Differential utilization of energy during wintering and spawning migration in Norwegian spring-spawning herring

The energy utilization in non-feeding Norwegian spring-spawning herring Clupea harengus was examined during periods of wintering in a fjord system in northern Norway and southward spawning migration (800 km distance) to the main spawning area off western Norway. At the onset of wintering the gonadosomatic index (IG) was higher and the amount of mesenteric fat was lower in male than in female herring, whereas the subsequent increase in IG and decrease in mesenteric fat occurred at a higher rate in females than in males. However, no differences were recorded between sexes in fat content (%) of whole fish at any time of the year. The absolute energy loss increased, whereas the relative energy loss decreased, with fish length. Furthermore, the absolute and relative length specific energy losses were two to three and three to four times higher, respectively, during spawning migration than during wintering. The distinct energy loss was attributed mainly to fat metabolism, whereas the storage protein only was transferred from the soma to the gonads and remained relatively constant in whole herring throughout the wintering and spawning migration. © 1999 The Fisheries Society of the British Isles     

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.     

Larval growth and drift pattern and the separation of herring spawning groups in the North Sea*

Data from two internationally co-ordinated survey programmes are used in an attempt to display the events in the early life that took place during the ‘recovery period’ of the North Sea herring Stock, 1979–1986. During the period, an increasing importance of the spawning grounds off Buchan and off Yorkshire is evident from distributions of newly hatched herring larvae. The dramatic changes in relative importance of spawning grounds are traced in the data on both late larvae (6 months old) and juveniles (18 months old). The onset of extended spawning off Buchan and off Yorkshire was followed by increase in distinct groups of late larvae of large mean length, and by enlargement of a group of medium-sized juveniles. It is hypothesized that the identified groups are interconnected. Thus, larvae spawned off Buchan and off Yorkshire are found to have growth rates superior to those attained by larvae from west of Scotland and around the Orkney/Shetland Isles, whereas the relationship is reversed in the juveniles. In the investigated period, larval drift routes exhibited the same trends from year to year, the drift of the northerly spawned larvae being the most variable. Thus, changes in the relative contributions from spawning grounds apparently influence the overall spatial distribution as well as size composition of larval and juvenile North Sea herring. The findings indicate that groups of larvae retain, to a large extent, separate distributions until metamorphosis, and point to larval drift pattern as a determinant of spawning group distinctness.     

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.     

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.     

Ticket to spawn: Combining economic and genetic data to evaluate the effect of climate and demographic structure on spawning distribution in Atlantic cod

Climate warming and harvesting affect the dynamics of species across the globe through a multitude of mechanisms, including distribution changes. In fish, migrations to and distribution on spawning grounds are likely influenced by both climate warming and harvesting. The Northeast Arctic (NEA) cod (Gadus morhua) performs seasonal migrations from its feeding grounds in the Barents Sea to spawning grounds along the Norwegian coast. The distribution of cod between the spawning grounds has historically changed at decadal scales, mainly due to variable use of the northern and southern margins of the spawning area. Based on historical landing records, two major hypotheses have been put forward to explain these changes: climate and harvesting. Climate could affect the distribution through, for example, spatial habitat shifts. Harvesting could affect the distribution through impacting the demographic structure. If demographic structure is important, theory predicts increasing spawner size with migration distance. Here, we evaluate these hypotheses with modern data from a period (2000–2016) of increasing temperature and recovering stock structure. We first analyze economic data from the Norwegian fisheries to investigate geographical differences in size of spawning fish among spawning grounds, as well as interannual differences in mean latitude of spawning in relation to changes in temperature and demographic parameters. Second, we analyze genetically determined fish sampled at the spawning grounds to unambiguously separate between migratory NEA cod and potentially smaller sized coastal cod of local origin. Our results indicate smaller spawners farther away from the feeding grounds, hence not supporting the hypothesis that harvesting is a main driver for the contemporary spawning ground distribution. We find a positive correlation between annual mean spawning latitude and temperature. In conclusion, based on contemporary data, there is more support for climate compared to harvesting in shaping spawning ground distribution in this major fish stock in the North Atlantic Ocean.     

