Biodiversity issues for the forthcoming tropical Mediterranean Sea

Present-day Mediterranean marine biodiversity is undergoing rapid alteration. Because of the increased occurrence of warm-water biota, it has been said that the Mediterranean is under a process of ‘tropicalization’. This paper analyses the main patterns of the Mediterranean Sea tropicalization and considers briefly its extent and consequences. As happened during previous interglacial phases of the Quaternary, Atlantic water, entering via the Straits of Gibraltar, carries into the Mediterranean species that are prevalently of (sub)tropical affinity. On the other side of the basin, Red Sea species penetrate through the Suez Canal, a phenomenon called lessepsian migration from the name of F. de Lesseps, the French engineer who promoted the cutting of the Canal. Also the many exotic species introduced by humans voluntarily or involuntarily are nearly always typical of warm waters. Climate change combines with Atlantic influx, lessepsian migration and the introduction of exotic species by humans to the establishment of tropical marine biota in the Mediterranean Sea. Present-day warming ultimately favours the spread of warm-water species through direct and indirect effects, and especially by changing water circulation. It is impossible at present to foresee to what extent the exuberance of warm-water species will affect the trophic web and the functioning of marine ecosystems in the Mediterranean Sea of tomorrow. While Mediterranean Sea communities are modifying their pattern of species composition, they do not seem to be acquiring a more marked tropical physiognomy: Mediterranean coastal marine ecosystems are still dominated by frondose algae (even if the species that are gaining ascendancy are of tropical origin) and not by corals as is normal in tropical seas.     

Seasonal variability of copepod abundance in the Balearic region (Western Mediterranean) as an indicator of basin scale hydrological changes

Spatial and temporal changes of the copepod community have been investigated and related to the environmental variability of the Balearic Sea (Western Mediterranean). The period studied spans from 1994 to 1999 during which we analyzed the abundance and structure copepod variability over a cross-shore transect. Results showed a close link between hydrological changes and the variations of copepod abundance. The synchronous variability of copepods and hydrography indicated the rapid response of this zooplankton group to the inflow of cold and warm water masses coming through the study area. Cluster analysis revealed four main copepod assemblages that distinguished the coastal from the oceanic species and those species with different water masses preference. The copepod assemblage composed of Calanus helgolandicus, Clausocalanus arcuicornis, C. pergens, C. paululus, Calocalanus tenuis and Pleuromamma gracilis was associated with cool salty waters, whereas the assemblage formed by Temora stylifera, C. pavo, C. styliremis, Centropages bradyi and Acartia danae was related to warmer less saline Mediterranean waters. Moreover, it is suggested that changes in sea water temperature and salinity are linked to large-scale changes likely occurring at a basin scale, which is reflected in the Western Mediterranean mesoscale hydrographic changes. Therefore, it is stressed that changes in the Balearic copepod community can be used as potential tracers of the western Mediterranean water masses. Guest editors: S. Souissi & G. A. Boxshall Copepoda in the Mediterranean: Papers from the 9th International Conference on Copepoda, Hammamet, Tunisia     

First record of a rare pelagic octopod,Ocythoe tuberculata (Cephalopoda: Ocythoidae), in Korea

The family Ocythoidae is found throughout the temperate and subtropical waters of the world’s ocean and contains a single species, Ocythoe tuberculata. Recently, a single female specimen of the rare pelagic species O. tuberculata was collected off Uljin in the East Sea/Sea of Japan of Korea by squid jigging for the first time. The appearance of this species could be considered as evidence of the changes in species distribution of sea surface. However, further studies are required to investigate whether this phenomenon is temporary or continuous.     

Spatial distribution and life‐history aspects of blackspot seabream Pagellus bogaraveo (Osteichthyes: Sparidae)

Spatial distribution and life history aspects of Pagellus bogaraveo in the eastern Ionian Sea were investigated using the data from 13 different studies carried out in the area from 1983 to 2010. The spatial patterns of the abundance, biomass and mean size showed that the species inhabits the shallow waters of the shelf (<170 m depth) as juveniles up to a certain size (<180 mm total length, L_T), moving to deeper waters of the slope (mainly 400–500 m depth) as adults. The spatial pattern of abundance indicated a continuous distribution of the species in deep waters, with hot‐spot areas of high values, whereas in shallow waters distribution was more discontinuous, with higher concentrations of juveniles in estuaries and brackish waters. The study of biological aspects of the species revealed (1) a difference in the increase in mass between males and females, (2) protandrous hermaphroditism, accompanied by the presence of primary females and males that do not change sex, (3) a sex ratio in favour of females >250 mm L_T, (4) the presence of hermaphrodites between 200 and 370 mm, (5) a long reproduction period from June to March, (6) a size at first maturity around 300 mm and (7) a diet composition of adults based mainly on fishes, and also on opportunistic behaviour in the food scarce environment of deep waters. The results suggest that the species' distribution and feeding strategies are the most appropriate for the oligotrophic eastern Ionian waters and that these conditions result in smaller sizes of the species in the east Mediterranean Sea compared to the west basin and the east Atlantic Ocean, with implications for the growth and reproductive biology of the species.     

