Planktic foraminifera from the mid-Cretaceous (Barremian–Early Albian) of the North Sea Basin: Palaeoecological and palaeoceanographic implications

A marine Cretaceous succession (Barremian–Albian) of a cored borehole (BGS 81/40), located in the Central North Sea Basin, has been examined with respect to its planktic and benthic foraminiferal content, as well as for calcareous nannofossils. The distribution patterns of foraminifera and calcareous nannofossils allow for a two fold division of the investigated interval. (1) The Barremian–earliest Aptian interval, which reflects a marine, temporary restricted setting. This is indicated by sporadic occurrences of planktic foraminifera with very rare planispiral forms suggesting short-term connections of the Boreal and Tethyan Realms. The benthic foraminiferal assemblages indicate aerobic, sometimes dysaerobic bottom-water conditions. High abundances of nannoconids in the Barremian suggest enhanced stratification and/or warm, oligotrophic surface water. (2) The late Aptian–early Albian interval, which was characterised by an open-oceanic environment with cool and aerobic surface water conditions. Planktic foraminifera are more abundant and diverse than in the lower interval. Trochospiral hedbergellids dominate the foraminiferal assemblages. The episodic occurrences of planispiral, clavate and trochospiral-flattened planktic morphotypes indicate the existence of a seaway between the Boreal and the Tethyan Realms. Aerobic to dysaerobic bottom-water conditions are suggested by the composition of the benthic foraminiferal assemblages. High abundances of cool-water taxa within the calcareous nannofossil assemblages indicate a cooling trend across the latest Aptian and earliest Albian.     

Experimental dissolution of a fossil foraminiferal assemblage (Paleocene–Eocene Thermal Maximum,Dababiya, Egypt): Implications for paleoenvironmental reconstructions

Dissolution experiments were carried out on a foraminiferal assemblage from the Paleocene–Eocene Thermal Maximum (PETM) at Dababiya, Egypt, in order to: 1) reveal the effects of differential dissolution on the composition of the foraminiferal assemblage and 2) develop objective criteria for the evaluation of dissolution in foraminiferal assemblages used in early Paleogene paleoenvironmental reconstructions, particularly with respect to neritic Midway-type assemblages from the Paleocene/Eocene transition. Our results confirm two general observations on modern foraminifera: 1) planktic foraminifera are much more vulnerable to dissolution than benthic foraminifera, leading to depressed P/B ratios and 2) dissolution susceptibility differs between size fractions, with the smaller specimens dissolving more rapidly than the bigger ones, leading to a larger average size of the remaining assemblage. Within a size fraction, wall structure and thickness are considered to be the main factors controlling differential dissolution susceptibility. We propose a ranking scheme for taxa with respect to dissolution resistance. Among the benthic taxa, Lenticulina is most resistant, followed by the agglutinated Gaudryina cf. ellisorae and Alabamina midwayensis. Biserial and triserial hyaline taxa and the porcelaneous Spiroloculina sp. are most susceptible to dissolution, whereas rotaliines, such as Cibicidoides and Anomalinoides have an intermediate susceptibility. This implies that mild dissolution of a Midway-type benthic assemblage leads to a relative enrichment in Lenticulina, Gaudryina and rotaliines. Amongst planktic foraminifera, the muricate taxa Acarinina and Morozovella are most resistant, followed by the cancellate Subbotina. The smooth and generally small Globanomalina and Zeauvigerina are least resistant to dissolution. Our data enable to objectively evaluate various degrees of dissolution in benthic and planktic foraminiferal assemblages retrieved from the lower Paleogene Tethyan outcrops. In this way taphonomic artifacts can be readily distinguished from paleoenvironmental signals affecting the primary composition of the assemblages. More generally, we propose that the combined use of foraminiferal numbers, P/B ratio and relative abundances of non-calcareous agglutinated taxa and Lenticulina may provide a powerful proxy for assessing dissolution in hemipelagic assemblages from Cenozoic and upper Cretaceous continental margins. In order to achieve more robust pre-Quaternary paleoenvironmental reconstructions based on quantitative foraminiferal data, application of dissolution proxies, like proposed here, or in slightly modified form, should become a more widely used micropaleontologic procedure. Particularly continental margin studies dealing with major biotic events (e.g. PETM) or employing P/B ratios for sea-level reconstructions should benefit from such an approach.     

