Diet of Atlantic cod in the southern Gulf of St Lawrence as an index of ecosystem change, 1959–2000

Major temporal changes in the importance of euphausiids and Atlantic herring Clupea harengus in the diet of Atlantic cod Gadus morhua , 10–75 cm L _F, occurred in shallow waters (<100 m depths) of the southern Gulf of St Lawrence between 1959 and 2000. Euphausiids represented 6–70% of prey mass for Atlantic cod 31–60 cm L _F from 1959 to 1987 but only trace amounts were detected in stomachs collected from 1990 to 2000. Mysids and gammarid amphipods (for Atlantic cod ≤45 cm L _F), and Atlantic herring (for Atlantic cod >45 cm L _F) largely replaced euphausiids in the Atlantic cod diets from 1990 to 2000. This diet change suggested there has been a major perturbation of the food web of the southern Gulf of St Lawrence. The importance of fishes (mostly Atlantic herring) in the diet of Atlantic cod >45 cm L _F increased significantly between the periods 1959–1980 and 1987–2000. Atlantic herring comprised 0–4% (mean 1.3%) of prey mass of Atlantic cod 46–60 cm L _F from 1959 to 1980 and increased to 2–42% (mean 19.6%) of the diet from 1987 to 2000. Atlantic herring comprised 0–25% (mean 9.4%) of the prey mass of Atlantic cod 61–75 cm L _F from 1959 to 1980 and increased to 42–81% (mean 54.2%) of prey mass from 1987 to 2000. This increased consumption of Atlantic herring was consistent with observed changes in abundance of Atlantic herring in the ecosystem between the late 1970s and 2000. The large changes in consumption of euphausiids and Atlantic herring represent highly significant changes and would need to be included in the development and interpretation of ecosystem-based management models for this ecosystem.     

The effect of temperature and somatic growth on otolith growth: the discrepancy between two clupeid species from a similar environment

Otolith growth rates of the early life stages of herring Clupea harengus ( n = 472) and smelt Osmerus eperlanus ( n = 348) collected in the Vistula Lagoon (Baltic Sea) during 1997–1999 were analysed. The larvae and early juveniles were not only collected in the same geographical area they were also of the same size (range 15–43 mm standard length, L _S), similar ages and were collected during the same seasons (May to July). Although the two clupeid species experienced very similar environmental conditions, there were significant discrepancies in the analysed relationships. The otolith growth of larval and juvenile smelt was very strongly related to somatic growth while temperature had a minor effect. In herring, the effect of somatic growth, although clearly visible and statistically highly significant, was of less importance than temperature. Furthermore, variation in the otolith size and L _S relationship was affected by temperature and somatic growth in both species, but the variance of otolith size at L _S was higher for herring than for smelt. Although growth backcalculation from otoliths can presently be recommended as an appropriate method for use with both smelt and herring (despite possibly lower precision and accuracy with the latter), other methods referring directly to short-term increment width changes ( e.g. marginal increment analysis) are recommended for smelt but not for herring.     

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.     

Parental effects on size variation in fish larvae

The growth and changes in the standard deviation about mean length were measured for sole, Solea solea L., and herring, Clupea harengus L., larvae from crosses of one or two males with one or two females. The sizes of sole larvae did not differ between crosses but the size variation increased faster with age when two females were used. Both maternal and paternal effects were detected for size at hatching in herring, but there was no parental influence on size variation. Size variation in herring larvae declined or was stable during the yolk sac stage and was not affected by the number of parents used for fertilizations.     

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.     

Drinking and water absorption by the larvae of herring (Clupea harengus) and turbot (Scophthalmus maximus)

The efficiencies of water absorption from the guts of the larvae of herring ( Clupea harengus L.) and turbot ( Scophthalmus maximus L.) were estimated by two methods, The first method was based on the differences in the rates of accumulation, by drinking, and clearance from the gut of radiolabelled inert markers. The second method used the equilibrium level of radioactivity in the larvae to measure the concentration of the markers in the gut above background as water is absorbed from the gut. Water absorption efficiencies for herring larvae drinking sea water were estimated to be 77% using both methods. When external salinity was reduced to 50% sea water, drinking rates and water absorption efficiency in herring larvae fell substantially.
Estimates of water absorption efficiency of seawater-adapted turbot larvae were similar (71–84%) to that of herring using both methods. Although temperature had a marked effect on both the rate of drinking and water absorption, there was no significant thermal effect on the efficiency of absorption from the guts of turbot larvae. The limitations of the techniques and the implications of the estimates in terms of water balance in fish larvae are discussed.     

