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.     

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.     

Recolonisation of the Robberg Peninsula (Plettenberg Bay,South Africa) by Cape fur seals

The Cape fur seal Arctocephalus pusillus pusillus colony at Robberg Peninsula, Plettenberg Bay, on the south-east coast of South Africa, was driven to extinction by indiscriminate harvesting by the late 1800s. Seals only began to recolonise this site in the 1990s. This study describes the recolonisation process from 2000 to 2009, exploring both within- and between-year count data of seals using the site. Counts increased over the study period from <300 animals to >3 100. Generalised linear models indicated the importance of year and month in explaining variability in the counts. Withinyear variability in the counts decreased over the study period, which may be related to an increasing proportion of resident (as opposed to transient) seals in the colony. However, the colony is currently still in a transition phase with a low ratio of breeding to non-breeding animals, based on the low numbers of pups born in the colony (currently still <100 per year). The influx of seals to the Robberg area may be associated with shifts in prey availability at the ecosystem level. The colony benefits from the protection afforded by the reserve status of the Robberg Peninsula and the existence of a marine protected area adjacent to it. However, human interference associated with fishing and/or ecotourism on the peninsula may inhibit development into a substantial breeding colony. Potential interventions for the conservation and management of this colony are discussed.     

A note on the stomach contents of fur seals Arctocephalus pusillus pusillus beached on the south-east coast of South Africa

The stomach contents of 36 South African fur seals beached over a 20 year period along the south-east coast of South Africa were examined. In all, 15 fish and nine cephalopod prey types were identified. Five species of fish (Merluccius capensis, Trachurus trachurus capensis, Cynoglossus zanzibarensis, Pterogymnus laniarius, Austroglossus pectoralis) and two cephalopods (Loligo vulgaris reynaudii, Octopus sp.) were of major importance in the diet. The proportion of demersal species in the diet is higher than previously reported, suggesting that the seals forage extensively on the sea floor. The most important fish species in the diet was M. capensis, and the most important cephalopod was L. v. reynaudii. Male and female seals showed distinct prey selection, suggesting possible intraspecific resource partitioning. There is no evidence of competition between seals and other top predators.     

Impact of predation by Cape fur seals Arctocephalus pusillus pusillus on Cape gannets Morus capensis at Malgas Island,Western Cape,South Africa

Cape fur seals Arctocephalus pusillus pusillus were estimated to kill some 6 000 Cape gannet Morus capensis fledglings around Malgas Island in the 2000/01 breeding season, 11 000 in 2003/04 and 10 000 in 2005/06. This amounted to about 29%, 83% and 57% of the overall production of fledglings at the island in these breeding seasons respectively. Preliminary modelling suggests this predation is not sustainable. There was a 25% reduction in the size of the colony, the second largest of only six extant Cape gannet colonies, between 2001/02 and 2005/06. There has been a large increase in predation by Cape fur seals on seabirds around southern African islands since the mid-1980s, coincidental with both an increase in the seal population, altered management of the islands and an altered distribution of prey for gannets and seals. At Malgas Island, most gannet fledglings were killed between 10:00 and 18:00, the period when most are in the water around the island, from mid-January to mid-March, the main fledging period. The Cape gannet is classified as Vulnerable.     

African penguins as predators and prey – coping (or not) with change

African penguins Spheniscus demersus live in the Benguela and western Agulhas ecosystems off southern Africa. Their numbers decreased throughout the 20^th century from at least 1.5 million to about 0.18 million adults, although different regional trends were apparent. They feed to a large extent on shoaling epipelagic fish, notably anchovy Engraulis capensis and sardine Sardinops sagax, and regional trends in the abundance of penguins are associated with trends in the abundance and distribution of these prey fish. Many first-time breeders emigrate from colonies where feeding or other conditions at the time are unfavourable to more favourable breeding localities. This has led to both the extinction and formation of colonies. Food now may limit colonies at relatively small sizes, a fact attributable to industrial fisheries reducing the densities of forage fish. African penguins share their habitat with several other predators, with which they compete for food and breeding space. One of these, the Cape fur seal Arctocephalus p. pusillus, increased through the 20^th century to 1.5 – 2 million animals at its close. Reported observations of predation by fur seals on seabirds have increased in recent decades and threaten the continued existence of small colonies of penguins. Stochastic modelling suggests that colonies of 10 000 pairs have a 9% probability of extinction in 100 years, so smaller populations should be regarded as "Vulnerable". However, in a period of prolonged food scarcity off southern Namibia, the regional population decreased from more than 40 000 pairs in 1956 to about 1 000 pairs in 2000, and many colonies numbering less than 1 000 pairs became extinct. The minimum viable population for African penguins is currently considered to be >40 000 pairs, likely of the order of 50 000 pairs, a figure equivalent to its level in 2000. The chance of survival of the species through the 21^st century is tenuous.     

