Christoph Hemleben

• Retired apl. Prof. Dr. Dipl. Geol. Akad.Dir. i.R.
• Eberhard-Karls-Universität Tübingen at University of Tübingen

The Atlas of Oligocene planktonic foraminifera followed a similar concept of wall texture that was applied to Eocene planktonic foraminifera (Hemleben & Olsson, 2006). Detail analyses of wall texture in normal perforate and microperforate planktonic foraminifera was used to determine phylogenetic groups important for their classification. Special a...

Ciperoella Olsson and Hemleben n. gen. is erected for Oligocene spinose species that have a neogloboquadrinid-type wall texture and 4½-5 similarly sized chambers in the final whorl. Four species are recognized as distinct, namely Ciperoella anguliofficinalis (Blow), Ciperoella angulisuturalis (Bolli), Ciperoella ciperoensis (Bolli), and Ciperoella...

The taxonomy, phylogeny and biostratigraphy of Oligocene and lower Miocene Globoturborotalita is reviewed. Globoturborotalita is a long-ranging genus appearing in the basal Eocene and still present in modern oceans with one living representative G. rubescens. Species attributed to this genus are generally common and cosmopolitan. The following spec...

The taxonomy, phylogeny, and biostratigraphy of Oligocene and lower Miocene Dentoglobigerina and Globoquadrina are reviewed. Because of the discovery of spine holes in various species assigned to these genera, the entire group is now considered to have been fully or sparsely spinose in life and hence part of Family Globigerinidae. One new species,...

Classification of modern and fossil planktic foraminifers is based on a morphological species concept (i.e. morphotypes) for practical reasons, i.e. a non-destructive enumeration from strew-mounted samples, and economical (i.e. time-saving) analyses. Detailed classification of each test, for example, using scanning electron microscopy or analysis o...

Calcification and dissolution of test CaCO3 cause changes in the surface water carbonate system. Deep-water chemistry affects and is affected by the dissolution of tests (e.g., Berger and Piper in JAMA 17:275–287, 1972; Dittert et al. in Use of proxies in paleoceanography. Springer, Berlin, pp. 255–284, 1999).

The temporal and spatial distribution of diet is presumably a major cause for the regional distribution of planktic foraminifer species, by providing the basis for growth and affecting their fecundity.

Continuity of the species in the vastness of the deep ocean is ensured by adaptive mechanisms characteristic of pelagic organisms to promote sufficient reproductive success.

Tests of varying complexity represent the most obvious and persistent visual expression of the overall planktic foraminifer morphology. Consequently, understanding the biological processes and environmental controls involved in the formation and deposition of the planktic foraminifer shell is of major interest to any related discipline, including m...

Analyses of planktic foraminifers are targeted towards three main goals.

The calcareous planktic foraminifer shell has been analyzed for its chemical composition, and assumed proxy of the chemical composition of seawater since the pioneering works of Samuel Epstein and Cesare Emiliani in the 1950s (e.g., Epstein et al. in Geol Soc Am Bull 62:417–426, 1951; Emiliani in J Geol 63:538–578, 1955).

Although the shell is one of the most obvious features of the living and dead planktic foraminifers, the cytoplasm is clearly of central biological importance as the site of the dynamic properties of life including transformation of the information stored in the genetic code into substantive morphology, metabolism, sensitivity, and activity, as wel...

Most of the about 50 extant planktic foraminifer morphospecies are ubiquitous in the global ocean (e.g., Bé in Oceanic micropaleontology. Academic Press, London, pp 1–100 1977; Hemleben et al. in Modern planktonic Foraminifera. Springer, Berlin, 1989). Single genotypes of those morphotypes are more limited to ocean basins and regions (e.g., Darling...

This book provides a comprehensive overview of the taxonomy, biology, sedimentation, and carbonate geochemistry of modern species. Students, early career and advanced scientists alike will profit from a broad synthesis of the current understanding of planktic foraminifers as an ecological indicator, biogeochemical factories, and proxies in paleocea...

Planktic foraminifers are marine protozoans with calcareous shells and chambered tests (Plate 1.1), first appearing in the mid-Jurassic approximately 170 million years ago, and populating the global ocean since the mid-Cretaceous (cf. Frerichs et al. in J Foraminifer Res 2:6–13, 1972; Caron and Homewood in Mar Micropaleontol 7:453–462, 1983).

