Phylogeny and Rates of Molecular Evolution of Planktonic Foraminifera: SSU rDNA Sequences Compared to the Fossil Record

Planktonic foraminifera are marine protists, whose calcareous shells form oceanic sediments and are widely used for stratigraphic and paleoenvironmental analyses. The fossil record of planktonic foraminifera is compared here to their molecular phylogeny inferred from ribosomal DNA sequences. Eighteen partial SSU rDNA sequences from species representing all modern planktonic families (Globigerinidae, Hastigerinidae, Globorotaliidae, Candeinidae) were obtained and compared to seven sequences representing the major groups of benthic foraminifera. The phylogenetic analyses indicate a polyphyletic origin for the planktonic foraminifera. The Candeinidae, the Globorotaliidae, and the clade Globigerinidae + Hastigerinidae seem to have originated independently, at different epochs in the evolution of foraminifera. Inference of their relationships, however, is limited by substitution rates of heterogeneity. Rates of SSU rDNA evolution vary from 4.0 × 10⁻⁹ substitutions/site/year in the Globigerinidae to less than 1.0 × 10⁻⁹ substitutions/site/year in the Globorotaliidae. These variations may be related to different levels of adaptation to the planktonic mode of life. A clock-like evolution is observed among the Globigerinidae, for which molecular and paleontological data are congruent. Phylogeny of the Globorotaliidae is clearly biased by rapid rates of substitution in two species (G. truncatulinoides and G. menardii). Our study reveals differences in absolute rates of evolution at all taxonomic levels in planktonic foraminifera and demonstrates their effect on phylogenetic reconstructions. Received: 21 January 1997 / Accepted: 17 April 1997     

Paleoceanographic implications of smaller benthic and planktonic foraminifera from the Eocene Pazin Basin (Coastal Dinarides,Croatia)

Summary Smaller benthic and planktonic foraminifera from the clastic sediments of the Pazin Basin (Istria, Croatia) were studied in order to obtain more data about paleoceanographic conditions that existed in the Middle Eocene Dinaric foreland basin. The succession investigated corresponds to the Middle Eocene planktonic foraminiferal zones Globigerapsis kugleri/Morozovella aragonensis (P11), Morozovella lehneri (P12), and Globigerapsis beckmanni (P13). Benthic foraminiferal assemblages from the clastic succession are dominated by epifaunal trochospiral genera suggesting oligotrophic to mesotrophic conditions and moderately oxygenated bottom waters. Planktonic foraminiferal assemblages indicate mesotrophic to eutrophic conditions of the surface waters, with increased eutrophication in the upper part of the section. Water depth, based on the ratio between planktonic and epifaunal benthic foraminifera and on the recognized species of cosmopolitan benthic foraminifera, was estimated to have been between about 900 and 1200 m. The basin was elongated and open to marine currents on both sides allowing good circulation and ventilation of the bottom water.     

Palaeoenvironmental conditions preceding the Messinian Salinity Crisis: A case study from Gavdos Island

The Messinian pre-evaporitic sedimentary succession of Gavdos Island (Metochia section) is a nearly uninterrupted succession of marine sediments, dominated by finely laminated diatomaceous marls, which are cyclically alternating with clayey diatomites and white diatomites. The qualitative and quantitative analysis of the planktonic foraminiferal fauna allowed the recognition of nine bioevents, which have been astronomically dated for the Mediterranean. The base of the diatomitic succession in Gavdos Island is dated at 6.722 Ma and the top at 6.015 Ma. The studied section contains benthic foraminiferal genera characteristic of an outer shelf to slope environment. The qualitative and quantitative analysis of this microfauna revealed three benthic foraminiferal fossil assemblages and the occurrence of allochthonous species transported into the bathyal environment by current activity. The cyclical pattern of the benthic foraminifera assemblages indicates that the studied sediments have been affected by repeated episodes of basin restriction characterized by low diversity benthic foraminifera populations, and a limited planktonic foraminifer association typified by shallow, surface-dwelling forms. This restriction was partly due to Antarctic cooling, which produced palaeo-Mediterranean sea-level oscillations during the Early Messinian, as a prelude to closure of the Atlantic connections. The relative impact of climatic versus tectonic control on sedimentation patterns within this basin is discussed.     

