Associations between wintering avian predators and schooling fish in the Skagerrak-Kattegat suggest reliance on predictable aggregations of herring Clupeaharengus

We investigated the distributional dynamics of piscivorous seabirds in relation to the distribution of young fish outside the breeding season in the Skagerrak and Kattegat, using synoptic bottom trawl and seabird surveys. Kittiwake Rissa tridactyla , Common Guillemot Uria aalge and Razorbill Alca torda were significantly correlated with herring Clupea harengus abundance at coarse spatial scales (>20 km). The association between the three seabird species and other abundant fish species like sprat Sprattus sprattus , whiting Merlangius merlangus and Norway pout Trisopterus esmarki was considerably weaker. In the Skagerrak and Kattegat, the location of aggregations of immature herring during winter is relatively stable and predictable. The correlation with herring and the lack of correlation with the abundant, but less stable occurrence of sprat of similar caloric value suggest that the tendency among piscivorous seabirds to depend on predictable concentrations of prey during the breeding season is a common feature during the non-breeding season as well. The results are discussed in relation to ecological and winter habitat characteristics of wintering piscivorous seabirds in the eastern North Atlantic, and in relation to the management and conservation of nursering herring in the Skagerrak and Kattegat.     

Wasp-waist interactions in the North Sea ecosystem

Background

In a “wasp-waist” ecosystem, an intermediate trophic level is expected to control the abundance of predators through a bottom-up interaction and the abundance of prey through a top-down interaction. Previous studies suggest that the North Sea is mainly governed by bottom-up interactions driven by climate perturbations. However, few studies have investigated the importance of the intermediate trophic level occupied by small pelagic fishes.

Methodology/Principal Findings

We investigated the numeric interactions among 10 species of seabirds, two species of pelagic fish and four groups of zooplankton in the North Sea using decadal-scale databases. Linear models were used to relate the time series of zooplankton and seabirds to the time series of pelagic fish. Seabirds were positively related to herring (Clupea harengus), suggesting a bottom-up interaction. Two groups of zooplankton; Calanus helgolandicus and krill were negatively related to sprat (Sprattus sprattus) and herring respectively, suggesting top-down interactions. In addition, we found positive relationships among the zooplankton groups. Para/pseudocalanus was positively related to C. helgolandicus and C. finmarchicus was positively related to krill.

Conclusion/Significance

Our results indicate that herring was important in regulating the abundance of seabirds through a bottom-up interaction and that herring and sprat were important in regulating zooplankton through top-down interactions. We suggest that the positive relationships among zooplankton groups were due to selective foraging and switching in the two clupeid fishes. Our results suggest that “wasp-waist” interactions might be more important in the North Sea than previously anticipated. Fluctuations in the populations of pelagic fish due to harvesting and depletion of their predators might accordingly have profound consequences for ecosystem dynamics through trophic cascades.     

Fatty acid composition of sprat (<Emphasis Type="Italic">Sprattus sprattus</Emphasis>) and herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>) in the Baltic Sea as potential prey for salmon (<Emphasis Type="Italic">Salmo salar</Emphasis>)

Sprat (Sprattus sprattus) and small herring (Clupea harengus) are the dominant prey fish of Atlantic salmon (Salmo salar) in the Baltic Sea. If the fatty acid (FA) proportions of sprat and herring differ, the dietary history of ascending salmon could be determined from their FA profiles. Therefore, we investigated the FA composition of several age groups of whole sprat and small herring, caught from the three main feeding areas of salmon in autumn and spring. Oleic acid (18:1n-9) was the most prevalent FA in sprat and characteristic of this species. In herring, palmitic acid (16:0) was the most common FA, but herring lipid was characterized by n-6 polyunsaturated FAs, and moreover, by palmitoleic acid (16:1n-7) and vaccenic acid (18:1n-7). Due to the higher lipid content of sprat, the concentrations of all other FAs, excluding these, were higher in sprat than in herring. The concentration of docosahexaenoic acid (DHA, 22:6n-3) increased with an increase in the lipid content and was consequently highest in the youngest specimens, being in young sprat almost double that of young herring, and 2.6 times higher in the sprat biomass than in that of herring. As a result of a decrease in the DHA concentration with age, the ratio thiamine/DHA increased with respect to age in both species, and was lower in sprat than in herring. It is concluded that an abundance of DHA in the diet of salmon most likely increases oxidative stress because of the susceptibility of DHA to peroxidation, and thus decreases thiamine resources of fasting, prespawning salmon. Because the FA composition of sprat and herring differs, and the relative abundancies of prey fish differ between the feeding areas of salmon, the feeding area of ascending salmon can most probably be derived by comparing their FA profiles.     

