The food of five gadoid species during the pelagic O-group phase in the northern North Sea

The diets of five gadoid species in the northern North Sea are described: the cod, Gadus morhua (L.); haddock, Melanogrammus aeglefinus (L.); whiting, Merlangius merlangus (L.); saithe, Pollachius virens (L.); and the Norway pout, Trisopterus esmarkii (L.). The diet of each species was shown to be a gradual progression to different prey species as the predators increased in length. There were differences in the food taken by each species with respect to numbers, species and size. These differences in diet suggest that the degree of competition between pelagic O-group gadoids is not great.     

Resource sharing among cod, haddock, saithe and pollack on a herring spawning ground

Cod Gadus morhua , haddock Melanogrammus aeglefinus , pollack Pollachius pollachius and saithe Pollachius virens were abundant fishes on herring Clupea harengus spawning grounds o. southwestern Norway. Their diets were species specific and ontogenetic changes were similar to patterns observed in other areas, indicating both inter- and intraspecific resource sharing. However, strong interspecific diet overlap occurred when cod and pollack preyed upon sandeel Ammodytes marinus , and when all the gadoids fed on herring eggs during the herring spawning season in the spring. These prey items were very abundant and the extensive diet overlap may not reflect strong competitive interaction.     

Observations on the food and diel feeding behaviour of pelagic O-group gadoids in the northern North Sea

The diets and diel feeding behaviour of six O-group gadoid species are described, the cod Gadus morhua (L.), the haddock Melanogramus aeglefinus (L.), whiting Merlangius merlangus (L.), saithe Pollachius virens (L.), Norway pout Trisopterus esmarkii (L.) and the blue whiting Micromesistius poutassou (R.). There were differences, in the main food groups selected by each species with respect to the type, size and number. The main prey consumed did not vary with depth. Cod, saithe and Norway pout had only one period of feeding activity whereas haddock and whiting apparently had two. The diets of haddock and whiting varied in association with these two periods. It is suggested that the differences between the diets were the result of each species having a different feeding behaviour which reflects that adopted in adult life.     

Does diet in Celtic Sea fishes reflect prey availability?

Feeding preferences of Celtic Sea fishes were investigated using a database of stomach content records, collected between 1977 and 1994. The diet of cod Gadus morhua , hake Merluccius merluccius , megrim Lepidorhombus whiffiagonis , whiting Merlangius merlangus and saithe Pollachius virens changed markedly as the animals grew larger, and although large predators generally chose larger bodied prey, the variability of prey sizes consumed also increased. Large predators continued to select small, low value, benthic prey ( e.g . Callionymus spp. and Trisopterus spp.) which were easier to catch, rather than larger, more energy lucrative pelagic prey ( e.g . mackerel Scomber scombrus ), even though these pelagic prey‐fishes were nearly always available and were often very abundant. Stock estimates of the International Council for the Exploration of the Sea and U.K. groundfish survey catches were used as indices of prey abundance. Blue‐whiting Micromesistius poutassou and other small pelagic fishes ( Argentina spp. and clupeoids) were identified as being particularly important, and were consumed by some predators more often than would be expected given the abundance of these prey in the environment. There was no evidence for density‐dependent feeding by predators on mackerel and only hake exhibited density‐dependent feeding on horse‐mackerel. Hake, cod and megrim consumed more blue‐whiting when this prey was at higher abundance in the environment. In choosing what prey to consume, predators must balance costs and benefits, considering the quality of prey and the energy expended during search, capture and handling.     

The trophic ecology of Atlantic cod: insights from tri-monthly, localized scales of sampling

