Feeding of hake and a first attempt at determining their trophic role in the South African west coast marine environment

During intensive biannual demersal fish-sampling cruises since January 1983 over the South African west coast shelf region (Orange River to Cape Agulhas, coast to 500-m isobath), feeding studies on both species of Cape hake (Merluccius capensis and M. paradoxus) were undertaken. These studies were of two forms, random sampling of the dietary preference and constituents of the hake caught during daylight only and intensive sampling of stomachs with food throughout a 28-hour cycle. From the index of relative importance, crustaceans were shown to be most important in the majority of length classes of hake studied, with mesopelagics (fish and cephalopods) and other hake also important. Differences by hake species, by predator size, by geographical location and by season were evident. The Cape hakes, which constitute some three-quarters of the demersal fish biomass in the area, are deduced to play a major role in the marine faunal environment, both as predators and prey, and some base-line figures of the quantities of different food items consumed by the resource are presented. Hake are largely opportunistic feeders and it is concluded that they can adapt to perturbations in the availability of prey, whether they be triggered by environmental cause or by, for example, purse-seining of particularly mesopelagic fish.     

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.     

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.     

Characterisation of the dietary relationships of two sympatric hake species,Merluccius capensis and M. paradoxus,in the northern Benguela region using fatty acid profiles

The two sympatric species of Cape hake, Merluccius capensis and M. paradoxus, have been the main targets of bottom-trawl fisheries off Namibia for several decades. The feeding ecology of these hakes has been studied mainly using stomach content analyses and thus there remain some gaps in our knowledge about food assimilated over the longer term. In this study, we used fatty acid (FA) profiles to characterise the dietary relationships of M. capensis and M. paradoxus. Muscle samples from hake (n=110) and their known prey (n=68) were collected during trawl surveys off Namibia during 2011. Significant differences between the neutral FA profiles of the hake populations were detected in December 2011 but not in January 2011, an indication of temporal variations in diet and resource partitioning. Comparisons of the neutral FAs in hake and the total FAs of potential prey showed no clear trophic connections, with the exception of flying squid Todarodes sagittatus, which had FA profiles very similar to those of M. paradoxus in December 2011. Our results highlight the complex and temporally shifting relationships that exist between hake and the large pool of prey available to them, and between the two hake species that overlap in their feeding habits and distribution within the highly productive Benguela Current region.     

The diet of Patagonian hake Merluccius australis polylepis and its daily ration of Patagonian grenadier Macrouronus magellanicus

The diet of the Patagonian hake Merluccius australis polylepis in 1986 was determined by analysis of stomach samples collected from commercial trawl catches. The major prey items were Patagonian grenadier Macrouronus magellanicus and, to a lesser extent, southern blue whiting Micromesistius australis. Large (81–100 cm) hake are mainly piscivorous and consume more grenadier than smaller (51–80 cm) hake. The smaller hake feed both on fish and invertebrates. There was no difference by sex in terms of stomach contents. The daily feeding cycle was also investigated from samples collected during 1991. Hake feed principally on 1-, 2-, and 3-year-old grenadier, which they seem to catch mainly at night. The daily ration of grenadier was between 0,043 and 0,26 per cent of body mass (1,692–9,849 g). The results are similar to those for other hake species in terms of composition of the diet, daily feeding cycle and daily ration.     

Greenland halibut diet in the Northwest Atlantic from 1978 to 2003 as an indicator of ecosystem change

The Northwest Atlantic marine community underwent dramatic changes during the last 30 years, including the collapse of many groundfish stocks and an increase in shrimp populations. Greenland halibut Reinhardtius hippoglossoides is an important commercial species and one of the top fish predators in this system. It is a large, wide-ranging flatfish that is found at depths up to 2200 m and it has an opportunistic diet which makes it a potential candidate for an ecosystem indicator. Analysis of stomach contents of Greenland halibut between 1978 and 2003 indicates that diet composition reflects the major changes in community structure. Over the entire period there was a clear increase in the importance of invertebrates, particularly after 1992. This change was associated with a higher importance of Pandalus shrimp and Gonatus squid and a protracted reliance on zooplankton by predators under 25 cm length. Capelin Mallotus villosus was the dominant prey between 1978 and 1992 for predators in the 12–63 cm range, but its importance dropped off drastically in the mid 1990s. Levels of main prey in the diet of Greenland halibut correlated well with fishery-independent surveys. Greenland halibut sample capelin well, compared to bottom trawl surveys and acoustic surveys. Greenland halibut consumed small shrimp which are not routinely caught by surveys and may be important in deriving information on year classes and growth of shrimp. Our results suggest that Greenland halibut's diet is a useful tracker of ecosystem change.     