Wie richtet eine Flußseeschwalbenkolonie (Sterna hirundo) ihr Abwehrverhalten auf den Feinddruck durch Silbermöwen (Larus argentatus) ein?

How a Common Tern (Sterna hirundo) Colony Defends itself against Herring Gulls (Larus argentatus) The subject of this study is the anti-predator behaviour of a small common tern colony near a large herring gull colony on the island of Mellum, West Germany (Fig. 1). In 1980 the number of gulls crossing this tern colony increased during the terns' chick-stage (Fig. 4). Observed predation of tern chicks was independent of tide and time of day (3., Fig. 5). The frequency of tern reactions corresponded to the number of herring gull crossings (Fig. 5, Table 1). The terns' responses increased between morning and evening (Fig. 8). Tern up-flights and attacks increased absolutely and as a percentage, with the advance of the breeding season (Fig. 3, 4). They were positively correlated with the observed chick predation and the number of pairs with chicks, most markedly with chicks older than 5 days (Figs. 3, 4; Table 1). This increased defence was maintained by fewer pairs as, by then, many had lost their own broods (Fig. 4). As the breeding season progressed, herring gulls increasingly became the main cause of tern up-flights and the object of the attacks (Figs. 9–11). The up-flights of the whole colony, which occurred frequently and spontaneously during incubation, were observed only rarely after hatching and were almost exclusively a response to herring gulls (Figs. 10, 12). The lower herring gulls flew over the colony, the more frequently common terns flew up or attacked and the more individuals were involved in these responses (Figs. 6, 13, 14). During the breeding period, communal up-flights and attacks by terns increased as a percentage (Figs. 12, 13, 15–17). Group-attacks effected changes in the gulls' flying-routes more often than did individual attacks (Fig. 18). Despite the defence behaviour and its adaptation to the predation pressure, herring gulls often succeeded in robbing chicks. This is why the breeding success of the common tern was poor (< 0.4 chicks/nest). Possible reasons for this are discussed.     

Predicting reproductive success from hormone concentrations in the common tern (<Emphasis Type="Italic">Sterna hirundo</Emphasis>) while considering food abundance

In birds, reproductive success is mainly a function of skill or environmental conditions, but it can also be linked to hormone concentrations due to their effect on behavior and individual decisions made during reproduction. For example, a high prolactin concentration is required to express parental behaviors such as incubation or guarding and feeding the young. Corticosterone level, on the other hand, is related to energy allocation or stress and foraging or provisioning effort. In this study, we measured individual baseline prolactin and corticosterone between 2006 and 2012 in breeding common terns (Sterna hirundo) using blood-sucking bugs. Reproductive parameters as well as prey abundance on a local and a wider scale were also determined during this period. Baseline prolactin and corticosterone varied significantly between years, as did breeding success. At the individual level, prolactin was positively and corticosterone was negatively linked to herring and sprat abundance. At the population level, we also found a negative link between corticosterone and prey abundance, probably reflecting overall foraging conditions. High prolactin during incubation was mainly predictive of increased hatching success, potentially by supporting more constant incubation and nest-guarding behavior. It was also positively linked to a lesser extent with fledging success, which could indicate a high feeding rate of young. Corticosterone concentration was positively related to high breeding success, which may be due to increased foraging activity and feeding of young. In general, our study shows that baseline prolactin and corticosterone levels during incubation can predict reproductive success, despite the presence of an interval between sampling and hatching or fledging of young.     

