Late Quaternary paleoceanography as recorded by benthonic foraminifera in Strait of Sicily sediment sequences

At specific times during the late Quaternary, a widespread low-salinity surface water layer was produced over the eastern Mediterranean which created stagnant conditions and associated deposition of sapropelic muds in the deep basins. The Strait of Sicily has been positioned above the anoxic deeper water masses and is therefore devoid of sapropel deposits. Sediment sequences in the region contain a valuable record of upper bathyal (200–1000 m) benthonic foraminiferal change during times of deep-basin stagnation.Quantitative examiniation of benthonic foraminiferal assemblages in Strait of Sicily Quaternary cores generally reveals changes coeval with sapropels S1, S2, S4, S5, and S6. The benthonic foraminiferal expression is seen as increases in frequency ofGlobobulimina affinis, Globobulimina pseudospinescens, Chilostomella mediterranensis, Bolivina dilatata, and otherBolivina species during intervals corresponding to the episodic abyssal stagnation events. Some cores also display a decrease inCassidulina carinata and an increase inArticulina tubulosa.Similar assemblage changes have been previously documented to be closely associated with the sapropel layers in the deep basins of the eastern Mediterranean (the actual sapropel layers usually do not contain benthonic foraminifera). However, in the Strait of Sicily area, the species exhibit less drastic changes than in the deep eastern basins because of the less severe anoxic conditions. In the deep eastern basins, benthonic foraminifera almost totally disappeared in sapropel layers; at the same time, there was a general reduction in foraminiferal numbers in the shallower Strait area.     

Late glacial to recent deep-sea benthic foraminifera from the northeastern Atlantic (Cadiz Gulf) and western Mediterranean (Alboran Sea): Paleooceanographic results

Deep-sea benthic foraminifera of the region of water exchange between the Mediterranean and the Atlantic Ocean, at the Gibraltar Strait, show significant water-mass relationships for the past 18,000 years.Cores used in this benthic foraminifera analysis were selected, according to the context of the distribution of various deep bottom water masses, on each side of the Gibraltar Strait:

• 1.(1) on the Atlantic side of the Strait, two cores (KS 8228, KS 8229) are located in the present day NADW, and one (KC 8221) in the upper part of the present Mediterranean outflow water,
• 2.(2) on the Mediterranean side of Gibraltar, two cores (KC 8241, SU 8107) are located in the deep water mass, and one (KS 8230) in the intermediate water mass.

In the two deep basins (1500–2800 m depth in the Gulf of Cadiz, 1200–1300 m depth in the Alboran Sea), the paleooceanographic changes appear to be in an opposite way for the past 18,000 years. The Alboran basin shows a paleooceanographic evolution from a well-oxygenated, nutrient-rich environment at about 18,000 yr B.P. to a nutrient-poor, oxygen-depleted environment from 13,000 to the present time; moreover, for the time-span synchronous with the well-known development of sapropels in the eastern Mediterranean basins between 10,000 and 7000 yr B.P., the faunal assemblage shows most unusual characteristics implying drastic environmental conditions. Conversely, in the Gulf of Cadiz, the environment pass as from a biotope occupied by an oxygen-depleted, nutrient-poor water mass at about 18,000 years B.P. to a biotope occupied by NADW since the Younger Dryas; this agrees with previous data obtained in the northeastern Atlantic Ocean (Caralp, 1987).In the epibathyal zones (550–800 m depth), on both sides of the Gibraltar Strait, paleohydrographic changes do not seem so important. According to the present and very late Holocene assemblages, which are similar on both sides of the Strait with a strong east-west flow, two other stratigraphic episodes have shown the same conditions of water exchanges: the end of the isotopic stage 2 and the Younger Dryas. Conversely, during the last glacial maximum, the Bølling-Allerød and the lower Holocene, westward water fluxes were probably lower. At no time, the hypothesis of a reversal or a stop in the east-west exchanges between Mediterranean and Atlantic Ocean may be justified.     

