Diel variation in feeding rate and prey composition of herring and mackerel in the southern Gulf of St Lawrence

Diel feeding patterns of herring Clupea harengus and mackerel Scomber scombrus in the southern Gulf of St Lawrence were examined based on samples obtained by midwater trawling between 19 and 26 June 2001. Within 3 h time periods, stomach contents tended to be more similar between fish from the same tow than between fish from different tows. Thus, in contrast to previous diet studies, which have used individual fish stomachs as independent observations, tow was used as the experimental unit in statistical analyses in this study. Diel patterns in stomach fullness were identified using generalized additive models. Two peaks in stomach fullness occurred for herring, one in the morning and the other in the evening. Mackerel showed an increase in feeding intensity throughout the day with a peak in mid‐afternoon. The diel changes in stomach contents suggested rapid gastric evacuation rates for both species, especially for herring. The estimate of the instantaneous evacuation rate for herring was twice that for mackerel. Calanus copepods (mainly C. hyperboreus ), fishes (mainly capelin Mallotus villosus ) and euphausiids were the main prey found in the stomachs of both species. Calanus copepods dominated the diet of herring regardless of time period. They also dominated the diet of mackerel during the late afternoon, evening and night while fishes and euphausiids were dominant during the morning and early afternoon. These diel patterns emphasize the need for sampling throughout the day and night in order to estimate ration and diet composition for bioenergetic and ecosystem models.     

Feeding Ecology of Northeast Atlantic Mackerel,Norwegian Spring-Spawning Herring and Blue Whiting in the Norwegian Sea

The Norwegian spring-spawning (NSS) herring (Clupea harengus), blue whiting (Micromesistius poutassou) and Northeast Atlantic (NEA) mackerel (Scomber scombrus) are extremely abundant pelagic planktivores that feed in the Norwegian Sea (NS) during spring and summer. This study investigated the feeding ecology and diet composition of these commercially important fish stocks on the basis of biological data, including an extensive set of stomach samples in combination with hydrographical data, zooplankton samples and acoustic abundance data from 12 stock monitoring surveys carried out in 2005–2010. Mackerel were absent during the spring, but had generally high feeding overlap with herring in the summer, with a diet mainly based on calanoid copepods, especially Calanus finmarchicus, as well as a similar diet width. Stomach fullness in herring diminished from spring to summer and feeding incidence was lower than that of mackerel in summer. However, stomach fullness did not differ between the two species, indicating that herring maintain an equally efficient pattern of feeding as mackerel in summer, but on a diet that is less dominated by copepods and is more reliant on larger prey. Blue whiting tended to have a low dietary overlap with mackerel and herring, with larger prey such as euphausiids and amphipods dominating, and stomach fullness and feeding incidence increasing with length. For all the species, feeding incidence increased with decreasing temperature, and for mackerel so did stomach fullness, indicating that feeding activity is highest in areas associated with colder water masses. Significant annual effects on diet composition and feeding-related variables suggested that the three species are able to adapt to different food and environmental conditions. These annual effects are likely to have an important impact on the predation pressure on different plankton groups and the carrying capacity of individual systems, and emphasise the importance of regular monitoring of pelagic fish diets.     

Movement,site fidelity and connectivity in a top marine predator,the killer whale

Movement, site fidelity and connectivity have important consequences for the evolution of population structure and therefore the conservation and management of a species. In this study photographs of naturally marked killer whales collected from sites across the Northeast Atlantic are used to estimate fidelity to sampling locations and movement between locations, expressed as transition probabilities, pt, using maximum likelihood methods. High transition probabilities suggest there is high inter-annual site fidelity to all locations, and large-scale movement between the spawning and wintering grounds of both Norwegian and Iceland stocks of Atlantic herring. There was no evidence of movement between the Norwegian herring grounds and Icelandic herring grounds, or between the mackerel fishing grounds and the herring fishing grounds. Thus the movement of predictable and abundant prey resources can lead to intrinsic isolation in this species We also find movement between the Northern Isles, Scotland and East Iceland. An association network indicates that killer whales predating seals around the Northern Isles, Scotland are linked to the community of killer whales that follow the Icelandic summer-spawning herring. This adds support to existing evidence of a broad niche width in some populations.     

