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 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.     

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.     

Diet of the South African fur seal (1974–1985) and an assessment of competition with fisheries in southern Africa

Almost 1 000 South African fur seals Arctocephalus pusillus pusillus were collected at sea (inshore and offshore) for examination of stomach contents and other biological parameters. The main prey of seals above the age of one year was teleost fish, of which at least 28 species were identified. Overall, 74,6 per cent of the diet was teleost fish, 16,7 per cent cephalopods, 4,8 per cent crustaceans such as shrimps, prawns, amphipods and isopods and 3,2 percent rock lobster. Within this overall dietary pattern there were marked regional differences. The importance of individual fish species in the diet was investigated from the numbers and the sizes of otoliths occurring in the stomachs, from which the identity and size classes of the prey species could be calculated. Again there were regional differences in the importance of different prey species. On the west coast of South Africa the most important species were anchovy and Cape hakes (each 23% of total diet), whereas on the south coast the important species were anchovy (17%), horse mackerel (14%), pilchard (12%) and Cape hake (14%). Off South West Africa the two dominant species were pelagic goby (52%) and horse mackerel (23%), anchovy and Cape hake constituting only 4 and 5 per cent respectively of total diet. The total annual consumption of anchovy and Cape hake by seals in South African waters was calculated at 125 000 and 120 000 tons respectively.     

Comparative analysis of the diets of smaller odontocete cetaceans along the coast of southern Africa

The diets of 20 species of smaller Odontoceti were investigated. The material was collected along the southern African coast between 1966 and 1990 and included 158 strandings, 29 animals killed incidentally and 49 animals caught under permit. In total, 57 species in 34 families of fish and 89 species in 24 families of cephalopods were identified from remains in the 236 stomachs investigated. Most Odontoceti fed on commercially unexploited cephalopods and fish, or on fish below commercially utilized sizes. However, the average sizes of anchovy and pilchard eaten by Delphinus delphis and Lagenorhynchus obscurus were larger than the sizes most frequently taken commercially. The species composition of the prey items reflected the onshore-offshore distribution (and possibly the foraging depths) of different Odontoceti. Tursiops truncatus and Cephalorhynchus heavisidii were closest to shore and D. delphis and L. obscurus over the continental shelf. The rest were distributed around the shelf-edge and just off the edge. A comparison of the stomach contents of stranded and non-stranded D. delphis, L. obscurus and C. heavisidii demonstrated that results from stranded animals are biased.     

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 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.     

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.     

Spatial, temporal and ontogenetic variation in diet of anchovy (Engraulis encrasicolus) on the Algerian coast (SW Mediterranean)

The diet of anchovy Engraulis encrasicolus was studied in three regions (Béjaia, Bénisaf and Ghazaouet) along the Algerian coast. Ontogenetic, spatial and seasonal variations in anchovy diet were investigated using multivariate analyses and analysed in relation with sea surface temperature and chlorophyll-a. 46 prey taxa of varying size between 0.57 mm (Euterpina acutifrons) and 6.8 mm (fish larvae) were recorded. Whatever the season, the region or the fish size, anchovy is exclusively zooplanktivorous and copepods were the most present prey, constituting 87% by number of the prey taken and found in 98% of the anchovy stomachs examined. However, their occurrence and number varied according to the different areas, seasons and fish size. During its first year of life, anchovy feeds almost exclusively on copepods (mainly small and medium size prey). As anchovy grows, copepods are gradually substituted by large crustaceans such as decapods and amphipods. Hierarchical cluster analysis, analysis of similarities (ANOSIM) and similarities percentage (SIMPER) indicated a distinct diet of anchovy of the bay of Bejaia from those of the bays of Bénisaf and Ghazaouet probably due to differences in hydrologic conditions. Diet differences also occurred between seasons. Summer and spring have distinct prey assemblages each and showed low diet similarities with the two other seasons. More prey species were found in the diet during winter (36) and autumn (30) and the vacuity index was lower in winter. Temporal variability in satellite-derived chlorophyll-a matched the seasonal variability in the diversity of the anchovy prey and feeding intensity as reflected by the vacuity index, suggesting further investigation of the potential use of satellite-derived chlorophyll-a data as a proxy for anchovy feeding intensity.     

