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.     

The Paleocene–Eocene Thermal Maximum as recorded by Tethyan planktonic foraminifera in the Forada section (northern Italy)

The Forada section in the Venetian Pre-Alps of northern Italy represents an expanded record of the Paleocene–Eocene Thermal Maximum (PETM) at a depositional paleodepth of about 1 km ± 0.5 km. High-resolution planktonic foraminiferal analysis of this section, in a time interval of approximately 1.3 Myr across the Paleocene/Eocene boundary, reveals striking faunal changes that allow the identification of eight phases (a–h). The late Paleocene was represented by stable, warm and oligotrophic surface water conditions (phase a). Unstable environmental conditions start well before the onset of PETM (ca. 150 kyr, phase b) and involved a change towards eutrophy, as marked by the increase of Subbotina and the concomitant decrease of Morozovella. This step is also characterized by enhanced fragmentation and dissolution.The interval corresponding to the main body of the carbon isotope excursion (CIE) is characterized by a marked increase of Acarinina, though with some differences in the species composition and relative abundance, both in high-and low-latitudes, particularly in the Tethyan area. Forada is no exception to this pattern. However, at Forada, two prominent peaks in abundance of acarininids are recorded ca. 30 kyr prior to the onset of the CIE, thus suggesting an increase in temperature heralding the onset of the PETM (phase c). Interestingly, the lower peak in abundance of Acarinina just precedes the 1‰ carbon isotope negative shift occurring below the onset of the main CIE. The basalmost Eocene, corresponding to the lower part of CIE curve, is represented by intense planktonic foraminiferal dissolution, implying an extraordinary rise of the CCD. This interval has an estimated duration of about 16 kyr (phase d).The dominance of acarininids in the lower part of the CIE (phase e, f; ca. 14 and 22.5 kyr) is interpreted as a consequence of the extreme warmth coupled with eutrophic conditions characterizing the Forada depositional environment at that time. These acarininids include at Forada also the temporally constrained Acarinina sibaiyaensis and A. africana. The morphological similarity between these peculiar species with the radially elongated chambered forms characterizing the Cretaceous anoxic events, suggests the hypothesis that depletion of oxygen in the upper water column might have been one of the factors causing their conspicuous occurrence at the PETM.The recovery in abundance of the specialized morozovellids and of other planktonic foraminiferal groups (e.g., biserials, globanomalinids, igorinids, planorotalids and pseudohastigerinids), occurring in the middle part of the CIE (ca. 30 kyr after the onset of the PETM), indicates an initial environmental recovery (phase g). A new stable state is definitely reached in the upper part of the Forada section where the relative proportions of the main component of planktonic foraminiferal assemblages move towards values similar to those of the late Paleocene conditions (phase h). However, the perturbation during the PETM produced significant changes in the ocean geochemistry that endured after the PETM event, as testified by the prominent high carbonate dissolution characterizing the marly levels, and the large variability in relative abundance among different components of the planktonic foraminiferal assemblages. These striking oscillations were not present in the latest Paleocene.     

Middle Eocene–Early Pliocene Subantarctic planktic foraminiferal biostratigraphy of Site 1090, Agulhas Ridge

The analysis of planktic foraminiferal assemblages from Site 1090 (ODP Leg 177), located in the central part of the Subantarctic Zone south of South Africa, provided a geochronology of a 330-m-thick sequence spanning the Middle Eocene to Early Pliocene. A sequence of discrete bioevents enables the calibration of the Antarctic Paleogene (AP) Zonation with lower latitude biozonal schemes for the Middle–Late Eocene interval. In spite of the poor recovery of planktic foraminiferal assemblages, a correlation with the lower latitude standard planktic foraminiferal zonations has been attempted for the whole surveyed interval. Identified bioevents have been tentatively calibrated to the geomagnetic polarity time scale following the biochronology of Berggren et al. (1995). Besides planktic foraminiferal bioevents, the disappearance of the benthic foraminifera Nuttallides truempyi has been used to approximate the Middle/Late Eocene boundary. A hiatus of at least 11.7 Myr occurs between 78 and 71 m composite depth extending from the Early Miocene to the latest Miocene–Early Pliocene. Middle Eocene assemblages exhibit a temperate affinity, while the loss of several planktic foraminiferal species by late Middle to early Late Eocene time reflects cooling. During the Late Eocene–Oligocene intense dissolution caused impoverishment of planktic foraminiferal assemblages possibly following the emplacement of cold, corrosive bottom waters. Two warming peaks are, however, observed: the late Middle Eocene is marked by the invasion of the warmer water Acarinina spinuloinflata and Hantkenina alabamensis at 40.5 Ma, while the middle Late Eocene experienced the immigration of some globigerinathekids including Globigerinatheka luterbacheri and Globigerinatheka cf. semiinvoluta at 34.3 Ma. A more continuous record is observed for the Early Miocene and the Late Miocene–Early Pliocene where planktic foraminiferal assemblages show a distinct affinity with southern mid- to high-latitude faunas.     