Comparison of radiolarian and sedimentologic paleoproductivity proxies in the latest Miocene–Recent Benguela Upwelling System

Estimating past ocean productivity from ocean sediments often gives different results depending on the measurement used. We have examined a suite of paleoproductivity proxies in latest Miocene–Recent sediments from DSDP Site 532 and ODP Site 1084, two deep-sea sections underlying the Benguela Upwelling System off the Atlantic coast of southern Africa. The productivity history of this system has been previously established via organic carbon concentration, diatom floras and alkenone based estimates of surface water temperature, and shows a change from low productivity in the early Pliocene to sustain high productivity in the late Pliocene–Recent. Each of our samples was split and simultaneously analysed for several proxies of ocean productivity, including organic carbon (TOC%), carbonate, abundance of opaline radiolarians, accumulation rate of benthic foraminifera (BFAR); the radiolarian faunal composition indices Upwelling Radiolarian Index (URI) and the Water Depth Ecology index (WADE); other proxies for opal and carbonate dissolution, plus stable isotopes of benthic foraminifera. Comparisons between proxies in the same measured samples, between sites in downcore plots and to the published productivity record for this region suggest that TOC and radiolarian faunal composition, particularly the WADE index, are good indicators of past productivity, albeit with different sensitivities (log–linear correlation WADE–TOC% r = 0.78, n = 65, p < 0.01). In contrast, carbonate, and carbonate-based proxies such as BFAR primarily reflect changes in dissolution. Radiolarian faunal composition indices do not appear to be affected by bulk opal accumulation or changes in opal preservation. WADE analysis of radiolarian faunas and TOC% measurements appear to be useful proxies for productivity in late Neogene sediments, particularly for sections where opal or carbonate dissolution is significant.     

Biostratigraphy and paleoecology of the Oligo-Miocene succession in Fars and Khuzestan areas (Zagros Basin,SW Iran)

Paleontological and biostratigraphical studies on carbonate platform succession from southwest Iran documented a great diversity of shallow-water benthic foraminifera during the Oligocene–Miocene. Larger foraminifera are the main means for the stratigraphic zonation of carbonate sediments. The distributions of larger benthic foraminifera in two outcrop sections (Abolhayat and Lali) in the Zagros Basin, Iran, are used to determine the age of the Asmari Formation. Four assemblage zones have been recognized by distribution of the larger benthic foraminifera in the study areas. Assemblage 3 (Aquitanian age) and 4 (Burdigalian age) have not been recognized in the Abolhayat section (Fars area), due to sea-level fall. The end Chattian sea-level fall restricted marine deposition in the Abolhayat section and Asmari Formation replaced laterally by the Gachsaran Formation. This suggests that the Miocene part of the formation as recognized in the Lali section (Khuzestan area) of the Zagros foreland basin is not present in the Abolhayat outcrop. The distribution of the Oligocene larger benthic foraminifera indicates that shallow marine carbonate sediments of the Asmari Formation at the study areas have been deposited in the photic zone of tropical to subtropical oceans. Based on analysis of larger benthic foraminiferal assemblages and microfacies features, three major depositional environments are identified. These include inner shelf, middle shelf and outer shelf. The inner shelf facies is characterized by wackestone–packstone, dominated by various taxa of imperforate foraminifera. The middle shelf is represented by packstone–grainstone to floatstone with a diverse assemblage of larger foraminifera with perforate wall. Basinwards is dominated by argillaceous wackestone characterized by planktonic foraminifera and large and flat nummulitidae and lepidocyclinidae. Planktonic foraminifera wackestone is the dominant facies in the outer shelf.     