Preliminary study of summer phytoplankton in coastal waters of the Gulf of Taranto (Ionian Sea)

Abstract

Phytoplankton abundances and species composition of coastal waters of the Gulf of Taranto (Ionian Sea) were investigated in July 1995. Abundances were lower than those of other Mediterranean coastal areas. Higher densities values were found in the central part of the studied area. In surface waters, small phytoflagellates (<10μm) and dinoflagellates, such as Heterocapsa niei and Oxytoxum variabile, were the dominant species. Diatoms, mainly belonging to the genus Chaetoceros, were also present.     

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 biological production of marine suspended barite and the barium cycle in the Western Mediterranean Sea

Suspended particulate barium was measured in the Western Mediterranean along 4 profiles sampled during the PHYCEMED 1 cruise in 1981. The non-terrigenous fraction of particulate barium (i.e. excess Ba; Ba_xs) can account for up to 96% of total barium. This fraction follows the barite settling and dissolution rate model we described earlier for the Atlantic Ocean, confirming barite as the carrier of excess barium. We propose that these Ba_xs concentrations, which are higher in the Western Mediterranean than in the overall Atlantic, may represent an adaptation of the microplankton to the high densities and density gradients of surface Mediterranean waters.A mass-balance for Ba in the Mediterranean Sea shows: (1) that the atmospheric source must be much more important than previously reported: (2) that contrary to the Atlantic situation, the internal recycling of barium is of little importance compared to the flow of barium through the system.     

The effects of northern italian rivers and Eastern Mediterranean ingressions on the phytoplankton of the Adriatic Sea

An analysis of the distribution of the phytoplankton standing crop during a period of high stability characterized the relative influence of northern Italian rivers and Mediterranean waters on the Adriatic Sea. The region influenced by northern Italian rivers was marked by low community diversities, relatively high chlorophyll a standing crops and cell densities, high microplankton to nannoplankton community volume ratios, and the dominance of Nitzschia seriata. To the contrary, the region influenced by Mediterranean waters exhibited high community diversities with prominent oceanic and warm water elements, low chlorophyll a standing crops and microplankton cell densities, low microplankton to nannoplankton ratios, and a relatively conspicuous coccolithophore flora.Nannoplankton populations remained relatively constant throughout the Adriatic. However eutrophication resulted in a marked increase in the microplankton component of the community, with the diatom Nitzschia seriata serving as a prime indicator of the process, even in regions several hundred kilometers downstream from nutrient sources.Even under relatively low rates of river discharge, the influence of northern Italian rivers could be traced along the entire western sode of the sea to the Otranto Strait, with low north to south and high west to east gradients resulting from the effect of a cyclonic surface circulation pattern on inflowing Mediterranean waters in the south and river discharge in the north.     

Phytoplankton biomass and species composition in a Mediterranean coastal lagoon

Phytoplankton populations were investigated weekly at a central station in the Fusaro lagoon (Mediterranean Sea) from 27 November 1989 to 18 June 1990 to assess species composition, temporal succession and standing stock of the different species. Chlorophyll concentrations varied from 1.2 to 73.2 µg 1^–1 in surface waters, and from 1.3 to 53.5 µg 1^–1 at the 4.5 m depth. Phytoplankton communities were dominated by Prorocentrum micans Ehrenberg in December and January, and by small-sized diatoms in the rest of the sampling period. In surface waters, a maximum biomass of 9.5 mg C 1^–1 was measured in January, in correspondence with high concentrations (8.1 × 10⁶ cells 1^–1) of P. micans, whereas an abundance peak of 159.9 × 10⁶ cells 1^–1 was registered on the last sampling date due to a massive bloom of a very small diatom, Minutocellus polymorphus (Hargraves & Guillard) Hasle, von Stosch & Syvertsen. On the whole, phytoplankton populations of the Fusaro lagoon showed distinct characters as compared to those of southern Tyrrhenian coastal waters and of other lagoons.     

