Christoph Hemleben

Assessing the reliability of magnesium in foraminiferal calcite as a proxy for water mass temperatures

Though many studies on the Mg contents in the calcitic tests of foraminifers exist, the processes controlling its uptake are still a matter of debate. Laboratory cultures offer an excellent opportunity to reveal these mechanisms. The Mg concentrations within single chambers of the planktic foraminifer Globigerinoides sacculifer (BRADY) maintained under controlled laboratory conditions were measured (1) at variable temperatures (19.5–29.5 °C) and constant salinity and (2) at variable salinity (22–45‰) and constant temperature. The experimental results suggest that under natural conditions, temperature is the leading mechanism controlling the Mg/Ca ratio. Temperature and magnesium are related proportionally. A temperature increase of ca. 10 °C gives rise to an increase of the magnesium concentrations of ca. 130%. Drastic (unnatural) salinity changes dominate the effects of temperature. A 110% change in the Mg/Ca ratio was observed when salinity was elevated or reduced by more than ca. 10‰. Specimens which underwent gametogenesis reveal significantly higher Mg concentrations than specimens that did not release gametes. Partition coefficients for Mg in foraminiferal calcite are orders of magnitude lower than values from inorganically precipitated calcite. When comparing observed Mg/Ca ratios of foraminiferal tests with predicted Mg/Ca ratios calculated according to empirical equations, it becomes evident that foraminiferal tests are undersaturated with respect to Mg for the water temperature they have experienced. Apparently, foraminifers are capable of controlling their Mg concentration. The physiological processes presumably responsible for such depressed Mg/Ca ratios appear to be temperature-controlled as deduced from the close relationship of the observed Mg/Ca ratios and water temperature. This study demonstrates that variations in temperature and salinity are definitely reflected in the Mg content of foraminiferal tests. Magnesium may thus serve as a paleo-proxy for past surface water temperatures, as long as postdepositional changes and salinity variations are of subordinate importance or can be excluded.

Phytodetritus on the deep-sea floor in a central oceanic region of the Northeast Atlantic

In a midoceanic region of the northeast Atlantic, patches of freshly deposited phytodetritus were discovered on the sea floor at a 4500 m depth in July/August 1986. The color of phytodetritus was variable and was obviously related to the degree of degradation. Microscopic analyses showed the presence of planktonic organisms from the euphotic zone, e.g., cyanobacteria, small chlorophytes, diatoms, coccolithophorids, silicoflagellates, dinoflagellates, tintinnids, radiolarians, and foraminifers. Additionally, crustacean exuviae and a great number of small fecal pellets, “minipellets,” were found. Although bacteria were abundant in phytodetritus, their number was not as high as in the sediment. Phytodetrital aggregates also contained a considerable number of benthic organisms such as nematodes and special assemblages of benthic foraminifers. Pigment analyses and the high content of particulate organic carbon indicated that the phytodetritus was relatively undegraded. Concentrations of proteins, carbohydrates, chloroplastic pigments, total adenylates, and bacteria were found to be significantly higher in sediment surface samples when phytodetritus was present than in equivalent samples collected at the same stations in early spring prior to phytodetritus deposition. Only the electron transport system activity showed no significant difference between the two sets of samples, which may be caused by physiological stress during sampling (decompression, warming). The chemical data of phytodetritus samples displayed a great variability indicative of the heterogeneous nature of the detrital material. The gut contents of various megafauna (holothurians, asteroids, sipunculids, and actiniarians) included phytodetritus showing that the detrital material is utilized as a food source by a wide range of benthic organisms. Our data suggest that the detrital material is partly rapidly consumed and remineralized at the sediment surface and partly incorporated into the sediment. Incubations of phytodetritus under simulated in situ conditions and determination of the biological oxygen demand under surface water conditions showed that part of its organic matter can be biologically utilized. Based on the measured standing stock of phytodetritus, it is estimated that 0.3–3% of spring primary production sedimented to the deep-sea floor. Modes of aggregate formation in the surface waters, their sedimentation, and distribution on the seabed are discussed.