Molecular evidence for an independent origin of modern triserial planktonic foraminifera from benthic ancestors

Gallitellia vivans is the only Recent representative of the triserial planktonic foraminiferal family Guembelitriidae. The origin and evolution of this interesting albeit poorly known family are enigmatic. To elucidate the phylogenetic relationships between G. vivans and other planktonic foraminifera, we sequenced the small subunit ribosomal DNA (SSU rDNA) for comparison to our extensive database of planktonic and benthic species. Our analyses suggest that G. vivans represents a separate lineage of planktonic foraminifera, which branches close to the benthic rotaliids Stainforthia and Virgulinella. Both genera resemble Gallitellia in general morphological appearance, having elongate triserial tests at least in their early ontogenic stages. The divergence time of G. vivans is estimated at ca. 18 Ma (early Miocene), suggesting an origin independent from the Cretaceous and Paleogene triserial planktonic foraminifera. Our study thus indicates that modern triserial planktonic foraminifera are not related to the Cretaceous–Paleogene triserial species, and that the sporadic occurrences in the fossil record are not the result of poor preservation, but reflect multiple transitions from benthic to planktonic mode of life.     

Molecular evidence for an independent origin of modern triserial planktonic foraminifera from benthic ancestors

Gallitellia vivans is the only Recent representative of the triserial planktonic foraminiferal family Guembelitriidae. The origin and evolution of this interesting albeit poorly known family are enigmatic. To elucidate the phylogenetic relationships between G. vivans and other planktonic foraminifera, we sequenced the small subunit ribosomal DNA (SSU rDNA) for comparison to our extensive database of planktonic and benthic species. Our analyses suggest that G. vivans represents a separate lineage of planktonic foraminifera, which branches close to the benthic rotaliids Stainforthia and Virgulinella. Both genera resemble Gallitellia in general morphological appearance, having elongate triserial tests at least in their early ontogenic stages. The divergence time of G. vivans is estimated at ca. 18 Ma (early Miocene), suggesting an origin independent from the Cretaceous and Paleogene triserial planktonic foraminifera. Our study thus indicates that modern triserial planktonic foraminifera are not related to the Cretaceous–Paleogene triserial species, and that the sporadic occurrences in the fossil record are not the result of poor preservation, but reflect multiple transitions from benthic to planktonic mode of life.     

Paleoenvironmental evolution of the eastern Mediterranean during the Messinian: Constraints from integrated microfossil data of the Pissouri Basin (Cyprus)

Integrated data of calcareous nannofossils, as well as planktonic and benthic foraminifera from the Pissouri Motorway section on Cyprus allow the reconstruction of surface- and bottom-water paleoenvironments of the eastern Mediterranean during the interval preceding the Messinian salinity crisis (MSC). Contrary to deeper-water locations, where benthic foraminifera faunas are suppressed or absent just after the Tortonian–Messinian boundary, sediments deposited at intermediate water depths do contain benthic assemblages. From the earliest Messinian onwards, a development towards increasingly unfavourable paleoenvironments is reflected in the planktonic and benthic microfossil records of the Pissouri section and proceeds with rather discrete time steps that can be correlated to sequences throughout the Mediterranean. Shortly after the Tortonian–Messinian boundary a transition is recorded in the sedimentology and the open marine, deeper-water taxa disappear from the benthic foraminifera assemblages; subsequently, the diversity of all fauna groups diminishes. The changes recorded at species level in both surface-water and sea-floor dwelling taxa suggest decreasing circulation of the bottom waters, associated with changes in the surface waters, most likely due to increasing stratification. From ∼6.73 Ma onwards, our data indicate a prominent change to more restricted conditions and increasing salinity at the sea floor together with intermittently rising surface water salinity. The dominance of oligotypic and monospecific assemblages and the frequent shifts in assemblage compositions of all microfossil groups indicate severely stressed environments after ∼6.4 Ma, probably related to increased salinity. The major changes in paleoenvironmental conditions, including oxygen deprivation due to stagnation and hypersalinity, can be explained by hydrographical changes in the Mediterranean basin, which are probably caused by tectonic movements in the Rif Corridor acting in concert with astronomical cyclicity. Evaluation of the paleodepth proxies indicates that the depth of the Pissouri Basin remained rather constant at ∼300–500 m, with a minimum of 200 m, until deposition of the “barre jaune”, the transitional interval towards the evaporites and that early shallowing to neritic depths, as was proposed before, is highly unlikely.     

