Modelling the presence of anchovy Engraulis encrasicolus in the Aegean Sea during early summer, based on satellite environmental data

Acoustic and satellite environmental data as well as bathymetry data were used to model the presence of anchovy, Engraulis encrasicolus during early summer in the northern Aegean Sea (Eastern Mediterranean). Generalized Additive Models (GAMs) were used for modelling and subsequently applied in a predictive mode to identify those areas in the Greek Seas and the entire Mediterranean basin that could support species’ presence. Model results were evaluated with the estimation of Receiver Operating Characteristic (ROC)-plots as well as qualitatively, based on (a) acoustic data from concurrent studies in certain areas of the northern Aegean Sea that were not included in the estimation of the GAM model and (b) historical acoustic data from the central Aegean and Ionian Seas. Mapping the estimated environmental conditions in the Mediterranean basin indicated areas that generally agree with the known distribution grounds of anchovy, such as the straits of Sicily and coastal waters of Tunisia, areas in the Tyrrhenian Sea, the Adriatic Sea, the Gulf of Lions and the Catalan Sea. Guest editor: V. D. Valavanis Essential Fish Habitat Mapping in the Mediterranean     

Fluctuating asymmetry and fitness correlations in two Engraulis encrasicolus populations

Correlations among several measures of fluctuating asymmetry (FA) and fitness‐related variables were assessed in two populations of the European anchovy Engraulis encrasicolus with fast growth (Aegean Sea) and slow growth (Ionian Sea), respectively. FA levels were borderline significantly higher in the Ionian than in the Aegean for some variables. Variation in otolith shape (deviation from population norm) was lower in the Ionian than the Aegean, contrary to expectation. Within the Aegean, there was no relation between any of the FA indexes and fitness estimators, while in the Ionian a composite otolith FA index was significantly negatively correlated to standard length at age only in 2 year‐old individuals. This difference between the Aegean and Ionian may have been related to the lower growth rate in the Ionian, as FA–fitness relations may be more apparent in less‐beneficial environments. The absence of significant correlations in the Aegean and the low correlation in one age group in the Ionian suggests that FA is not a sensitive indicator of individual fitness in adult E. encrasicolus.     

Anchovy (<Emphasis Type="Italic">Engraulis encrasicolus</Emphasis>) early life stages in the Central Mediterranean Sea: connectivity issues emerging among adjacent sub-areas across the Strait of Sicily

The combined use of field data on anchovy (Engraulis encrasicolus, Linnaeus, 1758) egg distribution in the Central Mediterranean Sea on both sides of the Strait of Sicily (Sicilian–Maltese and Tunisian waters) and Lagrangian simulations were used to assess the pattern of connectivity between these two sub-areas as a result of spawning activity. The field data were collected during ichthyoplankton surveys carried out in summer 2008 and 2010. The simulation runs showed considerable (up to 20%) rates of particle exchange in both directions (from Tunisian to Sicilian–Maltese waters and vice versa). However, considering the typical high mortality rates of anchovy early stages, the actual larval exchange rates across the Sicily Strait are supposed to be significantly lower (<1%), supporting the hypothesis that the anchovy population sub-units in the Strait of Sicily can be considered as separate fish stocks for the evaluation of their optimum exploitation rates.     

Pathways of unauthorized fish introductions and types of management responses

The study of the relationships among environmental factors, primary production, zooplankton community, and feeding behavior of fish species represents a key aspect to obtain a deeper knowledge of biological processes acting at ecosystem level. In this context, oceanographic data and biological samples were collected in two different surveys, carried out during late June 2011 and early July 2015 in the Strait of Sicily (Central Mediterranean Sea). Differences in abundance, biomass, and assemblage structure of zooplankton and anchovy (Engraulis encrasicolus Linnaeus, 1758) population were observed between the two surveys. Still, zooplankton biomass was significantly correlated to longitude, chlorophyll a recorded during the surveys (as a proxy of primary production), and oxygen concentrations. Such differences affected the isotopic composition of anchovy, which showed changes in δ¹⁵N and δ¹³C values between the two surveys. Mixing models identified different contributions of food sources in the two sampling periods, i.e., major consumption on large copepods and euphausiids in 2015 with respect to 2011, which was consistent with a greater availability of these preys in the environment in 2015. The obtained results evidenced that E. encrasicolus may be affected by any environmental change that influences the plankton community, which is the basis of the diet of these fishes. Such findings highlighted the importance to further investigate the trophic relationships among different compartments of the food web, as well as the possible environmental influences, in order to obtain a more complete picture of ecosystem functioning and also in the light of an ecosystem-based approach to fisheries management.     

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.     