Calcareous dinoflagellate cysts in surface sediments from the Mediterranean Sea: distribution patterns and influence of main environmental gradients

The distribution of calcareous dinoflagellate cysts in surface sediments from the Mediterranean Sea was quantitatively analysed. The samples contain 11 cyst species and the vegetative coccoid Thoracosphaera heimii. Cyst abundance increases towards the deeper parts of the basins and is generally higher in the eastern Mediterranean Sea. Three major distribution characteristics exist: (1) different assemblages in oceanic and neritic regions, (2) little agreement with the associations of areas studied so far like the Atlantic Ocean, and (3) a unique oceanic assemblage in the eastern Mediterranean Sea. A gradual change in cyst assemblages from the western to the eastern Mediterranean Sea was observed and statistically compared with the main environmental gradients in the upper water column. Temperature, nitrate concentration and possibly salinity appear to be the most important factors controlling cyst production. Three groups containing cysts with similar environmental preferences can be distinguished: (1) an eastern Mediterranean group related to relatively high temperature and salinity but low nitrate concentration, (2) a group of more or less consistently abundant cosmopolitan species tolerating or even preferring relatively low temperature and salinity but high nitrate concentration, and (3) a group containing species that are possibly adapted to neritic environments and have probably been transported from coastal areas into the studied regions. In contrast to other calcareous plankton, calcareous dinoflagellate cysts correlate strongly with the main environmental gradients in the Mediterranean Sea, bearing a high potential for palaeoenvironmental reconstructions.     

Paleoenvironmental evolution of the eastern Mediterranean during the Messinian: Constraints from integrated microfossil data of the Pissouri Basin (Cyprus)

Integrated data of calcareous nannofossils, as well as planktonic and benthic foraminifera from the Pissouri Motorway section on Cyprus allow the reconstruction of surface- and bottom-water paleoenvironments of the eastern Mediterranean during the interval preceding the Messinian salinity crisis (MSC). Contrary to deeper-water locations, where benthic foraminifera faunas are suppressed or absent just after the Tortonian–Messinian boundary, sediments deposited at intermediate water depths do contain benthic assemblages. From the earliest Messinian onwards, a development towards increasingly unfavourable paleoenvironments is reflected in the planktonic and benthic microfossil records of the Pissouri section and proceeds with rather discrete time steps that can be correlated to sequences throughout the Mediterranean. Shortly after the Tortonian–Messinian boundary a transition is recorded in the sedimentology and the open marine, deeper-water taxa disappear from the benthic foraminifera assemblages; subsequently, the diversity of all fauna groups diminishes. The changes recorded at species level in both surface-water and sea-floor dwelling taxa suggest decreasing circulation of the bottom waters, associated with changes in the surface waters, most likely due to increasing stratification. From ∼6.73 Ma onwards, our data indicate a prominent change to more restricted conditions and increasing salinity at the sea floor together with intermittently rising surface water salinity. The dominance of oligotypic and monospecific assemblages and the frequent shifts in assemblage compositions of all microfossil groups indicate severely stressed environments after ∼6.4 Ma, probably related to increased salinity. The major changes in paleoenvironmental conditions, including oxygen deprivation due to stagnation and hypersalinity, can be explained by hydrographical changes in the Mediterranean basin, which are probably caused by tectonic movements in the Rif Corridor acting in concert with astronomical cyclicity. Evaluation of the paleodepth proxies indicates that the depth of the Pissouri Basin remained rather constant at ∼300–500 m, with a minimum of 200 m, until deposition of the “barre jaune”, the transitional interval towards the evaporites and that early shallowing to neritic depths, as was proposed before, is highly unlikely.     

Distribution of the Eelpout Lycodes soldatovi(Pisces: Zoarcidae) over the Continental Slope of the Sea of Okhotsk

The eelpout Lycodes soldatovi was found at depths of 153 to 1005 m in the Sea of Okhotsk; however, it usually inhabits the depth range of 400 to 800. The mean and maximum abundance and the biomass of this species were 198.5 and 1037 individuals per km² and 173.7 and 1275 kg/km², respectively. In view of the data on the distribution of young fish (up to 30 cm in total body length) and the reports on the absence of Lycodes larvae, a hypothesis was developed that this species spawns predominantly in the waters off western Kamchatka and eastern Sakhalin at the depth range of 700 to 900 m. Large individuals (>50 cm) exhibit higher migratory activity and are more tolerant of environmental conditions. That is why they are encountered throughout the distribution area of this species, even in subzero temperature areas. Lycodes soldatovi seems to be a typical inhabitant of the Sea of Okhotsk, except for the waters off the South and Central Kuril Islands and the Pacific waters off the North Kuril Islands.     