Dietary variation within and between populations of northeast Atlantic killer whales,Orcinus orca,inferred from δ13C and δ15N analyses

Epidermal skin samples from eastern North Atlantic killer whales, Orcinus orca, were analyzed for carbon and nitrogen stable isotope ratios. From those, comparisons within a data set of 17 samples collected from Tysfjord, Norway, in November suggested that diet is relatively specialized during this time period at this location. There were significant differences between a small set of samples from Iceland and those collected from Norway, which had all been assigned to the same population by a previous population genetics study. The results would be consistent with matrilines feeding on either the Norwegian or Icelandic stocks of Atlantic herring (Clupea harengus). There was no significant difference within Icelandic samples between those assigned to the population known to feed upon herring and those assigned to a population hypothesized to follow Atlantic mackerel (Scomber scombrus). The greatest differences were between the epidermal samples analyzed in this study and tooth and bone collagen samples from the North Sea that were analyzed previously, which also showed significantly more variation in isotopic ratios than found for skin samples. These differences could reflect differences in turnover rate, differences in diet‐tissue fractionation and discrimination due to the amino acid composition of the different tissues, and/or greater competition promoting dietary variation between groups in the North Sea.     

Effects of alternative prey on predation intensity from herring Clupea harengus and sandeel Ammodytes marinus on capelin Mallotus villosus larvae in the Barents Sea

Stomach contents from lesser sandeel Ammodytes marinus and herring Clupea harengus caught at one and three stations, respectively, were analysed to investigate predation intensity on capelin Mallotus villosus larvae. Most capelin larvae were found in the anterior sections of the stomachs close to the oesophagus, indicating that rapid digestion rates of larvae reduced the abundance in the posterior sections. The anterior sections of the stomachs had alternating layers of either copepods or capelin larvae and krill. This shows that the individual predators switched between feeding on either copepods or on krill and capelin larvae. A total of 549 capelin larvae was found in 440 fish stomachs. Capelin larvae were found in 20% of the sandeel stomachs, and 24, 34 and 62% of the stomachs from the three herring stations, respectively. Many of the predator stomachs contained more than five larvae, and up to 28 larvae were found in one herring stomach. The herring appeared to prey selectively on the largest capelin larvae, and the number of larvae per stomach was much higher in stomachs where krill had recently been eaten than where copepods were dominant. The predation intensity measured in this study is much higher than reported in earlier studies, and it is suggested that the predators were feeding using a searching image, and that the concentrations of alternative prey (copepods and krill) varied around a switching threshold.     

Comparative feeding strategies of yellowfin tuna around St Helena and adjacent seamounts of the South Atlantic Ocean

Yellowfin tuna are the mainstay of the traditional tuna fisheries in St Helena waters, but there is limited knowledge of their ecology and feeding behaviour in the area. In this study yellowfin tuna stomach contents were used to assess spatio-temporal changes in feeding strategy and consider the role of tuna in the local ecosystem. Comparisons of the feeding spectra of yellowfin tuna between inshore regions of St Helena and oceanic seamounts demonstrated that in both areas the species was largely piscivorous. In inshore waters yellowfin consumed more neritic fauna, including significant numbers of crab megalopa, whereas around seamounts the diet included a greater diversity of epi- and mesopelagic fish and squids. The most important fish prey species in inshore waters was the St Helena butterflyfish Chaetodon sanctahelenae, and around seamounts was the pufferfish Lagocephalus lagocephalus. Results indicate that the diet spectrum of yellowfin tuna in St Helena waters is relatively similar to those of conspecifics living in waters with relatively low productivity, with strategies indicative of food-poor ecosystems. The availability of coastal fauna may make areas around islands and seamounts more attractive for feeding aggregations of yellowfin tuna, compared to the open ocean. The relatively unselective feeding of yellowfin tuna means that stomachs can provide valuable data on the species diversity, particularly in remote areas with limited opportunities for dedicated research expeditions.     