Faunal turnovers in central Pacific benthic foraminifera during the Paleocene-Eocene thermal maximum

Although it is well known that the Paleocene/Eocene thermal maximum (PETM) coincided with a major benthic foraminiferal extinction event, the detailed pattern of the faunal turnover has not yet been clarified. Our high-resolution benthic foraminiferal and carbon isotope analyses at the low latitude Pacific Ocean Shatsky Rise have revealed the following record of major faunal transitions: (1) An initial turnover which involved the benthic foraminiferal extinction event (BFE). The BFE, marked by a sharp transition from Pre-extinction fauna to Disaster fauna represented by small-sized Bolivina gracilis, expresses the onset of the PETM and the abrupt extinction of about 30% of taxa. This faunal transition lasted about 45-74 kyr after the initiation of the PETM and was followed by: (2) the appearance of Opportunistic fauna represented by Quadrimorphina profunda, which existed for about 74-91 kyr after the initiation of the PETM. These two faunas, which appeared after the extinction event, are characterized by low diversity and dwarfism, possibly due to lowered oxygen condition and decreased surface productivity. The second pronounced turnover involved the gradual recovery from Opportunistic Fauna to the establishment of Recovery fauna, which coincided with the recovery about 83-91 kyr after its initiation.     

The onset of the Paleocene–Eocene Thermal Maximum (PETM) at Sites 1209 and 1210 (Shatsky Rise, Pacific Ocean) as recorded by planktonic foraminifera

High-resolution biostratigraphic and quantitative studies of subtropical Pacific planktonic foraminiferal assemblages (Ocean Drilling Program, Leg 198 Shatsky Rise, Sites 1209 and 1210) are performed to analyse the faunal changes associated with the Paleocene–Eocene Thermal Maximum (PETM) at about 55.5 Ma. At Shatsky Rise, the onset of the PETM is marked by the abrupt onset of a negative carbon isotope excursion close to the contact between carbonate-rich ooze and overlying clay-rich ooze and corresponds to a level of poor foraminiferal preservation as a result of carbonate dissolution. Lithology, planktonic foraminiferal distribution and abundances, calcareous plankton and benthic events, and the negative carbon isotope excursion allow precise correlation of the two Shatsky Rise records. Results from quantitative analyses show that Morozovella dominates the assemblages and that its maximum relative abundance is coincident with the lowest δ¹³C values, whereas subbotinids are absent in the interval of maximum abundance of Morozovella. The excursion taxa (Acarinina africana, Acarinina sibaiyaensis, and Morozovella allisonensis) first appear at the base of the event. Comparison between the absolute abundances of whole specimens and fragments of genera demonstrate that the increase in absolute abundance of Morozovella and the decrease of Subbotina are not an artifact of selective dissolution. Moreover, the shell fragmentation data reveal Subbotina to be the more dissolution-susceptible taxon. The upward decrease in abundance of Morozovella species and the concomitant increase in test size of Morozovella velascoensis are not controlled by dissolution. These changes could be attributed to the species' response to low nutrient supply in the surface waters and to concomitant changes in the physical and chemical properties of the seawater, including increased surface stratification and salinity.Comparison of the planktonic foraminiferal changes at Shatsky Rise to those from other PETM records (Sites 865 and 690) highlights significant similarities, such as the decline of Subbotina at the onset of the event, and discrepancies, including the difference in abundance of the excursion taxa. The observed planktonic foraminifera species response suggests a warm–oligotrophic scenario with a high degree of complexity in the ocean structure.     

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.     