Maastrichtian-early Paleocene foraminiferal palaeobathymetry and depositional sequences at Gebel El Sharawna,south Luxor,Egypt

Maastrichtian-early Paleocene foraminiferal palaeobathymetry, palaeodiversity and vertical facies changes of Gebel El Sharawna, south Luxor, Egypt have been studied to determine the depositional sequences, their relationships to global records and/or tectonic signatures. Five benthonic assemblages are recorded and replicated in the present study reflect fluctuation in palaeo-water depth from restricted marginal marine to outer shelf palaeoenvironments. Four sequence boundaries that coincide with the Campanian/Maastrichtian, intra-early Maastrichtian, Early/Late Maastrichtian, Cretaceous/Palaeogene (K/Pg) and intra-Danian were recognized based upon sharp vertical facies changes, foraminiferal assemblage changes, hiatuses, mineral hard ground and reworking. The K/Pg unconformity reveals an unexpected ca. 4.2 Myr time gap as indicated by the absence of the CF2 Zone through lower part of the P1c Zone. It is easily distinguished in the field by conglomeration and winnowing of phosphate and glauconite in the lower Paleocene. These sequence boundaries defined five third-order depositional sequences mainly developed as the result of the eustatic sea-level changes, coupled with the Arabian–Nubian shield tectonic uplift at the southern edge of the Tethys Ocean.     

Palaeogene marine stratigraphy in China

Palaeogene deposits are widespread in China and are potential sequences for locating stage boundaries. Most strata are non‐marine origin, but marine sediments are well exposed in Tibet, the Tarim Basin of Xinjiang, and the continental margin of East China Sea. Among them, the Tibetan Tethys can be recognized as a dominant marine area, including the Indian‐margin strata of the northern Tethys Himalaya and Asian‐margin strata of the Gangdese forearc basin. Continuous sequences are preserved in the Gamba–Tingri Basin of the north margin of the Indian Plate, where the Palaeogene sequence is divided into the Jidula, Zongpu, Zhepure and Zongpubei formations. Here, the marine sequence ranges from Danian to middle Priabonian (66–35 ma), and the stage boundaries are identified mostly by larger foraminiferal assemblages. The Paleocene/Eocene boundary is found between the Zongpu and Zhepure formations. The uppermost marine beds are from the top of the Zongpubei Formation (~35 ma), marking the end of Indian and Asian collision. In addition, the marine beds crop out along both sides of the Yarlong Zangbo Suture, where they show a deeper marine facies, yielding rich radiolarian fossils of Paleocene and Eocene. The Tarim Basin of Xinjiang is another important area of marine deposition. Here, marine Palaeogene strata are well exposed in the Southwest Tarim Depression and Kuqa Depression. They comprise mostly neritic and coastal lagoon facies of the Tethyan realm. Palaeontological evidence suggests that the Paleocene/Eocene boundary here is in middle of the Qimugen Formation. The Tarim Basin was largely drained by Late Oligocene. To the east, the marine offshore Palaeogene strata are widespread in the North Taiwan and East Zhejiang depressions of the continental shelf basin of East China Sea. Abundant fossils including foraminifera, calcareous nannofossils, ostracods, pollen and bivalves occur in the marine environment. Biostratigraphically, the sequence is well correlated with the international planktonic foraminiferal and nannofossil zonations.     

Pleistocene agglutinated foraminifera from the Lomonosov Ridge and Amundsen Basin, Arctic Basin. Initial report on piston cores 2177-5 (KAL) and 2176-3 (KAL)

Noncalcareous Pleistocene sediments of the Central Arctic Ocean contain sparse benthic foraminiferal assemblages consisting entirely of agglutinated taxa. Deep water agglutinated foraminifera are studied from two piston cores collected from the Lomonosov Ridge and Amundsen Basin [Cores PS 2177-5 (KAL) and 2176-3 (KAL)]. Core PS 2177-5 (KAL) contains an assemblage of 10 species, dominated by Cyclammina pusilla Brady, and is interpreted to reflect a bathyal environment with variable organic flux and nutrition levels. Core PS 2176-3 (KAL) in the Amundsen Basin yielded a very depauperate benthic foraminiferal assemblage. It is assumed that the environment was inhospitable for agglutinated foraminifera.     

Deep-sea benthic foraminiferal recolonization of the 1991 Mt. Pinatubo ash layer in the South China Sea

A census count of Rose Bengal stained benthic foraminifera from the surface area on top of a 2 to 6 cm thick ashfall layer at three deep water stations along the western margin of the Philippines exhibits a unique assemblage composition of benthic foraminifera. The total number of benthic foraminifera is low and the ratio of living individuals to empty tests is high. Specific diversity is low, with a significant dominance of infaunal morphotypes including species of the genus Reophax (R. scorpiurus, R. bilocularis and R. dentaliniformis), which are regarded as successful recolonizers. Assemblages below the ash layers are diverse and contain many epifaunal suspension-feeding agglutinated and calcareous foraminifera. The 1991 Mt. Pinatubo eruption caused mass mortality of benthic foraminifera in a vast area of the eastern South China Sea followed by step-wise recolonization of the ash substrate. Three years after the eruption, the benthic foraminiferal community structure is still far from recovery to background levels.     

