Differential population structuring of two closely related fish species, the mackerel (Scomber scombrus) and the chub mackerel (Scomber japonicus), in the Mediterranean Sea

Population genetic structures of the mackerel (Scomber scombrus) and chub mackerel (Scomber japonicus) were studied in the Mediterranean Sea. Fragments of 272 bp (S. scomber) and 387 bp (S. japonicus) of the 5'-end of the mitochondrial control region were sequenced from spawning individuals collected off the coasts of Greece, Italy, Spain, and Portugal. High levels of mitochondrial control region haplotypic diversity (> 0.98) were found for both Scomber species. Nucleotide diversity was higher in the mackerel (0.022) than in the chub mackerel (0.017). Global F(ST) values were also higher and significant in the mackerel (0.024, P < 0.0001) as opposed to the chub mackerel (0.003, P > 0.05). Molecular variance analyses showed differential genetic structuring for these two closely related species. There is extensive gene flow between Mediterranean Sea and Atlantic Ocean populations of chub mackerel, which are organized into a larger panmictic unit. In contrast, Mediterranean Sea populations of mackerel show some degree of genetic differentiation and are structured along an east-west axis. The analysed eastern Mediterranean Sea mackerel populations (Greece, Italy) are clearly separated from that of the western Mediterranean Sea (Barcelona), which forms a panmictic unit with eastern Atlantic Ocean populations. The genetic structures of both species showed asymmetric migration patterns and indicated population expansion.     

Allozyme, mitochondrial-DNA, and morphometrie variability indicate cryptic species of anchovy (Engraulis encrasicolus)

Previous surveys of population structure in the Atlantic-Mediterranean anchovy Engraulis encrasicolus L. have reported heterogeneity in morphology, allozyme frequencies, and mitochondrial DNA haplotype frequencies at a regional scale. In particular, two stocks of anchovy have been detected in the Adriatic Sea. In this paper, the available data is reviewed with the aim to relate genetic variation to geography at the widest possible geographical scale, for investigating the evolutionary mechanisms underlying stock structure in anchovy. Correspondence analysis of allozyme frequencies (24 samples, three polymorphic loci) compiled from the literature indicates three distinct entities in the Mediterranean Sea. Open-sea or oceanic anchovy populations are genetically different from inshore-water populations within a region (Nei's ^ G _ST = 0.035–0.067), while broadscale geographical variation is weak for each of these two habitat-specific forms (^ G _ST = 0.005–0.006). Mitochondrial-DNA haplotype frequencies support the distinction between an inshore form and an oceanic form (^ G _ST = 0.067–0.107), with virtually no genetic differences among oceanic populations across the Gulf of Biscay, the western Mediterranean and the Ionian Sea (^ G _ST = −0.001). If natural selection on marker loci is unimportant, these results indicate the occurrence of two parapatric, genetically distinct, habitat-specific forms that are widely distributed throughout the Mediterranean Sea. Persistent allele and haplotype-frequency differences between these forms indicate reproductive isolation and the presence of an E. encrasicolus species complex in the Mediterranean. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society 75 : 261–269.     

Genetic structuring of European anchovy (Engraulis encrasicolus) populations through mitochondrial DNA sequences

Mitochondrial DNA sequence variation in 655?bpfragments of the cytochrome oxidase c subunit I gene, known as the DNA barcode, of European anchovy (Engraulis encrasicolus) was evaluated by analyzing 1529 individuals representing 16 populations from the Black Sea, through the Marmara Sea and the Aegean Sea to the Mediterranean Sea. A total of 19 (2.9%) variable sites were found among individuals, and these defined 10 genetically diverged populations with an overall mean distance of 1.2%. The highest nucleotide divergence was found between samples of eastern Mediterranean and northern Aegean (2.2%). Evolutionary history analysis among 16 populations clustered the Mediterranean Sea clades in one main branch and the other clades in another branch. Diverging pattern of the European anchovy populations correlated with geographic dispersion supports the genetic structuring through the Black Sea-Marmara Sea-Aegean Sea-Mediterranean Sea quad.     