Dead fish swimming: a review of research on the early migration and high premature mortality in adult Fraser River sockeye salmon Oncorhynchus nerka

Adult sockeye salmon Oncorhynchus nerka destined for the Fraser River, British Columbia are some of the most economically important populations but changes in the timing of their homeward migration have led to management challenges and conservation concerns. After a directed migration from the open ocean to the coast, this group historically would mill just off shore for 3-6 weeks prior to migrating up the Fraser River. This milling behaviour changed abruptly in 1995 and thereafter, decreasing to only a few days in some years (termed early migration), with dramatic consequences that have necessitated risk-averse management strategies. Early migrating fish consistently suffer extremely high mortality (exceeding 90% in some years) during freshwater migration and on spawning grounds prior to spawning. This synthesis examines multidisciplinary, collaborative research aimed at understanding what triggers early migration, why it results in high mortality, and how fisheries managers can utilize these scientific results. Tissue analyses from thousands of O. nerka captured along their migration trajectory from ocean to spawning grounds, including hundreds that were tracked with biotelemetry, have revealed that early migrants are more reproductively advanced and ill-prepared for osmoregulatory transition upon their entry into fresh water. Gene array profiles indicate that many early migrants are also immunocompromised and stressed, carrying a genomic profile consistent with a viral infection. The causes of these physiological changes are still under investigation. Early migration brings O. nerka into the river when it is 3-6° C warmer than historical norms, which for some late-run populations approaches or exceeds their critical maxima leading to the collapse of metabolic and cardiac scope, and mortality. As peak spawning dates have not changed, the surviving early migrants tend to mill in warm lakes near to spawning areas. These results in the accumulation of many more thermal units and longer exposures to freshwater diseases and parasites compared to fish that delay freshwater entry by milling in the cool ocean environment. Experiments have confirmed that thermally driven processes are a primary cause of mortality for early-entry migrants. The Fraser River late-run O. nerka early migration phenomenon illustrates the complex links that exist between salmonid physiology, behaviour and environment and the pivotal role that water temperature can have on population-specific migration survival.     

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.     

Biological features of the common fish species in Olyutorsky-Navarin region and the adjacent waters of the Bering Sea: 2. Families Macrouridae,Clupeidae, and Osmeridae

Biological features of the four common fish species, giant grenadier Albatrossia pectoralis (Macrouridae), Pacific herring Clupea pallasii (Clupeidae), Pacific rainbow smelt Osmerus mordax dentex, and Pacific capelin Mallotus villosus catervarius (Osmeridae), were studied under the 20-year dataset (1995?2015). These species inhabit the northwestern Bering Sea in the summer–autumn period and form the schoolings in the Olyutorsky-Navarin region. The size–age parameters of the fish caught by different sampling gear, as well as the peculiarities of the body length and body weight dynamics, spawning periods, spawning range, and conditions, were analyzed. The largest specimens of giant grenadier, Pacific herring, and Pacific rainbow smelt were observed in the catches performed by the bottom setline and the gill nets; the smallest fish were found in the trawl catches. The body length and body weight of Pacific herring were larger in the pelagic trawls compared to the bottom trawls; an opposite pattern was observed for the Pacific capelin. The abundant year-class in the species with short life cycle (capelin and herring) is well tracked on the longterm plots of the fish body size; this is accompanied by the decrease of their biological parameters. Herring stock covers large growing grounds; smaller body size was observed for the herring grazing in the coastal waters; young specimens dominate here.     

Scale analysis suggests frequent skipping of the second reproductive season in Atlantic herring

A common assumption in fish studies is that iteroparous fishes, once mature, normally reproduce in all consecutive seasons. An analysis of scales from Norwegian spring-spawning herring collected between 1935 and 1973 revealed a strong under-representation (47% of expected) of second-time spawners on the spawning grounds. This reduction is not satisfactorily explained by errors in scale-reading, suggesting that the second reproductive season is frequently skipped. Skipping a season may relate to trade-offs between growth, current and future reproduction, and survival, which are likely to be particularly strong for young adult herring.     

Life-history effects of migratory costs in anadromous brown trout

Mean size of sexually mature anadromous brown trout (sea trout) Salmo trutta in south-east Norway increased significantly with migratory distance ( D ) between the feeding area at sea and the spawning area in fresh water, from 32 cm for those spawning close to the river mouth to 43 cm at the spawning grounds 40 km inland. This was largely due to an increased size of the smallest anadromous spawners with increasing D . The raised mean size of the long-distance migrants is paralleled by an increased age at sexual maturity. Body mass at the same length of sea trout decreased with D in fresh water, meaning that the fish moving far inland was slimmer than those spawning near the coast. Gonadal mass of first-time spawning anadromous males declined significantly with D , and the fecundity and the ratio of fecundity over mean mass of the individual eggs adjusted for variation in fish mass, increased with D . There was no clear relationship between the ratio of anadromous to resident fish and D , probably because more variables than D , influence this relationship in the study streams.     