Evolution of Paleocene to Early Eocene larger benthic foraminifer assemblages of the Indus Basin,Pakistan

Afzal, J., Williams, M., Leng, M.J., Aldridge, R.J. & Stephenson, M.H. 2011: Evolution of Paleocene to Early Eocene larger benthic foraminifer assemblages of the Indus Basin, Pakistan. Lethaia, Vol. 44, pp. 299–320. The Paleocene–Early Eocene carbonate successions of the Indus Basin in Pakistan formed on the northwestern continental shelf margin of the Indian Plate in the eastern Tethys Ocean. Based on larger benthic foraminifera (LBF), eight Tethyan foraminiferal biozones (SBZ1–SBZ8) spanning the Paleocene to Early Eocene interval are identified. The base of the Eocene is identified by the first appearance of Alveolina sp. Other stratigraphically important LBFs that are characteristic of the earliest Eocene are Ranikothalia nuttalli, Discocyclina dispansa and Assilina dandotica. Stable isotope analysis through the Paleocene–Eocene (P–E) boundary interval identifies more positive δ¹³C values for the Late Paleocene (+3.4‰ to +3.0‰) and lower values (+2.7‰ to +1.6‰) for the earliest Eocene. However, there is insufficient sampling resolution to identify the maximum negative δ¹³C excursion of the Paleocene–Eocene Thermal Maximum. During Late Paleocene times LBF assemblages in the Indus Basin were taxonomically close to those of west Tethys, facilitating biostratigraphic correlation. However, this faunal continuity is lost at the P–E boundary and the earliest Eocene succession lacks typical west Tethys Nummulites, while Alveolina are rare; LBFs such as Miscellanea and Ranikothalia continue to dominate in the Indus Basin. The absence of Nummulites from the earliest Eocene of Pakistan and rarity of Alveolina, elsewhere used as the prime marker for the base of the Eocene, may imply biogeographical barriers between east and west Tethys, perhaps caused by the initial stages of India‐Asia collision. Later, at the level of the Eocene SBZ8 Biozone, faunal links were re‐established and many foraminifera with west Tethys affinities appeared in east Tethys, suggesting the barriers to migration ceased. □Biostratigraphy, Eocene, India‐Asia collision, larger benthic foraminifera, palaeoecology, Paleocene.     

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.     

Morphological abnormalities in Late Cretaceous and early Paleocene foraminifer tests (northern Patagonia, Argentina)

Morphological abnormalities are common in Late Cretaceous and early Paleocene foraminifer tests at two localities in northern Patagonia, Argentina. Protelphidium sp. in the Auca Mahuevo section (late Campanian–early Maastrichtian) exhibit abnormal size or shape of the later chambers, with the last chamber commonly larger than normal or inflated and variably extending onto one of the lateral sides of the test; modification of the coiling plane; protuberances near the proloculus or on one or more chambers; a double last chamber, and complex forms. Protelphidium hofkeri Haynes in the Cerro Azul section (Danian) exhibit abnormal size or shape of one or more chambers, producing peripheral irregularities. In addition, there are rare multiple tests in planktic species from the Cerro Azul section, probably teratological specimens. The sedimentology of the sections and the character of the accompanying faunas indicate that the abnormalities in the two benthic foraminiferal taxa were most probably caused by hypersalinity and/or fluctuations in salinity. The fossil occurrence of assemblages with abundant deformed specimens suggests that investigators should carefully look at many aspects of the environment before concluding that anthropogenic pollution is the only cause of deformations of living benthic foraminifera.     

Planktonic foraminiferal succession in late Cretaceous to early Palaeocene strata in Meghalaya, India

A roughly 10.5-m-thick succession within the Langpar Formation of the Um Sohryngkew River section, Meghalaya, India, constrained by the last occurrence of Globotruncanita stuarti and the first occurrence of Parasubbotina pseudobulloides , spans the K/T (Cretaceous–Tertiary) transition. The unit is divisible into three parts with the lower consisting of shaly limestone, weakly calcareous shale and silty shale with coal streaks. The middle part is dominated by calcareous shale with mud flakes, coprolites, burrows and pyrite nodules, followed by alternating limestone and marlite at the top. Planktonic foraminifera are rare to frequent within the unit. Based on the distribution of zonal indices, seven successive planktonic foraminiferal zones are recognized from across the K/T boundary. From base to top, these are CF4, CF3, CF2 and CF1 in the upper Maastrichtian part and Zone P0, Zone Pα and Subzone P1a in the lower Danian part. The biozones indicate that the section is biostratigraphically continuous across the K/T boundary. A similar foraminiferal succession and K/T transition is observed in the Langpar of the Cherrapunji-Mahadeo road section at a distance of over 5km. These K/T outcrops from Meghalaya provide the first record of a continuous K/T sequence in the Indian subcontinent with respect to planktonic foraminifera.     