Variation in the morphology of adult Apatemon gracilis Rudolphi, 1819 (Digenea: Strigeidae) reared in different avian hosts

Adult Apatemon gracilis (Rudolphi, 1819) were reared experimentally in three different avian hosts: herring gulls Larus argentatus Gmelin, domestic chicks Gallus gallus (L.) and eider ducklings Somateria mollissima (L.). Comparisons of size, body proportions and fecundity were made between these and specimens obtained from a naturally infected goosander Mergus merganser L. The eider duck proved to be a suitable experimental host, rearing adults of comparable size and fecundity to those from the natural host. The growth and development of the parasite in domestic chicks and herring gulls were significantly reduced. Similar conditions are likely to exist in the natural environment and this should be taken into consideration when choosing experimental hosts for use in the laboratory or when identifying species.     

Long-term changes in the prevalence of two helminth parasites (Cestoda: Trypanorhyncha) infecting marine fish

Changes in prevalence of the plerocerci of Grillotia angeli Dollfus, 1969 in mackerel, Scomber scombrus L., and of Lacistorhynchus sp. in herring, Clupea harengus L., were recorded over periods of eight years (1978–1985) and 11 years (1974–1984), respectively. Data were collected from 21 year classes of mackerel in an area to the south-west of Britain and Ireland and from seven year classes of herring in the eastern North Sea. Both sets of data showed sharp decreases in parasite prevalence from periods at relatively high levels to others at much lower levels. The changes in prevalence occurred at the same time in both host-parasite systems and coincided with the end of the hydrographic phenomenon known as the mid-70s salinity anomaly. Possible explanations for the changes which are discussed include changes in abundance of first intermediate and definitive hosts, variations in host year class strength, changes in hydrographic conditions and changes in host diet.     

Comparative analysis of alternative statistical models for differentiation of herring stocks based on meristic characters

A basic stock assessment problem is the mixing and separation of herring populations in their specific areas. Within the transition zone between the North and the Baltic seas (Skagerrak, Kattegat, The Sound) a mixing of two herring populations ( Clupea harengus L.) temporarily takes place. One major component stems from the Baltic Sea (spring spawner), and the other from the North Sea (autumn spawner); each stock exhibits different meristic characteristics. In order to separate the two herring populations, 'pure' learning samples of meristic characters were created in 1995 as differential variables. Mainly gained during two research surveys at two geographically remote areas where a mixing of herring components was relatively unlikely, these learning samples were considered as representative for one or the other herring population. This paper compares the current (ICES) separation approach for herring stock assessment with two alternative methods wherein vertebra counts (vc) are used as meristic characters. The two proposed methods are (a) an inverted variance weighted linear model, and (b) a separation rule based on a quadratic discriminant analysis. The paper summarizes and discusses the results of predicting fractions of the two herring components derived from all three separation models. The underlying example data set stems from a 1991–97 routine Swedish survey in the transition zone. In comparison, the quadratic discriminant analysis separation model was identified as superior to the two other methods. Furthermore, the model suggests a higher degree of mixing of the two herring stocks in the transition area than was previously thought; inter-annual changes in the geographical distribution of the two populations are suggested as being less variable than previously assumed.     

Geographical variation in the yellow‐legged gull: introgression or convergence from the herring gull?

