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.     

The influence of feeding behaviour on growth of Atlantic cod (Gadus morhua,Linnaeus, 1758) in the North Sea

The objective of this study was to resolve key mechanisms driving individual growth patterns of Atlantic cod (Gadus morhua). Growth dynamics were analysed by linking growth patterns with stomach content composition and environmental temperature. Samples were collected in August/September of the years 2009, 2010 and 2011 in the north‐eastern part of the central North Sea. Prey selection was assessed by identification of individual prey items in the stomach content to species. Ten feeding groups were identified consisting of individuals with one prey type dominating their stomach contents (≥75% by mass), of which six were used for growth analyses: “Sandeel”, “Clupeids”, “Norway pout”, “Flatfishes”, “Crustaceans” and “Brittle stars”. For each group, growth patterns were estimated based on measures of otolith growth increments. The stomach contents showed that cod as a species are opportunistic in their prey selection, but at the same time indicated that the total, broad feeding niche width of the population is dominated by individual diet specialization and that many individuals temporally show a preference for a particular prey type. The contribution of invertebrates and particularly crustaceans decreased with increasing cod size, whereas that of fish and predominantly herring increased. Prey type had a significant effect on growth, while temperature had no effect. Slowest growth was observed in the cod group preying on sandeel, while cod preying on Norway pout showed the fastest growth. No significant difference was observed between groups preying on brittle stars, crustaceans, flatfishes and herring. Growth in the year before capture did however not differ between any of these groups. Across sampling years, growth chronology patterns were similar but not significantly influenced by temperature.     

Stomach evacuation rate in the planktivorous Antarctic fish Pagothenia borchgrevinki

Summary Specimens of the Antarctic fish Pagothenia borchgrevinki were collected from just below the sea-ice in McMurdo Sound over the period 5–16 November 1987. The stomach contents of 54 freshly captured fish were analysed and showed very strong similarities with an earlier, more extensive, dietary analysis conducted at the same location (Foster et al. 1987). There was as much variation in stomach contents from fish within a school, as there was between schools, contrary to the expectation that fish within a school would have shared a more common feeding history. Fish were maintained in holding tanks for up to 48 h after capture; groups were sacrificed at 6 h intervals and the stomach contents analysed to determine evacuation rates. Evacuation rates of total stomach contents and individual prey species were fitted with an exponential model of decay. The exponential time constant for decay of stomach contents was 32 h (Table 2), while time constants for individual prey species ranged from 12 h for small prey species to 49 h for the larger crustacea. The feeding rate of P. borchgrevinki on important prey species was calculated from the above data (Table 2). Low feeding rates, and the low numbers of P. borchgrevinki argue against removal by predation as being the explanation for the observed reduction in plankton numbers seen in shallow waters (Foster 1989).     

Intra-annual variation in feeding of Atlantic cod Gadus morhua: the importance of ephemeral prey bursts

Seasonal prey bursts are important for the life cycles and energy budgets of many predators. This study documents the diet and, especially, the importance of the ephemeral occurrence of capelin as prey for Atlantic cod (Gadus morhua) in Godthaabsfjord, west Greenland, over an annual cycle. The cod showed clear differences in diet composition on the 11 sampling dates resulting in a spring–summer, late summer–autumn and winter cluster. Moreover, a single sampling date, 12 May, was defined by cod gorge feeding on spawning capelin, which led to average stomach contents 4.3 times higher than the average for the remaining sampling dates. Changes in nitrogen stable isotope values from 22 April to 7 July in cod liver and muscle tissue were used to calculate the consumption of capelin. Based on this, the consumption of capelin varied between 538 and 658 g wet weight for a 1.3 kg cod. Using published consumption/biomass estimates and observed growth rates, the capelin intake corresponds to 10.1%–33.3% of the annual food consumption and accounts for 28.1%–34.5% of the annual growth of the cod. The present study documents the omnivorous feeding mode of Atlantic cod but highlights the utilization and importance of ephemeral prey bursts for the annual energy budget of the cod. It is hypothesized that access to capelin is critical for the postspawning recovery of Godthaabsfjord cod.     

