Diet and gill morphology of the East Coast redeye round herring Etrumeus wongratanai off KwaZulu-Natal,South Africa

Stomach content analyses and measurements of gillraker morphology were used to assess the diet and feeding ecology of the East Coast redeye round herring Etrumeus wongratanai and provide data for comparisons with other small pelagic fishes off South Africa. Samples were collected by jigging from a kayak off Scottburgh, KwaZulu-Natal (KZN), over the period July 2013–May 2014. In total, 66 stomachs (containing 4 407 prey items) and 66 gill arches were examined. Stomach content analyses indicated that East Coast redeye round herring, at the time of sampling, fed on large (1 500–2 500?µm) particles, with ～2 500?µm particles making the greatest contribution to dietary carbon. The species feeds on larger particles than do anchovy Engraulis encrasicolus and sardine Sardinops sagax, but on particles of a similar size to those consumed by West Coast redeye round herring Etrumeus whiteheadi. There were significant differences in mean gillraker gaps between East Coast redeye round herring and both sardine and anchovy, but not between East Coast and West Coast redeye round herring. The differences in gillraker gaps between East Coast redeye round herring, sardine and anchovy are indicative of resource partitioning through differential feeding, based on zooplankton size. The data suggest that there is no resource partitioning between East and West coast redeye round herring, indicating that competition is likely to occur between them. However, such competition is minimised by limited spatial overlap between these two species.     

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.     

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.     

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.     

The potential effects of cannibalism and intraguild predation on anchovy recruitment and clupeoid fluctuations

To obtain a first estimate of the potential impact of density-dependent regulation by cannibalism and by inter-specific predation by a guild species (pilchard) on anchovy population growth, a simple model based on a Leslie matrix was constructed. Simulations in which the pilchard biomass was held at 0 and the anchovy biomass increased, demonstrated that the eigenvalue decreased linearly from 1,03 at 1 000 tons to 0,94 at 2 million tons, showing a potential density-dependent mechanism for population regulation. The impact of pilchard on anchovy population growth depended on the assumed clearance rate. At the maximum biomass tested (2 million tons of pilchard) the eigenvalue of the anchovy Leslie matrix varied from 0,91 to 0,36, depending on the anchovy biomass and the pilchard clearance rate used. Field evidence indicates that the fastest clearance rate tested may be the most realistic of the three options. The model assumed uniform distribution of eggs in the spawning area, but the results are sensitive to the simulated concentration of eggs. Therefore, the actual patterns of spatial distribution of both eggs and anchovy and pilchard adults would have a major impact on the extent of egg consumption. The results indicate that egg consumption is potentially an important intra- and interspecific mechanism causing population fluctuations.     

Comparison of the abundance of pilchard in Cape gannet diet and commercial catches off the Western Cape,South Africa

A hypothesis is presented that, in the absence of quota restrictions, commercial catches of pilchard Sardinops ocellatus in the western Cape and the abundance of pilchard in the diet of Cape gannet Morus capensis obtained at three South African colonies are directly related to pilchard biomass when the latter is low. There were significant correlations (p < 0,05) between the abundance of pilchard in gannet diet at Lambert's Bay and Malgas Island combined, and the total catch and CPUE at annual, though not monthly, time-scales. Trends in levels of correlation are discussed in relation to short-term variability, sampling intensity, statistical chance and the distribution of pilchard. Gannet diet and commercial catches reflect regional, but not local, changes in biomass. The 1983 pilchard year-class was relatively strong. Gannet diet is suggested to be a reliable monitor of the trend in pilchard biomass at low pilchard biomasses, a time at which direct surveys are least reliable.     

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.     

Comparison of anchovy biomass estimates measured by trawls,egg production methods and hydro-acoustics in the Chesapeake Bay and the Korea Strait

We compared estimates of anchovy biomass derived from trawl surveys, egg production method (EPM) and acoustic surveys, conducted in two remote regions. Biomass density of bay anchovy Anchoa mitchilli was estimated in Chesapeake Bay, USA, by trawls, EPM and acoustics from 1989 to 2000. Biomass density of Pacific anchovy Engraulis japonicus was estimated in the Korea Strait using EPM, simulation-based daily cohort analysis and acoustics from 1984 to 2006. Most of the existing estimates already had considered body-size-dependent gear selectivity, highlyvariable instantaneous natural mortality of anchovy eggs, and avoidance of trawl nets by adult anchovy. Despite great variability in the ratio of trawl to acoustic biomass estimates (0.034–8.35), annually-averaged biomass density of young-ofthe-year individuals derived by the two methods were similar for bay anchovy in Chesapeake Bay and Pacific anchovy in the Korea Strait (0.83 and 0.70 g m^?3, respectively). Results suggested that, despite substantial uncertainty, anchovy biomass estimates are generally compatible between EPM and acoustics. However, reported estimates of biomass density derived from the two acoustic surveys in the Korea Strait differed by a factor of 28, suggesting that further improvements in calibrations are required to reliably estimate anchovy biomass. The comparisons suggested that all biomass estimates could be biased and will require comparison and validation by other, independent sampling methods.     

