Deep-sea ecosystem variability of the Aegean Sea during the past 22 kyr as revealed by Benthic Foraminifera

Specific responses of the regional deep-sea ecosystems to climatic and oceanographic processes during the last 22 kyr are revealed by benthic foraminiferal faunas from two cores in the northern and southern Aegean Sea. Under glacial boundary conditions, high-diversity benthic foraminiferal faunas and elevated benthic foraminiferal numbers indicate enhanced organic matter availability and well-ventilated deep-water masses in the whole Aegean Sea. The glacial termination is accompanied by significant fluctuations in productivity and deep-water ventilation. In the northern Aegean Sea, meltwater inflow from mountain glaciers during the Bølling/Allerød warm period resulted in a restriction of local deep-water formation, as mirrored by a dominance of bolivinids. During the deposition of sapropel S1, drops in benthic foraminiferal number and diversity are more significant in the southern Aegean Sea when compared to the north. This suggests the persistence of local deep-water formation in the northern Aegean Sea during S1 deposition. In addition, faunal fluctuations within S1 at both sites suggest the repeated influence of short-term cooling events on the re-ventilation and re-colonization of Aegean deep-sea ecosystems. During the middle and late Holocene, benthic foraminiferal faunas document the establishment of oligotrophic and well-ventilated conditions in the southern Aegean Sea. The corresponding faunas from the northern Aegean Sea reflect generally mesotrophic conditions and variable deep-water oxygenation. During the entire Holocene, the deep-sea ecosystems of this region responded very sensitively to short-term changes in humidity and temperature. These abrupt climate changes controlled the inflow of nutrients from rivers and the Black Sea and the formation of local deep-water masses.     

General index

Abstract

Phytobenthos from Eastern Mediterranean Sea. — The A. states the results of an exploration, by diving techniques, in the Eastern Mediterranean Sea, along 13 stations located in the midle-western and southern region of the Aegean Sea. Besides to the values of various ecology elements (saltness, brightness, nutrition salts), informations about reologic regime in the Aegean Sea are reported. It has been collected 175 floral entities, with the following per cent composition: Chlorophyceae 14,5%, Phoeophyceae 17,0%, Rhodophyceae 68,3%. Eastern Mediterranean Sea underwater flora presents in the explored stations a R/P index equal to 4, which characterizes it as a subtropical one.     

Zooplankton distribution and diversity in a frontal area of the Aegean Sea

This study was carried out in the northern Aegean Sea during late summer and spring. The aim was to examine the spatial and temporal distribution of the zooplankton community across a dynamic frontal area and to investigate how the oceanographic heterogeneity structures the composition of the zooplankton assemblages. The low-salinity and cold Black Sea water coming from the Dardanelles Strait is modified by mixing with the underlying warm and saline Aegean water. These hydrological features result in a pronounced thermohaline front in the northern Aegean Sea throughout the year. In both seasons, zooplankton was collected using both 45 and 200 µm mesh plankton nets. A high abundance of zooplankton was observed in the surface layer at the stations closest to the Dardanelles Strait on the stratified side of the front. The zooplankton distribution and community structure in the northern Aegean Sea were strongly influenced by the hydrological features. The frontal structure acts as a boundary for the zooplankton community. The surface layer at the stratified stations had the lowest copepod diversity, from where it increased with depth and horizontally as the stratification weakened outside of the front. The total abundance of zooplankton collected with the 200 µm net was between two and 20 times lower than samples taken with the 45 µm net. The most pronounced differences were observed for the adults and copepodids of the small genera Oithona, Oncaea and Microsetella. Thus, to manage and understand the transfer of primary production up the food chain in the Aegean Sea, the smaller fraction of copepods should be taken into account in future investigations.     

Allozyme differentiation of bonito in the Mediterranean Sea

Bonito Sarda sarda samples collected in three areas from the Mediterranean Sea exhibited considerable genetic heterogeneity over all locations. This provided preliminary evidence for two different groups of bonito, one for the Ligurian and the Ionian Seas and the other for the Aegean Sea.     

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.     