Perturbation at the sea floor during the Paleocene–Eocene Thermal Maximum: Evidence from benthic foraminifera at Contessa Road, Italy

Detailed analyses of the benthic foraminiferal assemblages extracted with the cold acetolyse method together with high resolution geochemical and mineralogical investigations across the Paleocene/Eocene (P/E) boundary of the classical succession at Contessa Road (western Tethys), allowed to recognize and document the Paleocene–Eocene Thermal Maximum (PETM) interval, the position of the Benthic Extinction Event (BEE) and the early recovery of benthic faunas in the aftermath of benthic foraminiferal extinction. The stratigraphical interval spanning the P/E boundary consists of dominantly pelagic limestones and two prominent marly beds. Benthic foraminifera indicate that these sediments were deposited at lower bathyal depth, not deeper than 1000–1500 m. The Carbon Isotope Excursion (CIE) interval is characterized by high barite abundance with a peak at the base of the same stratigraphic interval, indicating a complete, although condensed record of the early CIE. A succession of events and changes in the taxonomic structure of benthic foraminifera has been recognized that may be of use for supra-regional stratigraphic correlation across the P/E boundary interval. The composition of the benthic foraminiferal assemblages, dominated by infaunal taxa, indicates mesotrophic and changing conditions on the sea floor during the last  45 kyr of the Paleocene. The BEE occurs at the base of the CIE within the lower marly bed and it is recorded by the extinction of several deep-water cosmopolitan taxa. Then, the lysocline/CCD rose and severe carbonate dissolution occurred. Preservation deteriorated, the faunal density and simple diversity dropped to minimum values and a peak of Glomospira spp. has been observed. Stress-tolerant and opportunistic groups, represented mainly by bi-and triserial taxa, dominate the low-diversity post-extinction assemblages, indicating a benthic foraminiferal recovery under environmental unstable conditions, probably within a context of sustained food transfer to the bottom. A three-phase pattern of faunal recovery is recognizable. At first the lysocline/CCD started to descend and then recovered. Small-sized “Bulimina”, Oridorsalis umbonatus, and Tappanina selmensis rapidly repopulated the severely stressed environment. Later on, Siphogenerinoides brevispinosa massively returns, dominating the assemblage together with other buliminids, Nuttallides truempyi, and Anomalinoides sp.1. Finally, a marked drop in abundance of S. brevispinosa is followed by a bloom of the opportunistic and recolonizer agglutinated Pseudobolivina that, for the first time, is recorded within the main CIE. A second interval of dissolution, but less severe than the previous one, has been recognized within the upper marly bed (uppermost part of the main CIE interval) and it is interpreted as a renewed, less pronounced shoaling of the lysocline/CCD that interrupted the recovery of benthic faunas. This further rise likely represents a response to persistent instability of ocean geochemistry in this sector of the Tethys before the end of the CIE. In the CIE recovery and post CIE intervals, the composition of the benthic foraminiferal assemblages suggests mesotrophic and unstable conditions at the sea floor. According to the geochemical proxy for redox conditions, the deposition of the PETM sediments at Contessa Road occurred in well-oxygenated waters, leading out a widespread oxygen depletion as major cause of the BEE. Changing oceanic productivity, carbonate corrosivity and global warming appear to have played a much more important role in the major benthic foraminiferal extinction at the P/E boundary.     

Benthic foraminiferal bioevents in lower to upper Toarcian strata of Southern Beaujolais (SE France)

This paper documents changes in benthic foraminiferal assemblages compared with high resolution ammonite biozonation along the lower Toarcian to upper Toarcian marine succession of Southern Beaujolais in southeastern France. Eight ammonite and three benthic foraminiferal zones including five subzones are distinguished based on the occurrence of twelve foraminiferal events. Each benthic foraminiferal subzone is characterized by its taxonomic and morphogroup composition, which represents the paleoecological response of these taxa and morphotypes of benthic foraminifera in the Early Jurassic and early Middle Jurassic. Major changes in abundance and diversity occur at the end of the Toarcian Oceanic Anoxic Event (T-OAE) and near the Early-Middle Jurassic transition. The low-abundance foraminiferal assemblages recorded in the Serpentinus ammonite Zone are interpreted as reflecting adverse environmental conditions after the T-OAE. The later recovery and development of the foraminiferal assemblages is documented in the Bifrons up to the Aalensis zones and is attributed to improved bottom water oxygenation. Common occurrences of agglutinated foraminifera represented mostly by Trochammina pulchra Ziegler in the Dispensum Zone point to an influx of cooler water masses during the late Toarcian. The morphogroup analysis carried out on the foraminifera and their paleoecological interpretations shed light on the changes in the stratigraphic record at the end of the T-OAE up to the Toarcian/Aalenian boundary.     