Benthic foraminiferal turnover across the Cretaceous/Paleogene boundary at Agost (southeastern Spain): paleoenvironmental inferences

We studied Upper Cretaceous and Lower Paleogene benthic foraminifera from the Agost section (southeastern Spain) to infer paleobathymetrical changes and paleoenvironmental turnover across the Cretaceous/Paleogene (K/P) transition. Benthic foraminifera indicate uppermost bathyal depths at Agost during the Abathomphalus mayaroensis Biochron (from about 400 kyr before the K/P boundary) through the early Plummerita hantkeninoides Biochron (about 120–150 kyr before that boundary). The depth increased to middle bathyal for the remainder of the Cretaceous, and remained so for the Danian part of the studied section (Parasubbotina pseudobulloides Biochron, at least 200 kyr after the K/P boundary). There were no perceivable bathymetrical changes at the K/P boundary, where 5% of the species became extinct, and the species composition of the benthic foraminiferal fauna changed considerably. Below the boundary, infaunal morphogroups constitute up to 65–73% of the faunas. Directly above the boundary, in the black clays of the lower Guembelitria cretacea Biozone, benthic foraminifera are rare. Several opportunistic taxa (e.g. the agglutinant Haplophragmoides sp.) have short peaks in relative abundance, possibly reflecting low-oxygen conditions as well as environmental instability, with benthos receiving food from short-lived, local blooms of primary producers. Above the clays through the end of the studied interval, epifaunal morphogroups dominate (up to 70% of the assemblages) or there is an even mixture or epifaunal and infaunal morphogroups. Infaunal groups do not recover to pre-extinction relative abundances, indicating that the food supply to the benthos did not recover fully over the studied interval (about 200 kyr after the K/P boundary). The benthic foraminiferal faunal changes are compatible with the direct and indirect effects of an asteroid impact, which severely destabilized primary producers and the oceanic food web that was dependent upon them.     

Benthic foraminiferal fauna during the time of the Indian-Asian contact, in southern Balochistan, Pakistan

Cretaceous and early Paleocene benthic foraminifera were studied from one section along the western Gaj River, southern Balochistan, Pakistan, to reconstruct the paleoenvironment of the Tethys Sea during the Indian-Asian contact. We recognize three lithostratigraphic units in ascending order: the Mughal Kot Formation, the Pab Sandstone, and the Jamburo Group. Both the Maastrichtian Mughal Kot Formation, which consists of shale with grey marly limestone, and the Maastrichtian Pab Sandstone, which consists of quartzose sandstone, indicate an open ocean environment as they have diversified planktic and benthic foraminiferal assemblages. The Maastrichtian-Paleocene Jamburo Group, consisting of dark grey, calcareous shale and marlstone with some sulfide grains, is characterized by low diversities of benthic assemblages. The change to the lower diversities may be associated with the development of poor circulation of deeper water that was caused by narrowing of the Tethys Sea.The Trochammina spp. Assemblage from the Jamburo Group, which can be correlated with flysch-type agglutinated foraminiferal assemblages, has a low benthic species diversity, indicating an unfavorable condition for calcareous foraminifera because of the development of oxygen-depleted water. The absolute abundance of agglutinated specimens shows a remarkable change from low numbers in the Maastrichtian to high ones in the Paleocene. The benthic foraminiferal evidence supports the hypothesis that the collision of the Asian and Indian plates occurred near the end of the Cretaceous.     

Response of calcareous nannofossils to the Paleocene–Eocene Thermal Maximum: Observations on composition,preservation and calcification in sediments from ODP Site 1263 (Walvis Ridge — SW Atlantic)

