Comparison of the feeding apparatus and diet of European sardines Sardina pilchardus of Atlantic and Mediterranean waters: ecological implications

In this study, the feeding apparatus (gill rakers, GR) and the diet composition of European sardine Sardina pilchardus populations living in two contrasting environments were compared: the upwelling area off western Iberia and the comparatively less productive region of the north‐western Mediterranean Sea. The importance of local adaptations in the trophic ecology of this species was estimated. Sardina pilchardus from the Atlantic Iberian coast and from the north‐western Mediterranean Sea have clear differences in the feeding apparatus and diet compositions. Those from the Atlantic Iberian coast have significantly more GRs than S. pilchardus of the same size range in the Mediterranean Sea. While S. pilchardus from the Mediterranean Sea mostly depend on prey ranging between 750–1500 and 3000–4000 µm, corresponding mostly to cladocerans, decapods and copepods, those from the Atlantic depend on smaller prey (50–500 and 1000–1500 µm) that include phytoplankton and copepods, particularly during summer months, and S. pilchardus eggs during the winter. The marked difference between the trophic ecology of S. pilchardus in the two areas studied appears to have originated from different dietary strategies that the two populations have adopted in contrasting feeding environments. These differences are shown to profoundly affect the size and quality of prey consumed, and the effect of cannibalism on the populations.     

Identification of subpopulations in pelagic marine fish species using amino acid composition

The spatial stock complexity of marine fish species requires that population structure is taken into account in fisheries management. The aim of this study was to determine whether the amino acid composition (AAC) of the adult fish allows the identification of subpopulations within the stock. During a cruise in November 2003 along the entire Mediterranean coast of Spain, individuals were collected of the following pelagic species: Sardina pilchardus, Sardinella aurita, Engraulis encrasicolus, Trachurus trachurus, Trachurus mediterraneus, Scomber scombrus and Scomber colias. Individuals of S. pilchardus and E. encrasicolus were also collected from the waters of the Strait of Sicily in 2002 and 2003. The AAC of the fish eyes was seen to be species specific, and therefore, the differences in AAC among species may be based on inherited characters. Moreover, a clear differentiation was seen between the Spanish and Sicilian populations of S. pilchardus and E. encrasicolus. Furthermore, in the Spanish waters of the Mediterranean Sea, discriminant analysis revealed a substantial separation between the northern and southern subpopulations of S. pilchardus, S. aurita and E. encrasicolus. Temporal variations in AAC within species in each area were lower than the spatial variations observed among areas for each species, probably reflecting the influence on the AAC of the contrasting environmental characteristics of each area. Our results indicate that the ACC of the eyes in adult fish is a good tool for discriminating among subpopulations in pelagic marine fish species.     

Cryptic diets of forage fish: jellyfish consumption observed in the Celtic Sea and western English Channel

To establish if fishes’ consumption of jellyfish changes through the year, we conducted a molecular gut-content assessment on opportunistically sampled species from the Celtic Sea in October and compared these with samples previously collected in February and March from the Irish Sea. Mackerel Scomber scombrus were found to feed on hydrozoan jellyfish relatively frequently in autumn, with rare consumption also detected in sardine Sardina pilchardus and sprat Sprattus sprattus. By October, moon jellyfish Aurelia aurita appeared to have escaped predation, potentially through somatic growth and the development of stinging tentacles. This is in contrast with sampling in February and March where A. aurita ephyrae were heavily preyed upon. No significant change in predation rate was observed in S. sprattus, but jellyfish predation by S. scombrus feeding in autumn was significantly higher than that seen during winter. This increase in consumption appears to be driven by the consumption of different, smaller jellyfish species than were targeted during the winter.     

Chaotic genetic patchiness in the pelagic teleost fish Sardina pilchardus across the Siculo-Tunisian Strait

ABSTRACT

This investigation aims at assessing patterns of spatial genetic structure of the teleost fish Sardina pilchardus across the Siculo-Tunisian Strait (a well-known discontinuous biogeographic area) and delineating putative genetic stocks within the species. For this purpose, a total of 180 specimens, collected from 11 locations stretching across the western and eastern Mediterranean coasts of Tunisia, were analysed genetically by means of 18 nuclear allozyme loci. The outcome of this study revealed strong genetic differentiation among populations, with the marked genetic distinctiveness of the central Tunisian population at Mahdia. Despite the delineation of seven well-defined genetic groups, no significant correlation was found between genetic and geographic distances. Besides, the recorded population subdivision did not align with biogeographic boundaries, suggesting the presence of chaotic genetic patchiness. Recent genetic bottlenecks were evidenced in S. pilchardus populations. Patchy migration patterns were recorded among the examined pairs of sardine populations. Among the recorded 16 polymorphic loci, GPI-2 and SOD appeared to be subject to natural selection. Patterns of population genetic differentiation and structuring were not found to be driven by outlier loci that appeared to be under selection. Furthermore, the detected neutral GPI-1 locus was found to be responsible for most of the genetic variation among identified genetic clusters. Hence, natural selection cannot cause the detected genetic heterogeneity among sardine samples. Different explanations to the origin of chaotic genetic patterns, observed within S. pilchardus, were discussed.     