Facies characteristic and paleoenvironmental reconstruction of the Fahliyan Formation,Lower Cretaceous,in the Kuh-e Siah area,Zagros Basin,southern Iran

The Lower Cretaceous Fahliyan Formation, part of the Khami Group, unconformably overlies the Hith Formation and is conformably overlain by the Gadvan Formation in the study area in southern Iran. The Fahliyan Formation is a reservoir rock in Zagros Basin. This formation was investigated by a detailed petrographic analysis in order to clarify the depositional facies and sedimentary environment in the Kuh-e Siah Anticline in Boushehr Province. Petrographic studies led to the recognition of 25 microfacies that were deposited in four facies belts: tidal flat, lagoon, and shoal in inner ramp and shallow open-marine in mid-ramp environment. An absence of turbidite deposits, reefal facies, and gradual facies changes indicate that the Fahliyan Formation was deposited on a carbonate ramp. Calcareous algae and benthic foraminifera are abundant in the shallow marine carbonates of the Fahliyan Formation. These skeletal grains have been studied in order to increase the understanding of their distributions in time and space. A total of ten genera belonging to different groups of calcareous algae and 16 genera of benthic foraminifera are recognized from the Fahliyan Formation at Kuh-e Siah section.     

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.     

Environmental perturbation in the southern Tethys across the Paleocene/Eocene boundary (Dababiya,Egypt): Foraminiferal and clay mineral records

Foraminiferal and clay mineral records were studied in the upper Paleocene to lower Eocene Dababiya section (Egypt). This section hosts the GSSP for the Paleocene/Eocene boundary and as such provides an expanded and relatively continuous record across the Paleocene/Eocene Thermal Maximum (PETM). Deposition of illite–smectite clay minerals is interpreted as a result of warm and arid conditions in the southern Tethys during the latest Paleocene. Benthic foraminiferal assemblages are indicative of seasonal variation of oxygen and food levels at the seafloor. A sea-level fall occurred in the latest Paleocene, followed by a rise in the earliest Eocene. Foraminiferal diversity and densities decreased strongly at the P/E boundary, coinciding with the level of global extinction of benthic foraminifera (BEE) and start of the Carbon Isotope Excursion (CIE) and PETM. In the lower CIE, the seafloor of the stratified basin remained (nearly) permanently anoxic and azoic. A sudden increase in mixed clay minerals (kaolinite and others) suggests that warm and perennial humid conditions prevailed on the continent. High levels of TOC and phosphathic concretions in the middle CIE are evidence for increased organic fluxes to the sea floor, related to upwelling and to augmented continental runoff. Low densities of opportunistic taxa appeared, indicating occasional ephemeral oxygenation and repopulation of the benthic environment. The planktic community diversified, although conditions remained poor for deep-dwelling taxa. An increase in illite–smectite dominated clay association is considered to mark the return of a seasonal signature on climatic conditions. During the late CIE environmental conditions changed to seasonally fluctuating mesotrophic conditions and diverse and rich benthic and planktic foraminiferal communities developed. Post-CIE planktic faunas consisted of both deep and shallow-dwelling taxa and buliminid-dominated benthic assemblages reflect fluctuating mesotrophic conditions.The frequent environmental perturbations during the CIE/PETM at Dababiya provided a rather specialized group of foraminiferal taxa (i.e., Anomalinoides aegyptiacus) the opportunity to repopulate, survive and subsequently dominate by a hypothesized capacity to switch to an alternative life strategy (population dynamics, habitat shift) or different metabolic pathway. The faunal record of Dababiya provides insight into the cause and development of the BEE: various severe global changes during the PETM (e.g., ocean circulation, CaCO₃-dissolution, productivity and temperature changes) disturbed a wide range of environments on a geologically brief timescale, explaining together the geographically and temporally variable character of the BEE. This allowed a number of specific but different foraminiferal assemblages composed of stress-tolerant and opportunistic taxa to be successful during and after the periods of environmental perturbations associated with the PETM.     

Benthic foraminiferal assemblages between two major extinction events: the Paleocene El Kef section,Tunisia

The development of benthic foraminiferal assemblages from the Paleocene outcrops of the El Haria Formation near El Kef, Tunisia is discussed qualitatively and quantitatively. The aim of the study is to reconstruct the paleoenvironmental evolution between the K/Pg boundary interval and the late Paleocene event, and to compare this evolution with results from other sites along the southern Tethyan margin. Eighty-four samples, covering virtually the entire Paleocene, provide a dataset that allows detailed qualitative and multivariate analysis. The benthic foraminiferal faunas indicate a complex pattern of environmental changes during the Paleocene, marked by the succession of different benthic associations. Following the K/Pg boundary event, community restoration was characterized by the gradual build-up of faunal diversity. Decreasing dominance and the entry of taxa common to normal marine, outer neritic to upper bathyal environments indicate the completion of the ecosystem restoration in Zone Plb. A highly diverse benthic foraminiferal assemblage persisted throughout the remainder of the early Paleocene into the earliest late Paleocene. At the P3a-P3b zonal transition relative sea-level lowering is evidenced by the sudden disappearance or decreasing abundance of deeper-water taxa (e.g. Anomalinoides affinis, A. susanaensis, Gavelinella beccariiformis). Neritic deposition continued into Zone P4, when trophic levels at the seafloor increased as indicated by the entry and increasing dominance of species such as Anomalinoides cf. aegyptiacus, Bulimina midwayensis, and B. strobila, which we consider to be sensitive to eutrophication. The combined effect of shallowing and the subsequent eutrophication led to the establishment of assemblages similar to late Paleocene benthic foraminiferal assemblages from Egyptian sections, some of which record the latest Paleocene extinction event. These assemblages were interpreted to be indicative of a middle neritic, highly eutrophic environment. Enhanced vertical fluxes of organic matter along the southern Tethyan margin may have resulted from intensified upwelling. This eventually led to oxygen deficiency at the seafloor. It appears that oxygen-deficient, high-productivity shelves were a common feature of the southern Tethyan margin during the latest Paleocene.     