Population genetics,gene flow,and biogeographical boundaries of Carcinus aestuarii (Crustacea: Brachyura: Carcinidae) along the European Mediterranean coast

Carcinus aestuarii Nardo, 1847 is a widespread coastal crab species throughout the Mediterranean Sea with a pelagic larval phase. This species tolerates a wide range of environmental conditions and typically inhabits fragmented habitats, such as embayments, lagoons and estuaries. It is therefore a good candidate species for studying and testing different phylogeographical hypotheses in the Mediterranean Sea. By contrast to its Atlantic sister species, Carcinus maenas, studies on the population genetic structure of C. aestuarii in its native range are still scarce. In the present study, specimens from along the European Mediterranean Sea were collected and DNA‐sequenced and analyses were applied to discriminate between present day and historical factors influencing the population genetic structure of this species. The results obtained demonstrate the existence of two genetically distinct geographical groups, corresponding to the eastern and western Mediterranean, with further subdivision within the East Mediterranean Basin. A strong asymmetric gene flow was recorded toward the Eastern Basin, which may play a crucial role in shaping the present day biogeographical patterns of this species and potentially other sympatric ones with pelagic larvae. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 771–790.     

Palaeoecology of Upper Pleistocene-Holocene bryozoan and foraminiferal assemblages from Ku?dili (Kadiköy, Istanbul, Turkey)

Bryozoan and foraminiferal assemblages from a core drilled in the Kusdili (Kadiköy, Istanbul) Upper Pleistocene-Holocene sequence were studied. Assemblages are absent from the very basal and top parts but they are well represented all along the middle part of the sequence. All the species found presently live in the Mediterranean. In the area sedimentation seemingly started about 13?000 years ago in very shallow near-shore environments with the deposition of pebbly sediments rapidly evolving to mud. Near the base, the presence of erosional surface allows to recognize three successive sedimentary cycles. Bryozoan and foraminiferal assemblages found in the middle part of the sequence testify to a shallow water, estuarine or lagoonal paleo-environment, in which effects of low salinity waters, more obvious toward the top, could be related to the reestablishment of a connection between the Marmara Sea and the Black Sea through the Bosphorus, about 7000 years BP.     

Tree species from contrasting hydrological niches show divergent growth and water-use efficiency

Tree species occupy different hydrological niches and climate warming may affect tree performance in those niches through increased drought stress. However, the effects of climate warming on growth, carbon and water fluxes would differ depending on species’ hydrological niche. Species from wet sites should show a lower growth dependence on precipitation and also lower intrinsic water-use efficiency (WUEi), as compared with species from dry sites which should improve more the WUEi. We test these ideas by comparing radial-growth rates (basal-area increment), climate- and drought-growth associations and WUEi of hackberry (Celtis australis) vs. Aleppo pine (Pinus halepensis) and maritime pine (Pinus pinaster) in two Mediterranean sites located in Spain. Species are subjected to similar regional climate conditions in each site but occupy contrasting local hydrological niches (hackberry in wet sites and pines in dry sites). Climate is warming in both study sites and drought-avoiding pines are responding by showing higher growth rates and improved WUEi. We also found a similar growth dependency on winter-spring precipitation and summer drought of all species and sites and comparable WUEi values and trends, excepting in hackberry from southern Spain which grew more, and showed a higher growth resistance to drought and lower and more stable WUEi values. Variables inferred from tree rings as growth rates and WUEi allow characterizing the hydrological niche of tree species, which may be contingent on site conditions and climate warming.     

Life history strategies of four small-size fishes in the Suez Canal, Egypt

Of 35 species collected from the shores of the Suez Canal and its lakes, five were abundant year round. Sihouettea aegyptia and Aphanius dispar are of Red Sea (warm water) origin, whereas Pomatoschistus marmoratus, Aphanius fasciatus and Atherina boyeri are Mediterranean species. S. aegyptia and A. dispar are distributed throughout the Suez system whereas A. fasciatus is restricted to the northern canal. P. marmoratus and A. boyeri have not spread southward beyond the Bitter Lakes. Salinity is the main limiting factor for the distribution of Mediterranean species. S. aegyptia and A. dispar are dominant in the Bitter Lakes (salinity 44‰), whereas P. marmoratus and A. boyeri are abundant in Timsah Lake (salinity 7·8–41·6‰). There was considerable interannual and monthly variation in the relative abundance of each species. The populations were dominated by a single age group, and life spans were no more than 2 years. Rapid growth was evident during the first spring in P. marmoratus and A. boyeri and during the first summer, early autumn in S. aegyptia and A. dispar . The relative abundance of each food item in the diet varied with fish size and season. S. aegyptia, P. marmoratus and A. dispar fed mostly on harpacticoid copepods, polychaetes, demersal eggs, diatoms and blue-green algae whereas A. boyeri fed mostly on planktonic copepods. The warm water species are summer spawners, whereas the temperate species are autumn-winter spawners.     

