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.     

Abundance,structure, growth and origin of inshore clupeid populations of the west coast of Scotland

The abundance, structure, growth, and origin of clupeid populations in inshore waters of the west coast of Scotland were studied from April 1970 to October 1972. Clupeid populations in the area comprise mainly young fish. 0-group autumn-spawned herring, probably of Minch origin, move into the area about April and spring-spawned ones (Clyde origin) about June. The timing and the body length at which each group arrives in the area during the different years is the same. After the initial immigration, the distribution of both 0-group clupeids becomes localized.Herring populations in the sea-lochs and associated inshore waters are mainly 0-group fish, which are replaced each year by a new incoming brood. The sprat populations of the sea-lochs are dominated by the 0-group; in the more ‘open’ areas the populations comprise older individuals. Year-class distribution in the ‘open’ areas resemble that of the commercial fishery.The rate of increase of 0-group autumn-spawned herring is 3.68 mm/week and that of spring-spawned 0-group herring is 2.83 mm/week. The curves of growth of 0-group sprats of the 1970 and 1971 year-classes are different; the rate of increase in length, however, averages 3.55 mm/week for both year-classes and the differences are not significant.In sprats, after their first year of life a rapid increase in length takes place in the spring. This increase is thought to enable the majority of the population to reach the minimum size (88–90 mm) for initial gonadal maturation and thereby make them capable of reproducing in their second year of life.     

The food of herring, Clupea harengus, in the North Sea, 1983–1986

The stomach contents of 5762 herring caught during North Sea trawl surveys in February 1983, February 1984 and February, May and August 1986 were analysed. The principal prey organisms were the copepods Calanus finmarchicus and Temora longicaudata , but Euphausiacea and the post-larval stages of Ammodytes spp. and clupeoids contributed a large percentage of the weight. Fish eggs were eaten, chiefly those of plaice, Pleuronectes platessa , but not in large numbers.     

Local and large-scale climatic variables as predictors of the breeding numbers of endangered Lesser Black-backed Gulls on the Norwegian Coast

In the 1970s and 1980s, the nominate subspecies of the Lesser Black-backed Gull (Larus fuscus fuscus) showed a dramatic drop in breeding numbers on the Norwegian Coast, and in 2000, the population in some colonies was only 10–20% of the population in 1980. This decline has been attributed to the collapse in the stock of Norwegian spring spawning herring (Clupea harengus). In this study, we examined whether local climate (sea and air temperatures), winter NAO (North Atlantic Oscilliation), and the year-class strength and size of 0-group herring could predict the relative changes in breeding numbers between years, mainly after the population collapse. Breeding birds were counted in 19 of the years between 1980 and 2007 in an archipelago on the coast of Helgeland, northern Norway. The best model predicting changes in breeding numbers for the period between 1980 and 2005 (for which data on 0-group herring was available) included mean local air temperature in winter (January–March) and winter NAO, explaining 57% of the variation between years, while the other factors had little effect. When also adding the years 2006–2007 (no herring data), the best model included only mean air temperature in winter, explaining 41% of the variation. In conclusion, the high positive correlation between breeding numbers and climatic factors probably resulted from a higher availability of important fish prey after mild winters, for which 0-group herring presently may only account for a limited proportion. However, this prey might have been of much more importance prior to the population decline.     

Interannual variations in condition indices of larval Norwegian spring-spawning herring

The sea water temperature off the Norwegian coast was lower in 1994 than in 1992 and 1993. RNA/DNA ratios of Norwegian spring spawning herring increased with increasing larval dry weight all years, except for larvae sampled south of 62°N in 1994. The RNA/DNA ratios indicated that each year, only a small portion ( 0.7%) of the larvae were starving. RNA/DNA ratios and temperature were negatively correlated in 1992, but in other years no significant correlations were found. Residuals of In RNA v . In DNA and In W v. L _s were poorly correlated in all years, but residuals of In RNA v. In DNA and In DNA v. In W were negatively correlated in all years. Principal component analysis showed that the RNA/DNA ratio and DNA (% of weight) were correlated with different axes. Abundance data for herring at the early larval and 0-group stages in 1992–1994 indicated higher mortality in 1994 compared with the other years. The data do not indicate that average larval condition was poorer in 1994 than in other years. However, the variability in larval condition was higher in 1994 than in other years, and the condition of later larval stages was relatively lower in 1994 than in other years.     