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.     

Dietary plasticity of the oceanic striped dolphin, Stenella coeruleoalba, in the neritic waters of the Bay of Biscay

The striped dolphin, Stenella coeruleoalba, is an oceanic species that occasionally occurs in neritic habitats; in the Bay of Biscay it is abundant offshore and erratic in occurrence over the shelf. Given that prey assemblages differ widely among these habitats both in terms of taxonomic composition and of ecology, this would suggest that striped dolphins are able to shift from vertically migrating meso-pelagic prey to neritic or coastal prey types. We investigated the striped dolphin's dietary plasticity by examining the stomach contents of individuals stranded along the French Atlantic coast. 1109 prey items were identified belonging to 30 distinct taxa and their biomass was calculated. Fish accounted for 91% of the diet by number and 61% by mass; the rest was mostly cephalopods, crustaceans being present as trace. Specific composition included both oceanic (myctophid and sternoptychid fish; histioteuthid, gonatid and brachioteuthid cephalopods), neritic (gadids and anchovy; loliginid, sepiolid and sepiid cephalopods) and even coastal (atherinid fish) prey types, showing that these animals had changed their diet as they moved over the shelf.     

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.     

Feeding habits of the ommastrephid squid Nototodarus sloanii on the Chatham Rise,New Zealand

The diet of the squid Nototodarus sloanii was determined from examination of stomach contents of 388 specimens of 14.3–41.0 cm dorsal mantle length sampled at depths of 211–760 m on the Chatham Rise, New Zealand. Prey items were predominantly mesopelagic fishes, with some crustaceans and cephalopods. The most important prey species identified was Maurolicus australis, followed by Lampanyctodes hectoris, and unidentified squids (Teuthoidea). Multivariate analyses using distance‐based linear models, non‐parametric multi‐dimensional scaling, analysis of similarities, and similarity percentages, indicated crustaceans were more important in the diet of smaller squid (83–480 g), the fish component of the diet was dominated by L. hectoris on the east Chatham Rise and M. australis on the west Chatham Rise, and there may be a difference in diet with sex, with crustaceans and cephalopods more important for females. The results indicated that N. sloanii, similar to other ommastrephid squids, foraged primarily in the mesopelagic layers.     

A preliminary investigation of the feeding ecology of squaloid sharks off the west coast of southern Africa

The diets of 15 species of squaloid sharks caught between Walvis Bay and Cape Agulhas over a depth range of 50–1 016 m are analysed. The most common prey items were fish and cephalopods. Hake Merluccius spp. were common in the diets of Centrophorus squamosus, Squalus acanthias, and S. cf. mitsukurii. Myctophids were extensively preyed on by Centroscyllium fabricii, Deania calcea, D. profundorum, Etmopterus cf. brachyurus, E. compagnoi, S. acanthias and S. megalops, among others. The most common cephalopods in shark stomachs were Histioteuthis miranda, Lycoteuthis ?diadema, Octopus vulgaris and Todarodes angolensis. Crustaceans were uncommon prey of most species, except for Centroscyllium fabricii and Etmopterus cf. granulosus. Centroscymnus coelolepis was the only squaloid with cetacean remains in the stomach. That species appears to parasitize its cetacean victims.     