The meso-scale trophic dynamics of cod Gadus morhua were examined based upon tri-monthly stomach sample collections from a nearshore, localized ( c. 800 km²) region off Cape Cod, Massachusetts, U.S.A. The major objective for this work was to relate any changes in cod diet and amount of food eaten to seasonal variations in prey availability, water temperature and spawning at a spatial scale between broad-scale and laboratory studies. Results suggested that the type and amount of food eaten by cod was generally consistent throughout a year and repeatable across years. Cod feeding was marked by two periods of increased feeding, corresponding to the arrival of small pelagic fishes in the area. This pelagic migration and subsequent increased feeding by cod occurred during important periods in the life history of cod ( e.g . spawning and overwintering). Similar annual patterns in food consumption and diet composition were remarkably consistent over the 2·5 years of the project, suggesting important feeding periods for cod that correspond to environmental and biological events. The diet of cod was composed primarily of several species of forage fishes [ e.g. herrings (predominantly Atlantic herring Clupea harengus ), sand lance Ammodytes sp. and Atlantic mackerel Scomber scombrus ], ophiuroids, Cancer sp. crabs and other small crustaceans. It was inferred that cod exhibited a maintenance diet on local forage fishes and benthic macroinvertebrates, augmenting their diet by seasonally gorge feeding upon migrating pelagic species.     

Feeding habits and gill raker morphology of three planktivorous pelagic fish species off the coast of northern and western Kyushu in summer

Feeding habits and gill raker morphology were examined for the three major planktivorous pelagic fishes, Japanese anchovy Engraulis japonicus , Pacific round herring Etrumeus teres and Japanese jack mackerel Trachurus japonicus , off the northern and western coasts of Kyushu, in the north‐eastern part of the East China Sea in the summer months of 2001. Using fishes in the same size range (80–140 mm, standard length), the stomach contents of the three fish species were compared. The diet of the Japanese anchovy mainly consisted of Oncaeidae copepods, while the diets of the Pacific round herring and Japanese jack mackerel were dominated by calanoid copepods at all stations. Comparisons between prey size in the stomach, zooplankton size in the water and gill raker morphology suggested that the stomach contents of the three species were characterized mainly by the difference in the feeding behaviour between Japanese anchovy (filter‐feeding) and the other two species (particulate‐feeding), rather than by the difference in the morphology of feeding apparatus only. It was concluded that behavioural adaptations in the feeding of these pelagic fishes brought about trophic partitioning to some degree in this pelagic ecosystem in summer. Although the diets of these three species overlapped to some extent, there was still little likelihood of competition between the Japanese anchovy and the other two species. The potential for competition between the Pacific round herring and the Japanese jack mackerel is discussed.     

Transitional ecological requirements for early juveniles of two sympatric stichaeid fishes,Cebidichthys violaceus andXiphister mucosus

Synopsis The distribution patterns, diets, and substratum (refuge) requirements of early juveniles of two sympatric stichaeid fishesCebidichthys violaceus andXiphister mucosus, were investigated in a rocky intertidal habitat at Diablo Canyon, California. Monthly investigations were conducted at low tide for four consecutive months, to assess ontogenetic differences in distribution, diet, and refuge requirements within and between the two species. Distinct differences in vertical zonation were exhibited by both stichaeids throughout the study. Interspecific zonation patterns were similar to those recorded for adults of both species. Diet analyses showed that early juveniles of both stichaeids were zooplanktivorous, differing markedly from the primarily herbivorous diets of adults. Changes in diet were largely due to the selection of larger prey taxa as both fishes, and their mouth size, grew over the study period. Predation by both fishes on water-column planktors (calanoid copepods, zoea and polychaete larvae) was greatest following initial intertidal settlement and habitat establishment by early juvenile fishes. Greater dependence on substrate-oriented and/or benthic prey (harpacticoid copepods, gammarid amphipods and mysid shrimp) was exhibited by both fishes as they grew in size. Affinities for sand, gravel, and pebbles during monthly field surveys were similar for both species throughout the study. As the fishes grew, their substratum preferences changed in relation to the substrata which provided the best refuge. Results from laboratory experiments indicated that young stichaeids select very specific substrata based on fish age (size) and substratum suitability (i.e. adequate refuge). Similarities in diet and substratum preferences, and changes in those preferences over time, appear to be the result of morphological similarities (body size and shape and mouth gape) for both species at a given age.     

Food of anadromous brown trout at sea

The food of migratory Salmo trutta in coastal marine waters along the Norwegian Skagerrak coast varied significantly with age, season and habitat of the fish. The main prey categories in terms of frequency of occurrence were fishes followed by crustaceans, surface insects and polychaetes. Seasonal variation in diet and within habitats was found, supporting the view that brown trout is an opportunistic feeder. An ontogenetic niche shift was observed with post-smolts feeding on inshore and shallow water prey communities, while larger brown trout fed mainly on pelagic fishes.     