Genetic variability of anchovy in the Azov-Black Sea basin

The intraspecific structure of the European anchovy (Engraulis encrasicolus) was studied on the basis of mitochondrial cytochrome b gene (cytb) fragment variability in 84 individuals from seven localities of the Black Sea and the Sea of Azov. The data on haplotype and nucleotide diversity and the values of neutrality tests suggested expansive growth of anchovy populations in the Azov-Black Sea basin. All samples from anchovy populations demonstrated a high level of haplotype diversity (Hd = 0.962). Two dominant haplotypes were identified, the frequencies of which were not directional, and they were present in all localities. Sequence analysis of the mitochondrial cytb gene fragment showed no differentiation between the Sea of Azov and the Black Sea subspecies.     

Modeling and predicting potential spawning habitat of anchovy (Engraulis encrasicolus) and round sardinella (Sardinella aurita) based on satellite environmental information

Anchovy and round sardinella are two important small pelagic species in the Mediterranean that spawn during the summer period. This is a first attempt to model and predict the two species’ potential spawning habitats in this area. Generalized additive models (GAMs) were constructed based on satellite environmental variables and presence/absence egg data, available from ichthyoplankton surveys conducted in the North Aegean Sea during early summer (June 2003–2006). These models were subsequently used to predict the probability of anchovy and round sardinella spawning in the Greek Seas as well as the entire Mediterranean and Black Sea during the same month of the year. The interaction of bottom depth and chlorophyll explained most of the deviance in the presence/absence GAMs of both species, indicating spawning over continental shelf areas with increased surface chlorophyll values. Round sardinella spawned closer to coast than anchovy. Predicted potential spawning areas for anchovy and round sardinella in unsampled areas of the Greek Seas and the entire Mediterranean and Black Sea were in good agreement with existing information on the distribution and extent of the spawning grounds, especially for anchovy. Modeling the species’ reproductive activity in relation to easily accessible environmental information and applying the models in a predictive way could be an initial, low-cost step to designate potential spawning fish habitats. Guest editor: V. D. Valavanis Essential Fish Habitat Mapping in the Mediterranean     

Identification of subspecies of European anchovy <Emphasis Type="Italic">Engraulis encrasicolus</Emphasis> (Engraulidae) in the wintering aggregations based on morphological parameters of otoliths

A new method of identification of the Sea of Azov and Black Sea subspecies of European anchovy Engraulis encrasicolus in mixed wintering aggregations is suggested based on discriminant analysis using a morphological parameter, the angle of the central groove of the otolith. This angle differs significantly in the subspecies, ranging between 92° and 148° and between 55° and 95° in the Azov and Black Sea subspecies, respectively. In the Azov anchovy, the difference between the eastern and western populations in the angle of the central groove is not registered. The classification equations obtained in the study can be used for subsequent assessment of the ratio between the abundances of each subspecies in mixed aggregations of European anchovy in the inshore zone of Crimea.     

Parallel genetic divergence among coastal–marine ecotype pairs of European anchovy explained by differential introgression after secondary contact

Ecophenotypic differentiation among replicate ecotype pairs within a species complex is often attributed to independent outcomes of parallel divergence driven by adaptation to similar environmental contrasts. However, the extent to which parallel phenotypic and genetic divergence patterns have emerged independently is increasingly questioned by population genomic studies. Here, we document the extent of genetic differentiation within and among two geographic replicates of the coastal and marine ecotypes of the European anchovy (Engraulis encrasicolus) gathered from Atlantic and Mediterranean locations. Using a genome‐wide data set of RAD‐derived SNPs, we show that habitat type (marine vs. coastal) is the most important component of genetic differentiation among populations of anchovy. By analysing the joint allele frequency spectrum of each coastal–marine ecotype pair, we show that genomic divergence patterns between ecotypes can be explained by a postglacial secondary contact following a long period of allopatric isolation (c. 300 kyrs). We found strong support for a model including heterogeneous migration among loci, suggesting that secondary gene flow has eroded past differentiation at different rates across the genome. Markers experiencing reduced introgression exhibited strongly correlated differentiation levels among Atlantic and Mediterranean regions. These results support that partial reproductive isolation and parallel genetic differentiation among replicate pairs of anchovy ecotypes are largely due to a common divergence history prior to secondary contact. They moreover provide comprehensive insights into the origin of a surprisingly strong fine‐scale genetic structuring in a high gene flow marine fish, which should improve stock management and conservation actions.     

Isolation of polymorphic microsatellite loci from the European anchovy,Engraulis encrasicolus

The European anchovy (Engraulis encrasicolus L., Engraulidae) is a widely spread fish resource which is overexploited along its area of distribution. A proper knowledge of the population genetic structure of this fish species is crucial to allow a sustainable management of fishery stocks. We developed five polymorphic microsatellite loci (expected heterozygosity per locus ranging from 0.46 to 0.97) for E. encrasicolus. These genetic markers can be applied to define the population genetic structure of European anchovy.     