Seasonal changes in species composition, numbers, mass, size, and isotopic composition of planktonic foraminifera settling into the deep sargasso sea☆

Abstract Planktonic foraminifera recovered from six successive sediment-trap samples collected over fourteen months at a depth of 3200 m near Bermuda show varied responses to seasonal changes in the near-surface hydrography. These include changes in species composition and within species, changes in number of individuals collected per unit time, average size and weight of individuals, and in oxygen and carbon isotopic composition. A total of seventeen species were identified and eleven were sufficiently abundant for isotopic analysis in at least some of the samples. The accumulations on the sea floor of some of these species can be expected to contain a strong seasonal component in their oxygen isotopic composition which may be utilized in paleoclimatic and paleo-oceanographic reconstructions. Best suited to this purpose are those species which form their tests in isotopic equilibrium with surface water and whose abundance is confined to a portion of the year only. The best examples are Pulleniatina o bliguiloculata, Globigerinoides conglobatus, and Neogloboquadrina dutertrei. Globigerinoides ruber is a good estimator of the average surface-water temperature because it occurs in rather even abundance throughout the year. Hastigerina pelagica, though very abundant in surface water, is practically eliminated, even before arrival on the sea floor, by partial resorption and structural weakening of the tests during gametogenesis. Orbulina universa, Globorotalia truncaltulinoides and Globorotalia hirsuta show morphological and istopic evidence for contibnued calcification in deeper water after initial test formation near ther surface. Globigerina bulloides represents the most striking example of oxygen isotopic disequilibrium with sea water.

Trophic control of benthic foraminiferal abundance and microhabitat in the bathyal Gulf of Lions, western Mediterranean Sea

Surface sediment was sampled at two bathyal sites in the southwestern Gulf of Lions in the western Mediterranean Sea in February and August 1997 to study the distribution and microhabitat of living (Rose Bengal stained) deep sea benthic foraminifera. Both standing stock and diversity of the faunas, and the microhabitat of distinct species mirror the trophic situation and the depth of the oxidised layer at the different sites. Our results suggest that the faunas do not comprise highly opportunistic species and are adapted to rather stable environments. In the axial channel of the Lacaze-Duthiers Canyon, organic matter fluxes are enhanced due to advective transport of organic matter resulting in elevated oxygen consumption rates in the surface sediment and a rather thin oxidised layer. The corresponding benthic foraminiferal fauna is characterised by rather high standing stock and diversity, and a well-developed deep infauna. In addition to freshly deposited phytodetritus, more degraded organic matter seems to be an important food source. In contrast, at the open slope, organic matter fluxes and oxygen consumption rates in the surface sediment are lower and the oxidised layer is much thicker than inside the canyon. The corresponding benthic foraminiferal fauna comprises mainly epifaunal and shallow-infaunal species with much lower standing stocks and clear differences between February and August. In August standing stocks are higher and the average living depths of most species shift towards the sediment surface. These differences can be attributed to patchiness or represent a seasonal trophic signal.

Forcing mechanisms for mid-Cretaceous black shale formation: evidence from the Upper Aptian and Lower Albian of the Vocontian Basin (SE France)

