Distinctive insular forms of threespine stickleback (Gasterosteus aculeatus) from western Mediterranean islands

Neutral and adaptive variation among populations within a species is a major component of biological diversity and may be pronounced among insular populations due to geographical isolation and island specific evolutionary forces at work. Detecting and preserving potential evolutionary significant units below the species rank has become a crucial task for conservation biology. Combining genetic, phenotypic and ecological data, we investigated evolutionary patterns among the enigmatic threespine stickleback populations from western Mediterranean islands, all of which are threatened by habitat deterioration and climate change. We find indications that these populations derive from different genetic lineages, being genetically highly distinct from the stickleback of mainland Europe and the northern Atlantic as well as from each other. Mediterranean island stickleback populations are also phenotypically distinct from mainland populations but interestingly stickleback from Iceland have converged on a similar phenotype. This distinctive island stickleback phenotype seems to be driven by distinct selective regimes on islands versus continents. Overall, our results reveal the status of western Mediterranean island stickleback as evolutionarily distinct units, important for conservation of biodiversity.

High-resolution marine record of sediment core MD99-2343 in the Balearic Sea

The IMAGES core MD99-2343, recovered from a sediment drift north of the island of Minorca, in the north-western Mediterranean Sea, holds a high-resolution sequence that is perfectly suited to study the oscillations of the overturning system of the Western Mediterranean Deep Water (WMDW). Detailed analysis of grain-size and bulk geochemical composition reveals the sensitivity of this region to climate changes at both orbital and centennial-millennial temporal scales during the last 50 kyr. The dominant orbital pattern in the K/Al record indicates that sediment supply to the basin was controlled by the insolation evolution at 40°N, which forced changes in the fluvial regime, with more efficient sediment transport during insolation maxima. This orbital control also modulated the long-term pattern of the WMDW intensity as illustrated by the silt/clay ratio. However, deep convection was particularly sensitive to climatic changes at shorter time-scales, i.e. to centennial-millennial glacial and Holocene oscillations that are well documented by all the paleocurrent intensity proxies (Si/Al, Ti/Al and silt/clay ratios). Benthic isotopic records (d13C and d18O) show a Dansgaard-Oeschger (D-O) pattern of variability of WMDW properties, which can be associated with changing intensities of the deep currents system. The most prominent reduction on the WMDW overturning was caused by the post-glacial sea level rise. Three main scenarios of WMDW overturning are revealed: a strong mode during D-O Stadials, a weak mode during D-O Interstadials and an intermediate mode during cooling transitions. In addition, D-O Stadials associated with Heinrich events (HEs) have a very distinct signature as the strong mode of circulation, typical for the other D-O Stadials, was never reached during HE due to the surface freshening induced by the inflowing polar waters. Consequently, the WMDW overturning system oscillated around the intermediate mode of circulation during HE. Though surface conditions were more stable during the Holocene, the WMDW overturning cell still reacted synchronously to short-lived events, as shown by increments in the planktonic d18O record, triggering quick reinforcements of the deep water circulation. Overall, these results highlight the sensitivity of the WMDW to rapid climate change which in the recent past were likely induced by oceanographic and atmospheric reorganizations in the North Atlantic region.

Nitrification rates in the NW Mediterranean Sea

During spring, ammonium oxidation and nitrite oxidation rates were measured in the NW basin of the Mediterranean Sea, from mesotrophic sites (Ligurian Sea and Gulf of Lions) to oligotrophic sites (Balearic Islands). Nitrification rates (average values for 37 measurements) ranged from 72 to 144 nmol of N oxidised/l/d, except in the Rhône River plume area where the rates increased to 264-504 nmol/l/d because of the riverine inputs of nitrogen. Maximal rates were located around the peak of nitrite within the nitracline at about 40 to 60 m and just above the phosphacline. At 1 station, relatively high values of nitrification (50 to 130 nmol/l/d) were also measured deep in the water column (240 m). Day-to-day variations were measured demonstrating the response within a few hours to hydrological stress (wind-induced mixing of the water column) and showing the role of hydrological characteristics on the distribution of nitrification rates. Because of the homogenous temperature (13°C) in the Mediterranean Sea, the spatial (geographical and vertical) fluctuations of nitrifying rates were linked to the presence of substrate due to mineralisation processes and/or Rhône River inputs. We estimate the contribution of nitrate produced by nitrification to the N demand of phytoplankton to range from 16% at mesotrophic to 61% at oligotrophic stations.