Late Quaternary planktonic foraminiferal biostratrigraphy,Strait of Sicily,Mediterranean Sea

The development of a detailed planktonic foraminiferal biostratigraphy for the Late Quaternary (latest 180,000 years) has allowed the dating and correlation of sediments from the Strait of Sicily. Principal component analysis of the assemblages has extracted a climatic history similar to that based upon oxygen isotopic oscillations. Intercore correlations are supported by tephrochronology and sapropel layer stratigraphy. Identification of Termination IA and IB in several high sedimentation rate cores indicates that the Mediterranean Sea experienced a two-stage warming during the last deglaciation. Cores collected between water depths of 200 and 600 m contain an erosional hiatus near their tops, reflecting an increase in the Levantine Intermediate Water flow sometime since 25,000 years B.P. and as recently as 7000 years B.P., during deposition of the youngest sapropel layer. A distinct and unusual planktonic foraminiferal succession is associated with sapropel layers. This begins in a protosapropel layer immediately beneath a sapropel horizon and continues upwards to the immediately overlying oxidized layer.     

A taphonomic study of modern pollen assemblages from dung and surface sediments in arid environments of Spain

Coastal plant communities in arid southeastern Spain are characterized by insect-pollinated scrub species, which fail to occur in Quaternary pollen sequences from valleys, marshlands and marine cores. We investigate pollen–vegetation relationships in samples from soil surfaces, animal dung, and sediments in depressions or basins that, in theory, should have pollen spectra that are comparable to those from sedimentary basins elsewhere. Pollen spectra from basins or depressions are very susceptible to long-distance wind and water transport. This can mask representation of pollen from the surrounding insect-pollinated vegetation, as can over-representation of basin-margin halophilous and hydrophilous pollen. Pollen spectra from biogenic materials of animal origin are the best analogues of local and regional vegetation, and show the best analytical potential in terms of pollen concentration and taxon diversity. Pollination properties of the species studied indicate they will rarely be found in most conventional pollen records. It cannot be stressed too strongly that insight into Quaternary vegetation of arid regions demands complementary pollen analysis of coprolites, urine-cemented deposits, and cave sediments with preserved biotic remains, in addition to water-lain sediments.     

Palaeoceanographic record of the last deglaciation in the Strait of Sicily

The palaeoceanographic evolution of the Levantine waters during the last deglacial time is investigated using the sedimentary record of a deep sea core, CS 70-5, from the Linosa basin (35° 44.4′N/13° 11.0′E, 1486 m water depth). Radiocarbon dating and oxygen isotope stratigraphy based on¹⁸O changes inGlobigerina bulloides allow us to recognize and to date the different steps of the deglaciation. These steps are synchronous with those reported in the North Atlantic, but correspond to a δ¹⁸O decrease of higher amplitude than in the Alboran sea or in the North Atlantic Ocean. Major faunal events permit the establishment of a local biozonation which differs from those reported either for the Alboran sea or the Eastern Mediterranean basin. Major breaks in the faunal assemblages occurred during Termination I_A, within the first step of the deglaciation around 14 kyr B.P., and near 10.6 kyr B.P. within the Younger Dryas. The onset of the last deglaciation induced important changes in the characteristics of the Levantine and Atlantic water masses which occupied the Strait of Sicily. The δ¹³C records ofGlobigerina bulloides andCibicidoides pachyderma indicate that the modifications observed in the assemblages of deep faunas are controlled by the oxygenation of the water column. δ¹³C records ofGlobigerina bulloides are similar throughout the West Mediterranean as well as in sediments located below the present Mediterranean outflow. The distributional pattern as well as its δ¹³C record suggest that this species could be a good recorder of the upper Levantine waters and, more precisely, of the mixing layer of the overlying Atlantic waters with the Levantine ones.Major influxes of Atlantic waters during Terminations I_A and I_B could have slowed down the vertical mixing of the different water layers present in the Strait of Sicily and caused a decrease in the oxygenation of the water column. During Termination I_B the effect of Atlantic influxes was reinforced by the occurrence of a low salinity layer in the eastern basin which led to the stagnation of the deep waters. The two episodes of decreased oxygenation in the Levantine waters also favored the precipitation of inorganic magnesium calcite.     