Host-parasite relationship between colonial terns and bacteria is modified by a mutualism with a plant with antibacterial defenses

Predator–prey and host-parasite interactions and mutualisms are common and may have profound effects on ecosystems. Here we analyze the parasitic and mutualistic associations between three groups of organisms: the plant Artemisia maritima, bacteria, and a colonial seabird (the sandwich tern Sterna sandvicensis) that breeds in dense colonies covered in feces produced by both adults and chicks. A disproportionately large fraction of colonies of the sandwich tern in Denmark were located in patches covered by A. maritima. This association was specific for the densely colonial sandwich tern, but was not present for four other sympatric species of terns that breed in much less dense colonies. A. maritima reduced the abundance of pathogenic Staphylococcus on chicken eggshells in a field experiment. Recruitment by sandwich terns breeding in patches of A. maritima was 18 % higher than for sandwich terns breeding in the absence of A. maritima. A. maritima benefitted from the association with sandwich terns due to the supply of nutrients from feces and uneaten food lost by young. These findings are consistent with sandwich terns exploiting the association with A. maritima and its antimicrobial properties to improve their reproductive success, while sandwich terns and A. maritima are involved in a mutualistic interaction.     

The biology of terns nesting at Aldabra Atoll, Indian Ocean, with particular reference to breeding seasonality

Five species of tern breed on Aldabra Atoll (09° 24' S; 46° 20' E). The Caspian tern Sterna caspia and Crested tern S. bergii feed exclusively in very shallow reef/lagoon water, the Fairy tern Gygis alba and Brown noddy Anous stolidus feed out at sea, and the Black-naped tern Sterna sumatrana is intermediate in its foraging. Both of the shallow-water species lay during the south-east monsoon season, the Caspian tern from April to August and the Crested tern from June to August, but the Crested tern also lays in December and January. The remaining three species have extended laying periods largely circumscribed by the north-west monsoon season from November to March. Breeding population size of the Caspian tern is in the low tens and of the Brown noddy in the low thousands, with the other species each numbering in the hundreds. The distribution and abundance of the nine species of tern breeding within the Seychelles ( sensu lato ) vary on the different island groups in a manner interpretable in terms of depth of the surrounding waters. Systematic differences between the central Seychelles and Aldabra groups in timing of breeding by terns which feed out at sea may be associated with seasonal latitudinal movement of the divergence zone between the South Equatorial Current and the Equatorial Counter-current, acting via correlated latitudinal shifts of prey species and game-fish abundance.     

Recruitment variability in North Atlantic cod and match-mismatch dynamics

Background

Fisheries exploitation, habitat destruction, and climate are important drivers of variability in recruitment success. Understanding variability in recruitment can reveal mechanisms behind widespread decline in the abundance of key species in marine and terrestrial ecosystems. For fish populations, the match-mismatch theory hypothesizes that successful recruitment is a function of the timing and duration of larval fish abundance and prey availability. However, the underlying mechanisms of match-mismatch dynamics and the factors driving spatial differences between high and low recruitment remain poorly understood.

Methodology/Principal Findings

We used empirical observations of larval fish abundance, a mechanistic individual-based model, and a reanalysis of ocean temperature data from 1960 to 2002 to estimate the survival of larval cod (Gadus morhua). From the model, we quantified how survival rates changed during the warmest and coldest years at four important cod spawning sites in the North Atlantic. The modeled difference in survival probability was not large for any given month between cold or warm years. However, the cumulative effect of higher growth rates and survival through the entire spawning season in warm years was substantial with 308%, 385%, 154%, and 175% increases in survival for Georges Bank, Iceland, North Sea, and Lofoten cod stocks, respectively. We also found that the importance of match-mismatch dynamics generally increased with latitude.

Conclusions/Significance

Our analyses indicate that a key factor for enhancing survival is the duration of the overlap between larval and prey abundance and not the actual timing of the peak abundance. During warm years, the duration of the overlap between larval fish and their prey is prolonged due to an early onset of the spring bloom. This prolonged season enhances cumulative growth and survival, leading to a greater number of large individuals with enhanced potential for survival to recruitment.     

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.     