Feeding by marine fish larvae: developmental and functional responses

Synopsis The relationship between prey consumption rate and prey concentration (functional response), and its change with growth (developmental response) were examined in the laboratory for three species of marine fish larvae: bay anchovy Anchoa mitchilli (Engraulidae), sea bream Archosargus rhomboidalis (Sparidae) and lined sole Achirus lineatus (Soleidae). The major objective was to determine relative predatory abilities of the larvae by fitting feeding rate data to developmental and functional response models. Feeding success, prey capture success, attack rates, handling times and search rates were estimated. Prey consumption rates and attack rates of bay anchovy usually were highest, but at the lowest prey level (50 per liter) first-feeding sea bream larvae had the highest consumption rate. Sea bream could consume prey at near-maximum rates at prey levels lower than those required by the other species. As larvae grew, time searching per attack decreased rapidly for all species, especially at low prey levels. Handling time also decreased, but most rapidly for bay anchovy. Search rates were highest for bay anchovy and lowest for lined sole. Bay anchovy had the best apparent predation ability, but when previous results on larval growth rates, survival rates and growth efficiencies were considered, sea bream larvae were the most efficient predators and the least likely of the three species to be limited by low prey levels.     

Habitat Selection Response of Small Pelagic Fish in Different Environments. Two Examples from the Oligotrophic Mediterranean Sea

A number of scientific papers in the last few years singled out the influence of environmental conditions on the spatial distribution of fish species, highlighting the need for the fisheries scientific community to investigate, besides biomass estimates, also the habitat selection of commercially important fish species. The Mediterranean Sea, although generally oligotrophic, is characterized by high habitat variability and represents an ideal study area to investigate the adaptive behavior of small pelagics under different environmental conditions. In this study the habitat selection of European anchovy Engraulis encrasicolus and European sardine Sardina pilchardus is analyzed in two areas of the Mediterranean Sea that largely differentiate in terms of environmental regimes: the Strait of Sicily and the North Aegean Sea. A number of environmental parameters were used to investigate factors influencing anchovy and sardine habitat selection. Acoustic surveys data, collected during the summer period 2002–2010, were used for this purpose. The quotient analysis was used to identify the association between high density values and environmental variables; it was applied to the entire dataset in each area in order to identify similarities or differences in the “mean” spatial behavioral pattern for each species. Principal component analysis was applied to selected environmental variables in order to identify those environmental regimes which drive each of the two ecosystems. The analysis revealed the effect of food availability along with bottom depth selection on the spatial distribution of both species. Furthermore PCA results highlighted that observed selectivity for shallower waters is mainly associated to specific environmental processes that locally increase productivity. The common trends in habitat selection of the two species, as observed in the two regions although they present marked differences in hydrodynamics, seem to be driven by the oligotrophic character of the study areas, highlighting the role of areas where the local environmental regimes meet ‘the ocean triad hypothesis’.     

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.     

Growth and length–weight relationship of the juvenile anchovy, Engraulis encrasicolus, in the nursery ground (Zrmanja River estuary–eastern Adriatic Sea)

Growth and length–weight relationships of 2564 juvenile specimens of anchovy, Engraulis encrasicolus (Linnaeus, 1758), were examined over a 2‐year period (August 1989 to April 1991) in the Zrmanja River estuary. Fish sizes ranged from 4.4 to 12.5 cm total length and 0.5 and 13.8 g, corresponding to age (in years) 0+ (98%) and 1+ (2%). Length–weight regression coefficient (b) varied between 2.95 and 3.42 and the regression constant (a) between 2.1 and 6.0 (×10^?3). The von Bertalanffy growth equation of juvenile anchovy was: l_t = 13.2 (1 ? e^?0.82(t+0.5)). Maturity was reached in June in males above 8.2 cm and females above 9.0 cm total length.     

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.     

Morphological parameters of abundant and threatened chondrichthyans of the northwestern Mediterranean Sea

The diversity of chondrichthyans in the Mediterranean Sea is relatively high; however, available data indicate that this group is declining in abundance and several species are becoming rare. As a result, the collection of biological data is a priority for demographic models, stock assessments, and food web analysis. In the present study, we report morphological parameters and length–weight relationships of several chondrichthyan species, both abundant and threatened, from the western Mediterranean Sea. Samples were obtained with commercial and scientific bottom trawl vessels between 2001 and 2013. A total of 893 individuals belonging to 11 families and 20 species were weighed and total lengths measured. In addition, seven species of large demersal sharks were measured and length–length relationships obtained to study the relationships between different body length measurements. All species showed positive allometric or isometric growth, except for Centroscymnus coelolepis. The results of the length–weight relationships reveal differences between the western Mediterranean and nearby areas, depending on the species studied.     