Conservation genetics of the short-beaked common dolphin (<Emphasis Type="Italic">Delphinus delphis</Emphasis>) in the Mediterranean Sea and in the eastern North Atlantic Ocean

Mediterranean Sea common dolphins have recently been listed as ‘endangered’ in the IUCN Red list, due to their reported decline since the middle of the 20th century. However, little is know about the number or distribution of populations in this region. We analysed 118 samples from the Black Sea, Mediterranean Sea and eastern North Atlantic at nine microsatellite nuclear loci and for 428 bps of the mtDNA control region. We found small but significant population differentiation across the basin between the eastern and the western Mediterranean populations at both nuclear and mtDNA markers (microsatellite F _ST = 0.052, mtDNA F _ST = 0.107, P values ≤ 0.001). This matched the differential distribution and habitat use patterns exhibited by this species in the eastern and the western parts of the Mediterranean Sea. The assignment test of a small number of samples from the central Mediterranean could not exclude further population structure in the central area of the basin. No significant genetic differentiation at either marker was observed among the eastern north Atlantic populations, though the Alboran population (inhabiting the Mediterranean waters immediately adjacent the Atlantic ocean) showed significant mtDNA genetic differentiation compared to the Atlantic populations. Directional estimates of gene flow suggested movement of females out of the Mediterranean, which may be relevant to the population decline. Phylogenetic analysis suggested that the observed population structure evolved recently.     

Molecular analysis of the recently described lizardfish Saurida lessepsianus (Synodontidae) from the Red Sea and the Mediterranean,with remarks on its phylogeny and genetic bottleneck effect

The opening of the Suez Canal in 1869 led to a massive influx of Red Sea species that invaded the Mediterranean; this was termed ‘Lessepsian migration'. Among these species was a species of lizardfish, identified by some authors as Saurida undosquamis and by others as S. macrolepis. Recently, the Red Sea and the Mediterranean populations were described according to external characteristics as a unique taxon, Saurida lessepsianus. Our molecular study confirms this finding and determines that all previous records of S. undosquamis and S. macrolepis in the Red Sea and the Mediterranean are misidentifications of S. lessepsianus. The Mediterranean population of S. lessepsianus exhibits a lower genetic variability than that of the Red Sea population, suggesting a bottleneck effect.     

Feeding habits of the swordfish (<Emphasis Type="Italic">Xiphias gladius</Emphasis> Linnaeus, 1758) in the subtropical northeast Pacific

Most of the studies carried out in the past on economically important fish species rely on single species approach. Ecosystem dynamics are characterized by complex interaction among species, sharing common habitat needs and thus forming characteristic assemblages. The analysis of spatio-temporal variability of fish community, coupled to the analysis of spatial indices, provides a synthetic view of the fish community status evidencing, if any, the way a community changes. Such considerations drive also to the development of ecosystem-based fishery management paradigm. In the present study changes in pelagic fish community structure in an upwelling ecosystem of the central Mediterranean Sea during the last 10 years was analysed, by focusing the attention on the five most abundant small pelagic species: Engraulis encrasicolus, Sardina pilchardus, Sardinella aurita, Trachurus trachurus and Boops boops. Our results evidenced a quite stable community structure, characterized by spatial occupation strongly driven by ecosystem characteristics and modulated according to specie-specific behaviour. Obtained results lead us to hypothesize that the observed stability of community could be linked to the presence of different environments leading to efficient space partitioning and resources utilization among species.     