Diet of harbor porpoises along the Dutch coast: A combined stable isotope and stomach contents approach

High stranding frequency of porpoises, Phocoena phocoena, along the Dutch coast since 2006 has led to increased interest in the ecology of porpoises in the North Sea. Stranded porpoises were collected along the Dutch coast (2006–2008) and their diet was assessed through stomach content and stable isotope analysis (δ¹³C and δ¹⁵N) of porpoise muscle and prey. Stable isotope analysis (SIAR) was used to estimate the contribution of prey species to the porpoises' diet. This was compared to prey composition from stomach contents, to analyze differences between long‐ and short‐term diet. According to stomach contents, 90.5% of the diet consisted of gobies, whiting, lesser sandeel, herring, cod, and sprat. Stable isotope analysis revealed that 70‐83% of the diet consisted of poor cod, mackerel, greater sandeel, lesser sandeel, sprat, and gobies, highlighting a higher importance of pelagic, schooling species in the porpoises' diet compared to stomach contents. This could be due to prey distribution as well as differences in behavior of porpoises and prey between the coastal zone and offshore waters. This study supports the need for multi‐method approaches. Future ecological and fishery impact assessment studies and management decisions for porpoise conservation should acknowledge this difference between the long‐ and short‐term diet.     

The feeding relationships of the larger fish species in a Scottish sea loch

Patterns within a fish assemblage have been examined from catches obtained over a period of 17 months using fixed nets and rod and line sampling. A total of 19 species were caught with summer samples dominated by dab and saithe and winter samples by whiting. Cod were caught regularly throughout the year. Catches at the study site of dab and whiting occurred mainly at night but for cod and saithe catches occurred throughout the day. There was no indication ofa crepuscular or tidal rhythm for any species. Differences between our own observations and those from other published studies lead us to conclude that no species-specific generalizations about feeding periodicities can be made. Feeding patterns are more likely to be determined by site-specific features of particular habitats and their associated prey and predator assemblages.
Examination of stomach contents data using a variety of methods indicates that conclusions are highly dependent on the form of the resource matrix used. However, our analyses indicate that during summer cod and dab feed on a much narrower range of prey than saithe and that there is significant overlap in their diets if a resource matrix based on percentage occurrence of prey species is used. Nevertheless, most diet overlap values for the species examined were low (between 0.1 and 0.2) suggesting that competition for food is unlikely. To date, no experiments have been performed to test the hypothesis that cod and dab compete for food.     

Diet diversity among five co-existing fish species in a tropical river: integration of dietary and stable isotope data

Diets were estimated from stomach contents and the MixSIR model using stable isotope values for five co-existing and abundant benthic fishes in relation to potential prey from a riffle habitat in a tropical river in eastern Thailand. Collectively, aquatic insects were of greatest dietary importance based on stomach contents and, except for one fish species, predicted from the MixSIR. The most prominent functional feeding insect groups in fish diets were collector-filterers and scrapers and, to a lesser extent, predators. MixSIR predicted shrimp to be the most important single dietary constituent for all fishes in contrast to stomach contents, which indicated they are a major item for only one species. MixSIR predicted plant material to be more important in fish diets than stomach contents where the composition of detritus was a concern. Differences in temporal feeding schedules, prey availability, species adaptations and others are important in understanding diet and in the construction of food webs. Stable isotope and dietary analysis provides a more accurate assessment of the food web structure and dynamics of tropical river ecosystems than either method alone.     

Feeding studies take guts – critical review and recommendations of methods for stomach contents analysis in fish

Studies on the feeding ecology of fish are essential for exploring and contrasting trophic interactions and population and community dynamics within and among aquatic ecosystems. In this respect, many different methods have been adopted for the analysis of fish stomach contents. No consensus has, however, been reached for a standardised methodology despite that for several decades there has been an ongoing debate about which methodical approaches that should be preferred. Here, we critically review and scrutinise methods, addressing their strengths and weaknesses and emphasising inherent problems and possible pitfalls in their use. Although our critical assessment reveals that no completely ideal approach exists, appropriate and reliable procedures can be adopted through careful considerations and implementation. In particular, we advocate that different objectives require different methodical approaches and the choice of method should therefore be closely linked to the research questions that are addressed. For a standardisation of methods, we recommend a combination of the relative-fullness and presence–absence methods as the optimal approach for the commonly applied feeding studies addressing relative dietary composition in terms of prey diversity and abundance. Additionally, we recommend the gravimetric method for objectives related to the quantification of food consumption rates and the numerical method for prey selection studies. DNA-based dietary analysis provides a new and promising complementary approach to visual examination of stomach contents, although some technical challenges still exist. The suggested method standardisation facilitates comparisons across species, ecosystems and time and will enhance the applicability and benefits of fish feeding studies in trophic ecology research.     

