State-dependent spawning migration in Norwegian spring-spawning herring

Norwegian spring-spawning herring Clupea harengus winter in fjords of northern Norway, whereas the subsequent spawning occurs at various locations along the coast with a main bulk off the south-western coast. The distance of the southward spawning migration tends to increase with the length and condition of the fish. The costs and benefits of the southward migration were modelled in terms of fitness (number of surviving offspring). The model assumes that larvae have increasing growth and survival rates further south as they pass through warmer water during the northward drift in the coastal current. In agreement with the observed spawning distribution, optimal spawning grounds were predicted off the southwestern coast and farther south with increasing fish length and condition. The present study suggests that homing is not a successful strategy for these herring, and instead the selection of spawning grounds depends on individual internal state (length, condition), the cost of migration and the probability of larval survival.     

Utilization of endogenous reserves and effects of starvation on the health of Prochilodus lineatus (Prochilodontidae)

Prochilodus lineatus Valenciennes (curimbatá) is an important migratory Neotropical fish. It does not feed during spawning migration, and often survives after spawning. The mobilization of energy reserves and some effects of starvation (zero to eight weeks) on fish health were experimentally evaluated. Hepatic glycogen and lipids from the perivisceral fat bodies were the main reserves mobilized during the first four weeks of fasting. During this period, somatic indices and blood parameters showed that fish health was not significantly affected. However, after five weeks of food deprivation, the main energy reserves were depleted and the fish became anaemic. The loss of muscle mass indicates that protein breakdown was an important energy source after the reduction of hepatic and perivisceral reserves. Mortality was increasingly observed from seven weeks of starvation. Prior accumulation of high amounts of reserves is essential to allow movements for long distances during spawning migrations in this species.     

Humpback whale (Megaptera novaeangliae) and killer whale (Orcinus orca) feeding aggregations for foraging on herring (Clupea harengus) in Northern Norway

Aggregations of predators on food patches have been documented for both terrestrial and marine animals. Here, we documented for the first time, and investigated, non-predatory aggregations occurring between humpback whales (Megaptera novaeangliae) and killer whales (Orcinus orca) while feeding on wintering Norwegian spring spawning herring (Clupea harengus) in Andfjord, northern Norway. Observational data were collected during 109 opportunistic surveys through three seasons 2013–2016. Killer whales were observed feeding on 59 occasions, with one to three humpback whales involved in 47 of these feeding events (79.7%), and there was an increased probability of finding feeding humpback whales when feeding killer whales also were observed. With killer whales identified as the initiating species in 94.4% of the feeding aggregations for which the first species was known, and with humpback whales joining and feeding on the fish ball afterwards, we suggest that humpback whales may benefit more from these aggregations than the opposite.     

The role of ``conservatism' in herring migrations

Herring (Clupea harengus) migrationstend to remain constant over periods of severalyears or even decades, despite environmentalvariation. When a migration pattern is changed,apparently in response to an environmentalstimulus, the change in migratory behaviorsometimes lasts longer than the environmentalstimulus itself. This paper reviews evidencethat the response of herring to environmentalchange is restrained by conservatism. Herringapparently develop certain migratory habitsduring the early stage of adult life and tendto adhere to these habits for life, even whenthe environment changes. Two elements ofconservatism are distinguished: the formationof habits within one generation, and thetransfer of habits between generations. Anumber of case studies on North Sea herring andNorwegian spring spawning herring are reviewedin order to find evidence for habit formationand tradition in the migrations of thesepopulations. By eliminating the possibilitythat homing to spawning, wintering and feedinggrounds is due to innate behavior orenvironmental constraints, it is shown that thereturn of these fish to the same areas eachyear is most likely a form of learned behavior.In many cases, new year-classes adopt the samemigration pattern as their predecessors, whichis explained by the social transfer of habitsfrom old herring to young ones. A change ofmigration pattern is usually initiated by arecruiting year-class that lacks the ``guidance'of older herring at the time it has to startits first migration. This may be the effect ofa special environmental condition that affectsthe distribution of the recruiting age groupmore than that of the adult stock component, andthereby results in a separation of the twostock components. After the recruitingyear-class has adopted the new migration route,it tends to repeat it in subsequent years. Thenew migration thus becomes habitual. Subsequentgenerations may copy the migration and continuethis behavior even after the originalenvironmental cause has disappeared.     