Demosponge distribution in the eastern Mediterranean: a NW–SE gradient

The purpose of this paper was to investigate patterns of demosponge distribution along gradients of environmental conditions in the biogeographical subzones of the eastern Mediterranean (Aegean and Levantine Sea). The Aegean Sea was divided into six major areas on the basis of its geomorphology and bathymetry. Two areas of the Levantine Sea were additionally considered. All available data on demosponge species numbers and abundance in each area, as well as their vertical and general geographical distribution were taken from the literature. Multivariate analysis revealed a NW–SE faunal gradient, showing an apparent dissimilarity among the North Aegean, the South Aegean and the Levantine Sea, which agrees with the differences in the geographical, physicochemical and biological characteristics of the three areas. The majority of demosponge species has been recorded in the North Aegean, while the South Aegean is closer, in terms of demosponge diversity, to the oligotrophic Levantine Sea. The number of studies in the Aegean and Levantine subareas was positively correlated with the number of species recorded within each Aegean subarea. Demosponge species with an Altanto-Mediterranean distribution prevailed in the Aegean and the Levantine. The reduced contribution of the endemic component, as compared to the western Mediterranean, is consistent with the general NW–SE decrease in the number of endemic species in the Mediterranean. Demosponge distribution at the order level showed also a NW–SE gradient, similar to that observed in the entire Atlantic–Mediterranean region, suggesting a warm water affinity. Sublittoral, circalittoral and bathyal zones were clearly distinguishable in the Aegean Sea on the basis of their sponge fauna. The total number of species was an exponential negative function of depth.     

The use of otolith shape and chemistry to determine stock structure of Mediterranean horse mackerel Trachurus mediterraneus (Steindachner)

Otolith shape and chemistry of Mediterranean horse mackerel Trachurus mediterraneus were simultaneously used to assess the feasibility of using these natural tags to discriminate populations throughout the Black, Marmara, Aegean and eastern Mediterranean Seas. Otolith shape and chemistry analyses showed a similar pattern of differentiation between T. mediterraneus stocks, revealing a clear discreteness of the middle Black Sea (Sinop) and Aegean Sea (Izmir) samples. Otolith upper side length and width, and Na, K, Mg and Ba, were the morphological traits and trace elements, respectively, differing most among groups. Overall assignment of individuals into their original sample was high for both otolith shape and chemistry. Highest reclassification rate was observed for the south-middle Black Sea and Aegean Sea samples for both analyses. Hierarchical cluster analyses also supported high differentiation of the south-middle Black Sea and Aegean Sea samples for both analyses. Mantel's test revealed that the Euclidean distance both for otolith shape ( r =−0·0917, P > 0·05) and chemistry ( r =−0·1248, P > 0·05) between these populations were not significantly associated with their geographical distances.     

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.     

Genetic diversity of sharpnose mullet <Emphasis Type="Italic">Liza saliens</Emphasis> introduced in southern Caspian Sea in comparison with one native Aegean Sea population

Sharpnose mullet, Liza saliens (Risso, 1810) is one of the valuable species in Caspian shoreline and it was first introduced to Caspian Sea from Black Sea between 1930 and 1934. In the present study, we used six microsatellite markers to obtain genetic information for L. saliens from four localities of southern Caspian Sea and for one native population from Greece coastline (northern Aegean Sea). Results showed lower number of alleles (Na = 5.7–6) and higher observed heterozygosity (Ho = 0.74–0.83) in Caspian Sea samples, related to the native Aegean population (Na = 8, Ho = 0.68). Significant deviations from Hardy-Weinberg expectations were detected in 19 out of 28 tests. Low genetic differentiation (provided Fst values) among samples was detected, but Behshahr sample was significantly differentiated from all the others. The UPGMA tree revealed three major clusters: the Behshahr sample and Greek population were clustered in two different clades, and the remaining three samples formed another cluster. The low genetic variation and spatial differentiation among Caspian Sea samples may be justified by the recent establishment.     

Pelagic-benthic coupling and diagenesis of nucleic acids in a deep-sea continental margin and an open-slope system of the Eastern Mediterranean

Downward fluxes of nucleic acids adsorbed onto settling particles play a key role in the supply of organic phosphorus and genetic material to the ocean interior. However, information on pelagic-benthic coupling, diagenesis, and processes controlling nucleic acid preservation in deep-sea sediments is practically nonexistent. In this study, we compared nucleic acid fluxes, sedimentary DNA and RNA concentrations, and the enzymatically hydrolyzable fraction of DNA in a bathyal continental margin (North Aegean Sea) and an open-sea system (South Aegean Sea) of the Eastern Mediterranean. The two systems displayed contrasting patterns of nucleic acid fluxes, which increased significantly with depth in the North Aegean Sea and decreased with depth in the South Aegean Sea. These results suggest that in continental margin and open-ocean systems different processes control the nucleic acid supply to the sea floor. Differences in nucleic acid fluxes were reflected by nucleic acid concentrations in the sediments, which reached extremely high values in the North Aegean Sea. In this system, a large fraction of DNA may be buried, as suggested by the large fraction of DNA resistant to nuclease degradation and by estimates of burial efficiency (ca. eight times higher in the North than in the South Aegean Sea). Overall, the results reported here suggest that the preservation of DNA in deeper sediment layers may be favored in benthic systems characterized by high sedimentation rates.     