Pseudanthocotyloides heterocotyle (van Beneden, 1871) Euzet & Prost, 1969 (Monogenea: Polyopisthocotylea: Mazocraeidae), a parasite of herring Clupea harengus L. and sprat Sprattus sprattus L. (Teleostei: Clupeidae)

A total of 690 herring Clupea harengus L. and 88 sprat Sprattus sprattus L. caught off the west coast of Sweden, in the North Sea and off the west and south coasts of the United Kingdom, were examined for gill parasites. The monogenean Pseudanthocotyloides heterocotyle (van Beneden, 1871) Euzet & Prost, 1969 was found in 38 (5.5%) herring and one (1.1%) sprat. The parasite was significantly (P>0.05) more common off the west coast of Sweden than elsewhere and most specimens (62.5%) were found on the pseudobranchs. Only the smaller herring were infected. P. heterocotyle is redescribed and its taxonomy discussed, together with the possibility of host and parasite misidentification in previous reports.     

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.     

Role of bay, fjord and seamount on the early life history of Lepidonotothen squamifrons from the Kerguelen Islands

Among the Kerguelen Islands' demersal fish, larvae of Lepidonotothen squamifrons are dominant during the summer over the island shelf and surrounding seamounts. Distribution of larval stages from different scientific surveys confirmed the position of the two known spawning grounds (south of the Kerguelen shelf and at the Kerguelen-Heard bank) and may indicate other spawnings on the Skiff bank and in bays and fjords. Different larval cohorts were observed for the shelf and the coastal zone whereas a unique one was observed for Skiff and Kerguelen-Heard seamounts. Larvae from the spawning ground south of the Kerguelen shelf appear in early summer and show a northwards dispersal to the northeastern nearshore zone, the Baleiniers Gulf, following the scheme of fish migration shown by Harden-Jones. Later on, larvae from the northern inshore spawning zone occur in the same area, which presumably then avoid intraspecific competition with the previous larvae. Larval distribution and migration strategy seemed to be in accordance with the island mass or seamount effect under a regular current. Accepted: 2 December 1999     

Migration route and spawning area fidelity by North Sea plaice

Data from plaice, Pleuronectes platessa L., tagged with electronic data storage tags, were used to test whether these fishes exhibited migration route and spawning area fidelity in successive spawning seasons. Depth and temperature data were recorded for each fish over 365-512 days in the central North Sea and this information was used to reconstruct movements based on tidal locations. We discovered highly directed seasonal migrations from the winter spawning area south of a major topographical feature, Dogger Bank Tail End, to summer feeding grounds 250 km to the north in deep, cold, thermally stratified water. Our results show synchronous timing of migration, repeated pre- and post-spawning migration routes and 100% spawning area fidelity, including two individuals that returned to within 20 km of their previous season's spawning location. This is the first study to provide a complete reconstruction of annual migrations by individual fishes, showing strong homing behaviour along consistent migration routes.     

Temporal stability of the population genetic structure of the White Sea herring <Emphasis Type="Italic">Сlupea pallasii marisalbi</Emphasis>

The length–age distributions and genetic variation of ten microsatellite loci were examined in samples of herring from the Chupa Inlet of the Kandalaksha Gulf of the White Sea. The samples were collected on the same spawning grounds over a period of 13 years and during the same spawning season. The obtained data demonstrate stability of the length structure and genetic characteristics (allele and genotype frequencies, allelic richness, and heterozygosity). The degree of temporal genetic differentiation was not statistically significant, θ = 0.0038 (CI,–0.0006 to 0.006). Spatial differentiation of the White Sea herring far outweighs the level of temporal variability, which makes it possible to suggest the important role of reproductive isolation in the formation of the population genetic structure of herring.     

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.     

Effect of spawning ground location on the transport and growth of chub mackerel (Scomber japonicus) eggs and larvae in the East China Sea