Distribution patterns of benthic foraminifera across the Ypresian–Lutetian Gorrondatxe section,Northern Spain: Response to sedimentary disturbance

We present a study of benthic foraminiferal assemblages from an Ypresian–Lutetian distal submarine fan system in the lower bathyal Gorrondatxe section (Basque-Cantabrian Basin, northern Spain). The objective of our study is to analyze the benthic foraminiferal distribution patterns and their response to sedimentary disturbance and related factors.Assemblages contain a high percentage of allochthonous taxa, such as asterigerinids and other shallow water taxa, which were transported downslope by turbidity currents.Detailed quantitative analyses, supported by R-mode cluster and Detrended Correspondence Analyses (after removing allochthonous taxa from the foraminiferal counts) allowed us to identify 6 assemblages that are divided into two groups related to the turbidite content in the Gorrondatxe section. Assemblages 1, characteristic of the turbidite-poor intervals with low sedimentary disturbance, include assemblage 1a (with highly diverse common middle–lower bathyal calcareous taxa) assemblage 1b (with common agglutinated taxa, mainly trochamminids), and assemblage 1c (characterized by calcareous taxa that are also common in the turbidite-rich interval).Assemblages 2, characterized by a high dominance, prevail in the turbidite-rich interval, and include assemblage 2a (characterized by the dominance of infaunal bolivinids and epifaunal cibicids), assemblage 2b (typified by moderate to low diversity and dominated by deep-infaunal Globobulimina species), and assemblage 2c (typified by very abundant suspension-feeding astrorhizids). The high abundance of bolivinids and Globobulimina species may be related to an enhanced input of low-quality organic matter transported by turbidity currents to the seafloor, representing different stages of recolonisation after disturbance and different energy regimes. High current activity was probably responsible for the abundance of cibicids, while moderate to low diverse and high dominance assemblages characterize the recolonisation of the substrate after disturbance.We conclude that sedimentary disturbance and other related factors such as current activity, resuspension of sediments at the seafloor, and supply of organic matter (and its quality) played an important role in the distribution of benthic foraminifera in the Gorrondatxe section. The identification of allochthonous taxa emerges as an essential aspect of the study of environments with sedimentary disturbance.     

The development of planktonic biogeography in the Southern Ocean during the Cenozoic