Microplankton respiratory activity and CO2 production rates in the Otranto Strait (Mediterranean Sea)

Respiratory activity and metabolic CO₂production of the microplankton in the Otranto Strait (Mediterranean Sea) were determined by monitoring the Electron Transport System activity. Ten stations were repeatedly investigated during two oceanographic surveys in February–March and August 1994. Respiratory activity and CO₂ production, estimated from the surface to the bottom, were higher in the euphotic layers (0-200 m) during summer (mean values: Winter = 0.024 μg C h⁻¹ dm⁻³; Summer = 0.042 μg C h⁻¹ dm⁻³); in the aphotic zone (deeper than 200 m), the rates were similar throughout different seasons (0.013 and 0.014 μg C h⁻¹ dm⁻³, respectively). A comparison with data collected by other authors from the euphotic layers of the Mediterranean Sea was made. Respiratory activities decreased from Western to Eastern Mediterranean Basins. The values of CO₂ production, integrated between 200 and 1000 m in the Otranto Strait (mean value 237.7 mg C m⁻² d⁻¹), were compared with other data collected from the Mediterranean Sea as well as from the Pacific, Atlantic and Indian Oceans. The comparison showed the Otranto Strait to be a site of organic matter oxidation. This revised version was published online in August 2006 with corrections to the Cover Date.     

Diagenetic stoichiometry and benthic nutrient fluxes at the sediment–water interface of Lake Illawarra, Australia

Benthic flux measurements of O₂, TCO₂ and inorganic nutrients were made at three stations (seagrass beds, shallow bare sand and deep mud) in Lake Illawarra (Australia) to compare the characteristics of diagenesis and benthic biogeochemical processes for different primary producers (seagrass or microphytobenthos, (MPB)) and/or sediment types (sand or mud).Seagrass beds exhibited the highest gross primary productivity while the lowest rates occurred at the deep mud station. At the shallow bare sand station only, the gross primary production (GPP) and respiration (R) were balanced, while at the other two stations, R exceeded GPP by as much as 2 fold, indicating more organic carbon was decomposed than produced at the time of sampling. In general, nutrient fluxes displayed typical diurnal variation.Organic carbon oxidation scenarios, evaluated by either calcium carbonate dissolution or sulfate reduction models, indicated that both models can represent organic matter mineralization. The difference of estimated total carbon oxidized in this lake using the two models was small, ranging from 0.2% at deep mud station to maximum of 21% at seagrass station. In addition, N₂ flux rates (net denitrification), estimated using carbon and nitrogen stoichiometry, were of similar magnitude as the rates estimated using LOICZ budget modeling or measured using the N₂/Ar technique.Finally, a comparison of calculated diffusive fluxes and measured fluxes using incubation cores indicated that the results were of similar magnitude at the deep mud station, but the incubation cores fluxes were much higher than the calculated diffusive fluxes at the other two stations. This may have been caused by bioturbation or bioirrigation.     

Quaternary calcareous nannofossils from Periantarctic basins: Paleoecological and paleoclimatic implications

Currently, coccolithophores have a widespread oceanic distribution and are reported from most latitudes, but not those higher than 65°S. Fifteen piston cores were sampled with the aim of investigating the distribution and abundance variation of Quaternary calcareous nannofossils of the Antarctic region, south of the Antarctic Divergence (> 65°S), particularly from Maud Rise, Bausan Bank, and from Weddell, Ross and Bellingshausen Seas. A calcareous nannofossil cold-taxa association is present in most of the cores examined and their discontinuous occurrence is thought to indicate key environmental relationships. The presence of calcareous nannofossils is correlated with interglacial intervals with warmer SSTs and may indicate high productivity and an open-ocean environment. Our results confirm that, during short periods of the late Quaternary, coccolithophorids occurred at southern high latitudes, in the western Antarctic basins, while in the eastern Antarctic basins they are nearly absent, suggesting more variable SSTs near West Antarctic Ice Sheet.     

