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.     

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”).     

Dinosaur sanctuary on the Chatham Islands, Southwest Pacific: First record of theropods from the K–T boundary Takatika Grit

Cretaceous–Tertiary (K–T) boundary (ca. 65 Ma) sections on a Southwest Pacific island containing dinosaurs were unknown until March 2003 when theropod bones were recovered from the Takatika Grit on the remote Chatham Islands (latitude 44° S, longitude 176° W), along the Chatham Rise. Tectonic and palaeontologic evidence support the eastward extension of a ca. 900 km land bridge that connected the islands to what is now New Zealand prior to the K–T boundary. The Chathams terrestrial fauna inhabited coastal, temperate environments along a low-lying, narrow, crustal extension of the New Zealand subcontinent, characterised by a tectonically dynamic, volcanic landscape with eroding hills (horsts) adjacent to flood plains and deltas, all sediments accumulating in grabens. This finger-like tract was blanketed with a conifer and clubmoss (Lycopodiopsida) dominated forest. The Chatham Islands region would have, along with New Zealand, provided a dinosaur island sanctuary after separating from the Gondwana margin ca. 80 Ma.     

Palynological indications of environmental changes during the Late Cretaceous–Eocene on the southern continental margin of Laurasia, Xizang (Tibet)

A palynostratigraphic and palynofacies study of a geological section at Cuojiangding in Zhongba County has provided a basis for discussing the palaeoenvironmental evolution of the southern continental margin of Laurasia in Xizang (Tibet) towards the end of the Cretaceous Period and during the Paleogene. It is the only record of fossil spores and pollen grains from collision- and convergence-related sediments close to the major Yarlung Zangbo Suture. Deposits of the Padana Formation at the base of the section through the Qubeiya, Quxia, and Jialazi formations to the Rujiao Zangbo Conglomerate at the top are considered to range in age from Santonian to Eocene or possibly Oligocene. During this period the India plate collided with, and continued to push against, Laurasia plate. Two spore and pollen zones and three palynofacies types are recognized. These are correlated with two transgressive–regressive cycles associated with the collision, periods of geologically rapid uplift being reflected by the molasse-like sediments of the upper Quxia Formation and the Rujiao Zangbo Conglomerate, which accumulated during the Late Paleocene and Eocene–Oligocene respectively. The first regression led to intermittent subaerial exposure and erosion of the Cuojiangding area. The second ended the marine history of the area and led first to the development of swamps in a subtropical climate, now preserved in the coal-bearing Qiuwu Formation, and later to the development of mountainous terrain, with a cooler climate at higher elevations.     

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.     

The early Ludfordian leintwardinensis graptolite Event and the Gorstian–Ludfordian boundary in Bohemia (Silurian,Czech Republic)

The late Gorstian to early Ludfordian hemipelagic succession of the south‐eastern part of the Prague Synform preserves a rich fossil record dominated by 28 species of planktic graptoloids associated with pelagic myodocopid ostracods, pelagic and nektobenthic orthocerid cephalopods, epibyssate bivalves, nektonic phyllocarids, rare dendroid graptolites, brachiopods, crinoids, trilobites, sponges and macroalgae. Faunal dynamics have been studied with particular reference to graptolites. The early Ludfordian leintwardinensis graptolite extinction Event manifests itself as a stepwise turnover of a moderate diversity graptolite fauna rather than an abrupt destruction of a flourishing biota. The simultaneous extinction of the spinose saetograptids Saetograptus clavulus, Saetograptus leintwardinensis and the rare S. sp. B. at the top of the S. leintwardinensis Zone was preceded by a short‐term acme of S. clavulus. Cucullograptus cf. aversus and C. rostratus vanished from the fossil record in the lower part of the Bohemograptus tenuis Biozone. No mass proliferation of Bohemograptus has been observed in the postextinction interval. Limited indigenous speciation gave rise to Pseudomonoclimacis kosoviensis and Pseudomonoclimacis cf. dalejensis. Egregiograptus (the only novel element of cryptic origin) made its earliest occurrence in association with the latest cucullograptids. A postextinction graptolite assemblage of moderate diversity, composed for the most part of long‐ranging taxa, persisted through the middle and upper tenuis Biozone until new rediversification in the Neocucullograptus inexpectatus Biozone. Unlike the graptoloids, the planktic, epiplanktic, nektonic and nektobenthic shelly fauna did not suffer significant extinction in the early Ludfordian. The Gorstian–Ludfordian boundary is placed at the lowest occurrence of S. leintwardinensis in spite of the very modest graptolite record available from the Ludfordian GSSP where no graptolites occur below the basal bed of the Lower Leintwardine Formation. Elsewhere, the Gorstian–Ludfordian boundary has been placed at the base of the Saetograptus linearis Biozone which has been considered roughly correlative with the leintwardinensis Biozone. Indeed, our morphometric study places the worldwide biozonal index species S. linearis in synonymy with S. leintwardinensis and thus considerably enhances the biostratigraphical utility of the latter index species. Pseudomonoclimacis antiqua sp. nov. is described.     