Calcareous nannoplankton, palynomorph, benthic foraminifera, and oxygen isotope records from the supraregionally distributed Niveau Paquier (Early Albian age, Oceanic Anoxic Event 1b) and regionally distributed Niveau Kilian (Late Aptian age) black shales in the Vocontian Basin (SE France) exhibit variations that reflect paleoclimatic and paleoceanographic changes in the mid-Cretaceous low latitudes. To quantify surface water productivity and temperature changes, nutrient and temperature indices based on calcareous nannofossils were developed. The nutrient index strongly varies in the precessional band, whereas variations of the temperature index reflect eccentricity. Since polar ice caps were not present during the mid-Cretaceous, these variations probably result from feedback mechanisms within a monsoonal climate system of the mid-Cretaceous low latitudes involving warm/humid and cool/dry cycles. A model is proposed that explains the formation of mid-Cretaceous black shales through monsoonally driven changes in temperature and evaporation/precipitation patterns. The Lower Albian Niveau Paquier, which has a supraregional distribution, formed under extremely warm and humid conditions when monsoonal intensity was strongest. Bottom water ventilation in the Vocontian Basin was diminished, probably due to increased precipitation and reduced evaporation in regions of deep water formation at low latitudes. Surface water productivity in the Vocontian Basin was controlled by the strength of monsoonal winds. The Upper Aptian Niveau Kilian, which has a regional distribution only, formed under a less warm and humid climate than the Niveau Paquier. Low-latitude deep water formation was reduced to a lesser extent and/or on regional scale only. The threshold for the formation of a supraregional black shale was not reached. The intensity of increases in temperature and humidity controlled whether black shales developed on a regional or supraregional scale. At least in the Vocontian Basin, the increased preservation of organic matter at the sea floor was more significant in black shale formation than the role of enhanced productivity.

High-resolution carbon isotope records of the Aptian to Lower Albian from SE France and the Mazagan Plateau (DSDP Site 545): a stratigraphic tool for paleoceanographic and paleobiologic reconstruction

High-resolution carbon isotope stratigraphy is established for the Aptian to Lower Albian of the Vocontian Basin (SE France), and correlated to the carbon isotope record of the Mazagan Plateau (DSDP Site 545). The carbon isotope stratigraphy of the Vocontian Basin is proposed as a standard reference curve for the Aptian to Lower Albian, due to the completeness and high temporal resolution of the stratigraphic succession, the good biostratigraphical time control, and the frequent occurrence of regional to global black shale horizons including Oceanic Anoxic Events 1a (OAE 1a) of the Lower Aptian and OAE 1b of the Lower Albian. The carbon isotope record appears better suited for long-distance short-term correlation of different marine and terrestrial environments than biostratigraphy because of the synchroneity of carbon isotope signals in a range of sediment types. However, the combination of both biostratigraphy and carbon isotope stratigraphy provides an effective tool to reconstruct biotic change and paleoceanography, and to correlate regional to global black shale horizons in different marine environments. This combined approach allows us to ascertain the synchroneities or diachroneities of first and last appearances of biostratigraphic marker species. Based on the demonstrated diachroneity of important biostratigraphic markers of the Aptian/Albian boundary, the globally observed break point between the end of the uppermost Aptian positive carbon isotope excursion and the onset of the pronounced negative shift of δ13C values, is an alternative criterion. The distinctive structure and amplitudes of the carbon isotope record are observed in both the inorganic and organic carbon and can therefore be recognized in all marine and terrestrial environments of the Aptian to Lower Albian.