The forgotten feeding ground: patterns in seasonal and depth-specific food intake of adult cod Gadus morhua in the western Baltic Sea

This study presents the diet composition of western Baltic cod Gadus morhua based on 3150 stomachs sampled year-round between 2016 and 2017 using angling, gillnetting and bottom trawling, which enhanced the spatio-temporal coverage of cod habitats. Cod diet composition in shallow areas (<20 m depth) was dominated by benthic invertebrate species, mainly the common shore crab Carcinus maneas. Compared to historic diet data from the 1960s and 1980s (limited to depth >20 m), the contribution of herring Clupea harengus decreased and round goby Neogobius melanostomus occurred as a new prey species. Statistical modelling revealed significant relationships between diet composition, catch depth, fish length and season. Generalized additive modelling identified a negative relationship between catch depth and stomach content weight, suggesting reduced food intake in winter when cod use deeper areas for spawning and during peak summer when cod tend to avoid high water temperatures. The results of this study highlight the importance of shallow coastal areas as major feeding habitats of adult cod in the western Baltic Sea, which were previously unknown because samples were restricted to deeper trawlable areas. The results strongly suggest that historic stomach analyses overestimated the role of forage fish and underestimated the role of invertebrate prey. Eventually, this study shows the importance of a comprehensive habitat coverage for unbiased stomach sampling programmes to provide a more reliable estimation of top predator diet, a key information for food web analyses and multispecies models.     

Feeding habits of horse mackerel, Trachurus trachurus (Linneaus, 1758), from the central Adriatic Sea

The feeding habits of the horse mackerel, Trachurus trachurus, from the central Adriatic Sea were investigated with respect to fish size and season. Stomach contents of 1200 specimens 12.9–37.6 cm total length (TL) taken at monthly intervals (January–December 1995) were analyzed. Of the total number of stomachs examined, 597 were empty (49.7%). This percentage varied significantly with season; the maximum number of empty stomachs was observed during January (80%) and February (82%) and a minimum number observed during July (15%) and August (19%). A total of 30 different prey species belonging to five major groups: Crustacea (Euphausiacea, Mysidacea, Decapoda), Cephalopoda, and Teleostei was identified in the stomach contents. Euphausiids were the most important ingested prey in all seasons as well as in small‐ and medium‐size classes (<28 cm TL). Two species of euphausiids, Nyctiphanes couchii (%IRI = 36.0) and Euphausia krohni (%IRI = 12.9), were the most frequent prey. The proportion of teleosts in stomach contents increased with the increasing size class of horse mackerel and was 15.5% IRI in >30 cm TL individuals. The mean weight of stomach contents increased significantly for fish larger than 24.1 cm TL, while the mean number of prey items significantly declined in the large‐size classes.     

Food and feeding habits of the African catfish Clarias gariepinus (Burchell, 1822) (Pisces: Clariidae) in Lake Koka,Ethiopia

The food and feeding habits of Clarias gariepinus (Burchell, 1822) were studied in Lake Koka, Ethiopia, in May 2011 (dry month) and September 2011 (wet month). Three hundred and thirty eight nonempty stomach samples were analysed using frequency of occurrence and volumetric methods of analysis. Detritus, insects, macrophytes, zooplankton and fish were the dominant food categories by occurrence, and they occurred in 79.6%, 63.6%, 63.0%, 56.2% and 15.4% of the stomachs, respectively. Volumetrically, the above food items comprised 24.3%, 14.1%, 14.5%, 19.3% and 21.8% of the total volume of food categories. Phytoplankton and gastropods were of low importance, and they occurred in 28.7% and 8.6% of the stomachs, respectively. Their volumetric contributions were 3.2% and 2.8% of the total volume of food categories. During the dry month, fish, zooplankton, insects and detritus were important food categories, while during the wet month detritus, macrophytes, insects and gastropods were important. Smaller catfish diets were dominated by detritus, macrophytes and insects, whereas larger catfish shifted to fish and zooplankton. Based on the results, C. gariepinus was found to be omnivorous in its feeding habits in Lake Koka.     