Genetic structure in the Mediterranean seagrass Posidonia oceanica: disentangling past vicariance events from contemporary patterns of gene flow

The Mediterranean Sea is a two‐basin system, with the boundary zone restricted to the Strait of Sicily and the narrow Strait of Messina. Two main population groups are recognized in the Mediterranean endemic seagrass Posidonia oceanica, corresponding to the Western and the Eastern basins. To address the nature of the East–West cleavage in P. oceanica, the main aims of this study were: (i) to define the genetic structure within the potential contact zone (i.e. the Strait of Sicily) and clarify the extent of gene flow between the two population groups, and (ii) to investigate the role of present water circulation patterns vs. past evolutionary events on the observed genetic pattern. To achieve these goals, we utilized SSR markers and we simulated, with respect to current regime, the possible present‐day dispersal pattern of Posidonia floating fruits using 28‐day numerical Lagrangian trajectories. The results obtained confirm the presence of the two main population groups, without any indices of reproductive isolation, with the break zone located at the level of the Southern tip of Calabria. The populations in the Strait of Sicily showed higher affinity with Western than with Eastern populations. This pattern of genetic structure probably reflects historical avenues of recolonization from relict glacial areas and past vicariance events, but seems to persist as a result of the low connectivity among populations via marine currents, as suggested by our dispersal simulation analysis.     

Grain-size controls on the occurrence of bioturbation

Grain size and grain-size related stresses impart a significant influence on the ichnological character of marginal-marine deposits. This is evident on the New Brunswick coastline of the Bay of Fundy, Canada, where three coarse-grained marginal-marine deposits are studied to assess grain-size controls on the occurrence and type of bioturbation. Firm mud and sand substrates exhibit the greatest diversity and density of bioturbation (i.e., bioturbation intensity). The types of organisms colonizing sands and firm-mud substrates are variable; however, the resultant trace assemblages are similar. Thixotropic muds exhibit significantly reduced trace diversity and density relative to firm mud, reflecting the additional stress placed on the organisms by the relatively soupy consistency of the sediment. A significant change in the trace assemblage occurs when sediment caliber passes the gradational sand—fine gravel boundary.Four main conclusions can be drawn from this study. First, for mixed sand and gravel, fine gravel, and coarse-gravel deposits, the degree of bioturbation (diversity ^? density) decreases more rapidly onshore (across the intertidal zone) than is noted in sand or mud deposits. Second, there is a decrease in the degree of bioturbation with increasing grain size for substrates composed of sand-sized and larger clasts. Third, burrows in gravels tend to be lined and/or robust, likely to maintain a stable environment within the burrow. Fourth, in coarse-gravel substrates or substrates with a significant component of coarse gravel, burrows are developed between the clasts and tend to be more permanent structures (than those developed in sand or mud), which are generally continuously occupied.The degree of burrowing noted in these modern gravel deposits contrasts with the relative paucity of biogenic structures reported in conglomerates preserved in the rock record. Based on the intensity of burrowing observed in the gravels, we hypothesize that ancient marginal-marine conglomerates are likely bioturbated, but that these burrows are likely distorted during burial and compaction.     

Glacial–interglacial deep-water changes in the NW Pacific inferred from single foraminiferal δO and δC

Oxygen and carbon isotope values of single benthic foraminiferal tests in a core from the Shatsky Rise, NW Pacific Ocean, show greater intra-horizon variance during the Holocene than during the Last Glacial Maximum (LGM). This greater variance is caused by the introduction of glacial specimens some 20 cm upward from their original deposition layer due to bioturbation. In contrast, foraminiferal populations belonging to glacial layers do not include Holocene specimens. The difference in direction of bioturbation greatly modifies climate information in horizons formed during and after deglacial events. After omitting glacial specimens from Holocene sediments, the glacial–interglacial difference in δ¹⁸O suggests that Pacific deep-water temperature changed by 2.4–3.8°C at the most. The δ¹³C values suggest that nutrient concentration was higher during the LGM than the Holocene. The glacial deep North Pacific Ocean apparently was influenced by cold deep waters of southern origin.     