Long-term changes in population genetic features of a rapidly expanding marine invader: implication for invasion success

Large blooms of Rhopilema nomadica, a highly venomous rhizostamatid scyphozoan species introduced to the Mediterranean through the Suez Canal, have become ubiquitous in the summer and winter months along the Israeli coasts since the mid-1980s. This species has since spread across the eastern Mediterranean and was sighted as far west as Tunisia and Sardinia. For the past 12 years, we have studied changes in the mitochondrial COI haplotypes diversity of R. nomadica to investigate small scale fluctuactions of genetic diversity and to reveal possible genetic structuring of the fast spreading invader in the Eastern Mediterranean. The 1091 COI sequences analysed, revealed a highly diverse population displaying 89 haplotypes, 46 of which appeared as singletons, low frequency haplotypes. All the specimens analysed throughout the period belong to a single unstructured population. Though lacking data from the source population in the Red Sea, the high within-population diversity and the high diversity of COI haplotypes support the hypothesis of multiple introductions events, or an open corridor with a continuous influx of propagules. Tajima’s D and Fu’s Fst negative values and the increased numbers of COI singletons from early to late sampling periods, have verified that the Israeli population is characterized by a rapid expanding population. Further research is needed for the evaluation of COI diversity and patterns in R. nomadica populations across the eastern Mediterranean Sea and Red Sea, as well as any correlation of the high variability between COI locus and phenotypic diversity.

     

First global approach: morphological and biological variability in a genetically homogeneous population of the European pilchard, Sardina pilchardus (Walbaum, 1792) in the North Atlantic coast

The European pilchard Sardina pilchardus represents the most commercially relevant fisheries resource in many countries bordering north Atlantic coasts and the Mediterranean Sea, being especially significant along the coast of Morocco. The continuous exploitation of this pelagic species for several decades places Morocco as the leader in sardine production. However, the conditions of exploitation of this resource underwent a great change during the last recent years. In order to identify the populations of the European pilchard sardine (Sardina pilchardus, Walbaum, 1792) in the Atlantic coast of Morocco and Spain, we have combined the truss network data to conduct multivariate analysis with biologic parameters and genetic analysis based on Microsatellite and mitochondrial control region data. Sardine morphometrics data truss variables from 10 samples spanning the Atlantic coast of Morocco were analysed by multivariate analysis. Thirteen morphometric measurements and some biological parameters such as the sex and the age of fishes were made for each individual. Discriminant analysis on size-corrected truss variables and cluster analysis of mean fishes shape using landmark data indicate, that the shape of north Moroccan sardines is distinct from the shape of sardines from south Morocco. However the analysis of the mitochondrial region and four microsatellites loci (Sp2, Sp7, Sp8 and SpI5) demonstrated an homogeneity population in the Moroccan Atlantic coast, with a low but significant genetic differentiation, which followed an isolation-by-distance pattern according to Mantel test.     

Fortes différences des ADN mitochonclriaux de populations de Sardinella aurita de la mer Méditerranée et de l'Atlantique Est

Digestion by restriction enzymes has been carried out on polymerase chain reaction products of the mitochondrial DNA control region of round sardine (Sardinella aurita) samples coming from the Eastern Atlantic and the Mediterranean Sea. The results show i) that, though the habitat is continuous, there is no gene flow between the two basins where two genetically differentiated groups can be recognized, ii) that each basin is genetically homogeneous and, iii) that the haplotypic diversity in the Mediterranean is between two and three times smaller than that observed in the Atlantic. These results can hardly be explained by a recent colonization from the Eastern Atlantic. This suggests that, for S. aurita and some other species, the Mediterranean Sea is genetically little influenced by the Eastern Atlantic.     

Temperature effects on asexual reproduction rates of scyphozoan species from the northwest Mediterranean Sea

In recent decades, many areas worldwide have experienced mass occurrences of jellyfish. To determine how temperature may affect jellyfish populations in the northwest (NW) Mediterranean Sea, we maintained polyps of three scyphozoan species, Aurelia aurita, Rhizostoma pulmo, and Cotylorhiza tuberculata in the laboratory at three temperatures (14, 21, 28°C) to test effects on survival and production of new polyps and ephyrae. Temperature significantly affected survival of all species, with longest survival of A. aurita and R. pulmo at 14°C and of C. tuberculata at 21°C. More polyps were budded by all species at temperatures above 14°C. A. aurita produced the most buds polyp⁻¹ (43.5) and R. pulmo the fewest (8.8). Strobilation occurred only at 14°C for A. aurita and at 21°C for C. tuberculata. For R. pulmo, fewer polyps strobilated and strobilated later at 14°C. These patterns of survival and asexual reproduction were seasonally appropriate for each species in the NW Mediterranean, where A. aurita medusae occur earliest (~April–May) in cool waters, followed by R. pulmo during May–June, and then by C. tuberculata in mid-summer. Comparisons among scyphozoan species suggested that many may be restricted by low temperatures, and that global warming may benefit temperate species, but not tropical or boreal species.     