Feeding ecology of central Baltic Sea herring and sprat

A unique dataset of stomach contents sampled between 1977 and 1999 in the central Baltic Sea was used to perform a comprehensive study of the feeding ecology of Central Baltic herring Clupea harengus and sprat Sprattus sprattus . Both fish species were mainly preying upon calanoid copepods with Pseudocalanus sp. dominating the diet of herring, whereas sprat generally preferred Temora longicornis . Sprat preyed upon older copepodite stages, indicating size‐selective particulate feeding, whereas herring additionally fed on smaller copepodite stages, indicating occasional low food supply inducing filter‐feeding. Additional food sources other than copepods were mysids in winter and autumn for medium to large herring, as well as cladocerans for sprat in spring and summer, determined by the seasonal occurrence of these plankton species. Seasonally the highest feeding activity of both fishes species occurred in spring and summer, the main reproductive periods of calanoid copepods. The most important food item for both predators in spring was Pseudocalanus sp. In summer sprat switched to T. longicornis and Acartia spp. Since the late 1970s, the total stomach fullness decreased and the fraction of empty stomachs increased. In parallel the amount of Pseudocalanus sp. in the diets of both fish species decreased. Further, a considerable dietary overlap between both species in spring indicated considerable competition for food resources, especially due to an enlarged sprat stock. The results of this study support the hypothesis that growth reductions observed in Baltic herring and sprat are due to combination of a change in food availability and an increase in density‐dependent competition.     

Diet composition of the juvenile chub mackerel (Scomber japonicus) in the Aegean Sea (Izmir Bay, Turkey)

Stomach contents of 296 juvenile chub mackerel (Scomber japonicus Houttuyn, 1782) specimens were examined based on samplings carried out in Izmir Bay (Aegean Sea, Turkey) during 2001. In terms of percentage weight (W%), fishes were the main food during summer and autumn. Thaliaceans (Salpa sp.) constituted the most important food source in winter, whereas planktonic crustaceans (Amphipoda, Copepoda) were the main prey during spring. According to the Bray–Curtis similarity index, diet of the chub mackerel was 64.1% similar during the summer, winter and autumn seasons.     

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.     

The response of gastric pH and motility to fasting and feeding in free swimming blacktip reef sharks, Carcharhinus melanopterus

In many fish and reptiles, gastric digestion is responsible for the complete breakdown of prey items into semi-liquid chyme. The responses of the stomach to feeding and to periods of fasting are, however, unknown for many lower vertebrates. We inserted data loggers into the stomachs of free-swimming captive adult blacktip reef sharks (Carcharhinus melanopterus) to quantify gastric pH, motility and temperature during fasting and following ingestion of food. Gastric acid secretion was continuous, even during long periods of fasting, with a mean pH of 1.66 ± 0.40 (± 1 SD) when the stomach was empty. Stomach contractions were greater following meals of mackerel than for those of squid. Gastric motility following feeding on mackerel, was positively influenced by ambient temperature, and followed a quadratic relationship with meal size, with maximum motility occurring after meals of 0.8-1.0% body weight. Diel changes in gastric motility were apparent, and were most likely caused by diel changes in ambient temperature. Gastric digestion in blacktip reef sharks is affected by both biotic and abiotic variables. We hypothesize that behavioral strategies adopted by sharks in the field may be an attempt to optimize digestion by selecting for appropriate environmental conditions.     