DIETS OF FIN, SEI, AND SPERM WHALES IN BRITISH COLUMBIA: AN ANALYSIS OF COMMERCIAL WHALING RECORDS, 1963–1967

Diets of fin ( Balaenoptera physalus ), sei ( Balaenoptera borealis ), and sperm whales ( Physeter macrocephalus ) were estimated from the stomach contents of individuals killed along the British Columbia coast from 1963 to 1967. The dominant prey types of fin whales were euphausiids, with minor contributions from copepods and fish. Sei whale stomachs contained primarily copepods in three years, whereas euphausiids or a variety of fish dominated the diet in the other two years. Sperm whales consumed primarily North Pacific giant squid ( Moroteuthis robusta ), but secondary prey differed between males and females. Female sperm whales frequently consumed ragfish ( Icosteus spp.) and other fish, whereas the male diet also contained rockfish ( Sebastes spp.). The high abundance of euphausiids along the British Columbia coast likely contributed to the presence of a summer resident population of fin whales. The high abundance of large copepods farther north probably influenced the migration of sei whales through the offshore waters of British Columbia. Sperm whale stomach contents differed by sex reflecting location and possibly breeding behaviors.     

Ecological significance of 0-group fish in the Barents Sea ecosystem

Investigations into the 0-group fish in the Barents Sea have been carried out since 1965, with the goal of estimating the abundance of 0-group fish. 0-group abundance indices have been used in the assessment of the recruitment level and in recruitment variability studies. However, the ecological importance of the 0-group fish in the Barents Sea has been less studied. Although 0-group capelin, herring, cod and haddock are widely distributed in the Barents Sea, the central area seems to be the most important, accounting for approximately 50–80% of the annual biomass. The total biomass of the four most abundant 0-group fish species can be up to 3.3 million tonnes, with an average of 1.3 million tonnes (1993–2009). Wide distribution and high biomass of pelagically distributed 0-group fish make these fishes an important element in the energy transport between different trophic levels and different geographical areas, having a critical impact on the entire Barents Sea ecosystem. In recent years, capelin have shown a pronounced northward shift in biomass distribution, and several successive strong year classes occurred during warm temperature conditions. Cod biomasses were unexpectedly low during warm years and were positively correlated with spawning stock biomass, while the correlation with temperature was not significant. Haddock and herring show, as expected, increasing biomass with increased temperature when the spawning stock is at a sufficiently high level.     

The microbial flora of herring fillets after storage in carbon dioxide, nitrogen or air at 2°C

Herring fillets from the Baltic Sea were stored in glass vessels in air, nitrogen or carbon dioxide (CO₂) at 2°C and the microbial development was studied. The microbiological shelf-life of the herring (the time to reach 10⁷ organisms/g) was prolonged by a factor of 3.5 in CO₂ as compared to air. The corresponding factor in nitrogen was 1.5. The microflora of fresh and spoiled herring was classified. The initial microflora was dominated by coryneforms, Flavobacterium spp., Moraxella -like organisms and Pseudomonas spp. The spoilage-flora in air (after 9 d) was dominated by Pseudomonas spp. and Moraxella -like organisms, and in nitrogen (14 d) Enterobacteriaceae, Vibrionaceae and Lactobacillus spp. were dominant. Homofermentative Lactobacillus spp. were the only organisms isolated from fillets stored in CO₂ (28 d). It was concluded that storing fresh fish in pure CO₂ at low refrigeration temperatures is a method with industrial potential. The method (1) improves the microbiological stability and (2) reduces the microbiological health hazards of the fish.     