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.     

Dietary analysis of harbour seals (Phoca vitulina) from faecal samples and overlap with fisheries in Erimo,Japan

The number of harbour seals in Japan has been rebounding since protection began in the mid‐1980s. With the increase in seal numbers, increased conflict with fisheries has occurred through depredation and the belief that seals compete with fisheries for prey. However, competition can only be determined if the prey species and quantities seals consume over time are known. We studied the diet of harbour seals in Erimo, site of the largest population of harbour seals in Japan, from 2011 to 2012 and assessed the degree of prey overlap with local fisheries. We used both hard parts and DNA techniques to identify prey items in seal scats, and compared these results to local fisheries data. A total of 46 prey occurrences was detected by both methods, of which 17 matched between techniques at least to the family level. Hard parts methods identified five incidences of prey undetected by DNA methods in five scats (one incidence per scat). DNA methods identified 24 additional prey occurrences in 13 scats, for which no hard part evidence for that prey had been found. This more than doubled the total number of prey occurrences across the 15 scats compared. Overall, the most frequently occurring harbour seal prey were walleye pollock (Theragra chalcogramma), sculpins and snailfishes. In contrast, the top three groups targeted by fisheries were codfishes, salmon and invertebrates. Many species common in the harbour seal's diet such as snailfishes and blennies were not targeted by fisheries. Fishes such as greenlings, sculpins, rockfishes and Japanese anchovy (Engraulis japonicus) were common in the diet of harbour seals, but made up a very small proportion of fisheries catches in Erimo. The importance of other prey species varied between seasons. Sculpins, greenlings and forage fishes were the top three prey groups for harbour seals in spring by percent modified frequency of occurrence, whereas the most important target groups by percentage mass caught by fisheries were codfishes, cephalopods and other invertebrates. Sculpins and rockfishes remained as important prey items for harbour seals in summer and autumn, in addition to codfishes, snailfishes and blennies. In contrast, the main groups caught by fisheries in summer were other fishes, particularly Japanese amberjack (Seriola quinqueradiata) and various species of sharks and eels, and invertebrates. By autumn, Erimo fisheries had focussed on catching salmon. Salmon were taken by harbour seals as well during this period, but at a relatively lower frequency compared to other prey groups. The results of our study show that although harbour seals consumed several of the prey species targeted by fisheries, the relative importance of these species to seals and fisheries and the seasons in which they were targeted were different.     

A preliminary investigation of the feeding ecology of catsharks (Scyliorhinidae) off the west coast of southern Africa

The diets of six species of catsharks (Carcharhiniformes: Scyliorhinidae) caught between Walvis Bay and Cape Agulhas at depths between 50 and 1 016 m were analysed. The most common prey items were teleosts and crustaceans. Lanternfish (Myctophidae) were common in the diets of Apristurus microps, A. saldanha, Apristurus spp. and Holohalaelurus regani, whereas the Cape anchovy Engraulis capensis was preyed on extensively by Scyliorhinus capensis. The most common crustaceans in the diet were Parapagurus spp., Mursia cristimanus and Euphausia lucens. Cephalopods were uncommon prey items, except in A. microps, A. saldanha, Apristurus spp. and H. regani.     