Spatially structured interactions between a migratory pelagic predator, the Norwegian spring-spawning herring Clupea harengus L., and its zooplankton prey

Spring-spawning herring Clupea harengus was patchily distributed over large parts of the Norwegian Sea in May 1995–2005, during the early phase of the annual feeding migration. Overall, herring tended to be found in areas with intermediate biomasses of zooplankton prey, intermediate water temperatures and relatively high salinities. Herring had more food in their stomachs in areas of relatively low water temperature and high herring abundance. Hydrographical conditions revealed that herring was feeding mainly within Atlantic water masses, and more intensely in western and northern regions of the Norwegian Sea. Zooplankton biomass was patchily distributed, and was generally higher towards the western parts of the Norwegian Sea. Here, zooplankton biomass in year _{i +1} was also negatively associated with herring spawning stock biomass in year _i , while there was no evidence for such an association in the eastern region; indicating that herring may have a geographically structured 'top–down' effect on the recruitment of its zooplankton prey. The fact that herring was not typically associated with the areas containing the greatest zooplankton biomasses may reflect that the fish had not yet reached the most profitable feeding grounds or alternatively that herring was depleting zooplankton biomass.     

The food and feeding habits of the silver pomfret, Pampus argenteus (Euphrasen), in Kuwait waters

The food habits of the silver pomfret, Pampus argenteus (Euphrasen 1788), in Kuwait waters were investigated by examining the stomach contents of 738 specimens collected from May 1996 to April 1997. Their diet consisted of a broad spectrum of food types, but Crustacea were dominant, with copepods and their eggs constituting 39 % other non-copepod Crustacea constituted 16 %. The next major food group was Bacillariophyta (21 %), followed by Mollusca (11 %), fish scales (10 %) and, finally, fish eggs and larvae (3 %). In summer the species fed on a wider variety of food items than in winter. Copepods, other non-copepod crustaceans, and molluscs tended to occur in the stomachs in higher frequencies with an increase in P. argenteus size (up to 18.5–20.4 cm), while the bacillariophytes tended to increase in stomachs at fish sizes between 22.5–24.5 cm. Analysis of monthly variations in stomach fullness indicated that feeding intensity fluctuated throughout the year, with a low during August and September, corresponding to the spawning period.     

The diet of Brycinus nurse (Pisces: Characidae) from Asa reservoir,Ilorin, Nigeria

From November 1991 to October 1993, 980 specimens of the characid Brycinus nurse were collected from Asa reservoir to examine its diet. The diet was analyzed using the frequency of occurrence, numerical and gravimetric methods. Two hundred and sixty nine (27.45%) of the stomachs examined were empty. The fish was an omnivore feeding extensively on a wide array of plant and animal food items. These consisted of 9 families, 10 genera and 10 species. The most extensively consumed plant food item was aquatic plant parts which occurred in 63.88% of the stomachs, and accounted for 6.06% by number and 12.10% by weight while the ephemeropteran, Povilla adusta was the most dominant animal food item, occurring in 50.92% of the stomachs, and accounting for 11.98% by number and 11.86% by weight. Conversely, the least consumed plant food item was Volvox occurring in 4.49% of the stomachs and accounting for 0.18% by number and 0.35% by weight, while the fish Barbus sp. was the least consumed animal food item occurring in 0.51% of the stomachs, accounting for 0.03% by number and 1.62% by weight. New food items not previously recorded such as a watermite. Aspatharia sinuata and Barbus callipterus were found in the stomach contents. The nonspecific feeding regime of the fish and its ability to utilize different food items effectively was what accounted for the prominence and wide distribution of the fish in the lake.     