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.     

Effects of temperature, prey type and prey size on gastric evacuation in small cod and whiting

Gastric lavage was used to investigate the effects of temperature, prey type and prey size on gastric evacuation in small cod Gadus morhua and whiting Merlangius merlangus . The fish were fed to satiation and subsequently the stomach contents were sampled to determine the rate at which food was evacuated. Satiation meal size was positively related to temperature and differed between prey types. The gastric evacuation rate (GER) also tended to increase with temperature and varied with prey type. GER at temperatures of 11.3–12.7° C averaged 1.5–1.7 times higher than at 6–9.5° C. There was no significant difference (P>0.05) in the evacuation of lugworm Arenicola marina , sandeel Ammodytes spp., and herring Clupea harengus , but the GER of brown shrimp Crangon vulgaris was much slower (P<0.05). No significant changes in GER were observed when fish were fed on three different size groups of either herring or brown shrimp. In whiting, there was no significant difference in the GER of individual herring or brown shrimp when they were fed as single species meals or incorporated in meals containing a mixture of prey species.     

OCCURRENCE, PHOTO-IDENTIFICATION AND PREY OF PACIFIC WHITE-SIDED DOLPHINS (LAGENORHYNCUS OBLIQUIDENS) IN THE BROUGHTON ARCHIPELAGO, CANADA 1984–1998

This study summarizes occurrence of Pacific white-sided dolphins ( Lagenorhyncus obliquidens ) in the Broughton Archipelago on the west coast of Canada, from October 1984 through December 1998. Dolphins were detected on 472 d. The annual percent of total occurrence rose from 0.4% in 1984 to 19% in 1994 and then declined to 2% in 1998. Seasonal occurrence peaked from 1 October through January. Dolphin group size ranged from 2 to 1,000; the most common range was 11-50. While unreported for the Broughton Archipelago prior to 1984, the species is represented by teeth distributed throughout the past 2,000 yr of First Nations midden sediment, suggesting sporadic long-term occurrence. Increased water temperature from the 1937–1984 mean of 8.6°C, to the 1985–1998 mean of 9.3°C and increased abundance of two fish populations in the study area are considered potential factors in the recent increase in occurrence. Of the 675 naturally marked dolphins that were photo-identified, 214 were resighted. A pair of dolphins was photographed swimming in tandem, fourteen months apart. Tight groups, of five or fewer extensively scarred dolphins with extremely falcate dorsal fins were seen within every aggregation of over 50 animals, suggesting the existence of all-male associations. Prey species were collected from 25 encounters with feeding dolphins; they included herring ( Clupea harengus ), capelin ( Mallotus villosus ), and Pacific sardine ( Sardinops sagax ). Predation on eulachon ( Thaleichthys pacificus ) is suspected. Unreported for the Broughton Archipelago, the capelin sampled in this study may belong to the Bering Sea population. Pacific sardines returned to commercial viability on the British Columbia coast in 1997 after a 60-yr population collapse. Dolphin frequency of occurrence declined following introduction of underwater acoustic deterrent devices into the study area.     

Temporal variability in marine feeding of sympatric Arctic charr and sea trout

The marine feeding pattern of anadromous brown trout (sea trout) Salmo trutta and Arctic charr Salvelinus alpinus was studied during June to August in 1992–1993 and 2000–2004 in a fjord in northern Norway. In general sea trout fed proportionally more on fishes than on crustaceans and insects (81, 1 and 18% by mass, respectively) by comparison with Arctic charr (52, 25 and 22% by mass, respectively). Herring Clupea harengus dominated the total fish diet of both species, but the Arctic charr also fed significantly on gadoids and sandlance Ammodytes spp. While sea trout became virtually all piscivorous at fork lengths ( L _F) ≥250 mm, the Arctic charr was ≥400 mm L _F before shifting totally to a fish diet. Despite annual variation in diet and forage ratios, there was a clear shift in diet from 1992–1993 to 2000–2004. Sandlance and different crustaceans constituted most of the diet during the initial period with a shift towards gadoids and especially herring during the latter period. This shift seemed to be associated with a high abundance of herring larvae during the latter sampling period, indicating a preferential selection on herring when present, particularly by sea trout. Furthermore, an index indicated dietary overlap in years with intensive feeding on herring of both species, and usually differences in the trophic ecology during years feeding mostly on other prey species. In combination, it was hypothesized that the two species reflect the type of marine prey present within a fjord system over time, and therefore provide an index of variation in the production and biological diversity of their potential prey within fjords.     