A new approach to graphical analysis of feeding strategy from stomach contents data—modification of the Costello (1990) method

A modification of the graphical Costello method is proposed for the analysis of stomach contents data. The new method allows prey importance, feeding strategy and the interand intra-individual components of niche width to be explored using graphical presentation. The analysis is based on a two-dimensional representation of prey-specific abundance and frequency of occurrence of the different prey types in the diet. The paper describes the new method and the parameters therein, and also present some examples of the utilization of the method. The method may be particularly well-suited for the examination of predictions made from optimal foraging, competition and niche theories.     

Stable isotope analysis as a tool to monitor dietary trends in little penguins Eudyptula minor

Long‐term dietary monitoring of seabirds can be used to relate population fluctuations to at‐sea events. Stomach flushing is a conventional dietary monitoring technique, but has a number of disadvantages. Stable isotope analysis (SIA) is a less invasive method that provides unbiased dietary information over a longer period. We evaluated stable isotope analysis as a potential tool for monitoring long‐term little penguin Eudyptula minor diet. We determined diet composition during the chick feeding stage using stomach flushing and SIA at three separate colonies, using spatial variation in diet as a surrogate for potential temporal variation. Bayesian isotopic mixing models were generated for blood and feathers to evaluate their ability to discriminate broad‐scale (fish, squid, crustaceans) and fine‐scale (individual prey species) diet composition. Differences in stable carbon and nitrogen isotope ratios were found between colonies: broad‐scale isotopic mixing models predicted different proportional contributions of broad taxa (fish, cephalopod, crustacean) to diet than was indicated by stomach samples, reflecting the bias incurred by one‐off stomach contents analysis. Fine‐scale isotopic mixing models predicted proportional contributions of prey items with less certainty. Blood isotopic mixing models had narrower confidence intervals than models for feathers, but trends in δ¹⁵N for feathers mirrored those for blood. Our results suggest that relying on stomach contents analysis to detect shifts in prey consumption in little penguins could be very misleading, resulting in a less‐than‐complete idea of total prey consumption. SIA of little penguin tissues could be used to monitor dietary shifts across dissimilar taxa that may affect population numbers, but would fail to detect shifts between fish species.     

Summer feeding ecology of harp seals (Phoca groenlandica) in relation to arctic cod (Boreogadus saida) in the Canadian high arctic

Summary The diet and feeding behaviour of harp seals, Phoca groenlandica, was examined in two high arctic locations. Fish otoliths were used to evaluate dietary composition and aspects of the population dynamics of the major prey species, arctic cod, Boreogadus saida. Harp seals, primarily adults, arrive in the high arctic in mid to late June and depart by early October. Their migration is undertaken specifically for feeding. Harp seals feed intensively on arctic cod, often occurring in dense multispecies aggregations in late summer. The average weight of harp seal stomach contents was high; glutted individuals contained as much as 6% of their body weight in food. Although arctic cod declined in abundance between years, size of cod ingested was similar between areas and years, and overlapped completely with cod taken by other marine mammals. Age/size segregation of arctic cod may account for poor representation of fish <3 years old in the seal diet. Widespread reproductive failure of arctic cod could have a profound influence on the energy balance of adult harp seals since there does not appear to be an alternate food source of equivalent energy value and abundance in arctic waters. Increasing harp seal populations will likely result in increased competition with a host of arctic cod predators, particularly ringed seals.     

Chick starvation in yellow‐eyed penguins: Evidence for poor diet quality and selective provisioning of chicks from conventional diet analysis and stable isotopes

The link between poor reproductive success and diet was investigated in yellow‐eyed penguins Megadyptes antipodes, by assessing diet at two localities separated by about 30 km: the north coast of Stewart Island where breeding success is low (0.38–0.67 chicks per pair in recent years), and Codfish Island where breeding success is higher (0.96–1.51 chicks per pair), and relating this to published data from South Island localities, where average breeding success was 1.1 chicks per pair. Diet composition, meal sizes and energetic content of meals and prey were determined from stomach contents, and stable isotope analyses of chick down, fledgling feathers and adult blood provided information on diet throughout the fledging period. The high proportion of stomachs that were empty or lacked diagnostic remains reduced sample size considerably, and variability between samples reduced the power to detect significant differences in meal size, proportions of empty stomachs and prey diversity of meals. Energetic content of Stewart Island meals was less than Codfish Island meals, and there was a non‐significant trend for smaller meal sizes and reduced prey diversity among Stewart Island samples. Both localities had lower prey diversity and smaller meals than South Island penguins. Blue cod Parapercis colias accounted for 99% of prey biomass in Stewart Island and 70% in Codfish Island stomach samples, where 27% of prey biomass was opalfish Hemerocoetes monopterygius. Isotopic mixing models carried out on larger sample sizes indicated that opalfish comprised a large proportion of the diet at both locations, with adults selectively provisioning chicks with opalfish while feeding mainly on blue cod themselves. We suggest the large blue cod consumed by Codfish Island and Stewart Island penguins, larger than those consumed by South Island penguins, is difficult to transfer to chicks by regurgitation. Oyster dredging around Stewart Island may have reduced the availability and abundance of alternative prey to Stewart Island penguins.     