Yellow-legged gulls Larus michahellis from the Atlantic Iberian coast exhibit some phenotypic similarities with the herring gull L. argentatus from Western Europe. To assess this phenomenon and its possible origin, we compared Mediterranean yellow-legged gulls, Atlantic Iberian yellow-legged gulls and herring gulls for several phenotypic traits (morphology, plumage), and used genetic data to determine the evolutionary history of the Atlantic Iberian yellow-legged gulls. Data from mitochondrial cytochrome b gene and microsatellite loci clearly indicate that Atlantic Iberian gulls are closely related to Mediterranean yellow-legged gulls, and do not show stronger signs of introgression with herring gulls relative to other populations of yellow-legged gulls. Atlantic Iberian yellow-legged gulls are more similar to herring gulls in body size and shape than to other yellow-legged gulls populations, but not in mantle colour and wing-tip pattern. Body size and other phenotypic and life history similarities with the herring gull ( L. argentatus argenteus ) such as voice, winter plumage and breeding phenology, previously described in several studies, might thus be interpreted as convergent characters. Within the yellow-legged gull, the high F _st-values obtained from four nuclear microsatellite loci indicate substantial population structure and reduced levels of gene flow between gull populations in Mediterranean France and Atlantic Iberia. Differences among these populations in breeding phenology and migration patterns, likely resulting from different local selection pressures, might contribute to this low level of gene flow.     

Population structure of Atlantic herring, Clupea harengus L.

During the last decade, genetic studies have been carried out on samples of Atlantic herring, CIupea harengus L., from the Baltic, North Sea, British Isles, Norwegian sea-waters and in the western part of the North Atlantic Ocean. Based on direct comparisons of samples drawn from several areas, the available information on allozyme variation was compiled into a comparable data set and analysed for population structure and genetic distance. The results are discussed in relation to the present stock model for herring.     

Thiaminase activity of Baltic salmon prey species: a comparision of net- and predator-caught samples

Thiaminase activity was determined for Gulf of Bothnia (GB) and Gulf of Finland (GF) Baltic herring Clupea harengus membras , sprat Sprattus sprattus and three-spined stickleback Gasterosteus aculeatus sampled from either trawl or gillnet catches or from Baltic salmon Salmo salar stomachs. The thiaminase activity in Baltic herring was about 10-fold higher than that in sprat, and there was almost no thiaminase activity in three-spined stickleback. Thiaminase activity of undigested Baltic herring found in Baltic salmon stomachs was significantly higher than that of trawl-caught Baltic herring from the same sea area, suggesting that there may be a higher risk of predation for Baltic herring with high thiaminase activity, possibly linked to their health. Thiaminase activity of the gastrointestinal contents of Baltic salmon, feeding almost entirely on Baltic herring in the GB, was significantly higher than for Baltic salmon feeding on both Baltic herring and sprat in the GF. Therefore, Baltic herring may be the major source of thiaminase for Baltic salmon. A tank experiment demonstrated that thiaminase activity in Baltic herring may vary, even within very short time periods. The results were consistent with the hypothesis that the thiaminase content in Baltic salmon forage fish may be an important link in the aetiology of the thiamine deficiency syndrome, M74, in Baltic salmon.     

The Dynamics of a Metapopulation: Changes in Life-History Traits in Resident Herring that Co-Occur with Oceanic Herring during Spawning

Different populations of Atlantic herring are regarded as forming a metapopulation, but we know little about the dynamics of the connectivity and degree of interbreeding between the populations. Based on data from three periods between 1962 and 2011, we identified the presence of two components of herring in a small semi-enclosed coastal marine ecosystem based on different somatic growth patterns and mean vertebrae sum (VS). The two components were interpreted as belonging to a resident herring population and the migratory, oceanic Norwegian spring spawning (NSS) herring population, and they co-occurred during spawning. In the 1960s, resident herring characterized by slow growth and low VS co-occurred with rapid growth, high VS oceanic NSS herring. Similar slow-growing resident and rapid-growing NSS herring were found in the 1970–80s, but both populations now had low VS suggesting similar origins. Finally, in the 2000s both populations showed rapid growth. The changes coincided with the NSS herring going from a state of high abundance and oceanic distribution to a collapse in the late 1960s that resulted in a coastal distribution closer to resident herring populations, before full recovery and resumption of the migratory, oceanic pattern in the 1990s. During all three periods, NSS herring were only present in the local system up to an age of about five years, but the synchronous spawning of the populations supports mixed spawning and interbreeding. During the investigation period both longevity, length at age (growth) and length-at-first maturity increased markedly for the resident herring, which then became more similar to the NSS herring. Genetic and/or cultural factors are believed to be the main causes of the observed changes in life history traits, although some effect of changes in environmental factors cannot be excluded. Our study suggests that relationships among populations in a metapopulation can be highly dynamic.     