Anchovy egg and larval distribution in relation to biological and physical oceanography in the Strait of Sicily

The European Anchovy (Engraulis encrasicolus, Linnaeus, 1758) represents one of the most important fishery resources in some areas of the Mediterranean. This short-lived, small pelagic fish is characterized by large interannual fluctuations, probably as a result of environmental variability. As part of the European Project Med 98-070, the main aim of which was the study of the anchovy population in the Strait of Sicily, icthyoplankton surveys were carried out between 1999 and 2001, during the peak spawning season for anchovy. Present work reports the relationship between meso-zooplankton biomass and the abundance of anchovy eggs and larvae in the Strait of Sicily. Data on anchovy egg abundance showed that the main spawning area was located in the north-western region of the study area. The branch of the Atlantic Ionian Stream, running parallel to the southern Sicilian coast, acts as a transport mechanism for anchovy eggs and larvae towards the southernmost end of the island, off Cape Passero. Observed distributions were largely consistent with local hydrographic features, which allow larvae to be retained in areas providing the necessary feeding conditions for recruitment success.     

Recruiting at the Edge: Kinetic Energy Inhibits Anchovy Populations in the Western Mediterranean

The Strait of Gibraltar replenishes the Mediterranean with Atlantic waters through an intense eastward current known as the Atlantic Jet (AJ). The AJ fertilizes the southwestern Mediterranean and is considered to be the ultimate factor responsible for the comparatively high fish production of this region. Here, we perform an analysis of the available historical catches and catch per unit effort (CPUE), together with a long series of surface currents, kinetic energy and chlorophyll concentration. We show that the high kinetic energy of the AJ increases primary production but also negatively impacts the recruitment of anchovy. We contend that anchovy recruitment in the region is inhibited by the advection and dispersion of larvae and post-larvae during periods of strong advection by the AJ. The inhibitory impact of kinetic energy on anchovy landings is not a transient but rather a persistent state of the system. An exceptional combination of events creates an outbreak of this species in the Alboran Sea. These events depend on the Mediterranean-Atlantic exchange of water masses and, therefore, are highly sensitive to climate changes that are projected, though not always negatively, for fish landings.     

Oogonial proliferation and early oocyte dynamics during the reproductive cycle of two Clupeiform fish species

Although oogonial proliferation continues in mature females in most teleosts, its dynamics and the transformation of oogonia to early meiotic oocytes during the reproductive cycle have received little attention. In the present study, early oogenesis was examined throughout the reproductive cycle in two Clupeiform fishes, the Mediterranean sardine, Sardina pilchardus, and the European anchovy, Engraulis encrasicolus. Observations using confocal laser scanning microscopy (CLSM) provided extensive information on markers of oogonial proliferation (mitotic divisions, oogonia nests) and meiotic prophase I divisions of oocyte nests (leptotene, zygotene, pachytene, diplotene) in ovaries of different reproductive phases. In sardine, oogonial proliferation persisted throughout the entire reproductive cycle, whereas in anchovy, it was more pronounced prior to (developing ovaries) and after (resting ovaries) the spawning period. Anchovy exhibited a higher rate of meiotic activity in developing ovaries, whereas sardine exhibited a higher rate in resting ovaries. The observed differences between the two species can potentially be attributed to different seasonal patterns of energy allocation to reproduction and the synchronization between feeding and the spawning season.     

Evaluation of a Mysis bioenergetics model

Direct approaches for estimating the feeding rate of the opossumshrimp Mysis relicta can be hampered by variable gut residencetime (evacuation rate models) and non-linear functional responses(clearance rate models). Bioenergetics modeling provides analternative method, but the reliability of this approach needsto be evaluated using independent measures of growth and foodconsumption. In this study, we measured growth and food consumptionfor M. relicta and compared experimental results with thosepredicted from a Mysis bioenergetics model. For Mysis rearedat 10°C, model predictions were not significantly differentfrom observed values. Moreover, decomposition of mean squareerror indicated that 70% of the variation between model predictionsand observed values was attributable to random error. On average,model predictions were within 12% of observed values. A sensitivityanalysis revealed that Mysis respiration and prey energy densitywere the most sensitive parameters affecting model output. Byaccounting for uncertainty (95% CLs) in Mysis respiration, weobserved a significant improvement in the accuracy of modeloutput (within 5% of observed values), illustrating the importanceof sensitive input parameters for model performance. These findingshelp corroborate the Mysis bioenergetics model and demonstratethe usefulness of this approach for estimating Mysis feedingrate.