Feeding ecology of dominant groundfish in the northern Bering Sea

We investigated current diets of the six most abundant benthic fish in the northern Bering Sea. Our objective was to explore feeding strategies and potential competition with other top predators as ecosystem changes occur in the northern Bering Sea ecosystem. Our approach used stomach content data collected from field sampling during spring 2006 and 2007. Calanoid copepods and ampeliscid amphipods were important prey of Arctic cod (Boreogadus saida) but in different proportions depending upon fish size, feeding location, and local environmental conditions. Snailfish (Liparidae) occupied a broad niche and fed on a variety of benthic amphipods. Arctic alligatorfish (Ulcina olrikii) and Arctic staghorn sculpin (Gymnocanthus tricuspis) consumed ampeliscid amphipods predominantly. Shorthorn sculpin (Myoxocephalus scorpius) had a less-diverse diet, with snow crab (Chionoecetes opilio) most important by weight. Finally, all Bering flounder (Hippoglossoides robustus) sampled had empty stomachs. Our results indicate that ampeliscid amphipods, which have high biomass in the central region of the northern Bering Sea, are the most important prey for the dominant groundfish in the Chirikov Basin. Generally, all dominant benthic fish in the northern Bering Sea had narrow feeding niches, except snailfish. High diet overlap was found among many of the fish species, including Arctic cod and snailfish, snailfish and Arctic alligatorfish, and Arctic alligatorfish and Arctic staghorn sculpin. These findings are consistent with a relatively short food chain for benthic fish that are for the most part specialized feeders with narrow preferences for food and may be affected by changes in benthic prey distributions.     

Feeding habits of harp seals (Phoca groenlandica) during early summer and autumn in the northern Barents Sea

The feeding habits of harp seals (Phoca groenlandica) in the Barents Sea were examined in studies conducted during June 1991, September 1990 and 1991, and October 1992. Analyses of stomach and intestinal contents were carried out and concurrent estimates of prey abundance were made using trawl gear. Harp seals appeared to feed at low intensity in the pack ice belt during the first half of June. There was little potential prey in the water column, but prawns (Pandalus borealis), capelin (Mallotus villosus) and polar cod (Boreogadus saida) were abundant close to the bottom. In September, the seals sampled in the northern pack ice areas of the Barents Sea fed on the pelagic amphipod Parathemisto libellula, krill (Thysanoessa spp.), prawns and, to a lesser extent, on fish species such as polar cod, sculpins (Cottidae) and snailfish (Liparidae). Trawling revealed that large quantities of Parathemisto libellala were present in the upper layers of the water column. Fish, mainly capelin and polar cod, were less abundant and occurred in deeper waters. In mid-October, the diet of seals in the northern Barents Sea consisted mainly of amphipods (Parathemisto sp.). Later in October, when increasing pack ice cover forced the harp seals to move south, the diet seemed to change from amphipods to fish prey, predominantly capelin and polar cod.     

A comparison of three methods to investigate the diet of breeding double-crested cormorants (Phalacrocorax auritus) in the Beaver Archipelago, northern Lake Michigan

In order to understand the role of waterbirds in aquatic food webs it is important to first get an accurate depiction of their diet. Three methods of dietary assessment (pellets, regurgitate and stomach contents) are compared here for breeding double-crested cormorants (Phalacrocorax auritus) of the Beaver Archipelago, northern Lake Michigan. By numerical frequency (percent number), each method yielded different depictions of the diet. However, in terms of presence and absence (percent frequency) of possible prey types, stomach content data did agree with both pellets and regurgitate data. However, differences were noted between regurgitate and pellets. In terms of biomass measured (percent biomass) in regurgitate and stomachs, data gathered agreed. In essence, pellets underestimate the importance of alewife (Alosa pseudoharengus) and overestimate the importance of crayfish (Orconectes sp.) in the diet when compared to both regurgitate and stomach analysis. The non-lethal method of regurgitate collection and analysis appears most practical in assessing cormorant diet in this system. In combination with information on avian foraging ecology and prey populations, these data may be used to investigate the relationships among cormorants and their prey, and lead to a better understanding of Great Lake food web dynamics.     

