Diet of the Cape fur seal Arctocephalus pusillus pusillus at the Robberg Peninsula,Plettenberg Bay,and implications for local fisheries

Cape fur seals Arctocephalus pusillus pusillus were harvested to extirpation on the Robberg Peninsula, Plettenberg Bay, on the south-east coast of South Africa, between the 17th and early 20th centuries. Seals returned to Robberg in small numbers during the early 1990s and their numbers subsequently increased. We studied the diet of this increasing population using faecal (scat) sampling to determine: the species composition and size of prey in the diet of Cape fur seals at Robberg; to explore temporal variation in the diet; and to investigate the potential for competition between seals and the fisheries around Plettenberg Bay. Of the 445 scats collected, 90% contained hard prey remains and 15 teleost prey species were represented in the 3 127 otoliths that could be identified. The seals’ most important prey species in terms of numerical abundance, frequency of occurrence and mass in the diet, were anchovy Engraulis encrasicolus, sardine Sardinops sagax, horse mackerel Trachurus capensis, sand tongue-fish Cynoglossus capensis and shallow-water hake Merluccius capensis (in decreasing order of importance for numerical abundance). The proportion of anchovy in the diet increased during the study period (2003–2008), whereas the proportion of sardine decreased. The estimated average annual consumption of sardine by seals was higher than the average annual catch made by purse-seine fisheries in this area, suggesting resource competition between seals and purse-seiners, especially in the light of continuing growth of seal numbers in the area. However, direct competition between seals and linefisheries appeared to be minimal. Scat sampling of Cape fur seals holds potential to serve a useful and cost effective indicator of temporal changes in sardine abundance.     

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.     

Fine‐scale diet of the Australian sea lion (Neophoca cinerea) using DNA‐based analysis of faeces

We applied DNA‐based faecal analysis to determine the diet of female Australian sea lions (n = 12) from two breeding colonies in South Australia. DNA dietary components of fish and cephalopods were amplified using the polymerase chain reaction and mitochondrial DNA primers targeting the short (~100 base pair) section of the 16S gene region. Prey diversity was determined by sequencing ~50 amplicons generated from clone libraries developed for each individual. Faecal DNA was also combined and cloned from multiple individuals at each colony and fish diversity determined. Diets varied between individuals and sites. Overall, DNA analysis identified a broad diversity of prey comprising 23 fish and five cephalopod taxa, including many species not previously described as prey of the Australian sea lion. Labridae (wrasse), Monacanthidae (leatherjackets) and Mullidae (goat fish) were important fish prey taxa. Commonly identified cephalopods were Octopodidae (octopus), Loliginidae (calamary squid) and Sepiidae (cuttlefish). Comparisons of fish prey diversity determined by pooling faecal DNA from several samples provided a reasonable but incomplete resemblance (55–71%) to the total fish diversity identified across individual diets at each site. Interpretation of diet based on the recovery of prey hard‐parts identified one cephalopod beak (Octopus sp.) and one fish otolith (Parapriacanthus elongatus). The present study highlights the value of DNA‐based analyses and their capabilities to enhance information of trophic interactions.     

Diet of mid-Atlantic Sowerby's beaked whales Mesoplondon bidens

The first mid-Atlantic diet of Mesoplodon beaked whales is presented, from ten Sowerby's Mesoplodon bidens stranded in the Azores region between 2002 and 2009. This doubles the worldwide number of stomachs sampled, and reveals new feeding habits for this species. The mean number of prey items per stomach was 85±89 (range: 12–238), with fish accounting for 99.3% and cephalopods contributing less than 1% of total prey. Fish otoliths from 15 families and cephalopod lower mandibles from three families were identified, representing 22 taxa. The diet consisted mainly of small mid-water fish, the most numerous being Diaphus sp., Lampanyctus sp. and Melamphaidae species. Myctophids were present in all stranded individuals, followed by Diretmidae, Melamphaidae and Opisthoproctus soleatus, while the remaining fish species were scarce or single occurrences. Consistency of diet in four different years reveals a divergence from all previous records in continental areas, where mainly neritic and shelf-break benthopelagic fish species have been reported. Mid-Atlantic Sowerby's beaked whales' showed dietary plasticity, feeding on the most abundant mid-water groups occurring between 0 and750 m. Trophic level from prey numerical frequency was estimated at 4.4±0.46.     

