Nanoflagellates (mixotrophs, heterotrophs and autotrophs) in the oligotrophic eastern Mediterranean

The vertical distribution (0 to 100 m) and abundance of nanoflagellates were examined in the oligotrophic Aegean Sea (east Mediterranean) in early spring (south basin) and late summer (north and south basins) of 1997 in the framework of the MATER project (Mass Transfer and Ecosystem Response). Different trophic types of nanoflagellates (mixotrophic, heterotrophic, and phototrophic) were identified based on the possession of chloroplasts and the consumption of Fluorescently Labelled Minicells (FLM). Bacterial production (leucine method) was compared with bacterivory estimated from FLM consumption. We found that mixotrophic nanoflagellates played a small role as bacterivores relative to heterotrophic nanoflagellates and total bacterivory roughly balanced bacterial production. In early spring with cool (14.2°C) well-mixed water columns, flagellate concentrations were lowest, phototrophic flagellates were the dominant group and concentrations varied little with depth. Average concentrations of mixotrophs, heterotrophs and autotrophs were 0.07, 0.34, and 0.64 x 103 cells/ml, respectively. Bacterial production in the 0 to 100 m layer averaged about 0.74 µg C/l/d. Estimated nanoflagellate bacterivory from FLM ingestion accounted for 40% of bacterial production with mixotrophic nanoflagellates consuming 5% of bacterial production. In late summer, total nanoflagellate concentrations were higher. Average concentrations of mixotrophs, heterotrophs and autotrophs were 0.09, 1.14, and 0.66 x 103 cells/ml, respectively, in the southern basin and 0.09, 1.1, and 0.98 x 103 cells/ml, respectively, in the northern basin. In September, bacterial production for both basins roughly balanced estimated nanoflagellate consumption. Similar to the March estimates, mixotrophic nanoflagellates accounted for about 5% of nanoflagellate bacterivory. In a nutrient enrichment experiment in March, treatments including phosphorus resulted in increased bacterial production and reductions in identifiable mixotrophs.

Calcareous dinoflagellate cysts in surface sediments from the Mediterranean Sea

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.

Stable isotope composition of Holocene benthic foraminifers from the Eastern Mediterranean Sea

Temporal and regional changes in paleoproductivity and paleoceanography in the eastern Mediterranean Sea during the past 12 kyr were reconstructed on the basis of the stable oxygen and carbon isotope composition of the epibenthic Planulina ariminensis and the shallow endobenthic Uvigerina mediterranea from three sediment cores of the Aegean Sea and Levantine Basin. The Younger Dryas is characterized by high d18O values, indicating enhanced salinities and low temperatures of deep water masses at all investigated sites. With the onset of the Holocene, d18O records show a continuous decrease towards the onset of sapropel S1 formation, mainly caused by a freshening and warming of surface waters at deep water formation sites. In the middle and late Holocene, the similarity of d18O values from the southern Aegean Sea and Levantine Basin suggests the influence of isotopically identical deep water masses. By contrast, slightly higher d18O values are observed the northern Aegean Sea, which probably point to lower temperatures of North Aegean deep waters. The epifaunal d13C records reveal clear changes in sources and residence times of eastern Mediterranean deep waters associated with period of S1 formation. Available data for the early and late phase of sapropel S1 formation and for the interruption around 8.2 kyr display drops by 0.5 and 1.5 per mil, indicating the slow-down of deep water circulation and enhanced riverine input of isotopically light dissolved inorganic carbon from terrestrial sources into the eastern Mediterranean Sea. The decrease in epifaunal d13C signals is particularly expressed in the southern Aegean Sea and Levantine Basin, while it is less pronounced in the northern Aegean Sea. This points to a strong reduction in deep water exchange rates in the southern areas, but the persistence of local deep water formation in the northern Aegean Sea. The d13C values of U. mediterranea records reveal temporal and regional differences in paleoproductivity during the past 12 kyr, with rather eutrophic and mesotrophic conditions in the North Aegean Sea and southeast Levantine Basin, respectively, while the South Aegean Sea is characterized by rather oligotrophic conditions. After S1 formation, increasing d13C values reflect a progressive decrease in surface water productivity in the eastern Mediterranean Sea during the middle and late Holocene. In the northern Aegean Sea, this time interval is marked by repetitive changes in organic matter fluxes documented by significant fluctuations in the d13C signal of U. mediterranea on millennial- to multi-centennial time scales. These fluctuations can be linked to short-term changes in river runoff driven by northern hemisphere climatic variability.