DNA-based identification methods of prey fish from stomach contents of 12 species of eastern North Pacific groundfish

Stomach content analysis of eastern North Pacific groundfish has been conducted routinely by researchers interested in understanding trophic interactions between key predator species and their prey. Identification of prey by traditional morphological methods has limitations however, due to the loss of identifiable characters from digestion and morphological similarities between taxa. Furthermore, some forage fish (e.g., osmerids, ammodytids, and juvenile gadids), common prey of Bering Sea and Gulf of Alaska groundfish, are difficult to distinguish because of their slender or fusiform shape, disarticulating easily during digestion. DNA-based identification methods were developed to differentiate among 18 fish species, some that are found at depths greater than 200 m, from four taxonomic families: Ammodytidae and Osmeridae (forage fish), Pleuronectidae (flatfish), and Gadidae (gadid fish). Polymerase chain reaction (PCR) amplification of a 739 basepair section of mitochondrial DNA cytochrome c oxidase I and an 862 basepair section of mitochondrial DNA cytochrome b was followed by restriction digest assays and resulted in species level resolution for 16 of 18 species of interest. PCR restriction digest assays applied to fish prey from stomach contents of groundfish indicated the presence of several target species, eulachon (Thaleichthys pacificus), walleye pollock (Gadus chalcogrammus), searcher (Bathymaster signatus), rock sole (Lepidopsetta bilineata), yellowfin sole (Limanda aspera) and either Bering flounder (Hippoglossoides robustus) or flathead sole (H. elassodon). The PCR restriction digest protocols improved the identification rate of predated fish from stomach contents compared to identification by conventional taxonomic methods alone, and DNA sequence analysis further resolved identification of unknown prey fish samples.     

Comparing trophic flows in the southern Benguela to those in other upwelling ecosystems

A balanced trophic flow model of the southern Benguela ecosystem is presented, averaging the period 1980–1989 and emphasizing upper trophic levels. The model is based largely on studies conducted within the framework of the Benguela Ecology Programme and updates the results of an expert workshop held in Cape Town in September 1989. Small pelagic fish other than anchovy Engraulis capensis and sardine Sardinops sagax, mainly round herring Etrumeus whiteheadi and mesopelagic fish, were important components of the food web in the southern Benguela. Severe balancing difficulties were encountered with respect to the semi-pelagic resources (hake Merluccius spp.) and demersal top predators (sharks), indicating the need for further research on the interaction of these groups with their ecosystem. The model is compared to other existing trophic flow models of ecosystems in major upwelling areas, i.e. the northern Humboldt Current (4–14°S), the California Current (28–42°N) and the southern Canary Current (l2–25°N), and to two independently constructed models of the northern Benguela ecosystem. These models are compared using network analysis routines of the ECOPATH software, focusing on the interactions between the five dominant fish species (anchovy, sardine, horse mackerel Trachurus trachurus capensis, chub mackerel Scomber japonicus and hake) that support important fisheries in all systems. The upwelling systems rank by size rather than species dominance. The ratio of catches and primary production differs between systems, partly because of differences in fishing regimes. Predation on the five dominant fish groups by other fish in the system was the most important cause of fish mortality in all models. Fishery catches are generally a larger cause of mortality for these groups than predation by mammals. The ecological cost of fishing appears to be comparatively low in the southern Benguela, because catches are low compared with the primary production, but also because the fishery is relatively low in the foodweb. However, in view of the very tight foodweb demonstrated in the model. it is likely that an increase in fishing pressure would cause severe trade-offs with respect to other components of the southern Benguela ecosystem.     

基于角质颚长度的头足类种类判别

本文根据2013 年8 月于舟山市沈家门东河菜场采集的5 种近海常见经济头足类(剑尖枪乌 贼、杜氏枪乌贼、金乌贼、曼氏无针乌贼和短蛸), 利用逐步判别和主成分分析法对其上、下角质颚 的各5 种长度指标(喙长、头盖长、脊突长、侧壁长、翼长)进行分析。结果显示, 这5 种头足类角质 颚长度差异显著(ANOVA: P<0.001), 其中以金乌贼的角质颚尺寸最大, 短蛸的角质颚尺寸最小。判 别分析显示, 角质颚长度适合用来划分头足类的种类, 综合判别成功率为96.2%, 其中以上颚侧壁 长、上颚头盖长和下颚脊突长对判定的贡献率最高。然而, 与之相比, 标准化后的角质颚长度则更适 合用来划分头足类的种类, 综合判别成功率达到100%, 其中以标准化下颚喙长和标准化下颚翼长对 判定的贡献率最高。主成分分析显示, 角质颚长度及标准化后角质颚长度对枪乌贼类和乌贼类的判 定成功率均达到100%.本文建立了一种基于角质颚长度判别分析法的头足类种类判定的新方法, 丰 富了头足类种类鉴定的技术手段, 为我国学者在相关领域的研究提供了基础。     