Sedimentary photosynthetic pigments of algae and phototrophic bacteria in Lake Hamana,Japan: temporal changes of anoxia in its five basins

We analyzed photosynthetic pigments of algae and bacteria (phototrophic sulfur bacteria: Chromatium and brown Chlorobium) in sediment cores and water samples obtained from five basins of Lake Hamana, a brackish, eutrophic, holomictic lake in Japan, and discussed our findings in relation to the distribution of the phototrophs. The four outer basins are connected to the central basin by narrow inlets. The prevalence of anoxia in Lake Hamana was demonstrated by the widespread presence of bacterial pigments in each core. The construction of training walls in 1954–1956 to direct tidal currents into the lake via Imagire-guchi Channel, the sole inlet for seawater, increased the lake water circulation, suppressed the development of anoxia, and caused Chromatium to disappear. Strong correlations (r ² 0.7) between total algal carotenoid (TAC) and total bacterial carotenoid (TBC) contents in each core were found in four basins. We ascribe this to the induction of anoxia by water stratification and algal proliferation, which precede the growth of phototrophic sulfur bacteria in the deeper layers of the water column. The slopes of the TBC–TAC correlations in the sediment cores, indicating the extent and stability of anoxia at each site, differed among basins (0.23–0.67) and were inversely related to the exchange rate of water by seawater intrusion in each basin.     

Ethological analysis of the trace fossil Zoophycos: hints from the Arctic Ocean

Löwemark, L. 2011: Ethological analysis of the trace fossil Zoophycos: Hints from the Arctic Ocean. Lethaia, Vol. 45, pp. 290–298. The distribution of the trace fossil Zoophycos in Quaternary marine sediments from the Arctic Ocean was studied in twelve piston and gravity cores retrieved during the Swedish icebreaker expeditions YMER80, Arctic Ocean‐96 and LOMROG I & II. The sampled cores span an area from the Makarov Basin to the Fram Strait. Zoophycos was only found in two cores taken at more than 2 km water depth on the slope of the Lomonosov Ridge, but was absent in cores obtained at shallower depth, confirming earlier observations of the trace maker’s bathymetric preferences. The two cores containing Zoophycos are characterized by quiet sedimentation and slightly enhanced food flux compared with the general Arctic. The occurrence of Zoophycos in these cores in a setting that is characterized by extreme seasonal variations in food flux due to the total ice coverage during winters and high primary productivity during the long summer days, is interpreted to be a cache‐behaviour response to pulsed flux of food to the benthic realm. □Arctic Ocean, ethology, Quaternary, spreiten, trace fossils, Zoophycos.     

Late Quaternary clay mineral record in Central Lake Baikal (Academician Ridge, Siberia)

We investigated the mineralogical composition of two cores recovered on the Academician Ridge (Central Lake Baikal, Siberia). Sedimentological features show that the cores are unaffected by turbidity currents. However, hemipelagic deposition is not continuous, but intermittently disturbed by syn- or post-sediment reworking (e.g., bioturbation, slumps, faulting). Such modes of deposition are consistent with the complex uplift history of the ridge. Bulk mineralogy suggests that terrigenous sediment supplies are constant through glacial/interglacial stages, and diluted by diatom-rich intervals related to warmer interglacial stages. The core stratigraphy is based on the correlation of the diatom zonation and opal abundance with the marine oxygen isotope reference curve SPECMAP. The ∼8-m cores partly recover the last four interglacial/glacial cycles, i.e., since oxygen isotope stage 8. We test the use of clay minerals as a proxy for paleoclimatic reconstruction. The clays are more weathered during the diatom-rich intervals in agreement with warmer climate conditions. However, the mean clay composition does not change significantly through glacial/interglacial stages. This observation implies that, in the Academician Ridge sediments, a simple smectite/illite ratio (S/I) does not alone provide a reliable indicator of climatic variation. It reflects the complex clay assemblages, especially the smectite group, delivered to Central Lake Baikal. Smectites include primarily illite-smectite mixed layers, made of a mixture of montmorillonite and beidellite. According to their behavior after cation saturation, the illite-smectite mixed layers are primarily transformed smectites, with some neoformed smectites intermittently observed. In addition, Al-smectites occur in minor proportions. We conclude that the S/I ratio has a climatic significance only if it evolves in parallel with the weathering stage of the clays and is confirmed by a change in the composition of the smectites.     