Breeding phenology and larval distribution of amphibians in a Mediterranean pond network with unpredictable hydrology

Understanding spatial and temporal breeding patterns in Mediterranean amphibian communities is urgent considering the rate of habitat loss. Breeding phenology and breeding habitat selection by amphibians were analysed through the monthly occurrence of larvae in a mosaic of 198 Mediterranean temporary ponds during three years. A generalized linear model (GLM) coupled with principal component analysis showed that, for almost all species, occurrence was significantly positively correlated to pond depth. In addition, pond openness negatively affected the presence of some species. Temporal breeding patterns varied among species. Some species exhibited flexibility in their breeding date (Pelobates cultripes, Pelodytes punctatus, Hyla meridionalis, Rana perezi), while others did not (Triturus marmoratus, Triturus helveticus, Bufo calamita). When faced with inter-annual hydrological variability, the first group had a more constant breeding success than the second. Variable hydrological conditions caused differential larval occurrence of species between years. These fluctuations might favour long-term persistence of the whole amphibian community. We finally discuss the implications of our results for the management of amphibian habitats in the Mediterranean region.     

A survey on bacteria inhabiting the sea surface microlayer of coastal ecosystems

Bacterial populations inhabiting the sea surface microlayer from two contrasted Mediterranean coastal stations (polluted vs. oligotrophic) were examined by culturing and genetic fingerprinting methods and were compared with those of underlying waters (50 cm depth), for a period of two years. More than 30 samples were examined and 487 strains were isolated and screened. Proteobacteria were consistently more abundant in the collection from the pristine environment whereas Gram-positive bacteria (i.e., Actinobacteria and Firmicutes) were more abundant in the polluted site. Cythophaga-Flavobacter-Bacteroides (CFB) ranged from 8% to 16% of total strains. Overall, 22.5% of the strains showed a 16S rRNA gene sequence similarity only at the genus level with previously reported bacterial species and around 10.5% of the strains showed similarities in 16S rRNA sequence below 93% with reported species. The CFB group contained the highest proportion of unknown species, but these also included Alpha- and Gammaproteobacteria. Such low similarity values showed that we were able to culture new marine genera and possibly new families, indicating that the sea-surface layer is a poorly understood microbial environment and may represent a natural source of new microorganisms. Genetic fingerprinting showed, however, no consistent differences between the predominant bacterial assemblages from surface microlayer and underlying waters, suggesting that the presence of a stable and abundant neustonic bacterial community is not a common trait of coastal marine environments.     

Comparative composition and dynamics of harmful dinoflagellates in Mediterranean salt marshes and nearby external marine waters

The taxonomic structure of phytoplankton populations in two Mediterranean coastal lagoons were compared with those of nearby marine waters (external waters). Mediterranean confined lagoons remain isolated for most the year and concentrate phytoplankton to a very high biomass. Coastal lagoons on the Mediterranean may, therefore, act as accumulators of neritic phytoplankton (including species related to harmful algal blooms). We examined whether coastal lagoons act as concentrators of marine toxic dinoflagellates during confinement periods, and the common environmental factors that favour growth of specific harmful species in the two ecosystems considered: coastal lagoons and external waters. An alternation between the dominance of diatoms and dinoflagellates was observed, coinciding with that described in Margalef's mandala, occurring in external waters as well as in coastal lagoons. Moreover, the temporal patter was different in the two ecosystems. Dinoflagellate species composition and their bloom period were highly variable in time and space, thus, species had to be analysed individually. Most of the dinoflagellate species found in this study were potentially harmful and high biomass producers. Harmful dinoflagellate species performed well in both, external waters and lagoons, but the specific species-dependent affinity to each of these environments determined which organisms bloom there. Thus, expansion of harmful algal blooms (HAB) to inland waters is not likely and some environmental factors such as the oxidised state of available nitrogen, became determinant to the success and bloom of a species in the coastal lagoon ecosystem.