黄东海近代厌氧沉积物中鳀鱼鱼鳞沉积信息的空间分布及重建潜力

Many studies have revealed that anchovy has exhibited large variability in population size on decadal tim-escales. However, such works concerning anchovy population are mainly based on short historical catch records. In order to understand the causes of variability in fish stocks （natural and/or anthropogenic） and calibrate the error between catches and standing stocks, it is essential to develop long-term time series of fish stocks from the time when human impacts are minimal or negligible. Well preserved fish scales from sediment record are regarded as useful indicators revealing the history of fish population dynamics over the last centuries. Anchovy scales was first analyzed over the Yellow Sea and East China Sea and the largest abundance was found in the central South Yellow Sea where is regarded as the largest overwintering ground for Japanese anchovy （Engraulis japonicas）. Thus in the central South Yellow Sea, two cores covering the last 150 years were collected for estimating fish scale flux. The scale deposition rate （SDR） records show that the decadal scale SDRs were obviously coherent between cores with independent chronologies. Thecalibration of downcore SDRs to the standing stocks of anchovy further validated that SDR is a reliable proxy to recon-struct the long-term anchovy population dynamic in the central South Yellow Sea where anoxic conditions prevail in the sediment. When assembled with other productivity proxies, it would be expected that SDR could be associated with changes in oceanic productivity and may make a contribution to determine the forcing factors and elucidate the mechanism of the process in future.     

Environmental factors associated with a distinct decline in pelagic fish abundance as detected by aerial fish spotting in February and March 1982 off northern South West Africa

Aerial fish-spotting flights were carried out between Cape Frio and Palgrave Point, off the northern coast of South West Africa, during February and March 1982. Results showed a pronounced drop in the number of shoals of pelagic fish (anchovy Engraulis capensis and pilchard Sardinops ocellata) within the area from one month to the next. Consideration of available hydro-acoustic survey data, environmental data and egg counts led to the conclusion that the decline in shoal abundance was brought about by a southward migration of shoals, and that the migration was possibly triggered by environmental changes between January and February 1982. The implied time lag of 14—32 days is difficult to explain, but it could have been a consequence of favourable spawning conditions which still prevailed during February, coupled with the development of dense phytoplankton to the south of the survey area during March.     

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.     

Aspects of the upwelling regime between Cape Point and Cape Agulhas,South Africa

A progressive increase in mean ovary mass (standardized by the cube offish length) over time was observed in specified length classes of pilchard Sardinops ocellatus sampled from purse-seine catches off the west coast of South Africa. This, in earlier investigations, was considered to have been a density-dependent response following the collapse of the stock in the mid 1960s, and it appeared to result mainly from the increasing frequency of occurrence of pilchard entering the batch-spawning cycle, particularly among fish of 16–20 cm L_c . The possibility that the observed trend was an artifact of changing distribution of fishing or shifting seasonality of spawning relative to the three-month period of sample collection was rejected following analysis of variance on spatially disaggregated data and inspection of seasonal patterns in ovary mass. Other causes of the increase in mean ovary mass could have been density-dependence, declining age structure, changes in the environment and selection for early maturity under prolonged high rates of mortality. The data are critically examined to evaluate the likelihood of each of these explanations.     

Occurrence and feeding of three shark species,Carcharhinus brachyurus,C. obscurus and Sphyrna zygaena,on the Eastern Cape coast of South Africa

The shark species Carcharhinus brachyurus, C. obscurus and Sphyrna zygaena, which attain more than 2 m when adult, occur in coastal waters of the Southern Cape as juveniles. They use these coastal waters as nursery areas and their relative abundance there varies through the year, possibly because of seasonally varying oceanographic conditions. Adult C. brachyurus occur in the same geographic area but usually deeper, although there is some overlap in juvenile and adult depth ranges. Adult C. obscurus prefer the warmer subtropical and tropical waters north of the study area. Adult S. zygaena are found over deep reefs at the edge of the continental shelf. There was considerable dietary overlap between the three species, the prey of all three being dominated by small schooling fish, such as pilchard, and squid. For the three species, animals of total length >2 m took larger prey, such as elasmobranchs, with greater frequency, although they continued to prey on squid and schooling fish. It is postulated that the nursery areas are in the study area because of the abundance of food there, and that avoidance of cannibalism would be enhanced by the use of such nurseries.     

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.     

Pilchard distribution in South African waters, 1983–1985

Seasonal movements of South African pilchard Sardinops ocellatus in the relatively abundant 1982/83 year-class are inferred by means of length frequency information from predator diet analyses and from commercial and research catches in various localities. Pilchard of approximately one year of age were most abundant in the vicinity of Algoa Bay on the south-eastern Cape coast in the summer of 1983/84. The pilchard length frequency distribution in gannet diet and research midwater trawl samples in this area provided a clear linkage, through the known juvenile growth rate, with fish of an average 6 months of age, which were abundant off the West Coast during May 1983. Pilchard in this year-class were subsequently found to be widely distributed over the Agulhas Bank, with a tendency for them to inhabit deeper water towards the shelf edge as they reached the age of first spawning (about 18 months). During 1985, a noticeable increase in the abundance of large, two-year-old and older pilchard in the diet of West Coast gannets and in West Coast commercial catches indicated a movement of 1982/83 year-class fish into that area. The changes in distribution of pilchard in this year-class are discussed in relation to seasonal differences in the marine environment off South Africa.     