Living benthic foraminifera of the Okhotsk Sea: Faunal composition,standing stocks and microhabitats

Live (Rose Bengal stained) benthic foraminifera were investigated in surface sediment samples from the Okhotsk Sea to reveal the relationship between faunal characteristics and environmental parameters. Live benthic foraminifera were quantified in the size fraction > 125 µm in the upper 8 cm of replicate sediment cores, recovered with a multicorer at five stations along the Sakhalin margin, and at three stations on the southwestern Kamchatka slope. The stations are from water depths between 625 to 1752 m, located close or within the present Okhotsk Sea oxygen minimum zone, with oxygen levels between 0.3 and 1.5 ml l^- 1. At the high-productivity and ice-free Kamchatka stations, live benthic foraminifera are characterized by maximal standing stocks (about 1700-3700 individuals per 50 cm²), strong dominance of calcareous species (up to 87-91% of total live faunas), and maximal habitat depths (down to 5.2-6.7 cm depth). Vertical distributions of total faunal abundances exhibit a clear subsurface maximum in sediments. At the Sakhalin stations, which are seasonally ice-covered and less productive, live benthic foraminifera show lower standing stocks (about 200-1100 individuals per 50 cm²), lower abundance of calcareous species (10-64% of total live faunas), and shallower habitat depths (down to 2.5-5.4 cm depth). Faunal vertical distributions are characterized by maximum in the uppermost surface sediments. It is suggested that 1) lower and strongly seasonal organic matter flux, caused by the seasonal sea ice cover and seasonal upwelling, 2) lower bottom water oxygenation (0.3-1.1 ml l^- 1), and 3) more pronounced influence of carbonate undersaturated bottom water along the Sakhalin margin are the main factors responsible for the observed faunal differences. According to species downcore distributions and average living depths, common calcareous species were identified as preferentially shallow, intermediate and deep infaunal. Foraminiferal microhabitat occupation correlates with the organic matter flux and the depth of the oxygenated layer in sediments.     

Evidence for calcification depth change of Globorotalia truncatulinoides between deglaciation and Holocene in the Western Atlantic Ocean

Measurements of the δ¹⁸O in tests of planktonic and benthic foraminifera in the Florida Straits are used to reconstruct the properties of the water column through time over the last 12 ka (Lynch-Stieglitz et al., in press). The isotopic composition of the foraminifera largely reflects the vertical density gradient. We use this reconstruction and δ¹⁸O measurements on Globorotalia truncatulinoides in a nearby core to track the depth habitat of this species from the last deglaciation to 1.6 ka B.P. Around 9 ka, G. truncatulinoides was calcifying in much shallower water than during the late Holocene. The downward migration toward its modern habitat is a regional phenomenon over the western tropical Atlantic continental slope. The cause is still unclear but we hypothesize that the shallower calcification depth may be a response to the presence of glacial melt water or to circulation changes. This study points to the value of further study of the ecology, life cycle and calcification depth for G. truncatulinoides and other planktonic foraminifera that are used to reconstruct the history of the thermocline and upper water column structure.     