Extinction of nanicellid foraminifera during the Frasnian–Famennian biotic crisis: some far‐reaching evolutionary consequences

This study evaluates the severity of the poorly known and mostly underestimated foraminiferal extinction during the Frasnian–Famennian biotic crisis and its evolutionary aftermath. During this global event, worldwide, truly plurilocular planispiral (Nanicellidae) and uniseriate, palmate (Semitextulariidae) foraminifera associated with metazoan reefs died out entirely. Highly advanced test morphology such as that of nanicellids did not reappear in the earth's history until the Late Triassic. Moreover, morphotype comparable to that of the Devonian bilaterally flattened and palmate semitextularids appeared again until the Middle Jurassic (Frondicularia, Lagenida). In terms of the degree of test septation and chamber arrangement as well as general test shape, these foraminifera were ‘very far ahead of their time’. In consequence, foraminifera suffered a significant collapse during the F‐F biodiversity crisis, leading to an amazingly long evolutionary time lag in the case of plurilocular foraminifera lasting at least 150 million years.     

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.     

Benthic foraminifera associated with a hydrocarbon seep in the Rockall Trough (NE Atlantic)

Recent benthic foraminiferal assemblages in surface sediments of the Rockall Trough (NE Atlantic) have been qualitatively and quantitatively studied in order to investigate the effects of hydrocarbon seepage on benthic foraminiferal populations. Species diversity and abundance data have been examined in samples of similar lithology collected from hydrocarbon seep and non-seep (control) areas at a water depth of about 1000 m. Three species groups with different environmental preferences can be recognized. Group 1 dominates seep samples, and includes species tolerant to hydrocarbon emission, especially Angulogerina bradyana. In contrast, the less tolerant Group 2 species are weakly represented at seeps but dominate control samples. Group 3 species occur in low frequencies in both seep and non-seep samples. Furthermore, the measurement of species diversity (Shannon-Wiener and Simpson indices) demonstrates a difference in foraminiferal occurrence and frequencies between the seep and non-seep sites. Thus, the benthic foraminiferal distribution pattern is guided by different sensitivities of the species to hydrocarbons, reduced bottom-water oxygen usually associated with seepage and/or to a relatively elevated organic matter content in the sediment.     

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.     

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.     

Benthic foraminifera as bioindicators of pollution: A review of Italian research over the last three decades

Since the 1950s, numerous studies have demonstrated the value of benthic foraminifera in detecting ecosystem contamination. The interest in benthic foraminifera has partly been driven by government policies and programs aimed at developing suitable, non-invasive bioindicators of marine environmental quality. This paper accomplishes two things: it reveals that Italian experience has significantly contributed to the advancement of our understanding of this topic and summarizes the most important results that have served to greatly improve our knowledge in this field. Although many issues are still a matter of debate, since it is difficult to separate natural vs human-induced pollution and a foraminiferal protocol has not yet been produced, foraminifera have been proven to be successful candidates as part of an integrated monitoring program.     

Distribution of live benthic foraminifera in the Ría de Vigo (NW Spain)

The spatial and vertical distributions of live benthic foraminifera from the muddy central axis of the Ría de Vigo were examined under downwelling and upwelling conditions. The spatial distribution of the main benthic foraminiferal species is controlled by salinity variations (San Simón Assemblage), food availability (Outer Ría Assemblage) and oxygen concentrations and/or reducing microenvironmental conditions in the sediment (Inner Ría Assemblage).The microhabitat distribution of benthic foraminifera is mainly dependant on the oxygenation/redox conditions in the sediment and the vertical segregation of food. In the inner areas of the ría, low oxygen and/or reducing microenvironmental conditions prevent the vertical segregation of microhabitats. In outer areas oxygen penetration is deep but microhabitat partitioning only occurs in the presence of additional food resources (mainly fresh organic carbon from phytoplankton blooms) which triggers the colonisation of the surface layer by new species and the migration upwards and reproduction of opportunistic species.     

Foraminiferal turnover across the Eocene–Oligocene transition at Fuente Caldera, southern Spain: No cause–effect relationship between meteorite impacts and extinctions

We studied planktic and small benthic foraminifera from the Fuente Caldera section, southern Spain, across the Eocene–Oligocene transition. Benthic foraminifera indicate lower bathyal depths for the late Eocene and earliest Oligocene. Detailed high-resolution sampling and biostratigraphical data allowed us to date precisely layers with evidence for meteorite impact (Ni-rich spinel), which occur in the lower part of the planktic foraminiferal Globigerapsis index Biozone and in the middle part of the small benthic foraminiferal Cibicidoides truncanus (BB4) Biozone (middle Priabonian, late Eocene). Major turnovers of foraminifera occur at the Eocene/Oligocene boundary, only. The impact did not occur at a time of planktic or benthic foraminiferal extinction events, and the late Eocene meteorite impacts did thus not cause extinction of foraminifera. The most plausible cause of the Eocene/Oligocene boundary extinctions is the significant cooling, which generated glaciation in Antarctica and eliminated most of the warm and surface-dwelling foraminifera.     