Plaice nurseries: effects on recruitment

The function of coastal nursery areas is discussed in relation to the variability in North Sea plaice recruitment. Since 60% of the recruitment of juveniles originates from the Wadden Sea, special attention is paid to this area. The concentration of juveniles in a restricted area seems to evoke only adverse effects: an increased risk of food limitation and hence reduced growth, and an increased vulnerability to predation. Despite these expectations, growth of most of the plaice in the Wadden Sea has always been optimal within the wide range of year-class strength observed and depends only on ambient water temperature. The same situation is indicated for some British bays, where the growth of 0-group plaice is far lower than in the Wadden Sea, because of lower temperatures. Mortality through predation seems to be relatively low in the Wadden Sea and restricted to only a short period, because of the absence of almost all potential predators. In the more open British bays higher mortalities are found, probably due to the presence of a number of predatory fish species. The low variability in the recruitment of plaice might be the combined result of optimal growth with the absence of between-year fluctuations in predator abundance in the Wadden Sea. As a result, observed mortality only depends on prey abundance and is therefore density-dependent both within one year and especially between years, reducing variations in recruitment. This suggestion is supported by the situation in more open British bays: here too, growth is maximal, but the abundance of predators shows larger fluctuations between years and, as a result, greater fluctuations in mortality are observed.     

Effects of individual life-history traits and weather on reproductive output of Black-headed Gulls Larus ridibundus breeding in the Wadden Sea, 1991–97

Capsule Annual breeding success was relatively constant and is shown to be related to clutch size and growth rate and to be positively affected by rain during egg-laying and advanced chick-rearing phase.

Aims To provide the first long-term data on breeding success of Black-headed Gulls in the Wadden Sea, to analyse its intrinsic and environmental co-variates, and to re-assess the significance of the severe winter 1995/96 on reproduction.

MethodsIn a relatively small colony at the Wadden Sea coast, clutches were selected randomly and enclosed to determine clutch size, egg biometrics, hatching and breeding success, and chick development in 1991 and from 1994 to 1997. Weather data (temperature, rain, wind) were related to life-history traits (clutch size, egg volume, chick development) and reproductive success.

Results Mean annual breeding success was 0.7 fledglings per pair. In 1996, breeding success tended to be lower and chick growth rate was significantly lower. Hatching success was lower in small clutches, chick mortality increased with decreasing chick growth rate. Rain during the egg-laying phase increased clutch size and decreased clutch predation. Rain during the early post-natal phase impaired chick growth. Rain during the phase of linear growth affected chick growth and fledging success positively and brood predation negatively.

Conclusions Breeding success of Black-headed Gulls breeding in the Wadden Sea is relatively constant between years, probably due to the use of terrestrial and marine feeding habitats. Rain may increase the availability of intertidal and terrestrial prey and thus may affect time budgets and food provisioning of parents positively. Lower breeding success in 1996 might have been caused by a relatively dry breeding season and possibly by the preceding severe winter.     

Diet overlap and resource partitioning among three forage fish species in Coorong,the largest inverse estuary in Australia

Protracted drought in southern Australia has degraded the system function of the Coorong estuary and intensified resource competition among forage fish species. The present study investigates the gut content, prey composition and dietary overlap of three forage fishes: smallmouth hardyhead (Atherinosoma microstoma), Tamar River goby (Afurcagobius tamarensis) and sandy sprat (Hyperlophus vittatus) influenced by environmental variation in the Murray Estuary and Coorong. The prey species identified in fish stomachs were dominated by crustaceans (amphipods, ostracods and harpacticoids), but nematodes and acanthocephalans were also common in all forage fishes. The diet of the sandy sprat and Tamar River goby highly overlapped (α = 0.8) in the Murray Estuary and all three forage fishes showed potential diet overlap (α ≥ 0.6) in the North Lagoon. Spatiotemporal variation of prey diversity was observed in smallmouth hardyhead while temporal variation of prey diversity was observed in sandy sprat and Tamar River goby. Overall, the prey abundance was temporally variable and predominantly regulated by salinity, pH, dissolved oxygen, water transparency and chlorophyll a in the Murray Estuary and Coorong. This study adds to our knowledge on dietary overlap and resource partitioning among small-bodied forage fishes mediated by environmental factors in the Murray Estuary and Coorong.     