Assessment of the diets of cod Gadus morhua and whiting Merlangius merlangus juveniles in a frontal region close to the Norwegian Trench: co‐existence or competition?

Over 200 pelagic juvenile cod Gadus morhua and whiting Merlangius merlangus were sampled in spring 2001 at stations inside and outside a frontal region close to the Norwegian Trench, and their stomach contents analysed in the context of zooplankton samples taken at the same time and same locations. Merlangius merlangus juveniles appeared to feed on a diverse zooplankton community, whereas G. morhua juveniles appeared to preferentially select Pseudocalanus elongatus, even though these prey items were not the dominant component of the prey community. The results suggest that during the pelagic juvenile stage, M. merlangus and G. morhua are direct competitors for zooplanktonic prey. In addition, because M. merlangus are more opportunistic, they may be more tolerant to changes in the species composition of the prey community, and therefore out‐compete G. morhua juveniles at times of low preferred prey abundance or under changing environmental conditions.     

Observations on the Diet of Some Slope Dwelling Fishes of Southern Portugal

The diets of 33 fish species from the southern Portuguese slope, at depths between 498 and 740 m were studied through the examination of stomach contents. Species of scyliorhinids, squalids, rajids, chimaerids, anguilliforms, notacanthids, macrourids, gadids and merluccids, with pelagic, benthope-lagic and benthic modes of life were examined. Nezumia sclerorhynchus feeding habits were studied in detail. Its diet its dominated by amphipods. No differences of prey preferences were recorded corresponding to predator size classes. The studied predators can exploit a wide range of prey, but they feed mainly on benthopelagic and epibenthic material. Consequently it is difficult to establish distinct feeding guilds, so confirming a generalist type of feeding for most of the upper slope dwelling fishes.     

Feeding habits of ocellate spot skate,Okamejei kenojei (Müller & Henle, 1841), in coastal waters of Taean,Korea

The feeding habits of Okamejei kenojei were studied using 592 specimens collected in the coastal waters of Taean, Korea from April 2008 to March 2009. O. kenojei is a bottom‐feeding carnivore that consumes mainly shrimp, fishes, and crabs. Its diet also includes small quantities of amphipods, mysids, cephalopods, euphausiids, copepods, isopods, and polychaetes. The total length (TL) of individuals in this study ranged from 8.2 to 49.0 cm. Cluster analysis based on %IRI (index of relative importance) identified three size classes. Group A (< 20 cm TL) ate primarily caridean shrimp and amphipods; group B (20–30 cm TL) ate exclusively shrimp; and group C (> 30 cm TL) ate penaeoidean shrimp, fishes, and crabs. O. kenojei showed ontogenetic changes in feeding habits. Although shrimps were the primary food consumed by all size groups, the proportion of shrimp in the total diet decreased and the consumption of fishes and crabs gradually increased with the body size of O. kenojei. Size of the prey organisms also increased. Smaller individuals fed mainly on small prey, such as amphipods, mysids, and small shrimp, whereas larger individuals preferred larger prey, such as larger shrimp, fishes, and crabs. The size‐related diet breadth and the percentage of empty stomachs were significant; the diet breadth gradually increased with body size, whereas the percentage of empty stomachs decreased. Seasonal changes in the O. kenojei diet were not significant, but shrimp constituted 97.3% of the summer diet by %IRI. Seasonal changes in diet breadth and the percentage of empty stomachs were not significant.     

Piscivore diet response to a collapse in pelagic prey populations

Pelagic fish populations in the upper San Francisco Estuary have experienced significant declines since the turn of the century; a pattern known as the pelagic organism decline (POD). This study investigated food habits of piscivorous fishes over two consecutive fall seasons following the decline of pelagic fish prey. Specifically, this study addressed the contribution of pelagic versus benthic prey to piscivorous fish diets, including the frequency of predation on special-status pelagic species, and the spatial variability in prey consumption. The piscivore community was dominated by Striped Bass and also included small numbers of Sacramento Pikeminnow and Largemouth Bass. Overall, pelagic prey items contributed less than 10% of the diet by weight in both years, whereas pre-POD studies gleaned from the literature found contributions of 39–100%, suggesting a major switch from pelagic to benthic prey resources. Between-year variation in piscivore diets reflected differences in environmental conditions associated with variation in freshwater outflow. No special status fish species were detected in any of the piscivore stomachs examined. The consequences of this pelagic to benthic diet shift warrants further investigation to understand its ecological relevance.     