In order to assess how insolation-driven climate change superimposed on sea level rise and millennial events influenced the Red Sea during the Holocene, we present new paleoceanographic records from two sediment cores to develop a comprehensive reconstruction of Holocene circulation dynamics in the basin. We show that the recovery of the planktonic...

In order to assess how insolation-driven climate change superimposed on sea level rise and millennial events influenced the Red Sea during the Holocene, we present new paleoceanographic records from two sediment cores to develop a comprehensive reconstruction of Holocene circulation dynamics in the basin. We show that the recovery of the planktonic...

We critically evaluate the applicability of Ca-isotope ratios in planktonic foraminifers as proxy for past sea surface temperatures (SST) and isotope composition of paleo-seawater (δ44Casw) reconstructions. Previous studies have shown discrepancies regarding the temperature sensitivity of Ca isotope fractionation in foraminifers of more than one or...

List of sequences newly obtained in the current study, including accession numbers and affiliations to morphospecies.

Contains all alignments in FASTA and all phylogenetic trees in Newick format inferred in the course of the study. The trees are not collapsed and appropriately annotated. The pseudocharacter matrix for the comparison with the current taxonomy is also contained, as well as the substitution matrix used for running poa and poaglo.

Comparative list of bootstrap support under ML and MP based on the six selected alignments; along with a qualitative rating of the listed bipartitions, and according phylogenetic relationships.

The high sequence divergence within the small subunit ribosomal RNA gene (SSU rDNA) of foraminifera makes it difficult to establish the homology of individual nucleotides across taxa. Alignment-based approaches so far relied on time-consuming manual alignments and discarded up to 50% of the sequenced nucleotides prior to phylogenetic inference. Her...

The chemical composition of shells of the planktonic foraminifer Globigerinoides ruber (white) is frequently used to determine past sea surface conditions. Recently, it has been shown that arbitrarily defined morphotypes within this species exhibit different chemical and isotopic signatures. Here, we investigate the occurrence through time and in s...

Published stable isotope records in marine carbonate are characterized by a positive δ18O excursion associated with a negative δ13C shift during the early Maastrichtian. However, the cause and even the precise timing of these excursions remain uncertain. We have generated high-resolution foraminiferal stable isotope and gray-scale records for the l...

The Red Sea is a marginal, oligotrophic basin of the Indian Ocean located at the interface between two climate systems: the Indian Monsoon and the Mediterranean climate. Whereas on glacial-interglacial timescales, the oceanography of the Red Sea is determined by eustatic sea level changes, at times of relatively stable sea level, the oceanography o...

The Red Sea is an extreme marine environment, with conditions limiting the application of standard geochemical proxies for the reconstruction of paleoclimate. In order to develop paleoenvironmental reconstruction methods which are not dependent on chemical signals, we investigated the distribution of planktonic foraminifera in the surface sediments...

We present SSU rDNA data resolving the seasonal and geographical distribution of 'cryptic' genetic types of the planktonic foraminifer morphospecies Globigerinoides ruber in the eastern Atlantic Ocean and the Mediterranean Sea. Analysis of 262 sequences revealed the presence of five genetic types belonging to two distinct lineages. Although the mor...

The Red Sea represents an extreme marine environment, with high salinity, high temperature and low level of nutrients, complicating the application of standard geochemical palaeotemperature proxies. In order to investigate the applicability of the TEX86 (TetraEther indeX of GDGTs with 86 carbons) proxy for sea surface temperature (SST) in the Red S...

Three multicores were studied in the Central Red Sea off Port Sudan at a W—E transect in order to reconstruct palaeoceanographic conditions of the past ~6000 years. Downcore fluctuations in the relative abundance of the epipelagic planktic foraminifera Globigerinoides ruber versus that of the deeper dweller G. sacculifer, together with the presence...

Ice cores from Antarctica record temperature and atmospheric carbon dioxide variations over the past six glacial cycles. Yet concomitant records of sea-level fluctuations-needed to reveal rates and magnitudes of ice-volume change that provide context to projections for the future-remain elusive. Reconstructions indicate fast rates of sea-level rise...

Virgulinella fragilis can be mainly observed in different, separated, oxygen-depleted and sulfide-enriched environments around the world and seems to be well adapted to such extreme habitats. Dispersal mechanisms behind this geographical distribution pattern are not yet understood. To analyze the genetic differentiation of geographically isolated p...