VARIABILITY IN THE DIET OF HARBOR PORPOISES (PHOCOENA PHOCOENA) IN SCOTTISH WATERS 1992–2003

Interannual, seasonal, and regional variation in the diet of porpoises, Phocoena phocoena , in Scottish (UK) waters was studied using stomach contents of animals stranded between 1992 and 2003. Most samples came from the east coast (including many porpoises killed by bottlenose dolphins), with smaller numbers from the west coast and from Shetland. The most important prey types, in terms of contribution by number and mass, were whiting ( Merlangius merlangus ) and sand eels (Ammodytidae). Multivariate analysis confirmed the existence of regional, seasonal, and interannual variation in diet, as well as differences ( i. e. , biases) related to cause of death. These differences were further explored using univariate analyses. Sand eels were more important in the summer months (quarters 2 and 3) and on the east coast, whereas gadids were more important in winter and in the Shetland area. Some, but not all, observed trends in the numerical importance and size of prey taken were consistent with trends in abundance and size of fish taken during research trawl surveys. There was some evidence that porpoises <1 yr old took more gobies (Gobiidae) and shrimps than did older porpoises. Clupeids (herring Clupea harengus and sprat Sprattus sprattus ) formed a relatively small proportion of the diet, but their importance varied from year to year. Although possible methodological biases prevent firm conclusions, it appears that the importance of clupeids in porpoise diet may have decreased since the 1960s, mirroring the decline in North Sea herring abundance. The recovery of the North Sea herring stock in recent years is not as yet reflected in porpoise diet.     

Inter‐annual variation in herring,Clupea harengus,and sprat,Sprattus sprattus,condition in the central Baltic Sea: what gives the tune?

The Baltic Sea ecosystem has undergone large changes during the last two decades, including a severe reduction in cod and herring biomass but, at the same time, a large increase in sprat abundance. The lower trophic levels of the Baltic Sea also changed due to environmental fluctuations, including variations in salinity and in volume of oxygenated water. In this apparently shifting environment, the conditions of herring and sprat have undergone large inter-annual variations during the past 15–20 years. In this study, we explore how abiotic factors (i.e. salinity and temperature) and biotic factors (biomass of the copepods Pseudocalanus elongatus , Temora longicornis , Acartia spp. and of cladocerans as well as clupeid abundance) in different seasons (May and August) affect clupeid body condition. Our analyses suggest that data of zooplankton biomass and abiotic factors in August have higher predictive power than May data. Although our analysis suggests that salinity (a bottom-up process) has an effect on sprat condition, total abundance of clupeids (a top-down process) is by far the most significant predictor of both herring and sprat condition. The strong correlation between clupeid abundance and total zooplankton biomass points to food competition and to top-down control by herring and sprat on common food resources. Furthermore, clupeid condition co-varied with the changes in the weight of zooplankton in the stomachs, which further suggest food competition being the main mechanism behind the changes in clupeid condition during the last two decades. Hence, our results are not in agreement with most of the current literature that has suggested that clupeid growth is regulated by environmentally mediated bottom-up processes acting on the abundance of copepods. This is, to our knowledge, the first evidence of food resources mediated density-dependent fish growth in a large marine ecosystem.     

Species interactions in a mixed colony of common terns (Sterna hirundo) and black skimmers (Rynchops niger)

Black skimmers (Rynchops niger) frequently nest among common terns (Sterna hirundo) on coastal islands from North Carolina to New York. Behavioural interactions among terns, skimmers, and several potential avian predator species were quantified during the 1973 breeding season at Fisherman Island, Virginia. Common terns were dominant to skimmers, chasing and dive-attacking neighbouring pairs, while skimmers rarely chased terns. Despite agonistic interactions, nesting success of both species was no different whether nearest neighbours were conspecifics or heterospecifics. Common terns chased away potential avian predators much more frequently than did skimmers. Black skimmers, then, appear to derive a ‘protective cover’ benefit from nesting among the terns. Neither nest-site limitation nor food ‘information-sharing’ appear to account for the species associations.     