In this study we present the results of a detailed analysis on calcareous nannofossil assemblages from sediment cores of ODP Site 1263 (Southern East Atlantic, Walvis Ridge). This section represents one of the few complete deep-sea sections that document the Paleocene–Eocene Thermal Maximum (PETM) in the pelagic realm. The PETM transient event was characterized by a brief, but intense interval of global warming, a global negative carbon isotope excursion (CIE), and widespread dissolution of seafloor carbonate sediments. Paired analysis at polarizing light microscope (LM) and Scanning Electron Microscope (SEM) documents the different “behavior” of nannofossils through the different phases of the PETM, at the onset of CIE, within the CIE, and during the recovery interval. The presence of anomalous specimens and morphotypes within some nannofossil taxa, recorded during previous LM high resolution analyses, has been further investigated in selected samples at the SEM. Besides the known representatives of the CIE-PETM “excursion nanno-flora”, as Rhomboaster calcitrapa group and Discoaster anartios, the analysis revealed the presence of peculiar morphotypes of Fasciculithus and deformed specimens of Discoaster nobilis group, Discoaster mediosus and Discoaster multiradiatus that are considered related to the anomalous amount of CO₂ in the ocean-atmosphere system during the early phase of PETM. Comparative analyses were performed in few selected samples from other PETM sections located at different latitudes in the Atlantic and Pacific oceans. Although the anomalous geochemical conditions during the PETM-CIE interval seem to have had some influence on the nannofossil production, calcification and assemblage composition, it results that local productivity together with post depositional (diagenetic) conditions were additional important controlling factors on nannofossil assemblages. Preliminary data from Eocene Thermal Maximum 2 (ETM2 or Elmo) suggest that nannofossil malformations are not exclusive of the PETM, and are associated to other episodes of perturbation of the C cycle.     

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.     

Catastrophic mass extinction and assemblage evolution in planktic foraminifera across the Cretaceous/Paleogene (K/Pg) boundary at Bidart (SW France)

A high-resolution biostratigraphic analysis of planktic foraminifers confirms that the Bidart section at the eastern margin of the Atlantic Ocean exhibits a continuous and complete Cretaceous/Paleogene (K/Pg) transition interval. The biozones and subzones recorded in this section are less expanded than their equivalent in Tunisian sections: El Kef (Global Stratotype Section and Point: GSSP for the K/Pg boundary) and Ellès (auxiliary section), but they are sufficiently thick to allow a detailed analysis of the evolution of the planktic foraminiferal assemblages across the K/Pg transition.Throughout the uppermost 4 m Maastrichtian, the planktic foraminiferal assemblages are highly diversified, containing up to 72 species. These Maastrichtian assemblages are rich in cosmopolitan taxa (70%), dominated by small biserial morphotypes which belong mainly to the genus Heterohelix which coexist with less abundant but highly diverse tropical and subtropical species.The extinction pattern at the Bidart section suggests a sudden catastrophic mass extinction at the K/Pg boundary which affected at least 53 out of 72 species. The species becoming extinct include globotruncanids (e.g. Contusotruncana spp., Globotruncana spp., Globotruncanita spp.) and complex heterohelicids (e.g. Racemiguembelina spp., Pseudotextularia spp., Gublerina spp.). At the Bidart section, only Archaeoglobigerina cretacea disappears 2 m below the K/Pg boundary event. Specimens of 18 small and even tiny Maastrichtian species, are found at the lowermost Danian. Only a few of these species belonging to the genera of Guembelitria, Hedbergella and Heterohelix are considered to be real “Cretaceous survivor species”, whereas the specimens belonging to the rest, are most probably reworked, because they differ in their preservation.Throughout lowermost Danian, the planktic foraminiferal assemblages are dominated by “opportunistic” species of the genus Guembelitria. These opportunists are associated to small and poorly diversified pioneer globigerinids (Palaeoglobigerina spp. and Parvularugoglobigerina spp.). These assemblages became progressively more diversified across the early Danian containing species with cancellate walls (Eoglobigerina spp., Parasubbotina spp., Subbotina triloculinoides and Praemurica spp.) and new taxa of biserial heterohelicids (Woodringina spp. and Chiloguembelina spp.) suggesting a paleoenvironmental recovery.     

Recent benthic foraminiferal assemblages from cold-water coral mounds in the Porcupine Seabight

Cold-water coral ecosystems are characterised by a high diversity and population density. Living and dead foraminiferal assemblages from 20 surface sediment samples from Galway and Propeller Mounds were analysed to describe the distribution patterns of benthic foraminifera on coral mounds in relation to different sedimentary facies. Hard substrates were examined to assess the foraminiferal microhabitats and diversities in the coral framework. We recognised 131 different species, of which 27 prefer an attached lifestyle. Epibenthic species are the main constituents of the living and dead foraminiferal assemblages. The frequent species Discanomalina coronata was associated with coral rubble, Cibicides refulgens showed preference to the off-mound sand veneer, and Uvigerina mediterranea displayed abundance maxima in the main depositional area on the southern flank of Galway Mound, and in the muds around Propeller Mound. The distribution of these species is rather governed by their specific ecological demands and microhabitat availability than by the sedimentary facies. Benthic foraminiferal assemblages from coral mounds fit well into basin-wide-scale distribution patterns of species along the western European continental margin. The diversity of the foraminiferal faunas is not higher on the carbonate mounds as in their vicinity. The living assemblages show a broad mid-slope diversity maximum between 500 and 1,300 m water depth, which is the depth interval of coral mound formation at the Celtic and Amorican Margin. The foraminiferal diversity maximum is about 700 m shallower than comparable maxima of nematodes and bivalves. This suggests that different processes are driving the foraminiferal and metazoan diversity patterns.     