Neoselachian (Chondrichthyes, Elasmobranchii) diversity across the Cretaceous-Tertiary boundary

Fishes are often thought to have passed through mass extinctions, including the Cretaceous-Tertiary (KT) event, relatively unscathed. We show that neoselachian sharks suffered a major extinction at the K/T boundary. Out of 41 families, 7 became extinct (17±12%). The proportional measure increases at lower taxic levels: 56±10% loss of genera (loss of 60 out of 107) and 84±5% loss of species (loss of 182 out of 216). However, the Maastrichtian and Danian are characterized by a high number of singleton taxa. Excluding singletons we have calculated a 34±11% loss of genera and a 45±9% loss of species. The simple completeness metric (SCM) for genera displays a decrease from the Maastrichtian (94%) to the Danian (85%) indicating a rather complete fossil record of neoselachian genera. The extinctions were heavy among both sharks and batoids (skates and rays), but most severe among batoids, which lost almost all identifiable species. There were equal losses among open marine apex predators (loss of Anacoracidae, Cretoxyrhinidae, and Scapanorhynchidae) and durophagous demersal forms from the continental shelf and shallow seas (Hypsobatidae, Parapaleobatidae, Sclerorhynchidae, Rhombodontidae). Benthopelagic and deep-sea forms were apparently little affected. New families with similar ecological roles (Carcharhinidae, Isuridae, Torpedinidae) replaced these families in the Danian, and full diversity of the different shark and batoid groups had been recovered by the end of the Paleocene or early Eocene. Sharks and rays suffered levels of extinction entirely in line with other groups of organisms at the K/T extinction event.     

Integrated magnetobiochronology of the Early/Middle Eocene transition at Agost (Spain): Implications for defining the Ypresian/Lutetian boundary stratotype

In this paper, we present an integrated study of a 115‐m‐thick section that spans the Ypresian/Lutetian boundary at Agost (Betic Cordillera, SE Spain). Our study includes magnetostratigraphic results and biostratigraphic and palaeoenvironmental data derived from planktic foraminifera, small and larger benthic foraminifera, and calcareous nannofossils. Our results demonstrate that the Agost section is continuous and spans from Zones P9 to P12 (E7 to E10), Zones CP11 to CP14a (NP13 to NP16), Zones SBZ11 to SBZ15, and Chrons C22n to C19r. The first occurrence (FO) of H. nuttalli (base of P10) and the FO of G. nuttalli (base of E8) are found within Chron C20r, at a much younger age (3–5 Myr) than previously considered in standard calibration schemes. Similarly, the boundary between SBZ12 and SBZ13 is located within Chron C21n, also at a younger age than previously considered. On the contrary, the FO of B. inflatus (base of CP12b) is found within Chron C21r, which conforms to the magnetostratigraphically calibrated age of ca. 48 Ma (middle part of C21r) considered in standard calibration schemes. These results corroborate earlier studies and indicate that all the events that have been proposed to mark the Ypresian/Lutetian boundary appear at different stratigraphic intervals and have different ages. Based on our results from Agost and on data from other sections elsewhere, we suggest that the Ypresian/Lutetian boundary might be approximated by the FO of B. inflatus (base of CP12b). The Agost section might be considered as a potential candidate to locate the Global Stratotype Section and Point (GSSP) of the base of the Lutetian Stage, because it includes all the events that might be selected as marker events for the Ypresian/Lutetian boundary and it fulfils most of the geological, biostratigraphic and infrastructure requirements demanded for definition of a GSSP.     

Stability and change in Tethyan planktic foraminifera across the Paleocene–Eocene transition

Examination of planktic foraminifera in the Tethys basin during the Paleocene–Eocene transition reveals two stasis intervals that are separated by a major saltation event coincident with the P–E short-term perturbation in global climate and oceanography. Changes occurred at many spatial and temporal scales as well as many taxonomic and ecologic hierarchical levels, though with various rates and magnitudes. The stasis intervals are marked by slow changes at the species level and account for 50% of the observed first and last appearances during a 2.5 Myr interval. The saltation event is marked by rapid changes at the species and morpho-guild levels and accounts for the remaining 50% of first and last appearances during an interval of about 100–200 kyr. Despite these changes, many taxonomic and ecologic units, such as the depth assemblages and genera, and faunal parameters, such as species richness and turnover rates, are stable with respect to the P–E perturbation. This coexistence of change and stability marks the crisis of Tethyan planktic foraminifera across the P–E transition and reveals the possible dynamics of ecological evolution.     

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.     

Habitat variables determining the occurrence of large benthic foraminifera in the Berau area (East Kalimantan, Indonesia)

Knowledge of the assemblage composition of large benthic foraminifera (LBF) in relation to environmental conditions is needed to interpret fossil records. In this study the assemblage composition of LBF is described for a carbonate shelf with a barrier reef system and some reefs outside the barrier. In a total of 140 samples, 34–35 species of LBF were found. Four clusters, roughly corresponding to substrate type were identified. Several small groups of samples were recognized that were collected locally inside, on, or outside the barrier. Microscale environmental gradients within the substrate or in the benthic boundary layer resulted in spatial differentiation of the microhabitat of each species.