Geochemistry of sediments cores in the Mediterranean Sea

We investigated five time-equivalent core sections (180-110 kyr BP) from the Balearic Sea (Menorca Rise), the easternmost Levantine Basin and southwest, south, and southeast of Crete to reconstruct spatial patterns of productivity during deposition of sapropels S5 and S6 in the Mediterranean Sea. Our indicators are Ba, total organic carbon and carbonate contents. We found no indications of Ba remobilization within the investigated core intervals, and used the accumulation rate of biogenic Ba to compute paleoproductivity. Maximum surface water productivity (up to 350 g C/m2/yr) was found during deposition of S5 (isotope stage 5e) but pronounced spatial variability is evident. Coeval sediment intervals in the Balearic Sea show very little productivity change, suggesting that chemical and biological environments in the eastern and western Mediterranean basins were decoupled in this interval. We interpret the spatial variability as the result of two different modes of nutrient delivery to the photic zone: riverderived nutrient input and shoaling of the pycnocline/nutricline to the photic zone. The productivity increase during the formation of S6 was moderate compared to S5 and had a less marked spatial variability within the study area of the eastern Mediterranean Sea. Given that S6 formed during a glacial interval, glacial boundary conditions such as high wind stress and/or cooler surface water temperatures apparently favored lateral and vertical mixing and prevented the development of the spatial gradients within the Eastern Mediterranean Sea (EMS) observed for S5. A non-sapropel sediment interval with elevated Ba content and depleted 18O/16O ratios in planktonic foraminifer calcite was detected between S6 and S5 that corresponds to the weak northern hemisphere insolation maximum at 150 kyr. At this time, productivity apparently increased up to five times over surrounding intervals, but abundant benthic fauna show that the deep water remained oxic. Following our interpretation, the interval denotes a failed sapropel, when a weaker monsoon did not force the EMS into permanent stratification. The comparison of interglacial and glacial sapropels illustrates the relevance of climatic boundary conditions in the northern catchment in determining the facies and spatial variability of sapropels within the EMS.

Paleoenvironmental changes in the western Mediterranean since the last glacial maximum: High resolution multiproxy record from the Algero-Balearic basin

The present study uses a multiproxy approach in order to further understand the evolution of climate responses in the western Mediterranean as of the Last Glacial Maximum. Sediments from ODP Site 975 in the Algero-Balearic basin have been analysed at high resolution, both geochemically andmineralogicallly. The resulting data have been used as proxies to establish a sedimentary regime, primary marine productivity, the preservation of the proxies and oxygen conditions. Fluctuations in detrital element concentrations were mainly the consequence of wet/arid oscillations. Productivity has been established using Ba excess, according to which marine productivity appears to have been greatest during cold events Heinrich 1 and Younger Dryas. The S1 time interval was not as marked by increases in productivity as was the eastern Mediterranean. In contrast, the S1 interval was first characterized by a decreasing trend and then by a fall in productivity after the 8.2 ky BP dry-cold event. Since then productivity has remained low. Here we report that there was an important redox event in this basin, probably a consequence of the major oceanographic circulation change occurring in the western Mediterranean at 7.7 ky BP. This circulation change led to reventilation as well as to diagenetic remobilization of redox-sensitive elements and organic matter oxidation. Comparisons between our paleoceanographic reconstruction for this basin and those regarding other Mediterranean basins support the hypothesis that across the Mediterranean there were different types of responses to climate forcing mechanism. The Algero-Balearic basin is likely to be a key area for further understanding of the relationships between the North Atlantic and the eastern Mediterranean basins.

Highly polymorphic microsatellite markers for the Mediterranean endemic fan mussel Pinna nobilis

Pinna nobilis is an endemic bivalve of the Mediterranean Sea whose populations have decreased in the last decades due to human pressure; as a consequence, it was declared a protected species in 1992. Despite its conservation status, few genetic studies using mitochondrial markers have been published. We report on the isolation and development of 10 microsatellite loci for the fan mussel, Pinna nobilis. All loci (2 di-nucleotide, 5 tri-nucleotide, 2 tetra-nucleotide and 1 penta-nucleotide) are characterized by high levels of polymorphism in 76 individuals tested from two populations in the Balearic Islands (Spain, Western Mediterranean Sea). The number of alleles ranged from 4 to 24 and expected heterozygosity ranged from 0.4269 to 0.9400. These microsatellites could be very useful for the assessment of the genetic diversity and connectivity patterns of P. nobilis and the establishment of new conservation strategies.