Impact and implications of large-scale environmental anomalies on the spatial distribution of spawning of the Namibian pilchard and anchovy populations

Trends in the spatial pattern of spawning of pilchard Sardinops ocellatus and anchovy Engraulis capensis during the period 1971–1985 were examined in relation to large-scale environmental anomalies outside the spawning area. Shifts in the locality of peak spawning of anchovy often coincided with that of pilchard, although sometimes anchovy were slower to respond to an environmental change. With negative sea surface temperature (SST) anomalies spawning shifted northwards, whereas with positive anomalies spawning took place both in the north and farther south. With a transition from negative SST anomalies early in the spawning season to zero or positive anomalies later in the spawning season, the impact on anchovy and pilchard spawning differed. Anchovy spawning was either reduced or absent during the negative phase of transitional seasons, thus avoiding unfavourable conditions. Pilchard, on the other hand, were spawning mainly in the north during the negative phase of the transitional seasons, but also in the south during the zero to positive phase. The shifts in spawning locality in response to environmental anomalies outside the spawning area imply that explanations for such shifts need not be sought in changes in biomass or age structure or in the fact that stocks may be genetically different, all theories previously advanced.     

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.     

Identification of and distinction between the scales of South African pilchard Sardinops ocellatus and Cape anchovy Engraulis capensis

Fossil fish scales hold potential for elucidating past fluctuations in fish populations. Only certain pilchard Sardinops ocellatus and anchovy Engraulis capensis scales can be distinguished easily from each other. Misidentification of the less-typical scales of the two species can introduce a bias in scale-based population studies and, in order to overcome this bias, scales removed from pilchard and anchovy reared in captivity were examined. A complete set of reference photographs of the scales of each species was compiled and study of these showed that both pilchard and anchovy have five distinctive scale types, each of which is found on different parts of the fish. It was also shown that the "typical" scales of each species, i.e. those referred to in the literature, constitute less than 50 per cent of the scales found on the fish.     

Variability in the diet of the Cape gannet at Bird Island,Algoa Bay,South Africa

Results are presented following a long-term study of the diet of the Cape gannet Morus capensis at Bird Island, Algoa Bay, examining 4 178 regurgitations representing 36 351 prey items collected during 12 breeding cycles from April 1979 to March 1991. In all, 32 teleost and two cephalopod species were recorded. The pelagic shoaling pilchard Sardinops ocellatus, anchovy Engraulis capensis and saury Scomberesox saurus scombroides were identified as the main prey species. The diet composition showed marked interannual fluctuations during the sampling period, which appear to be related to changes in the abundance of the main prey species. The relative abundance of pilchard, anchovy and saury was also found to change intra-annually, the first two species occurring more frequently in the diet during the breeding season and saury dominating the diet in the non-breeding season. These changes are thought to originate from temporal variation in the oceanographic regime within the foraging range of the gannets.     

Estimation of the annual consumption of food by Cape hake Merluccius capensis and M. paradoxus off the South African west coast

Information on hake stomach contents collected during research cruises off the west coast of South Africa between 1988 and 1990 is analysed. Estimates of the annual consumption and daily ration of the Cape hakes Merluccius capensis and M. paradoxus are obtained. Coefficients of variation (CVs) for these estimates are estimated by a bootstrap variance-estimation procedure. Cape hakes are opportunistic feeders. Juveniles feed mainly on crustaceans and the diet becomes increasingly piscivorous with age. Anchovy were the dominant dietary item of juvenile M. capensis, although this may be a reflection of the fact that anchovy were readily available at the time of the surveys. The large number of anchovy found in the diet during this study highlights the dangers of extrapolating consumption estimates for opportunistic feeders to other time periods. Pooling of data across geographic and seasonal strata introduces substantial bias in estimates of consumption and daily ration for only the pelagic prey species. Estimates of annual consumption are highly sensitive to the cruise selected to provide the estimates of numbers-at-length, because these latter estimates vary substantially between cruises. Furthermore, estimates of annual consumption and daily ration by prey species are very imprecise due to the effects of small sample sizes, the opportunistic nature of and natural variability associated with feeding. This implies that it is unlikely that model-estimation procedures which utilize these data will be able to provide particularly precise predictions. Therefore, before any larger-scale stomach collection exercise is undertaken, it is advisable to perform simulation studies to assess the sampling intensity required to achieve the desired levels of accuracy and precision for predictions from multispecies model-estimation procedures which make use of such data.     

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.     