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 diet of mesopelagic fish from the Pacific coast of Hokkaido,Japan

The diet of at least 28 species of mesopelagic fish from the Pacific coast of Hokkaido was examined. The dominant family was the Gonostomatidae (42%) which was represented by five species. The most abundant species wasCyclothone atraria which together with the other species of this genus preyed predominantly on copepods. Euphausiids and copepods were dominant in the diet ofGonostoma gracile. The next most abundant family was the Myctophidae (32%) which was represented by seven species. The dominant species,Stenobrachius nannochir, preyed mainly on copepods. Copepods were also the dominant food item of the other myctophids except forLampanyctus jordani which fed mainly on euphausiids. The other important family was the Bathylagidae (21%).Leuroglossus schmidti was the dominant species and its diet was more diverse with ostracods, copepods, molluscs and larvaceans being the most important food items.Bathylagus ochotensis had a similar diet. Copepods were the most important food items for all but a few species and their occurrence in the fish stomachs was related to the known vertical distribution of both predators and prey. Ostracods and euphausiids were also important prey items, the latter especially in large fish species. Molluscs and larvaceans were restricted to the two species of the family Bathylagidae.     

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.     

Predation patterns of demersal teleosts from the Cape south and west coasts of South Africa. 2. Benthic and epibenthic predators

The feeding patterns of 12 teleost species taken by demersal trawl from two areas of the Cape coast and which fed predominantly on benthic and epibenthic prey are described. Prey selection varied intraspecifically with growth. There was considerable overlap in prey taken by different predators, although the dominant food differed between the two coasts and according to habitat (soft substrata or hard reefs). Prey selection varied with depth, probably as a result of the influence of depth and other physical features on prey species distribution. Small crustaceans were taken by all the predators and constituted the single most important group, especially for small predators. Brachyurans were important as prey, particularly in those species inhabiting hard substrata. Fish were of minor importance as food, except in Chelidonichthys capensis, Malacocephalus laevis and Helicolenus dactylopterus. Commercially important species of fish and cephalopods were preyed on only occasionally. It appeared that prey were selected according to their abundance, size, behaviour and habitat use. The use of benthic and epibenthic prey constitutes an important pathway for energy to return from the benthos to the foodweb.     

Trophic relationships in the demersal fish community off Namibia

The benthic fish community off Namibia (between the Cunene River and Walvis Bay) in 50–650 m of water was studied during three bottom trawl surveys. The community was investigated on the basis of distribution, abundance and diet of 51 species, constituting 95 per cent of the demersal fish biomass. Dietary studies revealed the existence of three major trophic groups, one containing species that prey on pelagic and nektonic organisms, a second dominated by predators that feed on benthic polychaetes and copepods and a third group containing predators of benthic crabs, demersal shrimps and fish. Well differentiated from these groups are a few species that prey mainly on jellyfish and ophiuroids. Because for most of the species, their trophic level changes with growth, they were divided into size classes which are analysed independently. To describe interactions between the size classes, a similarity index combining diet affinity with spatial coincidence was applied. Cluster analysis showed that, for Merluccius capensis, Raja straeleni, R. clavata and Trachurus trachurus, there is a large difference in both geographic distribution and dietary preference between the size classes. For species such as Hoplostethus atlanticus, Austroglossus microlepis and Coelorinchus flabellispinis, the different size classes seem to share the same ecological niche. Variation in the number of species and the parameters mean abundance and niche width are described to enhance knowledge of the general structure of the community.     

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.     