Neogene stratigraphy,foraminifera, diatoms,and depositional history of Maria Madre Island,Mexico: Evidence of early Neogene marine conditions in the southern Gulf of California

Foraminifera and diatoms have been analyzed from an upper Miocene through Pleistocene(?) sequence of marine sediments exposed on Maria Madre Island, largest of the Trés Marias Islands off the Pacific coast of Mexico. The Neogene stratigraphic sequence exposed on Maria Madre Island includes a mid-Miocene(?) non-marine and/or shallow marine sandstone unconformably overlain by a lower upper Miocene to uppermost Miocene upper to middle bathyal laminated and massive diatomite, mudstone, and siltstone unit. This unit is unconformably overlain by lower Pliocene middle to lower bathyal sandstones and siltstones which, in turn, are unconformably overlain by upper Pliocene through Pleistocene(?) upper bathyal to upper middle bathyal foraminiferal limestones and siltstones. These beds are unconformably capped by Pleistocene terrace deposits. Basement rocks on the island include Cretaceous granite and granodiorite, and Tertiary(?) andesites and rhyolites. The upper Miocene diatomaceous unit contains a low diversity foraminiferal fauna dominated by species of Bolivina indicating low oxygen conditions in the proto-Gulf Maria Madre basin. The diatomaceous unit grades into a mudstone that contains a latest Miocene upper to middle bathyal biofacies characterized by Baggina californica and Uvigerina hootsi along with displaced neritic taxa. An angular unconformity separates the upper Miocene middle bathyal sediments from overlying lower Pliocene siltstones and mudstones that contain a middle to lower bathyal biofacies and abundant planktonic species including Neogloboquadrina acostaensis and Pulleniatina primalis indicating an early Pliocene age. Significantly, this Pliocene unit contains common occurrences of benthic species restricted to Miocene sediments in California including Bulimina uvigerinaformis. Pliocene to Pleistocene(?) foraminiferal limestones and siltstones characterize submarine bank accumulations formed during uplift of the Trés Marias Island area, and include abundant planktonic foraminifera such as Pulleniatina obliquiloculata and Neogloboquadrina duterteri. Common benthic foraminifera in this unit are indicative of upper bathyal water depths. The Neogene depositional history recorded on Maria Madre Island involves an early late Miocene subsidence event marking formation of the Trés Marias Basin with relatively undiluted diatomaceous sediment deposited in a low oxygen setting. Subsidence and deepening of the basin continued into the early Pliocene along with rapid deposition of terrigenous clastics. Uplift of the basinal sequence began in late Pliocene time accompanied by deposition of upper Pliocene-Pleistocene foraminiferal limestones on a rising submarine bank. Continued episodic uplift of the Neogene deposits brought the island above sea level by late Pleistocene time.     

Basin-scale paleobiogeography and paleoecology: an example from Karpatian (Latest Burdigalian) benthic and planktonic foraminifera and calcareous nannofossils from the Central Paratethys

We present the paleobiogeography and ecology of benthic and planktonic foraminifera and calcareous nannoplankton during the Karpatian (Latest  Burdigalian) in the Austrian part of the Central Paratethys. Results are based on quantitative data of assemblages from the Molasse and Styrian Basins. Our data suggest that, although adjacent, the two basins underwent different paleoceanographic evolution during the Karpatian. Dysoxic bottom waters characterized the Molasse Basin, together with reducing environments, which produced the formation of pyritized levels. In the Styrian Basin, intense volcanic activity enriched surface waters in nutrient and produced strong primary productivity. Reduced planktonic foraminiferal abundance and dominance of agglutinated benthic forms in the Styrian Basin can be attributed to paleoceanographic conditions. The planktonic-depleted assemblages bias the paleodepth estimation by 100P(P + B) in the Styrian Basin and new tools for assessing paleodepth are needed. This study shows that paleobiogeography of foraminifera and calcareous nannoplankton is strictly related to local ecological conditions in enclosed basins and demonstrates that quantitative ecological studies are the best way to monitor the biogeographic evolution of enclosed basins.     