Chub mackerel (Scomber japonicus) are an important pelagic fish species within the China Sea. Annual recruitment of this species is determined primarily by survival in the early life history stages. Minor changes in the physical marine environment can have a significant effect on the growth and survival of eggs and larvae, thereby affecting recruitment of population. To model this interaction, we constructed a bio-physical dynamic model of the early life history of chub mackerel in the East China Sea (ECS). The physical model was based on the unstructured grid Finite Volume Coast and Ocean Model (FVCOM) and simulated the 3-D physical fields. The biological model was based on individual-based models (IBMs) in which the early life stages of chub mackerel were divided into five stages based on age or length. The model was parameterized using functions describing spawning, growth, and survival for chub mackerel in the ECS. Using this coupled physical and biological model, driven by the March–July climatological forcing, we tracked super individuals from spawning grounds to the nursery grounds to evaluate the influence of the physical environment at each of the spawning locations (western, normal, eastern) on the transport and survival of chub mackerel. The model suggests that spawning location has a significant effect on larval transport, although the larvae were generally advected northeastward to enter the Japan/East Sea through the Tsushima/East Strait or southeastward with the Kuroshio Current which then flows along the eastern Japanese coast. Spawning to the west was highly influenced by the Taiwan Warm Current (TWC) during early transport when the larvae were advected northward and then northeastward. The speed of drifting during this period was relatively slow. The model predicted that a large number of eggs and larvae would enter and transit through China’s coastal waters (Changjiang River Estuary, Hangzhou Bay, and the Zhoushan Islands). Under this scenario, the majority of larvae were transported to the northern nursery grounds, 79% to the nursery at Jeju Island and 10% to the nursery at Tsushima Strait. In contrast, only 11% were transported to the southern nursery grounds in the Pacific Ocean and Kyushu. Larvae spawned at the eastern spawning ground were primarily influenced by the Kuroshio Current which transported the larvae southeastward. Kuroshio acts as a barrier, restricting larvae from being advected to the interior of the western Pacific Ocean. Under such circumstances, almost no eggs and larvae were retained in the coastal waters of China. Instead, the larvae were rapidly transported northeastward from the Chinese shelf towards the coast of Japan. The model predicted that a large number of larvae would be transported to the southern nursery grounds in the Pacific Ocean and Kyushu, before entering the Pacific Ocean and Japan Sea. In total, 36% of larvae were transported to the Pacific Ocean nursery, 45% to the northern nursery grounds of Jeju Island and Tsushima Strait, and 27% to the Jeju Island nursery. The three simulations assumed the same number of eggs were spawned (2.17 × 10¹²) and the survival of larvae at the western, normal, and eastern spawning grounds was 0.0306%, 0.0353%, and 0.0234%, respectively. The average length was 123.7, 126.0, and 123.5 mm, respectively. Our results suggest that larvae spawned in different regions encountered different physical environments and were subject to different transport processes. These differences explain the changes in survival and growth observed between larvae from the different areas. Survival and growth was highest for chub mackerel that were spawned at the normal spawning location and subject to suitable water depths and temperatures during transport.     

Hatch-period-dependent early growth and survival of Pacific herring Clupea pallasii in Miyako Bay, Japan

Considerable interannual variation in the abundance of larval and juvenile Pacific herring Clupea pallasii was detected in Miyako Bay, on the Pacific coast of northern Japan; abundances were high in 2001 and 2003 and low in 2000 and 2002. Hatch dates and growth rates for larval and juvenile survivors were estimated through otolith analysis. Water temperature and food availability were monitored on the spawning and nursery grounds in the inner part of the bay. The number of spawning females caught in nets set around the spawning ground was recorded during each spawning season (January to May) in 2000–2003. No correlation was found between the number of spawning females and the abundance of larvae and juveniles on the spawning and nursery grounds. The hatch dates of surviving larvae and juveniles were concentrated at the end of the spawning season in 2001 and in the middle of the season in 2003. The larvae experienced relatively high prey concentrations during the first-feeding period in 2001 but low concentrations in 2003. Survival of larvae during the first-feeding period may be a function of prey concentration as well as water temperature. In 2003, low water temperature would reduce starvation mortality during the first-feeding period. In contrast, unfavourable feeding conditions with higher temperatures during the first-feeding period seemed to result in low larval survival in 2000 and 2002. The 2001 larvae grew faster than those in 2003 because of the late hatch dates and the higher ambient temperatures that resulted. Temperature might be a major factor controlling growth rates of C. pallasii larvae in Miyako Bay.     

On the spatial relationship between the spawning and feeding parts of the range of lumpfish Cyclopterus lumpus (Cyclopteridae) in the Barents Sea and adjacent waters (according to results of analysis of the size composition similarity)

Results of analysis of similarity of size series of mature females of the lumpfish Cyclopterus lumpus collected on the spawning grounds in the Barents and Norwegian seas (Russian and Norwegian coasts), as well as in the areas of its feeding, are provided. The trajectories of feeding migrations of lumpfish in the southern part of the Barents Sea have been revealed. A clinal decrease in the average length of females in the range from west to east is shown. The area of feeding aggregations of lumpfish spawning off the Russian coast is distinguished. The boundaries of the census water area used to assess the commercial stock of lumpfish and to determine the values of its possible catch off the coast of Murmansk are specified.