The high sequence divergence within the small subunit ribosomal RNA gene (SSU rDNA) of foraminifera makes it difficult to establish the homology of individual nucleotides across taxa. Alignment-based approaches so far relied on time-consuming manual alignments and discarded up to 50% of the sequenced nucleotides prior to phylogenetic inference. Her...

The chemical composition of shells of the planktonic foraminifer Globigerinoides ruber (white) is frequently used to determine past sea surface conditions. Recently, it has been shown that arbitrarily defined morphotypes within this species exhibit different chemical and isotopic signatures. Here, we investigate the occurrence through time and in s...

Published stable isotope records in marine carbonate are characterized by a positive d18O excursion associated with a negative d13C shift during the early Maastrichtian. However, the cause and even the precise timing of these excursions remain uncertain. We have generated high-resolution foraminiferal stable isotope and gray-scale records for the l...

To settle debate on the timing of sea level fluctuations during marine isotope stage (MIS) 3, we present records of d18Oruber (sea level proxy) and magnetic susceptibility from the same samples within the single sediment archive (i.e., ‘‘coregistered’’) of central Red Sea core GeoTu¨ -KL11. Core-scanning X-ray fluorescence and environmental magneti...

Living (rose-Bengai-stained) foraminifera were investigated from the continental margin off the Pacific Coast of Costa Rica and Nicaragua. Assemblages from seven stations' including mounds and mud volcanoes, were compared in terms of foraminiferal density, distribution patterns, diversity and species composition and related to environmental factors...

To settle debate on the timing of sea level fluctuations during marine isotope stage (MIS) 3, we present records of d18O ruber (sea level proxy) and magnetic susceptibility from the same samples within the single sediment archive (i.e., "coregistered") of central Red Sea core GeoTü-KL11. Core-scanning X-ray fluorescence and environmental magnetic d...

To settle debate on the timing of sea level fluctuations during Marine Isotope Stage (MIS) 3, we present records of ?18Oruber (sea level proxy) and magnetic susceptibility from the same samples within the single sediment archive (i.e., ‘co-registered’) of central Red Sea core GeoTü-KL11. Core-scanning X-ray fluorescence and environmental magnetic d...

The last interglacial period, Marine Isotope Stage (MIS) 5e, was characterized by global mean surface temperatures that were at least 2 °C warmer than present1. Mean sea level stood 4–6 m higher than modern sea level2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, with an important contribution from a reduction of the Greenland ice sheet1, 14. Although some...

Specific responses of the regional deep-sea ecosystems to climatic and oceanographic processes during the last 22 kyr are revealed by benthic foraminiferal faunas from two cores in the northern and southern Aegean Sea. Under glacial boundary conditions, high-diversity benthic foraminiferal faunas and elevated benthic foraminiferal numbers indicate...

Strontium isotope ratios of the HCL-insoluble residue (“ISR”) and foraminifera of cores from the Red Sea and Gulf of Aden are used to monitor effects of hydrothermal, fluvial and desert dust transport to these regions during the past ∼0.5 Ma. While the Gulf of Aden was open-ocean, during low glacial sea levels the Red Sea was a semi-isolated basin,...

Living planktonic foraminifera (PF) samples from the Okinawa Trough of the northwestern Pacific Ocean were taken for DNA analysis. The SSU rDNA sequences of two PF species, Globigerina sp. and Pulleniatina obliquiloculata collected at Station WP01, were obtained and compared with those from the southwestern Pacific Ocean. Only small differences (<0...

Different source areas, oceanography and climate regimes influenced the clay mineral assemblages and grain size distribution of two sediment cores from the North and South Aegean Sea during the last glacial and the Holocene. In the North Aegean Sea, clay mineral composition is mainly controlled by sea level evolution, melting of southeastern Europe...

SYNOPSIS. During gametogenesis mother individuals of Hastigerina pelagica (d'Orbigny) undergo significant morphological changes. Thirty h before gamete release, the cytoplasm changes from pale orange to bright red, possibly due to transport of stored lipids from the inner region to more peripheral parts of the cytoplasm. During the next 10 to 15 h...

Well-preserved opal planktic assemblages containing diatom and silicoflagellate species were discovered in the finely laminated sapropels S5 and S7 from Site 67, core GeoTüKL51 during “RV Meteor”-Cruise 40, Leg 4 (34° 48 N; 27° 17 E). The siliceous microflora, composed quasi-exclusively of warm water species, is here studied for taxonomy, stratigra...