Field mortality estimates of anchovy larvae, Engraulis encrasicolus, in the western Mediterranean

Eggs and larvae of anchovy were obtained from plankton collections made along the Catalan coast (north-western Mediterranean) from 1983 to 1985 in order to determine spawning seasons and areas and other aspects of its life history. Previous studies have shown the existence of two main spawning areas, to the north and south of the Catalan coast, respectively. The northern one is characterized by a shorter spawning period related with the thermal cycle.
Mortality rates of larvae aged 4.9-15 days were calculated from the peak spawning month (June) of the three years, separately by spawning areas. Data on abundance by length classes were adjusted for escape and net avoidance and for duration of growth through a size class as derived from otolith readings. Mortality rates ranged from 0.17 to 0.58. Mortality was higher in 1983 than in 1984 and 1985, coinciding with a high production of anchovy eggs in that year. Mortality at the northern spawning area was lower than at the southern one.     

Characterization of swimming muscle in the Atlantic mackerel, Scomber scombrus L.

Both red and white muscle were removed from juvenile and adult Atlantic mackerel, Scomber scombrus L., for histochemical characterization of the muscle fibre types. Staining of white muscle for myosin ATPase, SDH, NADH diaphorase, GPDH and LDH revealed that these fibres are homogeneous. Red muscle was shown to be heterogeneous, of at least two fibre types recognizable on the basis of myosin ATPase staining with preincubation at a pH of 9·8. These two red types are dispersed throughout the red muscle and are present in both juveniles and adults. Red muscle is located both deep within the myotomes and as a superficial layer of muscle fibres. A third group of muscle fibres, intermediate in nature, was distinguished at the apex of the red muscle 'triangle,' between the epaxial and hypaxial muscle, using NADH diaphorase and myosin ATPase stains. This paper discusses the possibility that functionally different muscle fibres occur in the red swimming muscle of the Atlantic mackerel.     

A comparison of gut evacuation models for larval mackerel (Scomber scombrus) using serial photography

A novel technique is described, using serial photography of the gut contents of transparent living larval fishes, to generate individual gut evacuation time series. This technique was applied to Atlantic mackerel Scomber scombrus larvae to compare three widely used models of gut evacuation: linear, exponential and square-root. Regression r ² for the exponential model exceeded those for the linear and square root models in 20 of 21 time series, strongly supporting the exponential model. At the initial gut fullness for each time series, total gut evacuation rates calculated with the exponential model averaged 2·2 and 1·3 times greater than those calculated with the linear and square-root models, respectively, and would produce correspondingly higher estimates of feeding rates for field-collected larvae with similar levels of gut fullness. The results highlight the importance of choosing the appropriate evacuation model in feeding studies, particularly those intended to examine short-term changes in larval fish feeding rates, a contributing factor to the highly variable yearly recruitment of many marine fish species.     

New polymorphic microsatellite loci for population studies in the European anchovy, Engraulis encrasicolus (L.)

Eleven microsatellite loci were developed in the European anchovy, Engraulis encrasicolus, and tested in samples from two geographically distant populations (Atlantic and Mediterranean Sea). Number of alleles ranged from eight to 28 and observed heterozygosity from 0.440 to 0.920. There was no evidence of linkage disequilibrium, although two loci are indeed linked. All loci were in Hardy-Weinberg equilibrium, except for one locus in the Atlantic and two loci in the Mediterranean sample. These three loci plus two more showed evidence for null alleles.     