Insights into the diet of a poorly known species: pygmy killer whale Feresa attenuata from Cape Verde,West Africa

Despite having a worldwide distribution in tropical waters, knowledge on pygmy killer whales Feresa attenuata, including diet, is poor, with only a few studies carried out to date. The presence of otoliths and beaks in stomachs that have been examined indicate that the diet of F. attenuata includes squid, octopus and fish. In this study, the stomach contents of two F. attenuata recently stranded in Cape Verde, West Africa, were examined. A variety of fish hard parts and otoliths were recovered, in addition to the upper beak from an unidentified cephalopod and the remains of some marine arthropods. Lanternfish (Myctophidae) and deepbody boarfish Antigonia capros were identified from otoliths and comprised 95.4% of prey items. Whereas the sizes of the myctophids in the two whale stomachs were similar, those of other prey items differed. The findings provide important information on the diet of F. attenuata in the Cape Verde region.     

Feeding Habits of the Mesopelagic Fish Gonostoma gracile in the Northwestern North Pacific

The diet of Gonostoma gracile, a numerically abundant mesopelagic fish in the Subtropical Region and the Transition Domain of the northwestern North Pacific, was examined using 520 specimens collected during June–July 1988, June 1995 and November 1995. The prey included mainly copepods, ostracods, amphipods and euphausiids. Copepods and ostracods were the most abundant, comprising approximately 70% of the total diet. There was little evidence of an ontogenetic dietary shift; Pleuromamma copepods were the most abundant prey for all size classes of fish ranging from 19 to 116 mm in standard length. The size range of prey increased with growth, but all fish sizes examined fed mainly on 1–4 mm long prey. Luminescent copepods and ostracods were the most abundant prey, suggesting that G. gracile detects its prey visually. This revised version was published online in August 2006 with corrections to the Cover Date.     

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.     

Anchovy Engraulis encrasicolus diet in the North and Baltic Seas

The diet of anchovy (Engraulis encrasicolus) in the North and Baltic Seas was studied using stomach analysis from four sampling events in different areas. Zooplanktivory was confirmed; the most frequent prey items (in over 40% of stomachs) were copepods, malacostracan larvae and fish larvae. In the Baltic Sea, Paracalanus spp. and Pseudocalanus spp. were important in relative terms; in the German Bight, Temora spp. dominated the stomach contents. Relative abundances of prey items varied with area more than absolute abundance or presence absence of items. Moreover, the level of resolution of prey categories influenced which prey categories were considered to be most important in driving variability in stomach content. Anchovy diet is broad across the seasons, years and areas sampled, suggesting that it is not a specialist feeder in the North Sea. The similarity of diet between anchovy and other clupeids, as well as anchovy consumption of larval fish, makes the new increased anchovy population a potential intraguild predator of commercial species like herring.     

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.     

Feeding of the Norwegian spring spawning herring Clupea harengus (Linne) at the different stages of its life cycle

The aim of the research was to investigate the diet of herring at different stages of its life cycle. For that purpose feeding of 0-group and immature herring in the Barents Sea, as well as of mature fish from the Norwegian Sea, was studied. 0-Group herring was sampled in the Barents Sea in August–September 2002–2005 during the international 0-group and trawl-acoustic survey of pelagic fish, as well as during the trawl-acoustic survey of demersal fish in November–December 2003–2004. Stomach samples of immature herring (1–3 years) were collected in late May and early of June 2001 and 2005 in the south-western part of the Barents Sea during the trawl-acoustic survey for young herring. Stomach samples of mature herring were collected in the Norwegian Sea in 1996, 1998, 1999, 2001, and 2002 in the course of the international trawl-acoustic survey of pelagic fish. Feeding intensity of herring of all age groups varied considerably between years and this was probably associated with availability and accessibility of their prey. The 0-group herring was found to have the most diverse diet, including 31 different taxa. In August–September, copepods, euphausiids, Cladocera, and larvae Bivalvia were most frequent in the diet of 0-group herring, but euphausiids and Calanus finmarchicus were the main prey taken. In November–December, euphausiids and tunicates were major prey groups. It was found that C. finmarchicus in the diet of 0-group herring was replaced by larval and adult euphausiids with increasing fish length. C. finmarchicus was the principal prey of immature herring and dominated in the diet of both small and large individuals and mainly older copepodites of C. finmarchicus were taken. Larval and adult euphausiids were found in stomachs of immature herring as well, but their share was not large. The importance of different prey for mature herring in the Norwegian Sea varied depending on the feeding area and length of the herring. On the whole C. finmarchicus and 0-group fish were the most important prey for mature herring diet, but fish prey were only important in a small sampling area. Hyperiids, euphausiids, tunicates, and pteropods were less important prey, and in 2002 herring actively consumed herring fry and redfish larvae.     