Deep-sea drilling at high latitudes of the Southern Hemispheres has provided almost the only available data to evaluate the biogeographic development of the planktonic biota in the Southern Ocean during the Cenozoic (65 m.y. to Present Day). Paleontological investigations on Deep Sea Drilling Project (DSDP) materials have shown that the development of Cenozoic planktonic biogeography of the Southern Ocean is intimately linked with the evolution of the Southern Ocean water masses themselves. During the Cenozoic, this has included the development of the Circum-Antarctic Current system as obstructing land masses moved apart, the refrigeration and later extensive glaciation of the continent, and the development of the Antarctic Convergence (Polar Front) with related oceanic upwelling.Almost all evolution of calcareous planktonic microfossils has occurred outside of the Antarctic—Subantarctic region followed by limited migration into these water masses. Virtually no endemism occurs amongst calcareous microfossil groups at these latitudes. In contrast, conspicuous and widespread evolution has occurred within the siliceous microfossil groups especially during the Neogene. Low diversity and differences in stratigraphic ranges of Antarctic calcareous microfossils makes them only broadly useful for correlation. Relatively higher diversities within the Subantarctic provide a firmer basis for more detailed correlation, although the ranges of fossils are often different than at lower latitudes because of different paleoceanographic and paleoclimatic controls. Within the Antarctic water mass south of the Antarctic Convergence, siliceous microfossilsbiostratigraphy, oxygen isotopic stratigraphy and magnetostratigraphy, provide the only firm basis for correlation with low-latitude sequences.Eocene (55-38 Ma) sediments contain abundant calcareous microfossils even closely adjacent to the continent. Antarctic calcareous planktonic microfossils of this age exhibit relative high diversity, although this is lower than assemblages of equivalent age at middle and low latitudes. Within the Subantarctic region, Eocene planktonic foraminifera exhibit strong affinities with those in the temperate regions. Biogeographic differences exist between various sectors of the Southern Ocean related to biogeographic isolation preceding the development of the Circum-Antarctic Current. Subantarctic calcareous nannofossil assemblages of Paleocene and Eocene age exhibit higher diversity than Oligocene and Neogene assemblages. Siliceous microfossils are poorly represented or at best poorly known.One of the most dramatic changes in Southern Ocean planktonic biogeography occurred near the Eocene/Oligocene boundary (38 Ma). Since then, Antarctic planktonic foraminiferal assemblages have exhibited distinct polar characteristics, marked in particular by low diversity, and this event thus reflects the initiation of the Antarctic faunal and floral provinces. Profound paleoceanographic changes at this time, which triggered the biogeographic crisis, appear to be related to the initiation of widespread Antarctic sea-ice formation, and rapid cooling of deep and intermediate waters, in turn associated with increased Antarctic glaciation. During the Oligocene, planktonic microfossil diversity was low in all groups throughout the world's oceans. In Antarctic waters, the early Oligocene foraminiferal fauna is monospecific (Subbotina angiporoides), while in the later Oligocene two species (S. angiporoides and Catapsydrax dissimilis) were recorded. Calcareous nannofossil assemblages are of low diversity compared with the Eocene. Subantarctic foraminiferal faunas of Oligocene age display much higher diversity than those in the Antarctic, but early and middle Oligoceae faunas still exhibit the lowest diversities for the entire Cenozoic. Siliceous assemblages remain relatively inconspicuous in most regions of the Southern Ocean.The Paleogene-Neogene transition (22 Ma) is marked by a major change in the global planktonic biogeography, i.e. modern patterns developed in which permanent, steep faunal and floral diversity gradients existed between tropical and polar regions; a gradient which has persisted even during the most severe glacial episodes. Oligocene assemblages of low diversity and almost cosmopolitan distribution were replaced by distinctive belts of planktonic assemblages arranged latitudinally from the tropics to the poles. The establishment of the steep planktonic diversity gradients and latitudinal provinces near the beginning of the Neogene almost certainly were linked to the development of the Circum-Antarctic Current in the late Oligocene which effectively separated high- and low-latitude planktonic assemblages. These fundamental global circulation and biogeographic patterns have persisted through the Neogene.During the Neogene (22 Ma to Present Day), Antarctic calcareous microfossil assemblages exhibit persistent low diversity and high dominance, while Subantarctic assemblages are of much greater diversity. The beginning of the Neogene (= beginning of Miocene) heralded the development of the high-latitude siliceous microfossil assemblages towards their present-day dominant role. Siliceous biogenec productivity began to increase. These changes were linked to the initial development and later intensification of circulation associated with the Antarctic Convergence and Antarctic Divergence. The Antarctic Convergence sharply separates dominantly siliceous assemblages to the south from calcareous assemblages to the north. Radiolarian assemblages became more endemic. Relatively warm early and middle Miocene conditions are reflected by slightly higher diversity of planktonic foraminifera and by the presence, in the northern Subantarctic, of conspicuous discoasters in early Miocene sediments. In Antarctic waters, calcareous nannofossils become unimportant as biogenic elements after the middle Miocene.The latest Miocene ( 5 m.y. ago) was marked by northward movement of the Antarctic Convergence, corresponding expansion of the Antarctic water mass, and low diversity of calcareous assemblages. Pliocene planktonic foraminifera seem to be largely monospecific in Antarctic and southern Subantarctic sequences. During the Quaternary, Antarctic waters reached a maximum northward expansion and exhibit highest siliceous biogenic productivity for the Cenozoic. In the Subantarctic, Quaternary foraminiferal diversities are much higher than in Pliocene sequences. Although calcareous nannofossil diversity may be high, only a few species are abundant. Large northward shifts of Antarctic and Subantarctic water masses have occurred during the Quaternary although no southward penetrations have occurred much beyond that of the present day. Several radiolarian and foraminiferal species disappeared or appeared at or close to a number of paleomagnetic reversals during the last 4 m.y. These faunal events, which provide valuable datums, do not seem to be associated with major climatic changes.     