Size restricted carbon isotopes (δ13C) are used to track changes in the ontogenetic life strategies of two species of extinct planktonic foraminifera and demonstrate that the species Morozovelloides crassatus lost their photosymbiotic association prior to their extinction in the latest middle Eocene. M. crassatus exhibit a strong positive correlati...

We present new data on nuclear DNA sequences coding for the 3′ region of the small subunit ribosomal RNA (SSU rDNA), the 5.8 subunit and the internal transcribed spacers (ITS1 and ITS2). 56 SSU rDNA and 23 ITS clones were obtained from 12 specimens of the genus Chilostomella collected in the Pacific Ocean, the Atlantic Ocean and the Mediterranean a...

Benthic foraminiferal faunas from the Lower Maastrichtian (Globotruncana falsostuarti–Gansserina gansseri Planktic Foraminiferal Zone) of DSDP Site 390 (Blake Nose, western North Atlantic) were investigated in order to characterize changes in organic matter flux and bottom-water oxygenation and their relation to paleoceanographic changes. Benthic f...

Benthic foraminiferal assemblages are a widespread tool to understand changes in organic matter flux and bottom-water oxygenation and their relation to paleoceanographic changes in the Upper Cretaceous oceans. In this study, assemblage data (diversity, total number, and number per species and gram) from Deep Sea Drilling Project (DSDP) Site 390 (Bl...

Different source areas, oceanography and climate regimes influenced the clay mineral assemblages and grain size distribution of two sediment cores from the North and South Aegean Sea during the last glacial and the Holocene. In the North Aegean Sea, clay mineral composition is mainly controlled by sea level evolution, melting of southeastern Europe...

The termination of the penultimate glacial period (TII) shows both similarities and differences to the last termination (TI). Both terminations show significant cold reversals in the postglacial warming trend. TI consists of a continuously increasing sea-level trend, whereas TII may demonstrate a sea-level reduction midway through the termination....

Sediments from the western and southern part of the Arabian Sea were collected periodically in the spring intermonsoon between March and May 1997 and additionally at the end of the Northeast Monsoon in February 1998. Assemblages of Rose Bengal stained, living deep-sea benthic foraminifera, their densities, vertical distribution pattern, and diversi...

Sediments from the western and southern part of the Arabian Sea were collected periodically in the spring intermonsoon between March and May 1997 and additionally at the end of the Northeast Monsoon in February 1998. Assemblages of Rose Bengal stained, living deep-sea benthic foraminifera, their densities, vertical distribution pattern, and diversi...

Living (Rose Bengal stained) foraminifera in gas-hydrate-influenced sediments at the Cascadia convergent margin were investigated. Foraminiferal assemblages from the southern Hydrate Ridge and neighboring basins were compared in terms of abundances, vertical distribution, diversity, and species composition. At Hydrate Ridge, the presence of shallow...

Fluctuations in abundance, diversity and species composition of benthic foraminifera from two sites of the northern and southern Red Sea indicate strong variability of deep-sea ecosystems during the last four glacial to interglacial cycles. In total, five and four different benthic foraminiferal assemblages have been identified in the northern core...

Various proxy records from late Glacial and Holocene sediments of the eastern Mediterranean Sea reveal significant temporal and spatial variability suggesting that the different sub-basins were influenced by different oceanographic and climate regimes. In the North Aegean Sea, environmental conditions are controlled by sea level, exchange of water...

Benthic foraminiferal and calcareous nannofossil as-semblages, as well as stable isotope data from the Cam-panian/Maastrichtian boundary interval (71.4 to 70.7 Ma) of the Kronsmoor section (North German Basin), were investigated in order to characterize changes in surface-water productivity and oxygen content at the seafloor and their link to clima...

As one of the most important groups in micropaleontology, Foraminifera is traditionally described to have a membranous, agglutinated or carbonate shell according to its morphology, which resembles the marine granuloreticuloseans. However, recent molecular analyses on its ribosomal RNA gene have disclosed the existence of the naked, and also freshwa...

The Niveau Breistroffer black shale succession in the Vocontian Basin (SE France) is the regional equivalent of the widely distributed Late Albian Oceanic Anoxic Event 1d. The studied black shale-rich interval at the Col de Palluel section is 6.28 m thick and comprises four black shale units with up to 2.5 wt% total organic carbon (TOC) intercalate...