Greenland halibut diet in the Northwest Atlantic from 1978 to 2003 as an indicator of ecosystem change

The Northwest Atlantic marine community underwent dramatic changes during the last 30 years, including the collapse of many groundfish stocks and an increase in shrimp populations. Greenland halibut Reinhardtius hippoglossoides is an important commercial species and one of the top fish predators in this system. It is a large, wide-ranging flatfish that is found at depths up to 2200 m and it has an opportunistic diet which makes it a potential candidate for an ecosystem indicator. Analysis of stomach contents of Greenland halibut between 1978 and 2003 indicates that diet composition reflects the major changes in community structure. Over the entire period there was a clear increase in the importance of invertebrates, particularly after 1992. This change was associated with a higher importance of Pandalus shrimp and Gonatus squid and a protracted reliance on zooplankton by predators under 25 cm length. Capelin Mallotus villosus was the dominant prey between 1978 and 1992 for predators in the 12–63 cm range, but its importance dropped off drastically in the mid 1990s. Levels of main prey in the diet of Greenland halibut correlated well with fishery-independent surveys. Greenland halibut sample capelin well, compared to bottom trawl surveys and acoustic surveys. Greenland halibut consumed small shrimp which are not routinely caught by surveys and may be important in deriving information on year classes and growth of shrimp. Our results suggest that Greenland halibut's diet is a useful tracker of ecosystem change.     

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.     

Contrasting changes between the northern and southern Gulf of St. Lawrence ecosystems associated with the collapse of groundfish stocks

In order to have a global view of ecosystem changes associated with the collapse of groundfish species in the Gulf of St. Lawrence during the early 1990s, Ecopath mass-balance models were constructed incorporating uncertainty in the input data. These models covered two ecosystems (northern and southern Gulf of St. Lawrence; NAFO divisions 4RS and 4T), and two time periods (before the collapse, in the mid-1980s, and after it, in the mid-1990s). Our analyses revealed that the ecosystem structure shifted dramatically from one previously dominated by piscivorous groundfish and small-bodied forage species during the mid-1980s to one now dominated only by small-bodied pelagic species during the mid-1990s in both southern and northern Gulf. The species structure in the northern Gulf versus southern Gulf was different, which may explain why these two ecosystems did not recover the same way from the collapse in the early 1990s. Productivity declined in the northern Gulf after the collapse but increased in the southern Gulf. The collapse of groundfish stocks resulted in declines in the mean trophic level of the landings in both the northern and the southern Gulf. Even though fishing mortality was then intentionally reduced, this part of the total mortality was taken up by predation. The temporal changes in the internal structure of both ecosystems are reflected in their overall emergent properties.     

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 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.     

Prey spectrum of breeding sooty shearwaters (Puffinus griseus) in New Zealand

Prey remains from sooty shearwater (Puffinus griseus Gmelin, 1792) regurgitates, stomachs, and gizzards, were collected from four breeding colonies in southern New Zealand. We found a wide range of prey species (minimum 39 species of 35 genera), the most important of which were crustaceans (particularly euphausiid krill and hyperiid amphipods), cephalopods (notably arrow squid), fishes, and salps. Malacostracans (krill, amphipods, and decapods) were the predominant taxa of prey in both diversity and frequency of occurrence. Regurgitates were easily obtained from harvested chicks and provided the greatest range of prey remains (36 of the 39 species identified). However, these regurgitates comprised only 29% of the total number of samples collected. Fish, malacostracan, and salp prey ranged from 4 to 170 mm in size, whereas total lengths of squid ranged from 50 to 535 mm. Based on size and mass, the largest squid were undoubtedly scavenged, possibly in association with commercial fishers. The geographical distribution of prey species indicate that most sooty shearwaters breeding near Stewart Island forage in waters lying between the Subtropical and Polar Fronts. Our results suggest that the abundance of krill, the impact of fisheries and the influence of climate perturbations on prey species may play important roles in sooty shearwater breeding and survival.