Modern planktic foraminifera

Planktic foraminifers are marine protozoans with calcareous Shells and chambered tests. They first appeared in the mid-Jurassic and spread since the mid-Cretaceous over all the world’s oceans. Modern planktic foraminifers evolved since the early Tertiary, when the first spinose species occurred. Most species live in the surface to sub-thermocline layer of the open ocean, and in marginal seas like the Mediterranean, Caribbean, South China Sea, and Red Sea. Planktic foraminifers are absent in shallow marginal seas, for example, the North Sea. Planktic foraminifers respond to food, temperature and chemistry of the ambient seawater. Species abundance varies according to seasons, water masses, and water depths. Symbiont-bearing species depend on light and are restricted to the euphotic zone. Planktic foraminifers constitute a minor portion of total Zooplankton, but are major producers of marine calcareous particles (shells) deposited on the ocean floor where they form the so-called foraminiferal ooze.Planktic foraminifers contribute substantially to the fossil record of marine Sediments and are of high ecologic, paleoceanographic, and stratigraphic significance since the mid-Cretaceous. Radiocarbon (14C) gives an absolute age of shell formation within late Pleistocene and Holocene Sediments. Factors that determine the modern faunal composition are applied to Interpretation of the fossil assemblages, for example, by multiple regression techniques (transfer functions) to yield an estimate on ancient environmental parameters. The chemical composition of the calcareous shell (stable isotopes and trace elements) holds clues to the chemical and physical State of the ambient seawater and is useful in the reconstruction of temperature, chemical State, and biological productivity of the ancient marine environment.KurzfassungPlanktische Foraminiferen sind kalkschalige, marine Protozoen mit gekammerten Gehäusen. Sie sind seit dem mittleren Jura (∼170 Millionen Jahre) fossil überliefert. Seit der mittleren Kreide sind planktische Foraminiferen im marinen Pelagial weit verbreitet. Die meisten Arten sind an der Kreide / Tertiär-Grenze ausgestorben. Die modernen Arten, mit den spinösen -stacheltragenden- Arten, haben sich seit dem Tertiär entwickelt. Der Lebensraum der meisten Arten ist die euphotische Deckschicht bis knapp unterhalb der saisonalen Thermokline. Wenige Arten leben in der Tiefsee. Auch in tiefen Randmeeren leben planktische Foraminiferen, wie etwa im Mittelmeer, der Karibik, dem Südchinesischen Meer und dem Roten Meer. In flachen Randmeeren, wie der Nordsee, sind planktische Foraminiferen nicht heimisch. Das Artenspektrum variiert entsprechend des Futterangebotes, der Temperatur und des Chemismus des umgebenden Wassers. Symbiontentragende Arten sind lichtabhängig und an die euphotische Zone gebunden. Planktische Foraminiferen bilden nur einen geringen Teil der planktischen Biomasse, sind aber hauptsächlich an der Produktion und Sedimentation des marin-pelagischen, partikulären Karbonates beteiligt.Planktische Foraminiferen bilden den sogenannten Foraminiferen-Schlamm am Meeresboden und tragen substantiell zum fossilen Inhalt mariner Sedimente bei. Seit der mittleren Kreide sind planktische Foraminiferen stratigraphische Leitfossilien und wichtige ökologische und paläoozeanographische Indikatoren. Radiokohlenstoff (14C) der Foraminiferengehäuse wird zur absoluten Altersdatierung pleistozäner und holozäner Sedimente genutzt. Daten zur Ökologie rezenter Faunen werden mit multiplen Regressionsverfahren (Transferfunktionen) auf fossile Faunen übertragen und damit paläo-ökologische, -ozeanographische und -klimatologische Rekonstruktionen ermöglicht. Die chemische Zusammensetzung (stabile Isotope und Spurenelemente) der kalkigen Foraminiferenschale repräsentiert den chemischen und physischen Zustand des umgebenden Meerwassers und liefert wichtige Daten zur Rekonstruktion von Temperatur, chemischer Zusammensetzung und biologischer Produktivität vergangener Ozeane.

Planktic foraminiferal production stimulated by chlorophyll redistribution and entrainment of nutrients