Lipid biomarkers and trophic linkages between phytoplankton, zooplankton and anchovy (Engraulis encrasicolus) larvae in the NW Mediterranean

Identification and quantification of the main fatty acids inphytoplankton, zooplankton and Engraulis encrasicolus larvaewere used to study the relationship among the different trophiclevels. Three northwestern Mediterranean sea areas were studiedas representative of potentially heterogeneous environments.HPLC pigment analysis was performed to assess the contributionof different phytoplankton groups to total chlorophyll biomass.In spite of the physical variability, a homogeneous patternin phytoplankton composition among areas was found, with thePrymnesiophyceae representing more than half of the phytoplanktonchlorophyll biomass. This homogeneous spatial pattern was alsoreflected in the phytoplankton fatty acid proportions. Thus,significant amount of 14:0, 18:1(n-9) and 18:4(n-3) could berelated to the dominance of the prymnesiophytes; the low proportionof fatty acid markers of diatoms and dinoflagellates agreedwith the low abundances of those phytoplankton groups. Zooplanktonfatty acids also showed homogeneous proportions in the studiedareas, with a preferential accumulation of 16:1(n-7) and 20:5(n-3),and 22:6(n-3). This observation may indicate a preferentialgrazing on diatoms and dinoflagellates. Anchovy larvae displayeddiffering fatty acids contributions related to size. Large larvaecontained a higher proportion of polyunsaturated fatty acid(PUFA). Small larvae showed a high percentage of 18:1(n-9) and18:4(n-3) fatty acids, considered Prymnesiophyceae biomarkers.These results suggest an additional food resource for the anchovythat is complementary to a zooplankton diet, but probably ofprotozoan origin (i.e. microzooplankton). Significant logarithmicrelationships were found between dry mass and 20:5(n-3) and22:6(n-3) fatty acid proportions in E. encrasicolus larvae.These PUFA are considered essential for the fish-larvae development.     

Influence of environmental variability on anchovy early life stages (Engraulis encrasicolus) in two different areas of the Central Mediterranean Sea

The survival of early stages of small pelagic fish species (e.g. Engraulis encrasicolus) is highly dependent on environmental conditions in both spawning and nursery areas. Knowing the relationship between the mortality rates of the early stages and the environment may help to study and model recruitment fluctuations. During the summer of 2006, two consecutive oceanographic cruises were carried out in the Central Mediterranean sea (CMED) in two different areas: the western Libyan waters and the Sicilian?CMaltese waters. For the first time a nearly synoptic comparison between the two border areas of the CMED is performed. In spite of a higher overall egg density in the Sicilian?CMaltese waters, there was a higher density of anchovy larvae in Libyan waters. The comparison between the oceanographic datasets singled out different circulation patterns and different characteristics of water masses, which helped to explain the differences in density of the spawning products in the two areas. A Lagrangian transport model was used to evaluate the effects of major oceanographic features on offshore egg and larval transport. The results of the model simulations and the nutritional conditions, as indicated by lipid, carbohydrate and protein contents, support the hypothesis that the western Libyan waters may represent a more favourable nursery ground compared to the Sicilian?CMaltese waters in terms of environmental conditions and food availability.     

Late Pleistocene divergence and subsequent population expansion of two closely related fish species, Japanese anchovy (Engraulis japonicus) and Australian anchovy (Engraulis australis)

Climatic oscillations during the Pleistocene ice ages produced great changes in species' geographical distribution and abundance, which could be expected to have genetic consequences. Living in the temperate upwelling zones of the northwestern Pacific, Japanese anchovy (Engraulis japonicus) might have been affected by these severe climatic oscillations. To investigate the effects of Pleistocene climatic changes on the evolution in Japanese anchovy, fragments of 522 bp at the 5' end of mitochondrial DNA control region were sequenced for 241 individuals from 13 localities and 37 individuals of Australian anchovy. Japanese anchovy and Australian anchovy are reciprocally monophyletic and a late Pleistocene transequatorial divergence between the two species was indicated. High levels of haplotype diversity (>0.99) were found for all samples, indicating a high level of genetic diversity. Analyses of molecular variance and the conventional population statistic F(ST) revealed no significant genetic structure throughout the range of Japanese anchovy. Both mismatch distribution analyses and neutrality tests suggested a late Pleistocene population expansion for both Japanese anchovy (79,000-317,000 years ago) and Australian anchovy (45,000-178,000 years ago).     