Temperature,salinity tolerance,and buoyancy during early development and starvation of Clyde and North Sea herring,cod, and flounder larvae

Tolerance limits, at which 50% of larvae could survive high temperature and low salinity for 24 h, were determined for the yolk-sac larvae of Clyde and North Sea herring (Clupea harengus L.), cod (Gadus morhua L.) and flounder (Platichthys flesus L.) during early development and starvation. Clyde and North Sea herring, cod and flounder from hatching to the end of the yolk-sac stage, could withstand 21–23.5 °C, 20.5–23 °C, 15.5–18 °C and 21.5–24°C, respectively. The temperature tolerance was reduced by about 3.5–4 °C for Clyde herring and cod, 4–4.5 °C for North Sea herring and 8–8.5 °C for flounder when the larvae reached the point-of-no-return (PNR, when 50% of larvae, although still alive, are no longer strong enough to feed). The lowest salinity tolerance between hatching and the end of yolk-sac stage was 1–1.5‰ for Clyde and North Sea herring, 2–3‰ for cod and 0–1‰ for flounder. In no instance was there a loss of tolerance to low salinity during starvation. In fact, tolerance improved somewhat until the larvae became moribund. At hatching Clyde and North Sea herring larvae were negatively buoyant with a sinking rate of 0.35–0.4cm · s⁻¹ which steadily decreased until the larvae became moribund. Cod and flounder larvae, however, were positively buoyant at hatching but became progressively less buoyant and, by the end of the yolk-sac stage they were negatively buoyant with a sinking rate of 0.06–0.07 cm · s⁻¹. This sinking rate then decreased slightly until the PNR stage. The low salinity tolerance of all three species varied in a similar fashion to buoyancy.     

Selective predation on an introduced zooplankter ( Bythotrephes cederstroemi ) by lake herring ( Coregonus artedii ) in Harp Lake,Ontario

1. Bythotrephes cederstroemi (Crustacea: Onychopoda: Cercopagidae) invaded Harp Lake, Ontario in 1993, since when the zooplankton community has shifted from dominance of small-bodied to large-bodied species. During 1995 diets of adult lake herring (Coregonus artedii), Harp Lake’s primary planktivorous fish, were examined to determine the extent to which this conspicuous zooplankter has become integrated into the lake’s foodweb and whether fish condition has been affected in consequence. 2. Bythotrephes and Daphnia galeata mendotae were strongly preferred prey, whilst Holopedium gibberum and calanoid and cyclopoid copepods were negatively selected by lake herring. Predation on Bythotrephes and Holopedium was not size-selective, although D. galeata mendotae and calanoid copepods (Leptodiaptomus minutus and L. sicilis) consumed by herring were significantly larger than co-occurring conspecifics in the lake. 3. Caudal spines of Bythotrephes may form boluses in lake herring stomachs. However, the number of caudal spines in fish digestive tracts did not differ significantly from the number of Bythotrephes’ mandible pairs, indicating that the former were not differentially retained. 4. Lake herring weight-at-length relationships in lakes in Muskoka, Ontario, invaded by Bythotrephes did not differ from those in adjacent non-invaded lakes, indicating that Bythotrephes invasion of lakes apparently has not affected condition of lake herring.     