Pelagic-Benthic Coupling and Diagenesis of Nucleic Acids in a Deep-Sea Continental Margin and an Open-Slope System of the Eastern Mediterranean

Downward fluxes of nucleic acids adsorbed onto settling particles play a key role in the supply of organic phosphorus and genetic material to the ocean interior. However, information on pelagic-benthic coupling, diagenesis, and processes controlling nucleic acid preservation in deep-sea sediments is practically nonexistent. In this study, we compared nucleic acid fluxes, sedimentary DNA and RNA concentrations, and the enzymatically hydrolyzable fraction of DNA in a bathyal continental margin (North Aegean Sea) and an open-sea system (South Aegean Sea) of the Eastern Mediterranean. The two systems displayed contrasting patterns of nucleic acid fluxes, which increased significantly with depth in the North Aegean Sea and decreased with depth in the South Aegean Sea. These results suggest that in continental margin and open-ocean systems different processes control the nucleic acid supply to the sea floor. Differences in nucleic acid fluxes were reflected by nucleic acid concentrations in the sediments, which reached extremely high values in the North Aegean Sea. In this system, a large fraction of DNA may be buried, as suggested by the large fraction of DNA resistant to nuclease degradation and by estimates of burial efficiency (ca. eight times higher in the North than in the South Aegean Sea). Overall, the results reported here suggest that the preservation of DNA in deeper sediment layers may be favored in benthic systems characterized by high sedimentation rates.     

Genetic and morphological analyses of tub gurnard Chelidonichthys lucerna populations in Turkish marine waters

Genetic and morphological structure of tub gurnard Chelidonichthys lucerna populations in Turkish marine waters were investigated with mtDNA sequencing of 16S rRNA and morphological characters. C. lucerna samples were collected from the Black Sea, Marmara, Aegean and northeastern Mediterranean coasts of Turkey. The lowest genetic diversity was found in the northeastern Mediterranean (Iskenderun Bay) population, while the highest was in the Marmara population with overall average value of genetic diversity within populations. A total of 14 haplotypes was found, and the highest haplotype diversity was in the Black Sea whereas the lowest was in the northeastern Mediterranean population (Iskenderun Bay). The Black Sea and Iskenderun Bay populations showed the least genetic divergence (0.001081), while the highest was between the Marmara Sea and northeastern Mediterranean (Antalya Bay) populations (0.002067). Pairwise comparisons of genetic distance revealed statistically significant differences (P < 0.05) between the Marmara and both the Aegean and northeastern Mediterranean (Antalya Bay) samples. Neighbour joining tree analyses clustered the northeastern Mediterranean populations (Antalya Bay and Iskenderun Bay) as genetically more interrelated populations, whereas the Aegean Sea population was clustered as most isolated one. Discriminant function analysis of morphological characters showed that only the Black Sea population is differentiated from the other populations.     

The taxonomic status of the Black Sea and Marmara Sea populations of the Broadnosed Pipefish Syngnathus cf. argentatus Pallas (Teleostei: Syngnathidae) based on morphological and molecular characters

As the Black Sea and Marmara Sea population of the Broadnosed Pipefish Syngnathus cf. argentatus show some morphological differences from the Mediterranean Sea populations, some authors regard it as an endemic species Syngnathus argentatus Pallas, 1814, while others consider it as a synonym of S. typhle Linnaeus, 1758. The aim of this study is to compare the populations of the Black Sea and Sea of Marmara with the Aegean Sea population, using a combination of morphological and molecular characters, in order to clarify their taxonomic status. Sampling was carried out at three stations in the Black Sea, two in the Sea of Marmara and three in the Aegean Sea, and a total of 24 morphometric and 6 meristic characters were examined. Metric data were analysed by Principal Component Analysis (PCA) and phylogenetic relationships between the populations were analysed using both cytochrome oxidase subunit 1 (COI) gene and cytochrome b (cyt-b) gene sequences. Although constant differences were observed in snout depths between the Black Sea/Marmara Sea and the Aegean Sea populations, other morphological features and genetic analysis did not enable these populations to be differentiated. These findings indicate that S. argentatus is a synonym of S. typhle.     