Meta-ecosystems and biological energy transport from ocean to coast: the ecological importance of herring migration

Ecosystems are not closed, but receive resource subsidies from other ecosystems. Energy, material and organisms are moved between systems by physical vectors, but migrating animals also transport resources between systems. We report on large scale energy transport from ocean to coast by a migrating fish population, the Norwegian spring-spawning (NSS) herring Clupea harengus. We observe a rapid body mass increase during parts of the annual, oceanic feeding migration and we use a bioenergetics model to quantify energy consumption. The model predicts strong seasonal variation in food consumption with a marked peak in late May to July. The copepod Calanus finmarchicus is the most important prey and 23×10⁶ tonnes (wet weight) of C. finmarchicus is consumed annually. The annual consumption-biomass ratio is 5.2. During the feeding migration 17% of consumed energy is converted to body mass. The biomass transported to the coast and left as reproductive output is estimated from gonad weight and is about 1.3×10⁶ tonnes for the current population. This transport is to our knowledge the world’s largest flux of energy caused by a single population. We demonstrate marked temporal variation in transport during the last century and discuss the effects of NSS herring in the ocean, as a major consumer, and at the coast, where eggs and larvae are important for coastal predators. In particular, we suggest that the rapid decline of lobster Homarus gammarus landings in Western Norway during the 1960s was related to the collapse of NSS herring. We also discuss spatial variation in energy transport caused by changed migration patterns. Both climate and fisheries probably triggered historical changes in the migration patterns of NSS herring. New migration routes emerge at the level of individuals, which in turn determines where resources are gathered and delivered, and therefore, how meta-ecosystems function.     

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.     

Observations of the biology and parasites of postsmolt Atlantic salmon, Salmo salar, from the Norwegian sea

The main food items of postsmolt Atlantic salmon caught in the Norwegian sea during summer 1991 were Parathemisto spp. and herring ( Chipca harengus ). The most prevalent parasite species were Chloromyxion sp., Apatemon sp., H. adunction, A simplex and I. salmonis .     

Multi-mesh gillnets to estimate species composition and catch per unit effort of fish in a small water body in Zambia

A standard method comprising multi-mesh monofilament nylon gillnets and depth stratification, the Drottningholm method, was employed in test fishing of a small Zambian reservoir (32 ha) on eight occasions over 1 year. The catch per unit effort (cpue) from the 0–3 and 3–6 m zone was timated with benthic and pelagic nets, while in the 6–12 m zone only benthic nets were used. Results include species caught and cpue with coefficients of variation and confidence intervals, for net types and depth zones. A total of 11 species were caught, of which Burbus paludinosus , B. marequensis , Laheo cylindricus , and Oreochromis sp. dominated. Coefficients of variation did not indicate a preferable season for test fishing. During the cold season some species were not caught. The applicability of the method in estimation of species composition and cpue is discussed.     