Assessing the competitive effects of marine mammal predation on commercial fisheries

Commercial fishermen in many parts of the world perceive marine mammals as competitors for a scarce resource: fish. The importance of predation by marine mammals can be evaluated crudely by considering the marine mammal as a rival fishery whose operation is constrained by physiology and energetic requirements, rather than by catch limits and gear restrictions. Evidence collected from grey seals Halichoerus grypus in the United Kingdom, southern elephant seals Mirounga leonina and beluga whales Delphinapterus leucas shows that published information on distribution, based on the location of breeding sites and sightings in shallow water, may give an unrealistic picture of foraging distribution. Studies of diet and distribution can give an impression of the static nature of the inter-action between a marine mammal and the fishery. However, it is the dynamics of the interaction which are important. A critical question to be posed is how will the consumption of a particular prey species vary with its availability to the predator?     

Food and feeding of small fish in the Rakaia River,New Zealand

Benthic macroinvertebrates and fish species were collected at monthly intervals during a 12‐month period, from 3 areas of the lower Rakaia River. The composition and abundance of the benthos and stomach contents of the fish were analysed for seasonal trends, food niche breadth and overlap between pairs of fish species, and overlap between the benthos and the diet of each fish species. The diets of bluegilled bully (Gobiomorphus hubbsi), upland bully (G. breviceps), juvenile longfinned eel (Anguilla dieffenbachii), Galaxias brevipinnis, G. paucispondylus, and juvenile brown trout (Salmo trutta) were similar to the proportions of prey species in the benthos. Deleatidium and chironomid larvae dominated the benthos in all seasons and formed the majority of food items in the diet of these species. Food niche overlap between these species was high, indicating potential competition, but preferred habitat and feeding habit differences plus low fish population density and abundance of main prey items probably eliminate the occurrence of any serious competition. The diet of the other fish species differed from this pattern. The common bully (Gobiomorphus cotidianus) ate proportionally more chironomid larvae and fish eggs than occurred in the benthos; torrentfish (Cheimarrichthys fosteri) ate proportionally more chironomid larvae than occurred in the benthos; and quinnat salmon (Oncorhynchus tshawytscha) depended largely upon prey species of terrestial origin, such as adult Deleatidium and dipterans.     

The importance of cephalopods as prey for hake and other groundfish in South African waters

Literature on trophic relationships in the Benguela ecosystem has stressed the importance of cephalopods as prey of groundfish. The groundfish community of the shelf and upper slope of southern Africa is dominated by the Cape hakes, and the results presented (1984–1991) confirm that both species of hake are important predators of cephalopods, especially taking into consideration the abundance of hake in the ecosystem. However, geographic, seasonal and species variability are evident in the patterns observed. The main prey species are Sepia spp. (predominantly Sepia australis), Loligo vulgaris reynaudii, Todaropsis eblanae and Lycoteuthis ?diadema. The last-named is an important food organism for fish. Its systematic status needs revision, however. Qualitative results of studies of cephalopod predation are also provided for kingklip and monkfish.     

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.