Validation of growth zone deposition rate in otoliths and scales of flathead mullet Mugil cephalus and freshwater mullet Myxus capensis from fish of known age

Flathead mullet Mugil cephalus and freshwater mullet Myxus capensis are important components in South African estuarine fish communities and fisheries, but there is little information on their age and growth or age validation. This study validated the periodicity of growth zone formation in sectioned sagittal otoliths and scales of fish of known age that had been at liberty for 10 years. For both M. cephalus and M. capensis, the number of growth zones counted on otolith sections did not differ significantly from the known age of the fish. There were significantly fewer growth zones on scales than the known age of both M. cephalus and M. capensis. Growth zone deposition rate could therefore be validated as annual for otoliths but not for scales of the two species. It is recommended that future ageing studies focus on otoliths rather than scales.     

Spatial pattern of coastal fish assemblages in different habitats in North‐western Mediterranean

Variability of fish assemblages across habitat structures can depend on spatial scales. A hierarchical sampling design was used to assess the spatial variability of temperate fish assemblages in different habitats and at multiple scales. Underwater visual censuses were carried out along the coasts of Elba Island (NW Mediterranean) on Posidonia oceanica beds, rocky algal reefs and sandy habitat at three spatial scales, namely tens of metres (individual replicates), hundreds of metres (sites) and tens of kilometres (locations). At the assemblage level, there was a clear relationship between fish and habitat type and the observed habitat‐related differences were largely dependent on species identity. Fish assemblages on P. oceanica beds and rocky reefs shared a high number of species, whereas overlap with sandy assemblages was negligible. Multivariate analyses revealed significant differences in fish assemblages among habitats, although there was also a significant habitat × site interaction. These differences relied mainly upon assemblage composition and species richness. Assemblages on rocky reefs and P. oceanica meadows usually harboured a higher number of species and individuals compared with sandy assemblages. Nevertheless, the patterns of habitat‐related differences in species richness and, especially, in the total number of fish, changed significantly from site to site. Eight species showed significant differences over habitats, but they were not consistent due to the interaction of habitat with site. Predictability of fish at both assemblage and population levels decreased with the scale of observation, and the spatial pattern of fish observed at the smallest scale was likely dependent on factors other than habitat type.     

Diet shifts of Caribbean grunts (Haemulidae) and snappers (Lutjanidae) and the relation with nursery-to-coral reef migrations

The spatial size distribution of grunts and snappers have previously indicated the separation of juveniles in nursery habitats from the adults on the coral reef. This implies life cycle migrations from nursery habitats (such as seagrass beds and mangroves) to the coral reef. If diet shifts are related to such migrations, then the diets of these fish must change before or around the fish size at which such migrations take place. A wide size range of juveniles of two grunt species (Haemulon sciurus and Haemulon flavolineatum) and of two snapper species (Lutjanus apodus and Ocyurus chrysurus) were caught in seagrass beds and mangroves, and their gut contents identified and quantified. Regression analysis between fish size and dietary importance of small crustaceans showed a negative relationship in all four species. Positive relations were found for H. sciurus, L. apodus and O. chrysurus between fish length and the dietary importance of decapods, and for L. apodusand O. chrysurus between fish length and prey fish importance. Critical changes in the fish diets with fish size were examined by application of a Canonical Correspondence Analysis (CCA). The CCA yielded three clusters of size-classes of fishes with similar diets, and application of a Mantel test showed that each of these clusters had significantly different diets, and that each cluster diet was significantly specialised. The size at which a fish species ‘switched’ from one cluster to another was compared with size-at-maturity data and with the typical size at which these species migrate from the nursery habitats to the coral reef. H. sciurus and H. flavolineatum may be prompted to migrate from the nursery habitats to coral reef habitats because of dietary changes, or because of the development of the gonads. For L. apodus and O. chrysurus, a dietary changeover forms a more likely explanation for nursery-to-reef migrations than does sexual maturation because these species reach maturity at sizes much larger than the maximum size of individuals found in nursery habitats. Although other factors may theoretically initiate or promote the migration patterns, the results of this study indicate that ontogenetic dietary changes may crucially influence the nursery-to-coral reef migrations of these reef fish species.     

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.