Magnesium and strontium compositions of recent symbiont-bearing benthic foraminifera

Minor and trace elements in foraminiferal carbonates are potential paleo-proxies of climate, nutrient and seawater composition. There are very few reports of trace element composition of symbiont-bearing, larger foraminifera that are known to be important constituents of shallow-marine, modern and ancient carbonates. In this paper we examine the range of variation in Mg and Sr content of Recent species of these foraminifera from a lagoon of Lakshadweep Atoll (Indian Ocean) and Akajima Islands, Japan. Two hyaline species, Amphistegina lessonii and Neorotalia calcar,and two porcellaneous species, Amphisorus hemprichii and Marginopora vertebralis were collected live from Lakshadweep islands. Mg / Ca in these foraminifera is of an order of magnitude higher than the values reported for planktonic and symbiont-free benthic foraminifera. The Sr / Ca values are, however, comparable with the reported values in other foraminiferal taxa and they are found to vary within a narrow range. Electron-probe micro-analysis of three symbiont-bearing benthic species indicates spatial heterogeneity of high orders in Mg / Ca composition in all the species. The annual variation in temperature and pH of the lagoon water cannot explain the observed amplitude of the compositional variation. The photosynthesis and respiration of the symbionts and host foraminifera are possibly the major cause of compositional heterogeneity in individual tests, as has also been recently postulated for symbiont-bearing planktonic foraminiferal species. It highlights the need to isolate biological factors and necessitates species-specific paleotemperature scale in paleoclimatic analysis. We also analyzed δ¹⁸O, δ¹³C, Ca, Mg and Sr in carefully dissected chambers of a reef-dwelling, porcellaneous benthic foraminifer, Marginopora kudakajimaensis, collected live in four seasons. A moderate positive correlation is observed between Mg / Ca and temperature. However, large inter- and intra-test variation in Mg limits the precision of Mg / Ca as palaeotemperature proxy. The Sr / Ca of the test calcite is unrelated to temperature of the sea water. The δ¹³C of M. kudakajimaensis does not correlate with δ¹⁸O, Mg / Ca or Sr / Ca.     

Benthic origin and earliest evolution of the first planktonic foraminifera after the Cretaceous/Palaeogene boundary mass extinction

There are several hypotheses on the origin and evolution of the earliest Danian planktonic foraminifera. Most experts suggest that they descended from a few opportunist planktonic foraminifera species, mainly of the genera Guembelitria and Hedbergella, which are usually considered to be survivors of the Cretaceous/Paleogene (K/Pg) boundary mass extinction. Nevertheless, early Danian specimens of Guembelitria and Hedbergella remained morphologically well separated from the associated parvularugoglobigerinids (i.e. Parvularugoglobigerina and Palaeoglobigerina), the first trochospiral planktonic foraminifera appearing after the K/Pg boundary event. The most likely alternative is a benthic origin for the parvularugoglobigerinids, which would be consistent with molecular phylogenetic studies that have suggested several episodes of benthic-planktonic transitions in the evolutionary history of planktonic foraminifera. A review of material from the El Kef section and other Tunisian sections supports the previous hypothesis that the buliminid genus Caucasina is the ancestor of the first parvularugoglobigerinids (i.e. Parvularugoglobigerina longiapertura and Palaeoglobigerina alticonusa), on the basis of similarities in test and apertural morphologies and wall texture. The intermediate morphotypes between caucasinids and parvularugoglobigerinids, which appeared approximately 3–5 kyr after the K/Pg boundary, are assigned to Pseudocaucasina antecessor gen. nov. sp. nov.     

Experimental and field evidence of Antarctic foraminiferal tolerance to anoxia and hydrogen sulfide