Phylogeography of the Atlanto-Mediterranean sea cucumber Holothuria (Holothuria) mammata: the combined effects of historical processes and current oceanographical pattern

We assessed the genetic structure of populations of the widely distributed sea cucumber Holothuria (Holothuria) mammata Grube, 1840, and investigated the effects of marine barriers to gene flow and historical processes. Several potential genetic breaks were considered, which would separate the Atlantic and Mediterranean basins, the isolated Macaronesian Islands from the other locations analysed, and the Western Mediterranean and Aegean Sea (Eastern Mediterranean). We analysed mitochondrial 16S and COI gene sequences from 177 individuals from four Atlantic locations and four Mediterranean locations. Haplotype diversity was high (H = 0.9307 for 16S and 0.9203 for COI), and the haplotypes were closely related (p = 0.0058 for 16S and 0.0071 for COI). The lowest genetic diversities were found in the Aegean Sea population. Our results showed that the COI gene was more variable and more useful for the detection of population structure than the 16S gene. The distribution of mtDNA haplotypes, the pairwise FST values and the results of exact tests and AMOVA revealed: (i) a significant genetic break between the population in the Aegean Sea and those in the other locations, as supported by both mitochondrial genes, and (ii) weak differentiation of the Canary and Azores Islands from the other populations; however, the populations from the Macaronesian Islands, Algarve and West Mediterranean could be considered to be a panmictic metapopulation. Isolation by distance was not identified in H. (H.) mammata. Historical events behind the observed findings, together with the current oceanographic patterns, were proposed and discussed as the main factors that determine the population structure and genetic signature of H. (H.) mammata

Distribution pattern of Galeus atlanticus in the Alborán Sea (south western Mediterranean) and some sexual character comparison with Galeus melastomus

Galeus atlanticus is a small-sized deepwater catshark living on the slope bottom of the Alborán Sea. Given its external similarities with Galeus melastomus, both species are often confused, which makes G. atlanticus a very poorly understood species both in terms of catches and biological aspects. For this study, a total of 741 G. atlanticus specimens, caught during scientific surveys from 1997 to 2003, were analysed. The distributional patterns were described and the reproductive status estimated. Galeus atlanticus occurred throughout the Alborán Sea, between the Strait of Gibraltar and Cape Gata, including the slope of the Island of Alborán. Its bathymetric range extended from 330 to 790 m and no size depth trends have been observed. Mature specimens, both males and females, were caught in all seasons of the year. Size at first maturity was significantly different between sexes, with estimates of 32.9 cm for males and 36.9 cm for females. The differential growth of some secondary sexual characteristics, such as clasper length in males and oviducal gland diameter in females, were compared with those of G. melastomus.

Phytobenthic assemblages of Addaia Bay (Menorca, Western Mediterranean): composition and distribution

Study on the composition and distribution of Phytobenthic Assemblages of Addaia Bay (Menorca, Western Mediterranean)

Calcareous nannofossil bioevents of early-middle Pleistocene sediments from the Mediterranean Sea and the Noth Atlantic