Energy requirements of migrating Great White Pelicans Pelecanus onocrotalus

The Great White Pelican Pelecanus anocrotalus is the largest migrating bird in Israel and is an endangered species. The Palearctic populations of the Great White Pelican breed in eastern Europe and Asia and most of them pass through the ‘bottleneck’ of Israel to wintering grounds in Africa. Natural feeding sites for pelicans have diminished during recent decades due to human activities, and sites of extensive aquaculture have become the favourite feeding places for wintering and migrating Great White Pelicans. The fish industry has reported a significant impact on fish yield and the conflict between pelicans and fishermen has escalated so that hundreds of pelicans have died in recent years from shooting or accidental electrocution. We approached this management problem by studying the energy requirements of the Great White Pelican during migration and while wintering in Israel, under different feeding regimes (fish or chicks) and in different seasons, in captivity. The results show that a captive bird consumes 1.1 kg of fish per day. The basal metabolic rate and apparent metabolized energy of the Great White Pelican are both higher than predicted from allometric equations. Energetic demands were quite stable on both diets (fish and chicks) and during both seasons (winter and summer). The fat deposits of migrating pelicans averaged 313.5 g compared with 480 g in wintering birds (3.4% and 5.4% of body mass, respectively). Based on these fat contents and on the measured daily energy consumption, we calculated that birds that do not feed in Israel can fly only up to 1620 km from Israel southward and could not cover the distance to their likely wintering grounds in the Sudd area in southern Sudan. However, birds that replenish their fuel reserves could fly up to 2460 km and hence could reach this area. Therefore, we conclude that Great White Pelicans must feed in Israel during the autumn migration in order to complete their journey to Africa. One solution to the conflict between pelicans and fishermen could be to combine deterrents preventing pelicans from feeding in fish‐ponds with the provision of attractive alternative reservoirs, to ensure regular food supplies during autumn.     

Production of the 57 kDa major surface antigen by a non-agglutinating strain of the fish pathogen Renibacterium salmoninarum

Both the prevalence and tissue titer of viral hemorrhagic septicemia virus (VHSV) increased in Pacific herring Clupea pallasi following their introduction into net pens (pounds) used in the closed pound spawn-on-kelp (SOK) fishery in Prince William Sound, Alaska. VHSV was also found in water samples from inside and outside the SOK pounds after herring had been confined for several days; however, water samples taken near wild free-ranging, spawning herring either failed to test positive or tested weakly positive for virus. Little or no virus was found in tissue samples from free-ranging, spawning herring captured from the vicinity of the pounds, nor did the prevalence of VHSV increase following spawning as it did in impounded herring. The data indicated that increased prevalences of VHSV were correlated with confinement of herring for the closed pound SOK fishery and that infection was spread within the pounds through waterborne exposure to virus particles originating from impounded fish. In addition, pounds containing predominantly young fish had higher prevalences of VHSV, suggesting that older fish may be partially immune, perhaps as a result of previous infection with the virus. Operation of SOK pounds during spawning seasons in which young herring predominate may amplify the disease and possibly exacerbate the population fluctuations observed in wild herring stocks.     

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.     

Sexual dimorphism in the energetics of reproduction and growth of North Sea plaice, Pleuronectes platessa L.