Microsatellite DNA reveals genetically different populations of Atlantic bonito Sarda sarda in the Mediterranean Basin

Microsatellites were used to investigate population genetic structure of Atlantic bonito Sarda sarda from the Black Sea, Marmara Sea, Aegean Sea, north-eastern Mediterranean Sea and Adriatic Sea. Overall average observed heterozygosity was high (0.93). Average observed heterozygosity per locus ranged from 0.79 to 0.98. Pairwise F_ST estimates for all loci between populations ranged from 0 to 0.07626, and significant F_ST values (P < 0.001) were detected between populations; the Blacks Sea and Marmara Sea samples were not significantly different from each other, but significant different from the other samples, and Aegean Sea and north-eastern Mediterranean Sea samples were also not significantly different from each other, but significantly different from all other samples. The Adriatic Sea sample was significant different from all other samples. The Mantel test revealed a significant (P < 0.001, r = 0.68) isolation-by-distance for these 11 populations. Neighbour-joining analysis clustered the Black Sea and Marmara Sea samples together while collections from Aegean Sea and north-eastern Mediterranean Sea were clustered close to each other and far from the others. On the other hand, the Adriatic Sea collection presented very distinctive relationship from the others.     

Occurrence of the Erythrean invader Pteragogus pelycus Randall, 1981 (Teleostei: Labridae) in the eastern Aegean sea

The occurrence of the Erythrean fish Pteragogus pelycus, off Kusadasi Bay, Turkey, eastern Aegean Sea, is here reported for the first time. One specimen was collected and it is now maintained in aquarium. The finding shows that its population is expanding northwards, along the Anatolian coasts of the Aegean Sea, approaching the colonization of the whole south and central basin.     

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     

Habitat selection and temporal abundance fluctuations of demersal cartilaginous species in the Aegean Sea (eastern Mediterranean)

Predicting the occurrence of keystone top predators in a multispecies marine environment, such as the Mediterranean Sea, can be of considerable value to the long-term sustainable development of the fishing industry and to the protection of biodiversity. We analysed fisheries independent scientific bottom trawl survey data of two of the most abundant cartilaginous fish species (Scyliorhinus canicula, Raja clavata) in the Aegean Sea covering an 11-year sampling period. The current findings revealed a declining trend in R. clavata and S. canicula abundance from the late '90 s until 2004. Habitats with the higher probability of finding cartilaginous fish present were those located in intermediate waters (depth: 200-400 m). The present results also indicated a preferential species' clustering in specific geographic and bathymetric regions of the Aegean Sea. Depth appeared to be one of the key determining factors for the selection of habitats for all species examined. With cartilaginous fish species being among the more biologically sensitive fish species taken in European marine fisheries, our findings, which are based on a standardized scientific survey, can contribute to the rational exploitation and management of their stocks by providing important information on temporal abundance trends and habitat preferences.     

Genetic diversity and phylogenetic origin of brown trout Salmo trutta populations in eastern Balkans

The study focuses on the phylogenetic origin and genetic diversity of brown trout in the eastern part of the Balkan Peninsula. It further aims to reveal the impact of human-mediated transfers and stocking with non-indigenous trout on the populations in this area. For these purposes, mtDNA control region and microsatellite variation of 204 individuals from 16 populations were analysed. The results indicate that mtDNA haplotypes from the lower Danube basin and southern Black Sea basins differ substantially from a subclade of the Danubian lineage consisting of haplotypes found so far in the most of the Danube basin and in the Caspian and Aral Sea basins. Considering also the results of demographic analyses, this study evidences a complex evolutionary history of brown trout in the southern and western parts of the Black Sea basin. In the Aegean Sea basin, a high frequency of the central haplotype of Adriatic mtDNA lineage has been found. The other Adriatic lineage haplotypes found in this basin differ from the central haplotype by one mutational step only, indicating a recent evolution of the Adriatic lineage in the Aegean Sea basin. Substantial genetic differentiation among populations and basins was revealed. The hybridization with Atlantic brown trout was indicated in both sea basins, but especially in the Danube basin. Compared to other European regions, it can be inferred that the introgression of exogenous brown trout in the eastern Balkan populations is rather low.     

Extant silicoflagellates from the Northeast Aegean (eastern Mediterranean Sea): Morphologies and double skeletons

Silicoflagellate abundance, vertical distribution and morphology were studied during spring (March 2014) at three sampling stations located in the Northeast Aegean Sea adjacent to the Dardanelles Strait and characterized by a variable influx of cold, low-salinity Black Sea water. The silicoflagellate assemblage was dominated by Dictyocha stapedia and Stephanocha speculum with minor contribution of D. aculeata and Octactis pulchra. While specimens of D. stapedia were represented by the typical morphologies described in other areas of the Mediterranean Sea, populations of S. speculum displayed peculiar characters: they were large, predominantly 7-sided, with a small apical ring as well as apical ring spines, concave basal ring sides and non-rotated apical structure. Some of these features have been described for S. speculum at high latitudes, but the combined characters make these specimens slightly different from the high latitude populations. Similar morphologies have been observed in the western Black Sea, thus we can infer that the peculiar specimens detected in the Northeast Aegean are associated with the influx of Black Sea water masses.