Paleoenvironmental change at the Danian–Selandian transition in Tunisia: Foraminifera, organic-walled dinoflagellate cyst and calcareous nannofossil records

In the present study, we document paleoenvironmental change across the Danian–Selandian transition (planktic foraminiferal interval P2–P3b; calcareous nannofossil Zone NP4, Subzones NTp6–NTp8A; 61–59 Ma) in NW Tunisia. Diversifications of Paleogene planktic foraminifera with the evolution of the muricate and photosymbiotic lineages Morozovella, Acarinina and Igorina and of the biostratigraphically important nannofossils genus Fasciculithus are recorded within this interval. The present study aims to understand early Paleogene environmental changes in the southern Tethys, by analyzing the evolution of surface-water and–to a lesser extent–seafloor conditions. Three localities were investigated: Ain Settara, Elles and El Kef, all representing outer neritic deposition in the same basin, the Tunisian Trough. Paleoenvironmental changes are explored by combining planktic foraminiferal, organic dinocyst and calcareous nannofossils assemblages and several proxy parameters (planktic/benthic ratio, numbers of planktic foraminifera per gram, peridinioid/gonyaulacoid ratio; terrestrial/marine palynomorph ratio). In addition, also some geochemical parameters (calcite content and stable isotopes) are examined. Our records indicate that the environment evolved from an initially oligotrophic, open marine, deep outer neritic setting in P2–P3a towards a shallower and nutrient-rich setting from the base of Subzone P3b. This change is seen in the foraminiferal assemblages, with the substitution of Praemurica by Morozovella among the planktic foraminifera and an upward decrease in deeper benthic taxa. Also the organic-dinocyst assemblages show a peak of peridinioid cysts (Cerodinium and Lejeunecysta). Associated to these dinocyst assemblages, the lowest occurrence of Apectodinium is recorded, which seem to have evolved in this region, possibly in response to enhanced nutrient levels on the shelf. Additionally, a distinct change in calcareous nannofossil assemblages is also described, marked by the lowest appearance of Chiasmolithus edentulus, the lowest consistent occurrence of Fasciculithus and a slight increase in near-shore taxa (essentially Pontosphaera).This project provides an accurate understanding of paleoenvironmental change across the Danian–Selandian transition in Tunisia. Especially, integrating different proxies demonstrates a paleobathymetric shallowing from the Danian to the Selandian, associated to increase surface paleoproductivity. Furthermore, the results are compared with those from other localities along the Southern Tethyan margin (Egypt and Jordan) and a more regional paleoclimatic/paleoceanographic perturbation in the Southern Tethys is suggested.     

Palaeoenvironmental turnover across the Ypresian–Lutetian transition at the Agost section, Southastern Spain: In search of a marker event to define the Stratotype for the base of the Lutetian Stage

Marker events to define the stratotype for the base of the Lutetian Stage are poorly defined. To elucidate such markers and characterize palaeoenvironmental turnovers, we conducted an integrated study of the Ypresian–Lutetian (Y–L; early-middle Eocene) transition at the continuous Agost section (southeastern Spain). This 115-m-thick section, which consists of hemipelagic marls intercalated with hemipelagic limestones and turbidity sandstones, spans from planktic foraminiferal Zones P9 to P12 (E7 to E10) and calcareous nannofossil Zones CP11 to CP14a (NP13 to NP16). We report quantitative analyses of planktic and benthic foraminifera and characterization of trace fossil assemblages that are integrated with mineralogical analyses.Relative to benthic forms, planktic foraminifera constitute more than 80% of the foraminiferal assemblage. We found that the most abundant planktic species belong to the genera Acarinina, Morozovella, Subbotina, and Pseudohastigerina. Benthic foraminiferal assemblages are strongly dominated by calcareous taxa, with bolivinids being the most abundant group. Trace fossils showed the succession Nereites–Zoophycos–Cruziana ichnofacies throughout the Agost section. In addition to changes in palaeobathymetry, we deduced that quantity and quality of organic matter flux influenced by turbidity currents are the main factors controlling benthic assemblages. We distinguished several mineralogical boundaries at the Agost section, each associated with lithological facies changes suggesting a change in provenance rather than changes in weathering conditions. We made three observations that indicate an increase in sea water temperatures or a possible hyperthermal event related to the first occurrence (FO) of hantkeninids (i.e., the P9/P10 boundary): 1) a distinct peak in abundance of the benthic foraminifera Aragonia aragonensis; 2) the low-diversity of benthic foraminiferal assemblages; and 3) the occurrence of the planktic foraminifera Clavigerinella eocenica and Clavigerinella jarvisi. Benthic foraminiferal and trace fossil assemblages also suggest an associated relative fall of sea level from upper-middle bathyal to sublittoral depths. These characteristic indicators point to this boundary as a promising feature for defining the Global Stratotype Section and Point (GSSP) for the base of the Lutetian Stage. However, complementary magnetobiostratigraphic studies carried out at the Agost section point to the FO of calcareous nannofossil Blackites inflatus (base of CP12b), which occurred 3–5 Myr before the P9/P10 boundary, as the most suitable primary marker event. Whatever the marker event chosen, all the successive events recognized at the Agost section allow a complete characterization of the Y–L transition, and thus this section may be a suitable candidate to locate the GSSP for the Ypresian/Lutetian boundary.