographical regions. We therefore collected fish data from Namibia to compare with South Africa, because sardine from the two countries are considered to be separate stocks. Morphometric measurements (gill arch length and gill raker spacing) and meristic data (number of gill rakers) were collected from the left side of the first gill arch from a total of 377 sardine, approximately equally divided between the regions. A multivariate general linear model with caudal length as covariate was used to assess differences among fish from the four regions and significant differences were observed, although not always consistently across all fish size classes. Small South Coast sardine had shorter gill arches than small West Coast sardine, but adults had gill arches of similar length, longer than those from Namibia and the East Coast. Small sardine from the South Coast had fewer gill rakers than small sardine from the West Coast, but larger fish had similar numbers of gill rakers, significantly more than sardine from Namibia and the East Coast. Sardine from the West and South coasts had similar gill raker spacings, which were smaller than those of fish from the East Coast and Namibia. Despite spatially and particularly temporally unbalanced sampling, we consider that these differences provide evidence of spatial variation in Benguela sardine phenotype and that it would support the hypothesis of discrete sardine stocks off Namibia and South Africa. The results are consistent with the hypothesis of three sardine stocks within the southern Benguela.     

Spatial variation in meristic and morphometric characteristics of sardine Sardinops sagax around the coast of southern Africa

Spatial variability in phenotypic characteristics within a fish population may be used to infer the existence of multiple stocks, and knowledge of the population structure of exploited species is important for their sustainable management. In this study we investigated geographic variability in meristic (vertebral count) and morphometric (body shape) characteristics of sardine Sardinops sagax from three southern African regions: Namibia, the South African west coast, and the South African south coast. There were significant regional differences in vertebral counts, with Namibian fish tending to have more vertebrae than South African fish, but with no difference between sardine off the South African west and south coasts. Body shape was characterised using morphometric landmarks, and shape changes were investigated using geometric morphometrics, with influences on fish shape assessed using multivariate regressions. To remove the effects of allometry, the 22 shape variables (represented by Procrustes coordinates) were regressed against fish size (represented by centroid size). There was a significant allometric effect, which was removed by using the residuals of these regressions in further analyses. A multivariate multiple regression was applied to the 22 size-corrected shape variables and to three covariates: condition factor, stomach fullness and fat stage. The significant influence of these covariates was removed by using the residuals from this regression in a canonical variate analysis, where corrected shape variables were classified in relation to region, year and sex. The Mahalanobis distances from the canonical variate analysis differed significantly for most combinations of classifier variables, with clear separations among the three regions. Sardinops sagax off Namibia had thicker bodies and more contracted tail regions than South African fish, whereas S. sagax off the South African south coast had thinner bodies and smaller heads than those off the west coast. These results support the hypothesis of multiple sardine stocks around southern Africa.     

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.     

Ocean robotics in support of fisheries research and management

South Africa’s small-pelagic fishery is a socio-economically important component of the country’s commercial fisheries sector, second in value only to the demersal trawl fishery. Management of this sector relies on infrequent hydro-acoustic surveys, which provide measures of anchovy Engraulis encrasicolus and sardine Sardinops sagax biomass used in the assessments of stock status and in the development of management plans for the sustainable utilisation of these resources. We demonstrate how technological capabilities in ocean robotics at the Council for Scientific and Industrial Research (CSIR) could augment the current resource-intensive hydro-acoustic ship-based survey programme and create opportunities for expanding its spatial and temporal resolution. We successfully implement and demonstrate an autonomous wave glider, fitted with a hydro-acoustic sensor and compare the data to a collocated ‘traditional’ ship-based acoustics survey. In the future these autonomous systems approaches could be seen as a means to lessen the cost burden of the ship-based survey, while at the same time with the added advantage of continuous collection over much wider spatial and temporal domains. This could enable a more reflexive stock management approach taking into account the seasonal characteristics of the fishery and its ecosystem. Gliders thus have potential to increase dramatically the quantity of information available to fisheries managers, thereby reducing uncertainty and contributing to improved management of valuable fish resources. They are likely to contribute to improved knowledge of the ecology of small pelagic fish species off the coast of South Africa in a changing climate and should potentially also permit the collection of biomass data for other marine resources currently not routinely monitored.     