Geochemical analysis of sediments from three coring sites in the Mediterranean

In a gravity core from the eastern Mediterranean Sea, a chemically and mineralogically distinct, 5.5-cm-thick layer is present above sapropel S-1 and overlain by hemipelagic marls. Calcite is completely absent in this exotic layer, dolomite is present only in small amounts, and the Cr concentrations are significantly enhanced. The layer was deposited primarily under reducing conditions, but the distributions of redox-sensitive elements show that a large part of the exotic layer is now oxidised by a downward-progressing oxidation front. Sediments from within the nearby anoxic, hypersaline Urania Basin are similar to those from the exotic layer, in particular in S-, C-, and O-isotope distributions of pyrite and dolomite, as well as increased Cr concentrations. Mud expulsion due to expansion of gas-rich mud is proposed to explain the presence of the exotic layer outside the Urania Basin. The deposition of an anoxic layer above S-1 shielded the sapropel from oxidation which resulted in the rare occurrence of a complete preservation of S-1 and provides the first minimum age for the start of anoxic mud accumulation in the Urania Basin.

Geochemistry of Miocene Alborán Basin volcanism

We present new major and trace element and O-Sr-Nd-isotope data for igneous rocks from the western Mediterranean Alborán Sea, collected during the METEOR 51/1 cruise, and for high-grade schists and gneisses from the continental Alborán basement, drilled during the Ocean Drilling Programme (ODP Leg 161, Site 976). The geochemical data allow a detailed examination of crustal and mantle processes involved in the petrogenesis of the lavas and for the first time reveal a zonation of the Miocene Alborán Sea volcanism: (1) a keel-shaped area of LREE-depleted (mainly tholeiitic series) lavas in the central Alborán Sea, generated by high degrees of partial melting of a depleted mantle source and involving hydrous fluids from subducted marine sediments, that is surrounded by (2) a horseshoe-shaped zone with LREE-enriched (mainly calc-alkaline series) lavas subparallel to the arcuate Betic-Gibraltar-Rif mountain belt. We propose that the geochemical zonation of the Miocene Alborán Basin volcanism results from eastward subduction of Tethys oceanic lithosphere coupled with increasing lithospheric thickness between the central Alborán Sea and the continental margins of Iberia and Africa.

Sea surface temperature reconstruction for Mediterranean Sea samples

Alkenone unsaturation ratios and planktonic delta18O records from sediment cores of the Alboran, Ionian and Levantine basins in the Mediterranean Sea show pronounced variations in paleo-temperatures and -salinities of surface waters over the last 16,000 years. Average sea surface temperatures (SSTs) are low during the last glacial (averages prior to 13,000 years: 11-15°C), vary rapidly at the beginning of the Holocene, and increase to 17-18°C at all sites during S1 formation (dated between 9500 and 6600 calendar years). The modern temperature gradient (2-3°C) between the Mediterranean sub-basins is maintained during formation of sapropel S1 in the Eastern Mediterranean Sea. After S1, SSTs have remained uniform in the Alboran Sea at 18°C and have fluctuated around 20°C in the Ionian and Levantine Basin sites. The delta18O of planktonic foraminifer calcite decreases by 2 per mil from the late glacial to S1 sediments in the Ionian Basin and by 2.8 per mil in the Levantine Basin. In the Alboran Sea, the decrease is 1.7 per mil. Of the 2.8 per mil decrease in the Levantine Basin, the effect of global ice volume accounts for a maximum of 1.05 per mil and the temperature increase explains only a maximum of 1.3 per mil. The remainder is attributed to salinity changes. We use the temperature and salinity estimates to calculate seawater density changes. They indicate that a reversal of water mass circulation is not a likely explanation for increased carbon burial during S1 time. Instead, it appears that intermediate and deep water formation may have shifted to the Ionian Sea approximately 2000 years before onset of S1 deposition, because surface waters were as cold, but saltier than surface water in the Levantine Basin during the Younger Dryas. Sapropel S1 began to form at the same time, when a significant density decrease also occurred in the Ionian Sea.

Planktonic foraminifera and sea surface temperature reconstruction of sediments from the Mediterranean Sea

Water exchange between the Black Sea and the Mediterranean Sea has been a major focus of the paleohydrography of the eastern Mediterranean. Glacial melt water released from the Black Sea is a potential factor in the formation of sapropel S1, an organic-rich sediment layer that accumulated during the Early Holocene. A high-resolution study done on sediments from the Marmara Sea, the gateway between the Mediterranean and the Black Sea, sheds light on the Holocene exchange processes. Past sea surface temperature and sea surface salinity (SSS) were derived from stable oxygen isotope ratios (delta18O) of foraminiferal calcite and alkenone unsaturation ratios (Uk'37). Heavy delta18O values and high SSS in the Marmara Sea suggest absence of low salinity water from the Black Sea during S1. The comparison with data from the Levantine Basin and southern Aegean Sea outlines gradients of freshening in the eastern Mediterranean Sea, whereby the major sources of freshwater were closer to the Levantine Basin. It is thus concluded that the Black Sea was not a major freshwater source contributing to formation of S1. Given the absence of a low salinity layer, the deposition of organic-rich sediments corresponding to S1 in the Marmara Sea is likely the result of the global transgression and the concomitant re-organization of biogeochemical cycles, leading to enhanced productivity as shown by Globigerina bulloides.

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.