The chemical composition and energy content of North Sea plaice during the spawning period were examined in mature males and females and in immature fish, to study differences in the allocation of energy over reproduction and somatic growth between the sexes. At the beginning of the spawning period mature males and females had equal dry weights of lipid that were 70% higher than in immatures. Protein content in mature males was equal to that in immatures but was 23 % higher in mature females. Immature males and females did not differ in chemical composition. At the end of the spawning period, spent and immature fish had equal lipid contents, but protein content in spent females was 10% lower than in spent males, and 17% lower than in immatures. Gross energy content of the body decreased by 44% (65·2 to 36·3 J cm^-3) in mature females, 27% (55·0 to 40·OJ cm^-3) in mature males, and 9% (48·7 to 44·2J cm^-3) in immatures. Energy content of plaice eggs was estimated at 6·60 kJ per 1000 eggs. Reproductive investment was estimated from the energy loss during the spawning period and included the energy of sex products and spawning metabolism. Somatic growth comprised the annual increase in energy content of fish. The pattern of energy allocation over reproduction and somatic growth differed between males and females. Males started their reproduction at a smaller length and a younger age and allocated a higher proportion of the available energy into reproduction than females. Available energy resources for somatic growth and reproduction (surplus production) were equal between the sexes up to a length of about 30 cm. Beyond this length male surplus production levelled off whereas female surplus production continued to increase. The differences in surplus production and the allocation patterns are discussed. For female plaice the energy allocated into egg production was estimated as between 48 and 64% of the total amount of energy lost during spawning. The remaining energy is used for metabolism during the spawning period, yielding an estimate of the metabolic rate of mature females of between 6·4 and 9·1 kJ day^-1. A maximum estimate of the metabolic rate of mature males was 7·4 kJ day^-1.     

Climate change and condition of herring (Clupea harengus) explain long-term trends in extent of skipped reproduction

It is commonly assumed that iteroparous fish, once mature, normally reproduce in all consecutive seasons. Recent work has suggested, however, that in Norwegian spring-spawning herring—a population that undertakes extensive spawning migrations—almost one in two adults may skip their second spawning migration. Why should herring not return to spawn the year after first spawning, but instead wait an extra year? For herring, participation in distant, energetically costly, and risky spawning migrations will only pay off in terms of fitness if individuals are sufficiently large, and in sufficient condition, to both successfully migrate and spawn. Changes in the environment and individual condition should therefore affect the likelihood of skipped spawning. This paper describes long-term changes in the extent to which the second reproductive season is skipped in this herring population. These are shown to be linked to the size and condition of herring as first-time spawners, and to climatic factors possibly related to food availability. The findings corroborate the hypothesis that skipped reproduction results from trade-offs between current and future reproduction, growth and survival.     

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.     

Travel schedules to the high arctic: barnacle geese trade-off the timing of migration with accumulation of fat deposits

On their way from the wintering area to the breeding grounds in Spitsbergen, barnacle geese Branta leucopsis stage on islands off the coast of Norway. The aim of this study was to describe when the geese migrate in relation to the body stores deposited and explore questions related to the concept of optimal migration schedules and on the possible mechanisms involved. We estimated fat stores by repeated assessments of the abdominal profile index of individually marked females throughout staging. Reproductive success was derived from observations of the same individuals later in the annual cycle. Females arriving late, or with low fat stores at arrival, achieved higher fat deposition rates, probably by spending more time foraging. But they were unable to match final fat scores of birds that arrived earlier or with larger fat stores. Reproductive success was correlated with the timing of migration and individuals departing at intermediate dates achieved highest success. The exact date of peak reproductive success depended on the size of fat stores accumulated, such that low-quality birds (depositing less fat) benefited most from an early departure to the breeding grounds. Observations in the breeding colonies showed that these birds did not initiate a nest earlier but they spent a longer time in Spitsbergen before settling. The length of stay in Norway was close to the prediction derived from an optimisation model relating spring events to eventual breeding success. Poorest performing birds stayed longer than expected, perhaps depositing more fat to avoid the risk of starvation. Two possible mechanisms of the timing of migration were contrasted and it seemed that the geese departed for migration as soon as they were unable to accumulate any more fat stores.     

Comparison of fatty acid profiles of spawning and non-spawning Pacific herring, Clupea harengus pallasi