Structure and specific features of formation of aggregations of deepwater redfish <Emphasis Type="Italic">Sebastes mentella</Emphasis> (Scorpaenidae) in the Norwegian Sea pelagial

Results of studies of the structure and specific features of formation of aggregations of deepwater redfish Sebastes mentella in the Norwegian Sea pelagial are provided. Data on the biological characteristics of S. mentella from different regions of the Norwegian Sea pelagial do not allow us to consider its aggregations as separate populations. On the basis of comparative analysis of the structure of aggregations in the Norwegian Sea and of population parameters and specific features of the life cycle of S. mentella in different parts of the Atlantic Ocean, its complex intraspecies organization at sites of overlapping of two largest populations—North Atlantic and Norwegian-Barents Sea-was established. The formation of pelagic aggregations in the water area of the Norwegian Sea occurs due to migration of maturing and mature individuals from the adjoining water areas. In the southern areas of the Norwegian Sea, dominant recruitment comes from the pelagial of the northeastern part of the Irminger Sea, which is promoted by the direction of streams of the North-Atlantic Current; it is not excluded either that fish can migrate to the southern part of the sea from the slopes of the Faeroes, Faeroes-Icelandic threshold, and the western coast of Norway. Migration to the northern areas of the sea is mainly performed by rapidly maturing fish from the rearing region from the western slopes of the Barents Sea and Spitsbergen Archipelago. The presence of seasonal migrations hinders the formation in the pelagial of perch isolated groups and promotes their migration.     

Investigations into the biology of the garfish, Belone belone (L.), in Swedish waters

Garfish, Belone belone (L.), were collected from commercial fishermen between May and September of 1987 from around the Swedish coast. Fish belonging to the year classes of 1983, 1984 and 1985 dominated the catch, with very few fish from the year classes of 1980, 1981 and 1982. Maturity stages and condition were investigated, and estimates of fecundity calculated; these ranged from 2193 to 10804 eggs per ripe female. Examination of gut contents showed fish, especially juvenile clupeids and gasterosteids, as well as isopods and amphipods, to be the most important components of their diet. Garfish populations from both the Baltic and the west coasts were parasitized by copepods of the genus Caligus . A study of meristic characteristics showed a significant difference between the number of vertebrae in fish from the Baltic and those captured on the west coast of Sweden.     

Infection of wild fishes by the parasitic copepod Caligus elongatus on the south east coast of Norway

Natural Caligus elongatus Nordmann infections of wild coastal fishes on the Norwegian south east coast were monitored at various times of the year from 2002 to 2004. The prevalence for all coastal fish (n = 4427) pooled was 15%, and there were great differences between fish species and seasons. Lumpfish Cyclopterus lumpus L. spawners were the most infected fish, with a prevalence of 61% and a median intensity of 4 lice fish(-1), whereas gadids had a mean prevalence of 19% and a median infection of 1 to 2 lice fish(-1). Sea trout Salmo trutta L. and herring Clupea harengus L. carried C. elongatus at prevalence values of 29 and 21%, respectively. The results were compared with infection data for immature North Sea lumpfish. Lumpfish spawners caught on the coast in March to April had fewer lice than North Sea lumpfish in July. Spawners carried mostly adult lice, as did coastal fish hosts in May to June. The low development rates of lice at low spring temperatures and new genetic data suggest that the May to June adult lice could not have been offspring of the March to April lice, indicating transfer of adult lice to coastal fish. Most coastal fish species appeared to acquire new C. elongatus infections between May to June and September. The relatively high numbers of chalimii on North Sea lumpfish suggest that offshore fish sustain an oceanic population of this louse species.     

What is out there: diversity in feeding of gentoo penguins (Pygoscelis papua) around the Falkland Islands (Southwest Atlantic)

During austral spring 2000, the diet of Gentoo penguins, Pygoscelis papua, was studied and compared to prey availability, assessed by trawl and plankton surveys, in waters adjacent to a number of colonies along the coast of West Falkland. There was good agreement in size and abundance of key prey species in the penguin diet and from the plankton and trawl surveys. The diet of birds breeding adjacent to shallow sheltered waters was dominated by demersal species, in particular crustaceans, Munida spp. whilst birds breeding adjacent to deeper exposed waters generally preyed upon pelagic species of fish, Falkland herring Sprattus fuegensis and squid, Moroteuthis ingens. Gentoo penguins generally foraged at depths of up to 30 m, relatively close to the coast, and illustrated a certain degree of selection of prey species where possible, thus providing evidence that they are not fully opportunistic predators.