Island biogeography of the Mediterranean sea: the species–area relationship for terrestrial isopods

Aim We looked at the biogeographical patterns of Oniscidean fauna from the small islands of the Mediterranean Sea in order to investigate the species–area relationship and to test for area‐range effects. Location The Mediterranean Sea. Methods We compiled from the literature a data set of 176 species of Oniscidea (terrestrial isopods) distributed over 124 Mediterranean islands. Jaccard's index was used as input for a UPGMA cluster analysis. The species–area relationship was investigated by applying linear, semi‐logarithmic, logarithmic and sigmoid models. We also investigated a possible ‘small island effect’ (SIE) by performing breakpoint regression. We used a cumulative and a sliding‐window approach to evaluate scale‐dependent area‐range effects on the log S/log A regression parameters. Results Based on similarity indexes, results indicated that small islands of the Mediterranean Sea can be divided into two major groups: eastern and western. In general, islands from eastern archipelagos were linked together at similarity values higher than those observed for western Mediterranean islands. This is consistent with a more even distribution of species in the eastern Mediterranean islands. Separate archipelagos in the western Mediterranean could be discriminated, with the exception of islets, which tended to group together at the lowest similarity values regardless of the archipelago to which they belong. Islets were characterized by a few common species with large ranges. The species–area logarithmic model did not always provide the best fit. Most continental archipelagos showed very similar intercepts, higher than the intercept for the Canary island oceanic archipelago. Sigmoid regression returned convex curves. Evidence for a SIE was found, whereas area‐range effects that are dependent on larger scale analyses were not unambiguously supported. Main conclusions The Oniscidea fauna from small islands of the Mediterranean Sea is highly structured, with major and minor geographical patterns being identifiable. Some but not all of the biogeographical complexity can be explained by interpreting the different shapes of species–area curves. Despite its flexibility, the sigmoid model tested did not always provide the best fit. Moreover, when the model did provide a good fit the curves looked convex, not sigmoid. We found evidence for a SIE, and minor support for scale‐dependent area‐range effects.     

Facts and uncertainties about the genetic population structure of Atlantic bluefin tuna (<Emphasis Type="Italic">Thunnus thynnus</Emphasis>) in the Mediterranean. Implications for fishery management

The Atlantic bluefin tuna (Thunnus thynnus) is an extraordinary fish that has amazed humanity since ancient times. However, the continuous overexploitation of this fishery, particularly in the Mediterranean Sea, could result in a total collapse of this resource. Currently, this species is managed as two stocks: Western Atlantic and Mediterranean-Eastern Atlantic, with a recognized genetic differentiation between them. On the other hand, the population structure within the Mediterranean Sea is still unclear. The biological data supports the idea of two separate populations in the eastern and western Mediterranean basins. However, nuclear microsatellite and mitochondrial DNA (mtDNA) analyses of two samples representative of these two basins result in a lack of heterogeneity. A comparison of these results with previously published studies reveals some discrepancies. We have compared 59 genetic differentiation tests that include samples within the Mediterranean. Of these, about 60% gave significant differentiation while the remaining 40% were non-significant. But, when only nuclear-based loci were considered, genetic differentiation was detected in up to 73% of the cases with an average significant F _ST of only 0.018, whereas the average significant F _ST of the mtDNA-based studies was significantly higher (0.029). However, in some cases, it is difficult to reconcile the biology of the species with the results suggesting genetic differentiation. In conclusion, although it is not yet possible to reach a definitive conclusion about the population structure, but considering all biological and genetic data, we suggest an independent management approach for each basin to avoid the impact of a type II error that could lead to the possible loss of the regional subpopulations.     

Genetic and morphological variation in an ecosystem engineer,Lithophyllum byssoides (Corallinales,Rhodophyta)

Lithophyllum byssoides is a common coralline alga in the intertidal zone of Mediterranean coasts, where it produces biogenic concretions housing a high algal and invertebrate biodiversity. This species is an ecosystem engineer and is considered a target for conservation efforts, but designing effective conservation strategies currently is impossible due to lack of information about its population structure. The morphological and molecular variation of L. byssoides was investigated using morphoanatomy and DNA sequences (psbA and cox2,3) obtained from populations at 15 localities on the Italian and Croatian coasts. Lithophyllum byssoides exhibited a high number of haplotypes (31 psbA haplotypes and 24 cox2,3 haplotypes) in the central Mediterranean. The psbA and cox2,3 phylogenies were congruent and showed seven lineages. For most of these clades, the distribution was limited to one or a few localities, but one of them (clade 7) was widespread across the central Mediterranean, spanning the main biogeographic boundaries recognized in this area. The central Mediterranean populations formed a lineage separate from Atlantic samples; psbA pair‐wise divergences suggested that recognition of Atlantic and Mediterranean L. byssoides as different species may be appropriate. The central Mediterranean haplotype patterns of L. byssoides were interpreted as resulting from past climatic events in the hydrogeological history of the Mediterranean Sea. The high haplotype diversity and the restricted spatial distribution of the seven lineages suggest that individual populations should be managed as independent units.     