Crude lipid and fatty acid composition from liver, intestine, roe, milt and flesh of spawning and non-spawning Pacific herring (Clupea harengus pallasi) were examined to determine the relative effects of spawning on the nutritional value of herring. Depletion of lipid due to spawning condition was significant (P < 0.01) in all organ tissues and flesh of spawning herring. The lipid content ranged from an average of 1.9 to 3.4% (wet weight basis) in different organ tissues of spawning herring, to 10.5 to 16% in non-spawning fish. The fatty acid profile exhibited many differences in the relative distribution of individual fatty acids among organ tissues and between the two fish groups. Oleic acid (C18:1n-9), a major monounsaturated fatty acid (MUFA) found in all tissue lipids, decreased significantly (P < 0.01) in spawning fish. The two monoenes, C20:1n-9 and C22:1n-11, occurred at high concentrations in the flesh but at only minor proportion in the digestive organs and gonads. Spawning herring also had significantly (P < 0.01) higher polyunsaturated fatty acids (PUFA) content in the organ tissues, particularly in the milt and ovary, with docosahexaenoic acid (C22:6n-3, DHA) having the greatest proportion. Among the n-6 fatty acids, only C18:2n-6 and C20:4n-6 occurred at notable amounts and were present in higher proportions in spawning fish. We concluded that although relatively higher n-3 fatty acid content was found in the organ lipids of spawning herring, they are not an energy-dense prey food source due to the fact that both flesh and gonads contain a very low amount of lipid.     

Discrimination of Baltic herring populations with respect to Anisakis simplex larvae infection

Samples of herring Clupea harengus in the Baltic were collected during the spawning season in western, middle and eastern waters off the Polish coast. Two samples were taken per area, several morphometric characters were measured and some meristic characters (vertebrae and gill rakers) were counted. Infection with Anisakis simplex larvae, total length, age and maturity stage were recorded. Univariate and multivariate statistical tests were used to compare the samples, discriminant analysis was applied to separate herring populations and the fraction of herring populations in each area was estimated. The results obtained indicated that infected fish represented a mixed group of western and central Baltic herring. In the first samples collected from each area, the share of the western population decreased from 40% in the western to 18% in the middle and 13% in the eastern coasts. The share of western herring was higher in the second samples collected in the same area and decreased from 91% in the western coast to 14 and 32% in the middle and eastern ones, respectively.     

Spatial and temporal variations in parasite prevalence and infracommunity structure in herring (Clupea harengus L.) caught to the west of the British Isles and in the North and Baltic Seas: implications for fisheries science

Herring Clupea harengus L. viscera were examined for endoparasitic infections as part of a multidisciplinary stock identification project (WESTHER, EU Contract no. Q5RS-2002-01 056) which applied a range of stock discrimination techniques to the same individual fishes to obtain comparable results for multivariate analysis. Spawning and non-spawning adults, and juvenile herring were caught, over 3 years, by commercial and research vessels from numerous locations to the west of the UK and Ireland, along with control samples of spawning fish from the eastern Baltic Sea, and juveniles from sites in the eastern and western North Sea, and the north of Norway. The metacercariae of two renicolid digeneans (Cercaria pythionike and Cercaria doricha), one larval nematode (Anisakis simplex s.s.) and one larval cestode (Lacistorhynchus tenuis) were selected as tag species. Results were compared with those from herring collected between 1973 and 1982, which suggested remarkable stability in the parasite fauna of herring in the study area. These species were used to compare the parasite infracommunities of spawning herring. A significant variation in infracommunity structure was observed between different spawning grounds. These results suggest that the parasite fauna of herring are spatially variable but remain temporally stable in both the short and long term. Significant differences in prevalence and abundance of infections and comparisons of parasite infracommunity enabled the separation of putative herring stocks west of the British Isles. Distinctive patterns of parasite infection in two different spawning groups off the north coast of Scotland suggest that this area is occupied by two spawning populations, one recruiting from the west of Scotland, the other from outside this area, and most likely from the eastern North Sea. The distribution patterns of L. tenuis, C. doricha and C. pythionike suggest the potential for fish that spawn in three distinct International Council for the Exploration of the Seas (ICES) management units to be present in mixed aggregations found over the Malin Shelf, with significant implications for management in this area.     

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

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

The link between migration, the reproductive cycle and condition of Sardinella aurita off Mauritania, north-west Africa

The annual migration pattern of round sardinella Sardinella aurita up and down the north-western African coast between 12° N (Senegal) and 22° N (western Sahara) was shown to be associated with spawning activity and a distinct seasonality in fish condition, based on monthly sampling from commercial catches (2000–2003). Some S. aurita were found to spawn throughout the entire year, but a peak in spawning existed during the summer (June to September). The spawning cycle is apparent from seasonality in maturity stages, but is also demonstrated by the increase in gonad mass and fat content of the fish in springtime, the period preceding spawning. During the months after spawning, although feeding is maintained, the physical condition of the fish collapses, and fat content rapidly declines.     