Long-term observations on living benthic foraminifera in the laboratory were performed to investigate their behaviour and life style. We noticed that the formation of cysts or sedimentary envelopes is a common feature within many groups of foraminifera (organic walled, agglutinated and calcareous) in the laboratory. Several kinds of cyst were obser...

The behavior of deep-sea benthic foraminifera in response to seasonal inputs of food material was examined through feeding experiments. The experiments using sediments from Sagami Bay (1425 m water depth) containing Iiving foraminifera were carried out using microcosms (aquaria) with thin-walled glass sides. Three different nutrient levels were tes...

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 l...

The Early Albian Oceanic Anoxic Event 1b (OAE 1b) black shale is interrupted by one or more ventilation events that display significant changes in benthic and planktic popu-lations. Within the OAE 1b sections studied, at ODP Site 1049, DSDP Site 545, and the Vocontian Basin, the benthic foraminiferal repopulation events last between 500 and 1,250 y...

The Early Albian Oceanic Anoxic Event 1b (OAE 1b) black shale is interrupted by one or more ventilation events that display significant changes in benthic and planktic populations. Within the OAE 1b sections studied, at ODP Site 1049, DSDP Site 545, and the Vocontian Basin, the benthic foraminiferal repopulation events last between 500 and 1,250 ye...

The Early Albian Oceanic Anoxic Event 1b (OAE 1b) black shale is interrupted by one or more ventilation events that display significant changes in benthic and planktic populations. Within the OAE 1b sections studied, at ODP Site 1049, DSDP Site 545, and the Vocontian Basin, the benthic foraminiferal repopulation events last between ~500 and ~1,250...

Benthic foraminiferal and calcareous nannofossil assemblages, as well as stable isotope data from the Campanian/Maastrichtian boundary interval (~71.4 to ~70.7 Ma) of the Kronsmoor section (North German Basin), were investigated in order to characterize changes in surface-water productivity and oxygen content at the seafloor and their link to clima...

The Niveau Breistroffer black shale succession in the Vocontian Basin (SE France) is the regional equivalent of the widely distributed Late Albian Oceanic Anoxic Event 1d. The studied black shale-rich interval at the Col de Palluel section is 6.28 m thick and comprises four black shale units with up to 2.5 wt% total organic carbon (TOC) intercalate...

Up to now the benthic foraminiferal classification is based mainly on morphological characters leading to controversies and systematic rearrangements. Previous molecular phylogenetic studies using nuclear encoded small subunit (SSU) rDNA as a marker were able to resolve some of the major divergence points. However, they partly contradicted the morp...

Field studies of recent benthic foraminifera have shown that the vertical distribution in the sediment is controlled mainly by two parameters: oxygen concentration and food availability. The aim of the present paper is to determine the response of deep-sea foraminifera to changing bottom and pore water oxygen concentrations in the laboratory, under...

Here we outline a new, pragmatic methodology to derive relative sea-level estimates from central Red Sea oxygen isotope records based on a previously published model. In this paper, the methodology is described in detail, and it is shown that sea-level change is the dominant factor responsible for the recorded variability in Red Sea δ18O (PDB) for...

We propose that the observed short-term stable isotope fluctuations reflect changes in high- and low-latitude intermediate to deep water sources, based on a high-resolution stable isotope record of planktic and benthic foraminifera from the Early Maastrichtian (∼71.3 to ∼69.6 Ma) of Blake Nose (DSDP Site 390A, North Atlantic). Sources of these wate...

The distribution of diatoms, coccolithophores and planktic foraminifers mirrored the hydrographic and trophic conditions of the surface ocean (0–100 m) across the upwelling area off the Oman coast to the central Arabian Sea during May/June 1997 and July/August 1995. The number of diatoms was increased in waters with local temperature minimum and en...

Foraminiferal assemblages found at Great Meteor Seamount were studied in August 1998. Communities of living foraminifera in surface sediments from the plateau (290–325m water depth) and from the surrounding base (2,300–4,096m) were compared in abundances, diversity, and species composition. In this oligotrophic region, densities were very low, but...

The influence of microhabitat, organic matter flux, and metabolism on the stable oxygen and carbon isotope composition of live (Rose Bengal stained) and dead (empty tests) deep-sea benthic foraminifera from the Gulf of Lions (western Mediterranean Sea) have been studied. The total range of observed foraminiferal isotope values exceeds 1.0‰ for δ18O...

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...