Quantitative distributions of calcareous nannofossils are analysed in the early-middle Pleistocene at the small Gephyrocapsa and Pseudoemiliania lacunosa zone transition in deep-sea cores from the Mediterranean Sea and North Atlantic Ocean (Ocean Drilling Program [ODP] Sites 977, 964 and 967, Deep Sea Drilling Project [DSDP] Site 607). The temporal and spatial mode of occurrence of medium-sized gephyrocapsids and reticulofenestrids has been examined to refine biostratigraphic constraints and evaluate possible relationships of stratigraphic patterns to environmental changes during a period of global climatic deterioration. The timing of bioevents has been calibrated using high-resolution sampling and correlation to the delta18O record in chronologically well-constrained sections. Newly identified events and ecostratigraphical signals enhance the stratigraphic resolution at the early-middle Pleistocene. The first occurrence (FO) of intermediate morphotypes between Pseudoemiliania and Reticulofenestra (Reticulofenestra sp.) is proposed as a reliable event within marine isotope stage (MIS) 35 or at the MIS 35/34 transition. The distribution of Reticulofenestra asanoi is characterized by rare and scattered occurrences in its lowest range, but the first common occurrence (FCO) is consistently identified at MIS 32 or 32/31; the last common occurrence (LCO) of the species is a distinctive event at MIS 23. In the studied interval, Gephyrocapsa omega dominates among medium-sized Gephyrocapsa. The FO of G. omega and contemporaneous re-entry of medium-sized gephyrocapsids at the lower-middle Pleistocene transition are diachronous between the Atlantic Ocean and Mediterranean Sea and from the western to eastern Mediterranean. In the Mediterranean, the LO of G. omega falls at MIS 15, insolation cycle 54 and is isochronous among the sites. Abundance fluctuations of G. omega show notable relations to early-middle Pleistocene climate changes; they considerably increase in abundance at the interglacial stages, suggesting warm water preferences. Gephyrocapsa omega temporarily disappears during the glacial MIS 22 and MIS 20. Above MIS 20, an impoverishment in G. omega and in the total abundance of medium-sized gephyrocapsids occurs. A decrease in abundance of G. omega is observed between the western Site 977 and the easternmost Site 967 in the Mediterranean Sea, as a possible response to high salinity and/or low nutrient content. Possible environmental influences on the distribution of R. asanoi and of Reticulofenestra sp. are discussed.

Holocene sedimentation in the Strait of Otranto, Mediterranean Sea

An extensive radiograph study of 24 undisturbed, up to 206-cm long box and gravity cores from the western part of the Strait of Otranto revealed a great variety of primary bedding structures and secondary burrowing features. The regional distribution of the sediments according to their structural, textural, and compositional properties reflects the major morphologic subdivisions of the strait into shelf, slope, and trough bottom (e.g., the bottom of the northern end of the Corfu-Kephallinia Trough, which extends from the northeastern Ionian Sea into the Strait of Otranto): (1) The Apulian shelf (0 to -170m) is only partly covered by very poorly sorted, muddy sands without layering. These relict(?) sands are rich in organic carbonate debris and contain glauconite and reworked (?Pleistocene) ooids. (2) The slope sediments (-170 to -1,000 m) are poorly sorted, sandy muds with a high degree of burrowing. One core (OT 5) is laminated and shows slump structures. An origin of these slumped sediment masses from older deposits higher on the slope was inferred from their abnormal compaction, color, texture, organic content, and mineral composition. (3) Cores from the northern end of the Corfu-Kephallinia Trough (-980 to -1,060 m) display a few graded sand layers, 2-5 cm (maximum 30 cm) thick with parallel and ripple-cross-laminations, deposited by oceanic bottom or small-scale turbidity currents. They are intercalated with homogeneous lutite. (4) Hemipelagic sediments prevail in the more southerly part of the Corfu-Kephallinia Trough and on the "Apulian-Ionian Ridge", the southern submarine extension of the Apulian Peninsula. Below a core depth of 160 cm, these cores have a laminated ("varved") zone, representing an Early Holocene (Boreal-Atlanticum) "stagnation layer" (14C age approximately 9,000 years). The terrigenous components of the surface sediments as well as those of the deeper sand layers can be derived from the Apulian shelf and the Italian mainland (Cretaceous Apulian Plateau and Gargano Mountains, southern Apennines, volcanic province of the Monte Vulture). Indicated by the heavy mineral glaucophane, a minor proportion of the sedimentary material is probably of Alpine origin. If this portion is considered to be first-cycle clastic material it reaches the Strait of Otranto after a longitudinal transport of 700 km via the Adriatic Sea. The lack of phyllosilicates in the coarse- to medium-grained shelf samples might be explained by the activity of the "Apulian Current" (surface velocities up to 4 knots) which in the past possibly has affected the bottom almost down to depths of the shelf edge. The percentage of planktonic organisms, and also the plankton: benthos ratio in the sediments is a useful indicator for bathymetry (depth zonation).