Cathepsin D from Atlantic cod (Gadus morhua L.) liver. Isolation and comparative studies

The isolated cathepsin D-like enzyme from Atlantic cod (Gadus morhua L.) liver was shown to be a monomer with a molecular mass of approximately 40 kDa. It was inhibited by Pepstatin A and had an optimum for degradation of haemoglobin at pH 3.0. The purified enzyme had lower temperature stability than bovine cathepsin D. Antibodies raised against the purified enzyme and against two C-terminal peptides of cod cathepsin D recognized a 40 kDa protein in immunoblotting of the samples from the purification process. Both antisera showed cross reactivity with a similar sized protein in liver from cod, saithe (Pollachius virens L.), Atlantic herring (Clupea harengus L.) and Atlantic salmon (Salmo salar L.). A protein of same size was detected in wolffish (Anarhichas lupus L.) liver with the antibody directed against the purified enzyme. This antibody also recognized the native enzyme and detected the presence of cathepsin D in muscle of cod, saithe, herring and salmon. These antibodies may be useful in understanding the mechanisms of post mortem muscle degradation in fish by comparing immunohistochemical localization and enzyme activity, in particular in cod with different rate of muscle degradation. They may also be used for comparing muscle degradation in different fish species.     

Run timing of pelagic fishes in Gulf of St Lawrence: area and species effects

Run timings of four ubiquitous pelagic fishes in the Gulf of St Lawrence were compared for area and year effects. Run timing of alewife, Alosa pseudoharengus (Wilson), smelt, Osmerus mordax (Mitchill), and Atlantic salmon, Salmo salar L., was estimated at index rivers, and time of arrival of herring, Clupea harengus L., on spawning grounds was estimated from gillnet fisheries located around the Gulf, 1978–87. Time of pelagic fish migrations had strong species and area effects, but weak year effects. The species had their own unique migration times and durations, and could be divided into two groups: herring and smelt, which arrived in all areas of the Gulf at the same time; and alewife and salmon, which had significant area effects. Environmental data were not correlated with run timing. There was little annual variation in run timing among early-run species; only alewife had significant year effects. There was annual variation in the timing of late-run species, but it was not correlated with proximate environmental factors. Because run timing is an easily defined stock characteristic, observations over large areas and many years might be useful for understanding the impact of large-scale environmental changes on natural populations.     

Gastric emptying in cod, Gadus morhua L.: effects of food particle size and dietary energy content

Whole herring, Clupea harengus , were digested and emptied from cod stomachs much more slowly than meals of minced herring, showing that prey integrity is an important factor controlling emptying. Increasing the energy content of the minced herring by addition of fish meal and oil led to a reduction in the percentage of the meal emptied in the 24 h subsequent to feeding, but rates of energy throughput on the enriched herring diet were high. The physical form of the diet appears to be more important than energy content in the control of gastric emptying in cod. The results are discussed with respect to possible effects of dietary type on absorptive and metabolic processes.     

Evidence for Two Highly Differentiated Herring Groups at Goose Bank in the Barents Sea and the Genetic Relationship to Pacific Herring, Clupea pallasi

Genetic studies on Atlantic herring, Clupea harengus, have generally revealed a low level of genetic variation over large geographic areas. Genetically distinct herring populations in some of the Norwegian fjords are exceptions, and juvenile herring from the large oceanic herring, Norwegian Spring Spawners (NSS), are often found in mixture with local fjord populations as well as widely distributed in the Barents Sea. Research surveys in the eastern Barents Sea (Goose Bank) in 1993, 1994 and 2001 included collection of herring samples for allozyme analyses. As expected the results identified juveniles from NSS stock, but an additional unique group of herring (low vertebrae number), being almost fixed for alternative alleles at several allozyme loci, was detected. In some cases, the two groups of herring were taken in the same trawl catches as documented by highly significant departure from Hardy—Weinberg expectation with large excess of homozygotes providing evidence for population mixing. Large genetic differences (Nei's genetic distance = 1.53; F_ST = 0.754) were detected in pairwise comparisons based on five allozyme loci. The two herring groups were also compared with reference samples of Pacific herring, Clupea pallasi, including one sample from Japan Sea and three Alaskan samples. UPGMA dendrogram based on five allozyme loci revealed a close genetic relationship between the low vertebrae herring in the Barents Sea and the group of samples of Pacific herring. Although significant different in allele frequencies, one of the herring samples clustered together with the reference sample from Bering Sea with genetic distance of 0.008 and F_ST value of 0.032. The close genetic relationship found in this paper, suggest a re-evaluation of the taxonomic status of the Barents Sea herring populations investigated.