Seabirds in the diet of Cape fur seals Arctocephalus pusillus pusillus at three mainland breeding colonies in Namibia

Predation by Cape fur seals Arctocephalus pusillus pusillus on seabirds has previously been attributed to a few individuals, mainly males. Scat samples were collected at three mainland breeding colonies of seals in Namibia to determine the extent of seabird predation. Mainly females were sampled. No scats from the van Reenen Bay and Atlas-Wolf Bay colonies, and only two from Cape Cross, contained feather remains, giving an overall frequency of occurrence of feathers in scats of 0.1%. The feathers found were of African penguin Spheniscus demersus and a species of cormorant (Phalacrocorax sp.).     

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.     

An overview of the living marine resources of Namibia

This paper gives an overview of the main living marine resources of Namibia. It focuses on the scientific research conducted during the past decade as input to the management of these resources. The distribution and habitats of the most important harvested species and the main seabird populations are briefly described and discussed. The life histories of the major exploited species are summarized, with emphasis on spatial and temporal spawning patterns, dispersal of early life stages, migration patterns of recruits and adults, and diet, the latter particularly as it relates to potential competition between species. A number of commercially important species, such as the hake Merluccius capensis and M. paradoxus, deep-sea red crab Chaceon maritae, West Coast rock lobster Jasus lalandii, skipjack tuna Katsuwonus pelamis, southern albacore Thunnus alalunga and to a lesser extent Cape horse mackerel Trachurus capensis, southern African sardine Sardinops sagax and Cape anchovy Engraulis capensis, are distributed across national boundaries, requiring regional cooperation in research and management. The history and current status of the major fisheries is discussed. Over the past 30–40 years total annual catches have declined from a peak of around 2 million tons in the late 1960s to less than a million tons in the 1990s. This decline has been due, mainly, to a collapse in the sardine stock in the late 1960s and 1970s, and a reduction in the catches of hake and horse mackerel under a conservative management strategy in the past decade. Changes in the abundance and distribution of commercially important species, as determined by acoustic and trawl surveys and catch-based analytical methods, are presented. The effect of major environmental anomalies on the distribution and abundance of the resources in recent years is discussed. The most dramatic anomaly in recent years was the wide-scale advection of low-oxygen water into the northern Benguela from the Angola Dome in 1994, and the subsequent Benguela Niño of 1995, which appear to have severely impacted the Namibian sardine population and many other resources. The present socio-economic value of the Namibian fishing industry is given together with the broad policy, legislation and formal structures for managing the living marine resources.     

Cape cormorants decrease,move east and adapt foraging strategies following eastward displacement of their main prey

Numbers of Cape cormorants Phalacrocorax capensis breeding in South Africa decreased by nearly 50% from approximately 107 000 pairs in 1977–1981 to 57 000 pairs in 2010–2014. Although four colonies had >10 000 pairs in 1977–1981, there was just one such colony in 2010–2014. Almost all the decrease occurred after the early 1990s off north-west South Africa, between the Orange River estuary and Dassen Island. South of this, the number breeding in the two periods was stable, with some colonies being formed or growing rapidly in the 2000s. The proportion of South Africa’s Cape cormorants that bred south of Dassen Island increased from 35% in 1977–1981 to 66% in 2010–2014, with the opposite situation observed in the north-west. This matched a shift to the south and east in the distributions of two of the Cape cormorant’s main prey species, anchovy Engraulis encrasicolus and sardine Sardinops sagax. In 2014, an apparent scarcity of prey in the north-west resulted in Cape cormorants attempting to take bait from hooks of fishing lines over an extended period, a behaviour not previously recorded. The number of Cape cormorants breeding in the south may be constrained by the absence of large islands between Dyer Island in the west and Algoa Bay in the east. If so, it may be possible to bolster the southern population through the provision of appropriate breeding habitat, such as platforms, or restricting human disturbance at suitable mainland cliff breeding sites.     

"Predation" of anchovy by an Agulhas ring: a possible contributory cause of the very poor year-class of 1989

There was a marked decrease in abundance of anchovy Engraulis capensis off South Africa between November 1988 and November 1990 caused by formation of poor year-classes in 1989 and 1990. The percentage of anchovy in the diet of Cape gannets Morus capensis indicated that the most marked decrease in anchovy biomass was between June and July 1989. A filament from the Benguela upwelling front was in the process of being entrained by a passing ring of Agulhas Current water at that time. The ring extracted a large volume of frontal water over a period of 2–3 months, prevented its return to the shelf region and possibly removed anchovy larvae and pre-recruits from the Benguela system. It may have played a role in depressing the 1989 anchovy year-class, and hence the 1989 recruit biomass.     