Feeding ecology of silver hake larvae on the Western Bank, Scotian Shelf, and comparison with Atlantic Cod

The feeding ecology of the larvae of silver hake Merluccius bilinearis was examined during two time periods (October 1998 and December 1992) on the Western Bank, Scotian Shelf, north‐west Atlantic, and compared with the feeding ecology of Atlantic cod Gadus morhua larvae collected in the same samples in December 1992. During both time periods silver hake exhibited strong selection for late stage copepodids and adult copepods at a small size (>3·5 mm total length, L _T). The niche width measured as the diet breadth index ( I _DB) of silver hake declined rapidly as they increased in size and remained relatively constant from 3 to 11 mm L _T, during each time period. Atlantic cod larvae exhibited a broader niche width that was curvilinear over the same L _T. Atlantic cod were also less selective than silver hake, incorporating both naupliar and early stage copepodids in their diets throughout the length classes examined. Simple isometric relationships did not explain the differences in diet, as Atlantic cod larvae continued to feed on early stages of copepods at large size, while silver hake larvae quickly switched to large prey items. The strong selection and narrow I _DB observed for silver hake probably reflects adaptation to spawning during the periods between major secondary production peaks in temperate waters.     

An impact of seasonal squid migrations and fishing on the feeding spectra of subantarctic notothenioids Patagonotothen ramsayi and Cottoperca gobio around the Falkland Islands

Seasonal changes in the diet of two abundant notothenioid fishes, rock cod Patagonotothen ramsayi (Nototheniidae) and frogmouth Cottoperca gobio (Bovichthidae) were studied on the Falkland Island shelf. The rock cod is a near-bottom browser, feeding mostly on gelatinous and crustacean plankton. The diet of juveniles [10–16 cm total length (TL)] includes mostly copepods in summer and comb jellies Mnemiopsis leydyi in winter. Adult rock cod (17–34 cm TL) in summer consumed mainly plankton, with comb jellies and salps being of primary importance. In winter they mainly took benthos, primarily ophiuroids and lobster krill ( Munida spp.), and the comb jelly. During the fishing season, rock cod also scavenge on fishing discards. The frogmouth is a benthic ambush predator; both small (11–30 cm) and large (31–80 cm) representatives feed on the rock cod throughout the year. During the seasonal offshore migration of the squid Loligo gahi (June–August), this prey predominated in the large frogmouth diet, with crab, Peltarion spinosulum , of secondary importance. The region is seasonally exploited by the squid trawl fishery. As the fishing season commences (August–September), discarded rock cod also becomes an important food item for the frogmouth. Seasonal offshore migrations of L. gahi into the common depth range of both notothenioids have a significant direct (change in the C. gobio diet) and indirect (scavenging on fishing discards and targeting bottom scavengers by P. ramsayi and C. gobio during the squid fishing) impact on their diets over the Falkland shelf. The niche breadth of rock cod varied from 2.63 to 3.21 and that of frogmouth varied from 2 to 6.29, depending on fish size and season.     

Variation in the diet of Arctic Cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) in the Pacific Arctic and Bering Sea

Arctic cod, Boreogadus saida (Lepechin, 1774), is a nodal species in Arctic marine foodwebs as an important prey of many birds, marine mammals, and other fishes, as well as an abundant predator of zooplankton and epibenthic fauna. We examined the summer diet of Arctic cod across a latitudinal gradient extending from the southern limit of their distribution in the eastern Bering Sea to the northern margins of the eastern Chukchi Sea (ECS) continental shelf. Specimens were collected from demersal and pelagic trawls conducted between 1999 and 2012, and across a range of predator sizes (3–26 cm). Arctic cod diets vary with body size and between regions within the study area, and appear to vary between years in the eastern Bering Sea, indicating opportunistic feeding habits. Constrained Analysis of Principal Coordinates was conducted on ECS demersal samples and revealed consumption of fish and decapod crustacea were positively correlated with Arctic cod length while consumption of euphausiids and copepods had the opposite relationship. The demersal Arctic cod diet in the northern latitudes of the ECS was dominated by copepod consumption (47% by weight, %W), but copepods were less important (12–26%W) in the central and southern latitudes of the ECS and in the northern and eastern Bering Sea—areas where diets were more varied in their composition. High levels of variation in the diet of Arctic cod highlights the need to monitor Arctic cod diets to identify consistently dominant prey types and potential future changes to trophic relationships related to climate change or increasing anthropogenic activity.     