During September and October 1996 planktic foraminifers and pteropods were sampled from the upper 2500 m of the water column in the BIOTRANS area (47°N, 20°W), eastern North Atlantic, as part of the JGOFS program. Hydrography, chlorophyll fluorescence, and nutrient content were recorded at high spatial and temporal resolution providing detailed information about the transition time between summer and fall. At the beginning of the cruise a shallow pycnocline was present and oligotrophic conditions prevailed. Over the course of the cruise, the mixed layer depth increased and surface water temperature decreased by 1.5°C. Both chlorophyll-a dispersed in the upper 50 m by vertical mixing and chlorophyll-a concentrations at the sea surface increased. The nitracline shoaled and nutrient enriched waters were entrained into the mixed layer. Planktic foraminifers and pteropods closely reflected the changes in the hydrography by increased growth rates and changes in species composition. Three main groups of planktic foraminiferal species were recognized: (1) a temperate and low-productivity group dominated by Neogloboquadrina incompta characterized the shallow mixed layer depths. (2) A temperate and high-productivity group dominated by Globigerina bulloides characterized the period with wind-induced dispersal of chlorophyll-a and entrainment of nutrient-enriched waters. (3) A warm water group containing Globigerinoides sacculifer, Orbulina universa, Globigerinoides ruber (white), and Globigerinella siphonifera was most common during the first days of sampling. Synchronous with the hydrographic change from summer to fall, planktic foraminiferal and pteropod growth was stimulated by redistribution of chlorophyll-a and entrainment of nutrient-enriched waters into the mixed layer. In addition, the seasonal change in the eastern North Atlantic resulted in a transition of the epipelagic faunal composition and an increased calcareous particle flux, which could be used to trace seasonality in fossil assemblages and allow for better paleoceanographic interpretation of the boreal Atlantic.

Benthic foraminiferal record of ecosystem variability in the eastern Mediterranean Sea during times of sapropel S5 and S6 deposition

High-resolution benthic foraminiferal and geochemical investigations were carried out across sapropels S5 and S6 from two sediment cores in the Levantine Sea to evaluate the impact of climatic and environmental changes on benthic ecosystems during times of sapropel formation. The faunal successions indicate that eutrophication and/or oxygen reduction started several thousand years prior to the onset of sapropel formation, suggesting an early response of the bathyal ecosystems to climatic changes. Severest oxygen depletions appear in the early phases of sapropel formation. The initial reduction of deep-water ventilation is caused by a warming and fresh water-induced stratification of Eastern Mediterranean surface waters. During the late phase of S5 formation improved oxygenation is restricted to middle bathyal ecosystems, indicating that at least some formation of subsurface water took place. During S6 formation oxygen depletions and eutrophication were less severe and more variable than during S5 formation. Estimated oxygen contents were low dysoxic at middle bathyal to anoxic at lower bathyal depths during the early phase of S6 formation but never dropped to anoxic values in its late phase. The high benthic ecosystem variability during S6 formation suggests that water column stratification at deep-water formation sites was in a very unstable mode and susceptible to minor temperature fluctuations at a millennial time-scale. 5 2002 Elsevier Science B.V. All rights reserved.

The δ44Ca‐temperature calibration on fossil and cultured Globigerinoides sacculifer: New tool for reconstruction of past sea surface temperatures

We report direct δ44Ca-temperature calibration on cultured and fossil calcite foraminifera, showing that Ca isotopes are potentially a new proxy for past sea surface temperatures (SST). Samples have been analyzed using a 43Ca-48Ca double spike and thermal ionization mass spectrometry (TIMS). In order to avoid species-dependent isotope fractionation we focused our investigations on a single foraminifera species (Globigerinoides sacculifer), which is known to inhabit shallow euphotic waters in tropical and subtropical oceans. Ca isotope measurements were performed on cultured G. sacculifer that grew in seawater kept at temperatures of 19.5°, 26.5°, and 29.5°C. A δ44Ca change of 0.24 ± 0.02 per 1°C is defined by the weighted linear regression through reproduced δ44Ca data of the three temperatures (95% confidence level). Application of this new method to fossil G. sacculifer of an Equatorial East Atlantic sediment core (GeoB1112; 5°46.7′S, 10°45.0′W, 3125 m) indicates that the δ44Ca difference between marine isotope stage 1 (MIS-1) and MIS-2 is 0.71 ± 0.24. According to the current δ44Ca-temperature calibration this value corresponds to a temperature difference between MIS-1 and MIS-2 of ∼3.0 ± 1.0°C.