Diet composition and feeding of European anchovy Engraulis encrasicolus in Kiel Bight,western Baltic Sea

Diet composition of the expanding southern species European anchovy Engraulis encrasicolus in the western Baltic Sea was investigated. Results revealed an interesting case of bentho‐pelagic coupling with potential implications for local fish species through competition for food resources.     

Mitochondrial DNA reveals a mosaic pattern of phylogeographical structure in Atlantic and Mediterranean populations of anchovy (Engraulis encrasicolus)

This study extends the geographic coverage of a previous study of mitochondrial DNA restriction fragment length polymorphism in European anchovy. Both studies together include 24 samples representing 17 localities extending from the Black Sea, through the Mediterranean Sea to the eastern Atlantic as far south as Dakar, Senegal. Eighty-eight haplotypes define two clades (A and B) separated by 3.2% sequence divergence. Clade A has a star-like genealogy indicative of a recent population expansion. Clade B has a more complex genealogy, consisting of several haplotypes at intermediate frequencies. The distributions of these clades consist of a mosaic with abrupt changes between some areas and gradients between other areas. Clade A predominates the Black and Aegean seas, but is present throughout the Mediterranean. Unexpectedly, new data show that clade A is also at a high frequency in the Atlantic, from Portugal to at least Senegal. Overall, the level of genetic differentiation among populations is high (F(ST)=0.148, p<0.0001), with the greatest differences between basins. AMOVA reveals four main geographical groups: Atlantic, central Mediterranean, Aegean Sea, and Black Sea. Mismatch distribution clearly indicates historical bottleneck and population expansion for clade A, while for clade B such evidence is equivocal. This difference may reflect a range expansion for both clades, but with higher gene flow (Nm values) between demes for clade A. Both contemporary and historical processes are important in shaping the complex genetic population structure of European anchovy.     

The North Atlantic Oscillation affects landings of anchovy Engraulis encrasicolus in the Gulf of Cádiz (south of Spain)

The aim of this study was to explore the possible link between NAO (North Atlantic Oscillation) variations and the abundance of anchovy (Engraulis encrasicolus) stock in the Gulf of Cadiz. A significantly negative correlation was found between the NAO in the year previous to the anchovy landings. In the Gulf of Cadiz the mean anchovy landings after a negative NAO phase were significantly higher than after a positive NAO phase. A statistically significant negative relationship was obtained between the biennial NAO (estimated as the average monthly NAO values per two consecutive years starting with an odd‐numbered year) and the probability of having an anchovy landing value for a year higher than the average anchovy landings for the entire year. Results suggest that the NAO previous to the landing year is an ecological factor of European anchovy related to its abundance in the Gulf of Cádiz. This could have important applications for conservation strategies and species management.     

Spawning behaviour and diel periodicity of egg production in captive Atlantic mackerel,Scomber scombrus L.*

Spawning was observed in groups of captive Atlantic mackerel and occurred when a few fish swam away from the main school during a brief burst of high speed swimming. Counts of eggs from aquarium and field studies show that, in contrast to most pelagic fish species, spawning by Atlantic mackerel can occur at any time ofday or night and is not group-synchronized. Implications of the observed die1 periodicity of egg production to stock size estimation by the Daily Egg Production Method are discussed.     

Development of serotonergic neurons in the brain of the mackerel, Scomber scombrus. An immunohistochemical study

The ontogenetic development of serotonin immunoreactivity (5-HTir) neurons was investigated by immunocytochemistry in the brain of the mackerel, from fertilization to the complete resorption of the yolk sac (201 h). 5-HTir appears first in cell bodies in the diencephalon, on both sides of the third ventricle. At hatching time 5-HTir perikarya are also present in the mesencephalon and the inferior raphe but in small numbers. During larval stages, immunoreactivity appears consecutively in the area thalamo piaetectalis, the nuclei raphe pallidus and obscurus, the nuclei tegmenti dorsalis lateralis, recessus posterioris, lateralis hypothalami and paragigantocellularis lateralis. These results, together with previous findings, suggest a common pattern of distribution of the different 5-HTir neuronal populations in the brain of a teleost during ontogeny but a different sequence of appearance of initial 5-HTir in these populations between species.     