The common occurrence of foraminifera in strata deposited under anoxic environmental conditions throughout the geologic record raises the question of whether foraminifera actually survive anoxic or reducing conditions. To test this, benthic foraminifera from McMurdo Sound, Antarctica were exposed to anoxic or reducing conditions for 30 days. These Antarctic forms are known to be exposed to anoxic events in their natural environment. An adenosine triphosphate (ATP) assay was used to determine foraminiferal survival, and their ultrastructure was examined using transmission electron microscopy (TEM) to evaluate any possible effect that might have resulted from exposure to anoxic or reducing conditions. The experimental treatments did not have any statistically significant effects upon the foraminifers with regards to survival or average ATP content per individual including the four most common living species. Some ultrastructural evidence for encystment (fibrillar membrane-bound bodies in the cytoplasm of many specimens) was observed. Field data include the observation thatGlobocassidulina cf.G. biora resides in sediments to depths of 7 cm, as evidenced by ultrastructural investigations. A specimen ofGlobocassidulina cf.G. biora collected from sediments containing anoxic pore waters showed numerous bacteria within the confines of the organic lining. The occurrence of cytoplasmic inclusions similar in appearance to car☐ysomes within these bacteria suggest possible affinities to the chemolithotrophicThiobacillus. Bacteria of this type were not observed in specimens collected from oxygenated pore waters. These field and laboratory findings suggest that certain foraminifera may be facultative anaerobes able to withstand reducing conditions.     

Paleoceanographic interpretations of coccoliths and oxygen-isotopes from the sediment surface of the southwest Indian Ocean

The distribution of forty-four coccolithophore species in one hundred deep-sea core-tops from the southwest Indian Ocean is described. Three coccolith assemblages have been recognised (Maputo, Agulhas Current and deep water) by the relative abundances of four ecologically significant coccolithophore species (Gephyrocapsa oceanica, Emiliania huxleyi, Calcidiscus leptoporus and Umbilicosphaera sibogae). Their biogeographical distribution appears to be related to water temperature, nutrient concentration and dissolution.The degree of preservation of coccoliths and foraminifera indicates that the carbonate lysocline lies somewhere between 3500 and 4000 m, resulting in the concentration of dissolution-resistant microfossils below this depth.Stable oxygen isotope ratios of the planktonic foraminiferal species Globigerinoides sacculifer range between −1.5 to −1.0‰ PDB (equal to 22.8–25.1°C) and occur in a narrow band on the sea floor beneath the “A” route of the Agulhas Current.These values are about 0.5 per mil PDB lighter than samples analyzed on either side of this band and can be explained by the Agulhas Current's elevated temperature at the ocean surface of 2–3°C. Thus an oxygen isotope imprint of the Agulhas Current exists beneath it on the sea floor.The Agulhas Current is probably the major factor influencing sedimentation, sediment-distribution patterns and geological features in the study area. At present it is voluminous and fast flowing, possibly eroding sediments up to 2500 m below the surface.The oxygen-isotope ratios and nannoplankton counts obtained in this study indicate, however, that the majority of samples are most probably recent or at least not older than 85,000 years. This implies that sediments are accumulating on the ocean floor and that the Agulhas Current does not have a pronounced erosional influence, at least in areas from which cores were retrieved for this study.     

Calcareous nannofossil surface sediment assemblages from the Sicily Channel (central Mediterranean Sea): Palaeoceanographic implications

Quantitative analysis of 67 calcareous nannofossil assemblages from surface sediments recovered in a wide area across the Sicily Channel has been carried out in order to improve the interpretation of palaeontological data based on this planktonic group in a key area for Mediterranean palaeoceanographic studies. The investigation focused on three case studies that demonstrate the high potentiality of such a combined approach, taking into account the recent distribution of taxa or groups of taxa on the sea floor and the palaeontological record. The distribution of reworked specimens over the northern Sicily Channel sea floor validates the role of southern Sicily as a source region for reworked nannofossils and the role of rivers as their carrier. Eustatic sea-level fluctuations can be considered to be the main factor that influenced the abundance variations in sedimentary sequences of this area. The distribution of Florisphaera profunda can be explained in terms of topography (positive correlation) and mesoscale oceanographic circulation. In particular, its significant anti-correlation to the amount of chlorophyll-A deduced by satellite imagery validates the use of this species as a proxy for palaeoproductivity reconstructions. Finally, high abundance values of G. oceanica are confined to the westernmost part of the Sicily Channel, coinciding with a water mass salinity minimum. In particular, abundances of up to about 10% were observed in the westernmost part of the African Margin, suggesting the importance of the Atlantic Tunisian Current, whose activity is more pronounced in winter. The comparison of data of this species between 135 and 110 kyr BP, inside and outside the Channel, led us to deduce that the physical transport in almost unmodified waters of Atlantic origin might be the most important factor for its significant occurrence.     