How many can you catch? Factors influencing the occurrence of multi-prey loading in provisioning Greater Crested Terns§

Seabirds use several methods to transport food to their chicks; most species carry food in their stomachs or crops, but some terns and auks carry prey in their bills. Terns usually only carry one prey item at a time, limiting the rate at which they can provision their chicks, and restricting their effective foraging range. However, some terns do occasionally carry multiple prey, which should offer a selective advantage, but there are very few studies investigating the factors influencing the occurrence of multi-prey loading. We investigated the occurrence of multi-prey loads in provisioning Greater Crested Terns (Swift Tern) Thalasseus bergii bergii breeding on Robben Island, South Africa. Of 24 173 loads photographed, 1.3% comprised multiple prey items. Up to 11 fish were carried at once, but most multi-prey loads contained two Anchovies Engraulis encrasicolus, the most common prey item for this population of terns. Mixed species prey were recorded for the first time in a tern. Multi-prey loads occurred more frequently during mid- and late-provisioning, presumably because large chicks can cope with multiple prey, and have higher energetic requirements than small chicks. Mean standard length of Anchovies in multi-prey prey loads was less than Anchovies in single loads, possibly suggesting terns compensate for smaller prey sizes by bringing multiple prey back to their chick. The orientation of multiple Anchovies in a tern’s bill tended to be the same, suggesting that they were captured from polarised fish schools. At least some multi-prey loads were caught in a single dive.     

Intra- and interannual variation in the diet of the Magellanic penguin (Spheniscus magellanicus) at Martillo Island,Beagle Channel

Knowledge of seabirds’ diet at each breeding site and its temporal variation is key to understanding and evaluating how changes in marine resources affect each seabird population. In this study, we determined the diet of Magellanic penguins (MP, Spheniscus magellanicus) at Martillo Island, accounting for sex, breeding stage and year. We analyzed a total of 144 stomach contents during three consecutive breeding seasons (2006–2007, 2007–2008 and 2008–2009) and stages (incubation, early and late chick-rearing). MP fed mainly on fuegian sprat (Sprattus fuegensis), which represented 75 % of the biomass consumed by birds during the entire study. The next important prey was squat lobster (Munida gregaria), followed by Patagonian squid (Loligo gahi). Both sexes consumed similar prey items. We observed variation in diet relative composition among breeding years and stages. Fuegian sprat consumption decreased throughout the years whereas squat lobster increased. Penguins consumed a higher proportion of squat lobster and Patagonian squid during the incubation stage than in the chick-rearing stages, whereas fuegian sprat was almost the only prey item consumed during the late chick-rearing stage. MPs show certain flexibility in the use of resources probably as a response to changes in prey populations. Variability in the diet among different reproductive stages could be related to changes in the distribution and abundance of their main prey near the colony during the breeding season together with changes in the energy requirements of seabirds.     

Feeding ecology of pelagic fish species in the Gulf of Riga (Baltic Sea): the importance of changes in the zooplankton community

The feeding ecology of four pelagic fish species was studied in relation to their prey availability in the Gulf of Riga (Baltic Sea) during the summer 1999-2006. The zooplankton community was dominated by the cladoceran Bosmina longispina, rotifers Keratella cochlearis and K. quadrata and the copepod Eurytemora affinis, with the highest interannual variability in abundance recorded for B. longispina. The last influenced the diet of adult sprat Sprattus sprattus, juvenile smelt Osmerus eperlanus and three-spined stickleback Gasterosteus aculeatus as these were strongly selecting for B. longispina. The fish feeding activity did not match the abundance dynamics of their preferred prey, suggesting that fishes may switch to consume other prey in case the preferred diet was limited. A considerable dietary overlap indicated high potential competition between pelagic fish species. While herring Clupea harengus membras and G. aculeatus were relying on very different food, the diets of young O. eperlanus and G. aculeatus were very similar. Interannual variability in zooplankton composition and abundance significantly affected the diet composition of fishes, but those changes were insufficient to exert a consistent influence upon fish feeding activity and total amounts of zooplankton consumed.     

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.     

Diet comparison between pelagic and demersal whiting in the North Sea

Pelagic North Sea whiting Merlangius merlangus preyed upon (pelagic) sprat and sandeel, while demersal whiting preyed upon (demersal) Norway pout and herring. Values of diet breadth were low for both feeding groups using Levin's index. Diet overlaps within layers were low ( D <0·25), while the between layer food overlap was moderate ( D =0·25–0·74) to high ( D >0·74) using Schoener's index. Selection of prey was density dependent. However, prey size also played an important role. The diet of whiting shifted from amphipods and mysids to fish with increasing predator length, and the length of prey consumed increased significantly with length of whiting. The fact that the stomach contents differed between the feeding groups demonstrates the need for a sampling design that includes both pelagic and demersal habitats when trying to quantify the diet of whiting.