The Early Pliocene re-colonisation of the deep Mediterranean Sea by benthic foraminifera and their pulsed Late Pliocene–Middle Pleistocene decline

Ninety-five species and 19 genera of cosmopolitan, deep-sea benthic foraminifera belonging to the families Pleurostomellidae, Stilostomellidae and Nodosariidae, became extinct during the Late Pliocene–Middle Pleistocene. Only 50% of these (44 species) were present in the Pliocene or Pleistocene of the deep Mediterranean Sea (ODP Sites 654, 966, 967, 975, 976), being those which had successfully migrated in via the Strait of Gibraltar from the deep Atlantic following the annihilation of the Mediterranean deep-sea fauna during the Late Miocene Messinian Crisis. Most colonisation occurred within the first 0.8 myrs (5.3–4.5 Ma) after re-establishment of the Mediterranean–Atlantic link, with possibly a second lesser period of immigration in the Late Pliocene (3.4–3.0 Ma). We infer that colonisations may have been fortuitous and few in number, as some common members of the group in the Atlantic never succeeded in establishing in the Mediterranean Sea. There is no evidence of any new immigration events during the Pleistocene, implying that the present anti-estuarine circulation may have been in place throughout this period. Our studies suggest that these deep-water, low-oxygen-tolerant foraminifera survived the many periods of deep-water sapropel formation in the Pliocene–Early Pleistocene, possibly in somewhat shallower (~ 500 m) refuges with dysoxic, rather than anoxic conditions.The Pliocene–Pleistocene stratigraphic record of this group of elongate, cylindrical benthic foraminifera with constricted and specialised apertures is similar in the west and east Mediterranean basins. The group declined in abundance (flux) and diversity in two pulses, during the Late Pliocene (3.1–2.7 Ma) and the late Early Pleistocene (1.3–1.0 Ma) in concert with global, southern-sourced, deep-water sites (AABW, CPDW) and earlier than the single decline (1.0–0.6 Ma) in global, intermediate water sites (uNADW, AAIW). All species, with one possible exception, disappeared earlier in the Mediterranean than globally. The highest occurrence of any species of this group in Mediterranean sites was 0.8–0.43 Ma, comparable with 0.7–0.2 Ma outside with the youngest survivors being in abyssal, deep-water.Thus, despite the unusual oceanographic conditions and isolation, the deep Mediterranean Sea was in this case neither the centre for the evolution of new species nor a refuge where species survived after they had disappeared elsewhere.     

Diatom productivity peak and increased circulation during latest Quaternary: Alboran Basin (Western Mediterranean)

Diatoms were studied quantitatively in six latest Quaternary (⋍70 kyr B.P. to Recent) piston cores from the westernmost Mediterranean, the Alboran Basin, and the Atlantic region immediately to the west of the Straits of Gibraltar.The Atlantic cores completely lack diatoms. In the Alboran Basin, diatoms are common from late Stage 3 (⋍27.5 kyr B.P.) to Termination Ib (9 kyr B.P.) and in Recent core tops, but are absent in the other latest Quaternary intervals. Maximum accumulation of diatoms and highest abundance of species normally in sediments associated with increased productivity occurred during the latest Quaternary deglaciation, in the first phase of Termination I (⋍14.8 kyr B.P.).In the modern Alboran Basin, a region of high biological productivity occurs immediately east of the Gibraltar Straits. This high productivity results from upwelling associated with the interaction between the Atlantic inflow and the bottom topography near the Spanish coast. The upwelled nutrient-rich waters are then advected to the east and southeast by the surficial anticyclonic gyral circulation. Late Quaternary variations in diatom abundance are considered to reflect changes in this upwelling intensity with highest diatom abundances inferred to result from increased upwelling associated with an intensification of the anticyclonic gyral circulation. Highest inferred upwelling rates occurred during the first phase of latest Quaternary deglaciation. It is possible that an intensification of circulation within the Mediterranean Basin as a whole occurred from late Stage 3 to mid Termination I because widespread hiatus formation has been reported at this time in the Straits of Sicily due to an increase in the formation of intermediate waters. Diatoms were not preserved in other latest Quaternary intervals due to insufficient productivity to counterbalance their dissolution.     