Distribution of marine litter at bathyal and abyssal depths in the Mediterranean Sea

The distribution, type and quantity of marine litter accumulated on the bathyal and abyssal Mediterranean seafloor has been studied in the framework of the Spanish national projects PROMETEO and DOS MARES and the ESF-EuroDEEP project BIOFUN. Litter was collected with an otter trawl and Agassiz trawl while sampling for megafauna on the Blanes canyon and adjacent slope (Catalan margin, north-western Mediterranean) between 900 and 2700 m depth, and on the western, central and eastern Mediterranean basins at 1200, 2000 and 3000 m depth. All litter was sorted into 8 categories (hard plastic, soft plastic, glass, metal, clinker, fabric, longlines and fishing nets) and weighed. The distribution of litter was analysed in relation to depth, geographic area and natural (bathymetry, currents and rivers) and anthropogenic (population density and shipping routes) processes. The most abundant litter types were plastic, glass, metal and clinker. Lost or discarded fishing gear was also commonly found. On the Catalan margin, although the data indicated an accumulation of litter with increasing depth, mean weight was not significantly different between depths or between the open slope and the canyon. We propose that litter accumulated in the canyon, with high proportions of plastics, has predominantly a coastal origin, while litter collected on the open slope, dominated by heavy litter, is mostly ship-originated, especially at sites under major shipping routes. Along the trans-Mediterranean transect, although a higher amount of litter seemed to be found on the Western Mediterranean, differences of mean weight were not significant between the 3 geographic areas and the 3 depths. Here, the shallower sites, also closer to the coast, had a higher proportion of plastics than the deeper sites, which had a higher proportion of heavy litter and were often affected by shipping routes. The weight of litter was also compared to biomass of megafauna from the same samples. On the Blanes slope, the biomass of megafauna was significantly higher than the weight of litter between 900 and 2000 m depth and no significant differences were found at 2250 and 2700 m depth. Along the trans-Mediterranean transect, no significant differences were found between biomass and litter weight at all sites except in two sites: the Central Mediterranean at 1200 m depth, where biomass was higher than litter weight, and the Eastern Mediterranean at 1200 m depth, where litter weight was higher than biomass. The results are discussed in the framework of knowledge on marine litter accumulation, its potential impact on the habitat and fauna and the legislation addressing these issues.

Subsurface time series of chlorophyll a at the PhytoCly station, Bay of Calvi, Corsica, Mediterranean Sea

The development of the winter-spring phytoplankton bloom was investigated in the Bay of Calvi (Corsica, Ligurian Sea, northwestern Mediterranean) in 1979, 1986, 1988, 1997 and 1998. A drastic reduction of phytoplankton biomass was evidenced over the last 2 decades, in relation to long-term changes in climatic and environmental conditions. Between 1979 and 1998, the monthly averaged chlorophyll a concentrations at 1 m decreased by about 80% during February, March and April. Simultaneously, major changes to hydrodynamic conditions include warmer water, overall decrease of salinity at 10 m depth, longer periods of bright sunshine and lower wind stress. The changes in environmental conditions were large enough to affect the vertical stability of the water column during the winter-spring period and to reduce nutrient replenishment of the surface layer prior to the usual period of phytoplankton growth. Until 1986, the main factor driving nutrient replenishment was the winter upward mixing of nutrient-rich deep waters, while the progressive reduction of mixing from 1988 induced nutrient limitation of surface waters in the last decade. The following hypotheses on changes in the development of the winter-spring phytoplankton bloom are made: (1) Until 1986, phytoplankton peaks took place in relatively high-nutrient waters and were diatom-dominated. (2) Between 1986 and 1988, decreasing Si availability led to Si limitation which caused a reduction in diatom abundance. This resulted in the disappearance of the diatom-dominated pulses and in lower phytoplankton biomass and was accompanied by a shift toward non-siliceous phytoplankton. (3) In 1988, 1997 and 1998, decreasing nitrate availability led to nitrate limitation, thus explaining the progressive reduction in non-siliceous phytoplankton biomass. Other, associated changes in benthos assemblages and ichthyofauna are documented. The conclusions from the Bay of Calvi are extended to the whole western Corsican coast. This confirms that the Mediterranean reacts rapidly to external perturbations, which are driven by climate change in that particular area.