Food consumption of yellowfin tuna,Thunnus albacares, in Sri Lankan waters

Synopsis A total of 4181 stomachs of yellowfin tuna,Thunnus albacares (22–164 cm FL), mainly caught by gillnets in the period from July 1984 to June 1986 were analyzed. Food consumption of yellowfin tuna in nature was determined using the values of the average stomach content, incorporating laboratory measurements of gastric evacuation rates in a feeding model. The diet of yellowfin tuna around Sri Lanka comprised a variety of macro zooplanktonic and nektonic organisms. Juveniles < 39 cm FL, are planktivores. Tuna > 40 cm FL gradually increase their consumption of fishes with increasing size. Among fish speciesAuxis sp. are the most important. The daily food consumption of juvenile yellowfin tuna (22–59 cm FL) range from 1.8 g to 136.2 g, increasing to about 284.7 g to 551.9 g for the medium size tuna (60–99 cm FL). The adult tuna (100–>130 em FL) consume around 513 g to 538.8 g of prey per day. Daily ration estimates increase from 2.1% to 5.5% of body weight with increasing size up to 70 cm FL, beyond which it decreases. The predatory pressure of yellowfin tuna on commercially important fishes and other species is discussed.     

Feeding habits of the king soldier bream,Argyrops spinifer (Forsskål, 1775) (Perciformes: Sparidae), in the northern Persian Gulf

The feeding habits of the king soldier bream Argyrops spinifer (Forsskål, 1775), were examined in the northern part of the Persian Gulf (coastal waters of Bushehr, Iran). The stomach contents of 622 specimens, 13.5–64.2 cm in total length, collected by a trawl boat from June 2010 to May 2011, were analysed to assess the influence of size, season and sex on their diet composition. Amongst the examined stomachs, 57 were empty (vacuity index = 9.2%), which varied significantly (P < 0.05) throughout the year. According to the results, this species is a generalist carnivorous feeder with a diet consisting of 52 different prey items belonging to nine main categories. Bivalvia, Gastropoda, Malacostraca, Crustacea and Teleostei were the most important prey items. Fish size and gender showed no significant effects on diet composition. Moreover, in the main food catagories no seasonal variations were detected.     

Invasion of the Atlantic rock crab (Cancer irroratus) at high latitudes

With the increase in global oceanic trade the establishment of non-indigenous marine organisms has become a major environmental and economic problem worldwide. Recently, the Atlantic rock crab (Cancer irroratus) was reported in Icelandic waters, Eastern North Atlantic. This is the first record of this relatively large crab species outside its natural range, i.e. the east coast of North America. The crab was most likely transferred to Iceland as larvae in ballast water and has successfully established a reproducing population in Icelandic waters. The species is distributed along the southwestern- and western-coast of Iceland. Adult specimens are now common in Faxaflói Bay, Southwest Iceland, but with sporadic occurrences in western and northwestern Icelandic waters. The green crab (Carcinus maenas) and the spider crab (Hyas araneus) are the only native brachyuran decapod species commonly found in its new habitat, but despite its recent colonization the rock crab was the most abundant brachyuran in the areas studied in southwest Iceland. Egg bearing rock crab and green crab females were found from June to October, while egg bearing spider crab females were seen from July to December. In Southwest Iceland both rock crab and green crab larvae were abundant in mid-summer but rare in both spring and autumn, which is opposite of what was observed for the spider crab. The size and abundance of adult crabs, their reproductive conditions, and occurrence of all larval stages, indicate that the Atlantic rock crab has successfully colonized Iceland.     

The Food and Feeding Chronology of Yellow Perch (Perca flavescens) in Lake Washington

The food habits of yellow perch (Perca flavescens) in Lake Washington, Seattle, U.S.A. were studied over a period of nine months. Altogether 549 yellow perch were examined for the stomach contents. The daily activity pattern of the yellow perch in Lake Washington appears to correspond roughly with the feeding pattern. The maximum numbers of perch caught by the nets during 24 hour study periods correspond well with degree of fullness of stomachs noted for the different times of the day. The food consisted mainly of cottids, mysid shrimps (Acanthomysis awatchensis) and chironomid pupae and larvae. There was no difference in the food consumed between the different sexes and between the different size groups. Maximum feeding by perch during the day was observed just before dark and the daily ration was calculated to be about 1.4% of their wet body weight.