Diatoms in surface sediments of the Indonesian Archipelago and their relation to hydrography

Diatoms from 53 surface sediments (water depth 350–7200 m) of the Indonesian Archipelago were studied to determine the recent distribution of important assemblages. Three significant assemblages were distinguished, each related to hydrographic parameter(s) of the overlying water mass. The first assemblage is related to the warm saline surface waters of Pacific and Indian Ocean origin; the second assemblage reflects the low-salinity lobe in Makassar Strait; and the third corresponds to major seasonal upwelling areas in the Arafura Sea and south of Java.The normalized ratio of typically Pacific Ocean and Indian Ocean species reflects the fluctuations of the Pacific versus Indian Ocean inflow during the Late Quaternary.Absolute diatom abundances (ADA; in valves g^–1 carbonate-free dry wt) and diatom accumulation rates (DAR; in valves CM^–2 ka^–1) of autochthonous species did not correspond to daily rates of primary production in the photic zone and consequently cannot be used for paleoproductivity estimates.Besides vertical transport some lateral transport of diatoms occurs, as was demonstrated by the presence of three groups of allochthonous species. They are indicators of productivity in the littoral environment, bottom currents and river outflow.     

Analysis of the hatching site and migratory behaviour of the swordtip squid (Uroteuthis edulis) caught in the Japan Sea and Tsushima Strait in autumn estimated by statolith analysis

In contrast to the swordtip squid (Uroteuthis edulis) caught in the spring and summer, those caught in the autumn in the Japan Sea and the Tsushima Strait are characterized by thick bodies with large clubs, long tentacles and large suckers. In addition, most females and many males are immature. However, the reasons for these characteristics, as well as the hatching site and migratory behaviour, are not well known. The empirical water temperature of juveniles was estimated at ～17°C through strontium:calcium (Sr:Ca) ratios in statoliths and information on seawater temperature distribution. The ontogenetic variation in Sr:Ca ratios decreased from hatching to 60 days after hatching and then remained stable. The estimated water temperature was ～21°C after 60 days. Thus, individuals of the autumn-migrating group most likely hatch in the southern East China Sea in the winter and early spring and then move north-eastwards with currents and pass through the Tsushima Strait into the southern Japan Sea during or after July. However, the squid could be present in these waters without strong currents northeast of Tsushima Island, where thermoclines occur during summer, and become confined in limited layers until vertical mixing occurs in the autumn. The distribution of the squid would thereafter extend to fishing grounds in the southern and south-eastern Japan Sea and then to the Tsushima Strait. This migratory behaviour could be the cause of the observed characteristics because bottom seawater temperatures are much lower in the Japan Sea than in the East China Sea.     

Influence of the mode of macrofauna-mediated bioturbation on the vertical distribution of living benthic foraminifera: First insight from axial tomodensitometry