Use of particle-size data to predict potential pelagic-fish yields of some southern African areas

A particle-size spectrum model is used to estimate standing stocks of some pelagic fish from measurements of phytoplankton chlorophyll in two exploited regions in southern Africa (southern Benguela and off South West Africa/Namibia) and in two unexploited regions (the Agulhas Bank and off the eastern Cape Province). The model is based on the assumption that equal biomasses occur in logarithmically equal size classes in the pelagic marine environment. Phytoplankton, with an equivalent spherical diameter ranging from 1 to 128 μm, occupy 21 size classes on the logarithmic scale. Two different size ranges are assumed for some commercial, pelagic fish species, equivalent to exploitation with two different purse-seine mesh sizes. A mesh of 12,7 mm would catch 8 size classes of pilchard, horse mackerel and anchovy whereas a mesh of 32 mm would catch only 3,3 size classes of pilchard and horse mackerel. From the model, the potential biomass of these commercial pelagic fish is estimated, after allowing for the presence of other commercial and non-commercial fish and other taxa in the exploited size range. Total pelagic fish production is estimated by assuming constant turnover rates of 1·y^?1 and 1,5·y^?1 when exploited with 32 and 12,7 mm mesh nets respectively. Consideration of the maximum and mean reported catches in the exploited areas indicates that only some 25 per cent of pelagic fish production is exploitable by man. On this basis, the unexploited Agulhas Bank region may yield some 400 000 metric tons (wet) of pelagic fish of the species considered, and the East Coast region some 90 000 tons. Exploitation in these regions cannot be recommended, because the Agulhas Bank is an important spawning ground for many pelagic species, and the fish in both regions probably act as a reserve buffer for the heavily exploited pelagic resource of the Western Cape.     

Distribution of and predation on saury Scomberesox saurus scombroides in continental shelf waters off the Cape Province,South Africa

Although the size distribution of larvae and early juveniles of the saury Scomberesox saurus scombroides in continental shelf waters off the Cape Province, South Africa, is consistent with a south-north passive dispersal by known currents, the size of late juveniles and adults increases from north-west to south-east. Occurrence of these stages is highly seasonal: they are found in summer off the Western Cape and mainly from late summer to winter in Algoa Bay in the Eastern Cape, possibly reflecting longitudinal migration. Large numbers of late juveniles and adults are often found downstream of upwelling plumes off the Western Cape when warmer waters lie close inshore. They are sometimes taken in purse-seines in association with adult round herring Etrumeus whiteheadi and pilchard Sardinops ocellatus. Important predators of late juvenile and adult saury are fast-moving, surface-feeding species: yellowfin tuna Thunnus albacares, Cape gannet Morus capensis and Cory's shearwater Calonectris diomedea. Cape gannets consume an estimated 2 277–6 044 tons of saury annually in South African waters. Predator diets provide important time-series of occurrence, abundance and length-frequency distributions of late juvenile and adult saury.     

Age and growth of redeye round herring Etrumeus whiteheadi off the South African coast

Ages of redeye round herring Etrumeus whiteheadi were estimated with associated errors and used to infer life-history information, such as age composition, age-at-maturity and instantaneous mortality rate. Samples were collected in November 2005 during a research survey aimed at estimating the biomass of spawning pelagic fish off South Africa’s west and south coasts. Replicate age estimates obtained from sagittal otoliths were collected with slight bias and relatively high precision. A von Bertalanffy model describing growth of the combined sexes, including juveniles, was L_t = 20.41(1 ? e^{?0.41(t ? 1.92)}). Kimura’s likelihood ratio test revealed no statistically significant differences between growth parameters of males and females. Results suggested that otolith length is a better predictor of age than otolith weight. Maturity estimates for E. whiteheadi were similar to those previously documented.     

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.     

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.     

Food and population variability in five regions supporting large stocks of anchovy,sardine and horse mackerel

The Benguela, California, Humboldt and Canary currents and the coastal waters of Japan support large stocks of sardine or pilchard Sardinops spp. and Sardina sp., anchovy Engraulis spp., horse mackerel Trachurus spp. and chub mackerel Scomber japonicus, with hake Merluccius spp. also abundant in each of the systems except off Japan. In these systems many of the more numerous fish, birds and mammals are opportunistic feeders having catholic diets. As a result species often have overlapping diets, and when a particular food item is plentiful it may be consumed by a wide variety of organisms at higher trophic levels. Frequently the performances of predator populations have been related to the availability of forage species, but predators have not always been influenced similarly by gross changes in the systems. Predators with the most generalized feeding strategies appear best buffered against collapses of particular prey resources. Although seabirds are generally opportunistic feeders, limitations in their foraging range may make them more susceptible than most other predators to system perturbations. Opportunistic feeding by species at lower trophic levels suggests that they could be advantaged by an increased food supply, and it may not be unreasonable to interpret species replacements in catches as reflecting shifts in the dominance of species. Overall catches from the systems have usually been more stable than the catches of individual species, which have shown wide fluctuations. Shifts in the dominance of species in the catches have often been sustained over a number of years. Generalized feeding creates the possibility that more than one species may benefit from energy made available by the collapse of an abundant resource, and there is evidence that species more abundant in former years have in some instances been partially replaced by a number of organisms. The likelihood of a particular organism becoming the main replacing species will probably be influenced by the degree to which it is utilized during and after the collapse of the originally dominant resource.