Food- and habitat-segregation in sympatric grayling and brown trout

Grayling in the large reservoir Aursjøen, Norway, did not use the pelagic habitat and only trout larger than 187 mm and older than 4+ were caught there. Both species preferred the upper 8 m in the benthic habitat, but grayling were caught deeper than trout in August and September. Grayling were more closely associated with soft substratum than trout. Association with soft substratum decreased with increasing size in grayling, whereas this relationship was reversed in trout. A low number of empty stomachs indicated a low degree of food competition, but despite this the two species segregated in different key prey items—probably due to differences in mouth shapes. The key prey item for grayling was the benthic Eurycercus lamellatus , with zooplankton dominating the diet of the smallest individuals. Large trout (>300 mm) preferred the benthic Lepidurus arcticus , whereas medium trout (160–300 mm) ate predominantly the limnic-pelagic Bythotrephes longimanus , irrespective of habitat. The distribution of key prey items seemed to be the major factor controlling the habitat use, except in small trout which showed a generalist feeding pattern throughout the period. A generalist feeding pattern, a close association to coarse substratum and absence from the pelagic habitat indicated that small trout were negatively influenced by large individuals. Small grayling seemed less affected by aggressive encounters as they were less associated with coarse substratum and ate predominantly high-risk food such as zooplankton. These individuals probably did shoal in order to reduce predation risk.     

Variation in the diet of Genypterus blacodes(Ophidiidae) around the Falkland Islands

The diet change with size, season and area was investigated using the stomachs of 496 kingclip Genypterus blacodes collected around the Falkland Islands (south‐west Atlantic) between August 2001 and September 2002. The key prey species were rockcod Patagonotothen spp., benthic isopods and Patagonian grenadier Macruronus magellanicus . Kingclip <50 cm total length ( L _T) fed mainly on crustaceans and small fishes. With size the diet shifted away from crustaceans towards Patagonotothen spp. in kingclip 50–100 cm L _T, and finally towards larger fishes such as M. magellanicus and Micromesistius australis australis in kingclip >100 cm L _T. The niche breadth was highest in fish >100 cm L _T and the lowest in fish <50 cm L _T. The larger kingclip generally selected larger individuals of the same prey species, with the exception of the Patagonian squid Loligo gahi , where all ingested squid were of similar size, regardless of the predator length. The importance of the main prey species varied substantially between five consequent seasons studied, and appeared to follow the seasonal abundance and availability of prey. The spatial variability in the diet was found in kingclip caught in regions occupied by transformed temperate and sub‐Antarctic waters. The rockcod, which is available throughout the year around the Falkland Islands, was the most important prey in the kingclip diet. Kingclip takes advantage of other seasonally abundant prey species during their seasonal migrations ( e.g . L. gahi ) and also scavenge on discards from fishing vessels when available.     

Diet composition,feeding niche partitioning and trophic organisation of large pelagic predatory fishes in the eastern Arabian Sea