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.     

Size variations of the calcareous nannofossil taxon Discoaster multiradiatus (Incertae sedis) across the Paleocene–Eocene thermal maximum in ocean drilling program holes 690B and 1209B

Size measurements of the calcareous nannofossil taxon Discoaster multiradiatus were carried out across the Paleocene–Eocene Thermal Maximum (PETM) in Ocean Drilling Program Holes 690B (Maud Rise, Weddell Sea) and 1209B (Shatsky Rise, Pacific Ocean). Morphometric investigations show that D. multiradiatus specimens are generally larger at ODP Site 1209 than at ODP Site 690. A limited increase in size of D. multiradiatus is recorded at ODP Site 1209, whereas significant enlargements characterize ODP Site 690. Preservation is comparable at both sites: nannofossils are moderately preserved with some evidence of etching/overgrowth in the PETM interval. Yet, D. multiradiatus variations do not correlate with preservation state and morphometric data most likely represent primary signals rather than diagenetic artifacts. There is a direct relationship between D. multiradiatus size and paleotemperatures: largest specimens are coeval with global warming associated with the PETM, inferred to result from excess atmospheric CO₂ due to (partial) oxidation of massive quantities of methane. Size increases and largest specimens of D. multiradiatus occur at different stratigraphic levels within PETM at ODP Sites 690 and 1209. A marked shift in diameter size was observed at the onset and peak of the Carbon Isotopic Excursion (CIE) at ODP Site 690, but only at the end of CIE and initial recovery interval at ODP Site 1209. This diachroneity is puzzling, but indeed correlates well with reconstructed changes in surface and thermocline water masses temperature and salinity in the PETM interval at low and high latitudes. The presumed high concentrations of carbon dioxide seem to have not influenced the morphometry of D. multiradiatus. The major size increase of D. multiradiatus in the CIE of ODP Site 690 could represent the migration of larger-sized allochtonus specimens that moved from peri-equatorial/subtropical areas to higher latitudes during the warmest interval of the PETM, although no direct evidence of distinct populations/subpopulations has been obtained from the frequency diagrams. As a result, we infer that D. multiradiatus is a proxy of water masses stratification and might be used for deriving temperature–salinity–nutrient conditions in the mixed layer and thermocline and their dynamics.     

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.     

Food supply to the seafloor in the Pacific Ocean after the Cretaceous/Paleogene boundary event

Deep-sea benthic foraminifera show important but transient assemblage changes at the Cretaceous/Paleogene (K/Pg) boundary, when many biota suffered severe extinction. We quantitatively analyzed benthic foraminiferal assemblages from lower bathyal–upper abyssal (1500–2000 m) northwest Pacific ODP Site 1210 (Shatsky Rise) and compared the results with published data on assemblages at lower bathyal (~ 1500 m) Pacific DSDP Site 465 (Hess Rise) to gain insight in paleoecological and paleoenvironmental changes at that time.At both sites, diversity and heterogeneity rapidly decreased across the K/Pg boundary, then recovered. Species assemblages at both sites show a similar pattern of turnover from the uppermost Maastrichtian into the lowermost Danian: 1) The relative abundance of buliminids (indicative of a generally high food supply) increases towards the uppermost Cretaceous, and peaks rapidly just above the K/Pg boundary, coeval with a peak in benthic foraminiferal accumulation rate (BFAR), a proxy for food supply. 2) A peak in relative abundance of Stensioeina beccariiformis, a cosmopolitan form generally more common at the middle than at the lower bathyal sites, occurs just above the buliminid peak. 3) The relative abundance of Nuttallides truempyi, a more oligotrophic form, decreases at the boundary, then increases above the peak in Stensioeina beccariiformis. The food supply to the deep sea in the Pacific Ocean thus apparently increased rather than decreased in the earliest Danian. The low benthic diversity during a time of high food supply indicates a stressed environment. This stress might have been caused by reorganization of the planktic ecosystem: primary producer niches vacated by the mass extinction of calcifying nannoplankton may have been rapidly (<10 kyr) filled by other, possibly opportunistic, primary producers, leading to delivery of another type of food, and/or irregular food delivery through a succession of opportunistic blooms.The deep-sea benthic foraminiferal data thus are in strong disagreement with the widely accepted hypothesis that the global deep-sea floor became severely food-depleted following the K/Pg extinction due to the mass extinction of primary producers (“Strangelove Ocean Model”) or to the collapse of the biotic pump (“Living Ocean Model”).     