Regionally coherent trends in colonies of African penguins Spheniscus demersus in the Western Cape,South Africa, 1987–2005

From 1987 to 2005, numbers of African penguins Spheniscus demersus breeding in South Africa's Western Cape Province increased by about 50%. Numbers decreased at the four northernmost colonies in the region: Lambert's Bay and the three colonies in Saldanha Bay, although at Jutten Island the decrease is inferred from an estimate for 1987, derived from interpolation. Numbers also decreased at Geyser Rock and Dyer Island on the South Coast. At five colonies between Saldanha Bay and Dyer Island there were large increases. At a sixth colony in that region, Seal Island, where Cape fur seals Arctocephalus pusillus pusillus limit breeding space, numbers remained stable. At two colonies that were initiated in the early 1980s, Robben Island and Boulders, increases were initially rapid (>20% per annum) and matched growth of the South African stock of sardine Sardinops sagax. Strong growth at Dassen and Vondeling islands, between Robben Island and Saldanha Bay, was observed from about 1996–2002, when there was a large increase in the biomass of pelagic fish off South Africa. Increases at colonies between Saldanha Bay and Boulders slowed after 2002, whereas the colony at Dyer Island stabilised at that time. In 2003, a new colony was initiated east of Dyer Island at De Hoop Nature Reserve. These latter trends followed an eastward shift in the distribution of sardine. Small penguin colonies may act as foci for growth in a period when the distribution of prey is changing. Hence, it is important that they be maintained, especially those that, if lost, would increase the isolation of regional populations. Some of the small colonies are less susceptible to oil spills than colonies in the proximity of harbours, and for that reason also are important.     

Seasonal fluctuations in occurrence of inshore Bryde's whales in Plettenberg Bay,South Africa,with notes on feeding and multispecies associations

Seasonal fluctuations in the occurrence of inshore South African Bryde's whales Balaenoptera edeni were investigated between November 2005 and June 2008. Sighting data were collected in Plettenberg Bay on the south-east coast of South Africa. Bryde's whale occurrence was modelled in relation to the following environmental covariates: sea surface temperature, chlorophyll a concentrations and wind speed. Seasonal increases in encounter rates (sightings per day) were observed during summer and autumn, with a peak in April that corresponded to increased feeding activity and above average aggregation sizes. All three environmental covariates were significant factors in terms of explaining variability in the occurrence of whales. Multispecies associations with common dolphins Delphinus capensis and Cape gannets Morus capensis were most common in summer and autumn, when feeding activity was highest.     

Dusky dolphins Lagenorhynchus obscurus and Cape fur seals Arctocephalus pusillus pusillus: fatty acid composition of their blubber and prey species

The fatty acid composition of the blubber of five dusky dolphins Lagenorhynchus obscurus and five Cape fur seals Arctocephalus pusillus pusillus from the northern Benguela ecosystem (South-East Atlantic) and their main prey was determined. Differences in fatty acid composition of the inner and outer blubber layer of the dolphins were substantial, with higher relative amounts of monoenic fatty acids with 14, 16 and 18 carbons in the outer layer and higher relative amounts of saturated, long-chain monounsaturated and polyunsaturated fatty acids in the inner layer. This stratification is similar to the general pattern that has been observed in a variety of marine mammals, but the degree of stratification is the highest yet observed, with the long-chained (n3) fatty acids being more than 20 times more abundant in the inner than in the outer layer. On the other hand, the seals’ blubber consisted of only one uniform layer, densely supported by connective tissue. The whole-body fatty acid composition of the prey was species specific. The dolphins and seals had significantly different fatty acid composition of their blubber. In both species, the blubber fatty acid composition was different from the composition of the prey, indicating that a phylogenetic component is involved in the determination of the fatty acid composition of the blubber.     

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.     

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.     

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.