We investigated the influence of bioturbation by macrofauna on the vertical distribution of living (stained) benthic foraminifera in marine intertidal sediments. We investigated the links between macrofaunal bioturbation and foraminiferal distribution, by sampling from stations situated on a gradient of perturbation by oyster-farming, which has a major effect on benthic faunal assemblages. Sediment cores were collected on the French Atlantic coast, from three intertidal stations: an oyster farm, an area without oysters but affected by oyster biodeposits, and a control station. Axial tomodensitometry (CT-scan) was used for three-dimensional visualization and two-dimensional analysis of the cores. Biogenic structure volumes were quantified and compared between cores. We collected the macrofauna, living foraminifera, shells and gravel from the cores after scanning, to validate image analysis. We did not investigate differences in the biogenic structure volume between cores. However, biogenic structure volume is not necessarily proportional to the extent of bioturbation in a core, given that many biodiffusive activities cannot be detected on CT-scans. Biodiffusors and larger gallery-diffusors were abundant in macrofaunal assemblage at the control station. By contrast, macrofaunal assemblages consisted principally of downward-conveyors at the two stations affected by oyster farming. At the control station, the vertical distribution of biogenic structures mainly built by the biodiffusor Scorbicularia plana and the large gallery-diffusor Hediste diversicolor was significantly correlated with the vertical profiles of living foraminifera in the sediment, whereas vertical distributions of foraminifera and downward-conveyors were not correlated at the station affected by oyster farming. This relationship was probably responsible for the collection of foraminifera in deep sediment layers (> 6 cm below the sediment surface) at the control station. As previously suggested for other species, oxygen diffusion may occur via the burrows built by S. plana and H. diversicolor, potentially increasing oxygen penetration and providing a favorable microhabitat for foraminifera in terms of oxygen levels. By contrast, the absence of living foraminifera below 6 cm at the stations affected by oyster farming was probably associated with a lack of biodiffusor and large gallery-diffusor bioturbation. Our findings suggest that the effect of macrofaunal bioturbation on the vertical distribution of foraminiferal assemblages in sediments depends on the effects of the macrofauna on bioirrigation and sediment oxidation, as deduced by Eh values, rather than on the biogenic structure volume produced by macrofauna. The loss of bioturbator functional diversity due to oyster farming may thus indirectly affect infaunal communities by suppressing favorable microhabitats produced by bioturbation.     

Deep-Sea Biodiversity in the Mediterranean Sea: The Known,the Unknown,and the Unknowable

Deep-sea ecosystems represent the largest biome of the global biosphere, but knowledge of their biodiversity is still scant. The Mediterranean basin has been proposed as a hot spot of terrestrial and coastal marine biodiversity but has been supposed to be impoverished of deep-sea species richness. We summarized all available information on benthic biodiversity (Prokaryotes, Foraminifera, Meiofauna, Macrofauna, and Megafauna) in different deep-sea ecosystems of the Mediterranean Sea (200 to more than 4,000 m depth), including open slopes, deep basins, canyons, cold seeps, seamounts, deep-water corals and deep-hypersaline anoxic basins and analyzed overall longitudinal and bathymetric patterns. We show that in contrast to what was expected from the sharp decrease in organic carbon fluxes and reduced faunal abundance, the deep-sea biodiversity of both the eastern and the western basins of the Mediterranean Sea is similarly high. All of the biodiversity components, except Bacteria and Archaea, displayed a decreasing pattern with increasing water depth, but to a different extent for each component. Unlike patterns observed for faunal abundance, highest negative values of the slopes of the biodiversity patterns were observed for Meiofauna, followed by Macrofauna and Megafauna. Comparison of the biodiversity associated with open slopes, deep basins, canyons, and deep-water corals showed that the deep basins were the least diverse. Rarefaction curves allowed us to estimate the expected number of species for each benthic component in different bathymetric ranges. A large fraction of exclusive species was associated with each specific habitat or ecosystem. Thus, each deep-sea ecosystem contributes significantly to overall biodiversity. From theoretical extrapolations we estimate that the overall deep-sea Mediterranean biodiversity (excluding prokaryotes) reaches approximately 2805 species of which about 66% is still undiscovered. Among the biotic components investigated (Prokaryotes excluded), most of the unknown species are within the phylum Nematoda, followed by Foraminifera, but an important fraction of macrofaunal and megafaunal species also remains unknown. Data reported here provide new insights into the patterns of biodiversity in the deep-sea Mediterranean and new clues for future investigations aimed at identifying the factors controlling and threatening deep-sea biodiversity.