Endemic and Indo-Pacific plankton in the Mediterranean Sea: a study based on dinoflagellate records

Aim To investigate biogeographical patterns based on published dinoflagellate records from the Mediterranean and Black Seas, and to provide a tentative list of endemic and Indo‐Pacific dinoflagellates in the Mediterranean Sea. Location Mediterranean Sea, Black Sea. Methods Checklists of dinoflagellates of the Mediterranean and Black Seas were compared with worldwide literature records. Only species reported in the Indo‐Pacific Ocean or exclusively known in the Mediterranean Sea were selected for biogeographical analysis. Results Dinoflagellates in the Mediterranean Sea comprised c. 43% of the world marine species and c. 88% of the dinoflagellate genera. Species richness among the Mediterranean sub‐basins showed marked differences due to the less reliable records of unarmoured (athecate) and rare dinoflagellates. These differences disappeared when only the more easily identifiable taxa were considered. Of the 673 dinoflagellates cited in the Mediterranean, 87% were also reported in the Atlantic Ocean. Only 40 taxa (6% of the total) were considered to be potential Indo‐Pacific species. Most were reported from the Ligurian Sea (21), and only two species from the Levantine basin. The other 48 taxa (7% of total) were known exclusively from the Mediterranean Sea, mainly from the Ligurian Sea. Half of these taxa were reported by a single author. Main conclusions Substantial dinoflagellates species richness can be attributed, in part, to the historical tradition of taxonomic studies in the Mediterranean Sea. The list of species of both Indo‐Pacific and exclusively Mediterranean species included taxa of dubious taxonomic validity or that were insufficiently known. The exclusion of these questionable taxa revealed the near absence of endemic dinoflagellates in the Mediterranean Sea compared with macroscopic organisms. This could be related to: (1) continuous replenishment of the plankton populations by the inflow of Atlantic water through the Strait of Gibraltar, (2) the possibility that species introduced during the Pliocenic flooding after the Messinian salinity crisis have not had enough time to diverge from their Atlantic ancestors, and/or (3) the reliance on traditional taxonomy based on morphological characters, which precludes the detection of cryptic speciation.     

Craniometrical variations among eastern Brazilian marmosets and their systematic relationships

The craniometrical variations among marmosets on the eastern coast of Brazil or theCallithrix jacchus group (C. jacchus, C. penicillata, C. kuhli, C. geoffroyi, C. flaviceps, andC. aurita) were analyzed to test various hypotheses for the arrangement of the species and phylogenetic relationships ofC. kuhli andC. flaviceps. Q-mode correlation coefficients and principal component analysis were employed to extract important shape factors from the craniometrical data. On this basis, the shape distances among the eastern Brazilian marmosets were found to be larger than those between the marmosets which are recognized as distinct species. This evidence supports the hypothesis that the members of theC. jacchus group are distinct species.C. kuhli shows the strongest resemblance toC. geoffroyi within theC. jacchus group.C. flaviceps is the most similar toC. aurita in its cranial shape. These findings indicate thatC. kuhli is most closely related toC. geoffroyi, whileC. flaviceps is toC. aurita.     

Morphometric differentiation of Pseudocalanus minutus populations in the Barents Sea

Dvoretsky VG. and Dvoretsky AG. 2011. Morphometric differentiation of Pseudocalanus minutus populations in the Barents Sea. —Acta Zoologica (Stockholm) 00 : 1–12. We investigated spatial variations in the morphometric characteristics (total length of body, lengths of cephalothorax, abdomen and antennules, and their relative proportions) of Pseudocalanus minutus, an abundant copepod species across the Barents Sea in August–September 2007. Females were found to have higher values for the measured parameters than males. The average absolute morphometric characters of both sexes increased from the south to the north. In most cases, parameters were similar in the southern, central, and eastern regions delineated by cluster analyses of oceanographic variables. The morphometric characteristics were strongly correlated with environmental variables in both males and females. Multiple regression analysis showed that temperature together with salinity explained 72–85% (in females) and 38–91% (in males) of the total variations in the log₁₀‐transformed morphometric parameters. According to principal component analysis and discriminant function analysis, two distinct groups could be separated in the Barents Sea. The first group included the copepods from the northern region; the other included populations from the southern, central, and eastern regions. The observed morphological variation can be interpreted as geographical variation connected with hydrological variability.