Spring migration of Atlantic mackerel,Scomber scombrus,in relation to water temperature through Cabot Strait (Gulf of St. Lawrence)

Synopsis Sea surface temperatures across Cabot Strait (Gulf of St. Lawrence) ranged from 6 to 9°C on June 3, 1989, but only from 3 to 5°C on June 2, 1990. Periods of peak commercial landings of mackerel in eastern Cape Breton Island extended from May 22 to June 3 in 1989, and from May 28 to June 2 in 1990. In late May 1990, Atlantic mackerel were captured with a purse-seiner during exploratory fishing in Cabot Strait; commercial quantities were caught in water as cold as 2.8°C. The presence of mackerel in water a full 4°C colder than its reported lower tolerance limit indicates that the development of the 7°C isotherm is not a requirement for the vernal appearing of mackerel. The overlap of the periods of peak commercial landings between 1989 and 1990, despite: marked differences in warming chronology, suggests that the movements of mackerel are not as closely linked to water temperature as previously reported. The fish's thermal preferences could be subordinate to their reproductive requirements at this stage of their spawning migration.     

Ultrastructure of atrial and ventricular myocardium in the pike Esox lucius L. and mackerel Scomber scombrus L. (Pisces)

Summary Atrial and ventricular muscle in the pike and mackerel hearts consists of narrow, branching cells. The atrial cells in the two species are similar whereas the ventricular cells differ. The sarcolemma is attached to the Z and M lines of the sarcomere. Intercalated discs are common, and the transverse parts display desmosomes and intermediate junctions. Nexuses are uncommon and only occur in the longitudinal parts of the intercalated discs. The sarcoplasmic reticulum forms a regular hexagonal network on the myofibrillar surface. Subsarcolemmal cisternae form peripheral couplings at the I-A level. Junctional processes are usually inconspicuous, but an electron dense substance is present between the sarcolemma and the Junctional sarcoplasmic reticulum. Specific heart granules are common in atrial cells of both species and in ventricular cells of the pike, but are very scarce in mackerel ventricular muscle.This work was supported by grants from the Norwegian Research Council for Sciences and the Humanities     

Genetic differences between populations of sardine, Sardina pilchardus, and anchovy, Engraulis encrasicolus, in the Aegean and Ionian seas

Genetic distances based on electrophoretic variation and multivariate analysis of several morpho-metric and meristic characters suggest that populations of sardine and anchovy from the Aegean and Ionian seas do not form one panmictic population. The distinction between these two classes of populations emerges only as a statistical property, and for some characters the within-sea variation is larger than the between-sea variation. Thus, the reproductive isolation between populations inhabiting the two seas appears to be only partial. Our findings do not support the' pure' or' discrete' stock concept. Instead, they provide evidence for the dynamic population structure model according to which physical (e.g., hydrographic) or biological (e.g., predation or behaviour) factors impose a population structure maintained in a semi-equilibrium state under the opposing influences of migration and selection. Such a dynamic state may not be stable in the long run, and it may not allow for the accumulation of genetic divergence necessary for the emergence of higher taxa. It must, nevertheless, be taken into account in the management of exploited natural populations.     

Long-term changes in growth and age at maturity of mackerel, Scomber scombrus L., from the North Sea

Mean length and mean weight at age of Atlantic mackerel in the North Sea increased significantly between 1960 and 1979, then, decreased markedly during the 1980s. In contrast, median age at maturity decreased (c. 1-1.5 years) from 1960. Median length at maturity remained stable until the 1980s, when it declined sharply. The significance of these changes is discussed in relation to variation in sea temperature, food availability and stock size. It is speculated that the changes during the 1980s resulted from immigration of western stock mackerel.