The onset of spring and timing of migration in two arctic nesting goose populations: the pink‐footed goose Anser bachyrhynchus and the barnacle goose Branta leucopsis

An earlier onset of spring has been recorded for many parts of Eurasia in recent decades. This has consequences for migratory species, both in changing the conditions encountered by individuals on reaching migratory sites and in affecting cues regulating the timing of migration where decisions to migrate are influenced by local environmental variables. Here we examine the timing of spring migration for two arctic goose populations, the pink‐footed goose Anser brachyrhynchus (during 1990–2003) and barnacle goose Branta leucopsis (during 1982–2003), which both breed on Svalbard. The satellite‐derived Normalised Difference Vegetation Index (NDVI) was used to express the onset of spring at their wintering and spring staging sites. Pink‐footed geese use several sites during spring migration, ranging from the southernmost wintering areas in Belgium to two spring staging areas in Norway, and distances between sites used along the flyway are relatively short. There was a positive correlation in the onset of spring between neighbouring sites, and the geese migrated earlier in early springs. Barnacle geese, on the other hand, have a long overseas crossing from their wintering grounds in Britain to spring staging areas in Norway. Although spring advanced in both regions, there was no corresponding correlation in the timing of onset of spring between their wintering and spring staging sites, and little evidence for barnacle geese migrating earlier over the whole study period. Hence, where geese can use spring conditions at one site as an indicator of the conditions they might encounter at the next, they have responded quickly to the advancement of spring, whereas in a situation where they cannot predict, they have not yet responded, despite the advancement of spring in the spring staging area.     

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.     

Seasonal changes in the chemical composition of fish body and nutritional value of the muscle of the pollan (Coregonus pollan Thompson) from Lough Neagh,Northern Ireland

Studies on the chemical composition of adult pollan were carried out monthly from January to December 1977. Sex, weight of scaled fillet, and gonads were examined in 184 fish. Chemical composition which includes water, fat and ash content was analysed in the whole fish, muscle and gonads. This paper attempts to account for the fat and protein flow within fish of both sexes during gonad maturation. Extreme differences between males and females have been recorded in the fat deposition process during summer and spawning period. In spite of our fragmentary knowledge of spawning behaviour in pollan, an explanation is offered for these phenomena. The changes in fat content of female gonads were followed closely throughout the year and the cost of egg production was estimated as a percentage of the amount of fat deposited in the body. Since a similar growth pattern was found for both sexes of pollan, differences in feeding intensity might account for the difference in energy expenditure. Amino acid content was analysed in muscle and gonads and an Essential Amino Acid Index (EAAI) was calculated. There were some changes in EAAI which depended on sex and season. Finally, a new method for evaluation of muscle production expressed as fat and protein is used to determine the most suitable time for fish harvesting.     

Variation in the calorific value and total energy content of the lesser sandeel (Ammodytes marinus) and other fish preyed on by seabirds

Wet and dry calorific values (energy densities) and total energy content of lesser sandeel Ammodytes marinus , sprat Sprattus sprattus , Atlantic herring Clupea harengus , whiting Merlangius merlangus , saithe Pollachius virens and cod Gadus morhua , were measured. Calorific values varied both within and between species. Larger fish tended to have higher calorific values than small ones, particularly when considered in terms of wet weight, but there was considerable temporal variation. The calorific values and body weights of sandeels larger than 10 cm showed marked seasonal trends and in consequence the total energy content of a sandeel of given length in summer is approximately double the spring value. The calorific values of herring and sprat also varied from month to month but seasonal cycles were less obvious. Whiting varied least. Both calorific values and total energy content of individual sprat were very variable in summer (the spawning season of this species). Because there is so much intraspecific variation, care must be taken when assessing the relative merits (in terms of energy) of different species as prey. The practical difficulties of obtaining reliable data on wet calorific values are discussed and the use of dry calorific values and dry weight/length relationships is recommended.