From mass of body elements to fish biomass: a direct method to quantify food intake of fish eating birds

The winter diet of great crested grebe (Podiceps cristatus) was studied in the major lakes of Insubria region, N Italy, in 2000–2003 by analysing stomach and oesophagus contents of birds found dead. Winter diet of cormorant (Phalacrocorax carbo) was studied using pellet analysis. Diet was expressed in terms of numerical abundance, frequency and biomass of prey. A detailed methodology is described of how to prepare a reference collection of otoliths, pharyngeal bones and chewing pads (key-elements), essential for species-specific identification of fish remains in stomachs and pellets, and to estimate ingested fish biomass. Correlations between mass (mg) of key-elements and mass of corresponding fish were significant, and exponential regression models were used to calculate biomass of each single prey item directly from the mass of its key-elements. Advantages of this method are compared with the more traditional two step approach in which size of elements is used to estimate fish length, followed by calculating fresh mass from fish length. Cyprinids formed the largest part of the diet for both predators. Great crested grebes selected small prey, feeding mainly on bleak (Alburnus alburnus alborella), while diet composition of cormorants varied significantly in relation to prey availability. Handling editor: K. Martens     

Distribution and diet of 0-group cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) and haddock (<Emphasis Type="Italic">Melanogrammus aeglefinus</Emphasis>) in the Barents Sea in relation to food availability and temperature

Distribution of 0-group cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) in August–September 2005 and 2006 was mainly restricted to the Atlantic waters of the western and central areas of the Barents Sea. The main distribution of 0-group fish overlapped largely with areas of high biomass (>7 gm⁻² dry weight) of zooplankton. The copepod Calanus finmarchicus and krill Thysanoessa inermis, which are dominant zooplankton species in both Atlantic and boreal waters of the Barents Sea, were the main prey of 0-group cod and haddock. The main distribution, feeding areas and prey of 0-group cod and haddock overlapped, implying that competition for food may occur between the two species. However, though their diet coincided to a certain degree, haddock seems to prefer smaller and less mobile prey, such as Limacina and appendicularians. As 0-group fish increased in size, there seems to be a shift in diet, from small copepods and towards larger prey such as krill and fish. Overall, a largely pelagic feeding behaviour of 0-group cod and haddock was evident from this study.     

Fish eggs and larvae in the diet of herring Clupea harengus membras Linnaeus, 1758 and the sprat Sprattus sprattus balticus (Schneider, 1904) (Clupeidae) in the Southeastern Baltic Sea

The qualitative and quantitative composition of fish eggs and larvae in the stomach contents of Baltic herring and sprat was examined. The occurrence of fish eggs and larvae in the diet of herring was markedly higher, compared to sprat. Eggs of four and larvae of nine fish species were found in the herring stomachs. The prey species composition of the herring and sprat diets was related to the ichthyoplankton composition and to the degree of overlapping of the prey and predator distributions. Sprat eggs were the predominant prey in the herring diet in May and sprat larvae prevailed in August. The daily ration estimates indicate that the sprat larvae consumption by herring in August exceeded the sprat egg consumption in May by many times. Cod eggs in the herring stomachs were only found at the optimal oxygen content and salinity for the cod eggs in the near-bottom water layer.     

Feeding patterns of the dominant benthic and demersal fish community in a temperate estuary

The diet and feeding relationships of the 12 most abundant benthic and demersal fish species of the Mondego Estuary, Portugal, were studied between June 2003 and May 2006. Fishes were caught during the night using a 2 m beam trawl. The stomach contents were analysed for prey identification, counting and weighing. According to prey importance in diets, three main feeding guilds were identified: (1) invertebrate feeders, (2) invertebrate and fish feeders and (3) plankton and invertebrate feeders. Besides these main feeding guilds, some fishes also presented fractions of algae and zooplankton in their stomach contents. The most abundant prey items were macroinvertebrates, with several polychaetes ( Nephtys spp., Capitellidae, Spionidae and Eunicidae), Corophium spp. and Crangon crangon among the dominant prey. Pomatoschistus spp. were the most preyed on fishes. Several fish species showed a tendency for a specialized diet, but almost all also showed some degree of opportunistic feeding by preying on other food resources. High diet overlap was found between some fish species, yet exploitative competition could not be concluded.