Importance of foraminifera for the formation and maintenance of a coral sand cay: Green Island, Australia

 CaCO₃ production by reef-building organisms on Green Island Reef in the Great Barrier Reef of Australia is estimated and compared with the contribution of benthic foraminifera to the sediment mass of the vegetated sand cay. Major constituents of the cay are benthic foraminifera (mainly Amphistegina lessonii, Baculogypsina sphaerulata, and Calcarina hispida), calcareous algae (Halimeda and coralline algae), hermatypic corals, and molluscs. Among these reef-building organisms, benthic foraminifera are the single most important contributor to the sediment mass of the island (ca. 30% of total sediments), although their production of CaCO₃ is smaller than other reef-building organisms. Water current measurements and sediment traps indicate that the velocity of the current around Green Island is suitable for transportation and deposition of foraminiferal tests. Abundant foraminifera presently live in association with algal turf on the shallow exposed reef flat, whose tests were accumulated by waves resulting in the formation and maintenance of the coral sand cay. Accepted: 30 June 1999     

Benthic foraminifera and their role to decipher paleoenvironment during mid-Cretaceous Oceanic Anoxic Events - the “anoxic benthic foraminifera” paradox

Oceanic Anoxic Events (OAEs) are a widespread, recurring and well-studied feature of the mid-Cretaceous greenhouse climate, reflecting severe perturbations in Earth's climate and oceanography. To investigate bottom-water environments before, during and after these OAEs and to understand the dynamic fluctuations of seafloor environments, benthic foraminifera are a commonly employed tool. Here, a review of the response of benthic foraminiferal assemblages to OAEs (i.e., OAEs 1a, 1b, 1d, 2, and 3) will be given. I will discuss in detail how benthic foraminifera can be used to identify the paleoclimatic and paleoceanographic mechanisms responsible for OAE formation. The main feature of benthic foraminiferal assemblages during oceanic anoxic events is a dramatic decrease in both diversities and abundances or their total absence. However, some studies record the permanent occurrence of benthic foraminifera throughout OAEs or short-term repopulation events. These patterns clearly indicate that mid-Cretaceous OAEs are complex events. The paradox of anoxic (and even euxinic) conditions during OAEs coincident with the occurrence of benthic foraminifera (here called the “anoxic benthic foraminifera” paradox) is most readily explained by the sampling (and therefore temporal) resolution of typical foraminiferal studies that have homogenized hundreds to thousands of years in a single sample.     

Origin of the Eocene planktonic foraminifer Hantkenina by gradual evolution

Hantkenina is a distinctive planktonic foraminiferal genus characterized by the presence of tubulospines (robust hollow projections) on each adult chamber, from Middle and Upper Eocene marine sediments worldwide. Here we illustrate its evolutionary origin using c. 150 specimens from 30 stratigraphic intervals in two sediment cores from Tanzania. The specimens, which span an estimated time interval of 300 ka, show four intermediate steps in the evolution of the tubulospines that amount to a complete intergradation from Clavigerinella caucasica, which does not possess them, to Hantkenina mexicana, which does. Stable isotope analyses indicate that the transitional forms evolved in a deep planktonic habitat not occupied at that time by other species of planktonic foraminifera. We discuss the morphogenetic constraints involved in the evolutionary transition and propose an ecological/adaptive model for the selective pressures that resulted in the evolution of tubulospines. We compare our record with similar, recently described assemblages from Austria and Italy, and we update the biostratigraphy and systematic taxonomy of the key morphospecies involved in the transition.