Information on the ecology and feeding behaviour of the large oceanic predatory fishes is crucial for the ecosystem approaches to fisheries management models. Co-existing large pelagic predators in the open oceans may avoid competition for the limited forage by resource partitioning on spatial, temporal or trophic levels. To test this, we studied the prey species composition, diet overlap, trophic level, and trophic organisation of 12 large predatory fishes co-existing in the eastern Arabian Sea. Stomach contents of 1,518 specimens caught by exploratory longline operations in the Indian Exclusive Economic Zone during the years 2006–2009 were analysed. Finfishes were dominant prey of all species except blue marlin (Makaira nigricans) and yellowfin tuna (Thunnus albacares), which fed mainly on cephalopods, and long-snouted lancetfish (Alepisaurus ferox) and pelagic stingray (Pteroplatytrygon violacea), which fed mainly on crustaceans. Common dolphinfish (Coryphaena hippurus) and yellowfin tuna fed on a wider variety of prey than the other species, while the diets of lancetfish and black marlin (Istiompax indica) were narrowest. Pelagic stingray and great barracuda (Sphyraena barracuda) fed on species occupying epipelagic waters, whereas the contribution of mesopelagic prey was higher in the diets of swordfish (Xiphias gladius) and pelagic thresher (Alopias pelagicus). Trophic levels of these fishes ranged from 4.13 to 4.37. Diet overlap index revealed that some of the large pelagic predatory fishes share common prey species. Cluster analysis of the diets revealed four distinct trophic guilds namely ‘flyingfish feeders’ (common dolphinfish and great barracuda); ‘mesopelagic predators’ (pelagic thresher and swordfish); ‘crab feeders’ (lancetfish, pelagic stingray and silky shark) and ‘squid feeders’ (yellowfin tuna, Indo-Pacific sailfish (Istiophorus platypterus), skipjack tuna (Katsuwonus pelamis), black marlin and blue marlin). Large predatory fishes of the eastern Arabian Sea target different prey types, and limit their vertical extent and time of feeding to avoid competing for prey.     

Fish feeding guilds in shallow rocky and soft bottom areas on the Swedish west coast

Multivariate analyses of the diet relationships within fish assemblages resulted in the recognition of five major feeding guilds in shallow (0-9 m) rocky- and soft-bottom habitats on the Swedish west coast. The two habitats had similar guilds that could be broadly described as piscivores, decapod feeders, copepod feeders and amphipod and mollusc feeders. The last guild on the rocky bottom was more strictly one of amphipod feeders, whereas the last guild on soft bottoms consisted of infauna feeders (mainly polychaetes and bivalves). A comparison of the diet of the entire fish assemblages showed that amphipods and gastropods were more prevalent as prey taxa on rocky bottoms, whereas bivalves and teleosts were more common on soft bottoms. A large proportion (12 out of 27 species) of the fish species investigated was found in both habitats. These fish species generally seemed to rely on the same major prey groups in the two habitats, but also included prey taxa characteristic for the habitat in which they resided. Sea trout Salmo trutta , mackerel Scombrus scombrus , saithe Pollachius virens , whiting Merlangius merlangus and cod Gadus morhua were found to be the major piscivores in both the investigated habitats. Herring Clupea harengus and members of the family gobiidae were the most prevalent prey species for piscivores within the rocky- as well as the soft-bottom fish assemblage. A comparison of feeding modes suggested that the fish assemblage on rocky bottoms predominantly (60% of the food items) relied on food found in vegetation. On soft bottoms, the average distribution of food categories among fish species was 44% benthic, 35% pelagic and 21% vegetation-associated food items.     

Deep-sea mystery solved: astonishing larval transformations and extreme sexual dimorphism unite three fish families

The oceanic bathypelagic realm (1000–4000m) is a nutrient-poor habitat. Most fishes living there have pelagic larvae using the rich waters of the upper 200m. Morphological and behavioural specializations necessary to occupy such contrasting environments have resulted in remarkable developmental changes and life-history strategies. We resolve a long-standing biological and taxonomic conundrum by documenting the most extreme example of ontogenetic metamorphoses and sexual dimorphism in vertebrates. Based on morphology and mitogenomic sequence data, we show that fishes currently assigned to three families with greatly differing morphologies, Mirapinnidae (tapetails), Megalomycteridae (bignose fishes) and Cetomimidae (whalefishes), are larvae, males and females, respectively, of a single family Cetomimidae. Morphological transformations involve dramatic changes in the skeleton, most spectacularly in the head, and are correlated with distinctly different feeding mechanisms. Larvae have small, upturned mouths and gorge on copepods. Females have huge gapes with long, horizontal jaws and specialized gill arches allowing them to capture larger prey. Males cease feeding, lose their stomach and oesophagus, and apparently convert the energy from the bolus of copepods found in all transforming males to a massive liver that supports them throughout adult life.