Abundance and distribution of avian and marine mammal predators of sardine observed during the 2005 KwaZulu-Natal sardine run survey

Opportunistic observations to determine the relative abundance and distribution of marine mammal and seabird predators of sardine Sardinops sagax were carried out during a dedicated multidisciplinary research survey off the South African east coast in June and July of 2005 that was timed to coincide with the annual sardine run. Associations between different predator species, between predators and clupeoids, and between predators and oceanographic variables, were examined. Species’ distributions were primarily separated by latitude and distance from shore. Bryde's whale Balaenoptera edeni, African penguin Spheniscus demersus, Cape cormorant Phalacrocorax capensis and West Coast round herring Etrumeus whiteheadi were predominantly found in the cool southern part of the survey region. Peak sardine run activity occurred within 4 km of shore at the northward limit of a strip of cool water (<21 °C) stretching along the East Coast. The principal predators associated with this activity were common dolphins Delphinus capensis and Cape gannets Morus capensis, and their nearshore distribution was associated with sardine and East Coast round herring E. teres. Few clupeoids were encountered along the KwaZulu-Natal continental shelf, although patches of high sardine abundance were recorded near the shore immediately south of Durban. It is clear that during the 2005 survey the sardine run terminated in this region, probably as a result of the nearshore intrusion of warm water from the Agulhas Current.     

Acoustic study on the abundance and distribution of anchovy spawners and recruits in South African waters

Direct surveys of Cape anchovy Engraulis capensis spawner and recruit biomass in South African coastal waters were made with a 38-kHz calibrated echo-sounder and custom-built digital echo-integrator. The spawning surveys were conducted each November between 1983 and 1985, predominantly on the Agulhas Bank. Recruit surveys were conducted in autumn 1985 and 1986, mainly on the West Coast. The spawning fish were widely distributed across the continental shelf, whereas recruit concentrations were highest close inshore. Employing a target strength per kilogramme expression for herring Clupea harengus, the spawning biomass in 1984 was estimated at 1,06 million tons and that in 1985 at 0,98 million tons. Similarly, recruit biomass in May/June 1985 and June 1986 was estimated at 310 and 466 thousand tons respectively. From an analysis of error in the latter two estimates, it was considered approximately 80 per cent certain that the recruitment in 1986 was greater than in 1985. The estimates of spawning biomass are 3–4 times higher than earlier estimates made by Virtual Population Analysis (VPA) of commercial catch data, and suggest that the stock is less seriously threatened by present catch levels than was indicated by VPA. It is concluded from the distribution and age structure of the spawning population that the VPA estimates are invalid because of the failure of commercial vessels to sample the major part of the adult population.     

Broad-scale distribution patterns of sardine and their predators in relation to remotely sensed environmental conditions during the KwaZulu-Natal sardine run

The annual movement of South African sardine Sardinops sagax up the east coast of South Africa, known as the ‘sardine run’, was investigated using data from aerial surveys for the period 1988–2005 and compared with remotely sensed sea surface temperature (SST) and chlorophyll a data. Sardine sighting rates were highest within the Waterfall Bluff Bight off the Eastern Cape Coast, where conditions appeared to be most favourable. Sardine and predator sightings decreased significantly northwards of Mdoni on the KwaZulu-Natal (KZN) coast, whereas the proportion of nearshore sightings increased. The causal mechanism for this inshore concentration is suggested to be the influx of warm Agulhas Current water from the Durban Eddy that forces sardine shoreward. Cape gannet Morus capensis, common dolphin Delphinus capensis and sardine distributions were associated, and there was an association between SST and sardine and predator distributions. There was a marked increase in bottlenose dolphin Tursiops aduncus sightings upon commencement of the sardine run, with these dolphins being consid-ered to be a ‘migratory’ stock that enters KZN waters every winter.     

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.     

Ecosystem considerations of the KwaZulu-Natal sardine run

The annual winter sardine run along the South African east coast impacts the KwaZulu-Natal (KZN) coastal system in a variety of ways. These include ecological impacts, such as enrichment of a largely oligotrophic environment, competition between migrant sardine Sardinops sagax, other migrant and resident small pelagic fish species, and interactions with predators, as well as the socio-economic impacts of the sardine run on the local people. Enrichment of KZN coastal waters with organic nitrogen contained within the sardine is compared with alternative sources of nitrogen such as upwelling, river, sewage and stormwater runoff, and groundwater discharge. The sardine run appears to contribute most nitrogen to this system—96 000 t compared to 500–3 300 t for each of the other significant sources at trophic level 2, although upwelling estimates are extremely wide. Nonetheless, the majority of surviving sardine, their young and predators return southwards, suggesting that the nett export of nitrogen to KZN waters during the run is likely to be of a similar order of magnitude as that from other sources. Further, whereas the sardine supply of nitrogen is exclusively during winter, the bulk of the riverine input is in summer, thus ensuring that nitrogen supply in the region is maintained at fairly constant levels throughout the year. Competition for food between small pelagic fish is minimised by resource partitioning, but further dietary data are needed for resident species. Although interactions between sardine and top predators must exist, further studies are needed to confirm links between top predator life cycles and the sardine run. The estimated value of sardine as a tourist spectacle is compared to that from a seasonal beach-seine or boat-based purse-seine fishery for this species. Whereas the estimated value of the sardine as a tourist attraction appears substantially higher than could be derived from catching them, the small-scale beach-seine fishery itself draws tourists and also provides limited, seasonal employment opportunities.