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.     

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.     

Larger benthic foraminiferal assemblages and their response to Middle Eocene Climate Optimum in the Kohat Basin (Pakistan,eastern Tethys)

The Middle Eocene Climatic Optimum (MECO) at ～40 Ma is a significant global warming event associated with pronounced changes in the hydrosphere, atmosphere, and biosphere. The Kohat Formation in the Kohat Basin (eastern Tethys, Pakistan) is studied for identifying the response of larger benthic foraminifera (LBF) to MECO. The LBF assemblages in the Kohat Formation, covering from the Shallow Benthic Zones (SBZ) 15 to 17, suggest middle Lutetian to early Bartonian in age. Microfacies analyses indicate a lagoonal (inner carbonate ramp facies belt) to open marine (middle carbonate ramp facies belt) paleodepositional environment of the Kohat Formation. A distinct positive δ¹³C shift marks the stratigraphic position of the MECO in this formation. At the Peak-MECO event that is marked by the onset of the positive carbon isotope excursion (CIE), no evident compositional variation in the LBF assemblages is observed. However, significant changes in the LBF assemblages with the local first and last occurrences of some LBF genera can be observed in the Post-MECO and CIE recovery phase. These changes are verified by the sudden disappearance of Alveolina and orthophragminids and initial dominance of larger shell-size Nummulites fabianii, Heterostegina, and Linderina species accompanied by an increase in the species diversity. Here, we argue that the change in the observed LBF assemblages in the uppermost part of the Kohat Formation might be related to a larger foraminiferal turnover occurring during the Post-MECO event and corresponds to the CIE recovery phase.     

The effect of bioturbation by polychaetes (Opheliidae) on benthic foraminiferal assemblages and test preservation

Biological activity such as burrowing can alter benthic foraminiferal shell preservation and may also modify benthic foraminiferal assemblages by vertical mixing, inducing sediment homogenization. Here, we analyse benthic foraminiferal assemblages and taphonomy of upper Miocene marine deposits from Conil de la Frontera (Cádiz, south‐western Spain). The deposits consist of marls displaying a pervasive alternation of intensively bioturbated beds dominated by Macaronichnus segregatis traces (ichnofabric index 4–5) and non‐bioturbated beds. Benthic foraminiferal assemblages are dominated by Cibicidoides mundulus and Cibicides refulgens, indicating that the marls were deposited on an oligotrophic, well‐oxygenated upper slope. The impact of burrowing on the preservation of benthic foraminiferal tests was tested using Q‐mode cluster analysis, which found two well‐differentiated groups of samples, one including the non‐bioturbated beds and the other encompassing the bioturbated ones. Fragmentation and recrystallization account for the differentiation of these groups, both being higher in the bioturbated sediments. Aggressive chemical digestion by the Macaronichnus trace‐makers, assumed to be a polychaete worm of the family Opheliidae, etched the microfossil shells, making them more vulnerable to fragmentation. Intense bioturbation favoured the circulation of pore fluids, encouraging recrystallization. Pervasive burrowing resulted in significant vertical reworking of microfossils. As a consequence, benthic foraminiferal assemblages in the bioturbated beds were homogenized in the mixed layer; that is, the uppermost layer of the substrate totally burrowed. The alternation of bioturbated and non‐bioturbated beds reflects episodic transfer of food particles down slope from shallower parts of the shelf as well as from the continent due to storms under otherwise homogeneous oligotrophic marine conditions.