Benthic foraminiferal ecomarker species of the terminal Cretaceous (late Maastrichtian) deep-sea Tethys

Benthic foraminiferal distribution patterns throughout the late Maastrichtian Tethyan deep sea are analyzed. Many species are ubiquitously distributed throughout this region and therefore it is hard to assess their ecological preferences. However, five species show distribution patterns, which suggest that they may have distinctive paleoenvironmental preferences. These preferences are interpreted from hypothesized surface circulation and upwelling patterns. Additional information comes from Recent benthic foraminiferal ecology and from responses to the Cretaceous/Paleogene (K/Pg) boundary event. This enables us to assess the ecological preferences of these late Maastrichtian taxa, and establish them as ecological-marker (ecomarker) species for paleoenvironmental interpretation of the late Maastrichtian bathyal-abyssal Tethyan realm.(1) Eouvigerina subsculptura is suggested to be indicative of reasonably oxygenated upper-middle bathyal environments, though with high abundance of utilizable organic matter. (2) Sliteria varsoviensis is linked to areas of late Maastrichtian upwelling and seems to have been an epibenthic species with an opportunistic life mode. (3) Gavelinella beccariiformis and (4) Nuttallides truempyi are considered to be indicative of oligotrophic conditions unless they occur with a large proportion of endobenthic morphotypes. (5) Gavelinella pertusa is proposed to indicate neritic-middle bathyal environments of the ‘boreal’ realm, which might be influenced by more seasonal food-fluxes and by higher oxygen levels than similar settings in the (sub)tropics.Finally, the anomalous high abundances of the buliminid species Sitella cf. plana in deep open ocean environments is discussed in terms of possible mechanisms permitting such a (morphologically) opportunistic species to thrive in such an assumedly oligotrophic environment.     

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.     

Benthic foraminiferal paleoenvironmental reconstruction from the Upper Coniacian-Lower Campanian succession in north-western Tunisia

Quantitative analysis of benthic foraminifera is used to characterize the paleoenvironments of the Upper Coniacian-Lower Campanian succession in the Jbil section of north-western Tunisia. Foraminiferal parameters and benthic foraminiferal assemblages show that the studied section includes four distinct paleoenvironmental phases. From oldest to youngest, these are as follows: (1) an interval with a Praebulimina reussi assemblage with infaunal ratios as high as 96.1%. High abundances of P. reussi, reflecting an increase in organic matter flux to the seafloor (meso-to eutrophic) under oxygenated bottom-water conditions. (2) An interval characterized by a Gavelinella costulata assemblage with mixed infaunal/epifaunal foraminifera with higher Fisher's alpha values (ranging from 4 to 15.2), reflecting mesotrophic conditions in an outer shelf environment. (3) An interval with a Gaudryina laevigata assemblage indicative of a middle to outer shelf environment; there is a considerable increase in infaunal agglutinated foraminifera, as well as a relatively abundant and moderately diversified oxic/suboxic foraminifera. (4) The final interval occurs in the lower Campanian (the Globotruncana ventricosa Zone) and includes a Bolivinoides decoratus assemblage reflecting an outer shelf to upper bathyal environment. It contains a higher planktonic percentage and biodiversity with a slight increase in dysoxic species; the mixed infaunal/epifaunal content (57.6 to 73.3%) reflects mesotrophic conditions. Four well-recognized major sea-level falls are matched by the dual signatures of eustatic sea-level changes. These are coincident with the results of this study, which represent the first documentation of these events in Tunisian faunal and paleoenvironmental changes, at the following boundaries: Coniacian/Santonian, intra-Santonian, Santonian/Campanian, and intra-early Campanian.     

Benthic foraminifera across the Cretaceous–Tertiary (K–T) boundary: a review

The response of the Earth’s biota to global change is of fundamental interest to paleontologists, but patterns of change in paleontologic data are also of interest to a wider spectrum of Earth scientists in that those patterns are of great significance in constraining hypotheses that attempt to explain physical changes in the Earth’s environment. The Cretaceous–Tertiary (K–T) boundary is a case in point. Some paleontologists have criticized the bolide impact hypothesis, not because they deny the impact but because the proposed effects of that impact do not always conform to the available paleontological data. Benthic foraminifera are of particular interest in this context because it has been suggested for over 20 years that shallow-water benthic foraminifera were affected more severely than deep-water benthic foraminifera by events at the K–T boundary. This observation adds to the fact of planktonic foraminiferal extinction and indicates that K–T boundary environmental effects were largely restricted to shallow waters. In this paper I review all published works on smaller benthic foraminifera at the K–T boundary and conclude the following. (1) Shallow-water benthic foraminifera were not more severely affected than deeper dwelling species. True extinction, as opposed to local extinction and/or mass mortality, is generally quite low no matter what the water depth. (2) The data are not sufficient in quality, quantity and geographic range to conclude that there is a latitudinal pattern of extinction. (3) In general, biotic changes (such as they are) begin before the boundary in shallow and intermediate depth waters and at the boundary in deep water. Disagreements about the placement of the boundary and the presence, absence and duration of hiatuses hinder more precise conclusions. (4) There appears to be preferential survivorship of epifaunal species into the early Danian with a short interval dominated by infaunal taxa in the earliest Danian. This pattern can best be explained by short-lived input of increased amounts of organic matter at the boundary followed by a sudden collapse of primary productivity and, hence, major reduction or cessation of organic flux to the seafloor. In summary, based on the current dataset, smaller benthic foraminifera, no matter whether they lived in shallow or deep waters, high or low latitudes, or infaunal or epifaunal microhabitats, survived the environmental events across the K–T boundary quite well. Mass extinction does not characterize this group of organisms at this time.     

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.     

Planktonic foraminiferal turnover, diversity fluctuations and geochemical signals across the Eocene/Oligocene boundary in Tanzania

A major turnover in planktonic foraminifera occurred across the Eocene/Oligocene (E/O) boundary. New drill holes through the E/O boundary in southern Tanzania contain extremely well-preserved and diverse assemblages of planktonic foraminifera. Here we document a 1.2 million year record of assemblages, diversity and stable isotope fluctuations through this critical interval, which is often dissolved and/or recrystallised in carbonate-rich facies. The E/O boundary is marked by the abrupt extinction of all five remaining species of the family Hantkeninidae and a distinct size reduction in the genus Pseudohastigerina. The boundary is preceded over a short stratigraphic interval by the extinction of Turborotalia cerroazulensis, Turborotalia cocoaensis and Turborotalia cunialensis. Quantitative analysis of planktonic foraminiferal assemblages reveals significant changes in the abundance of certain species and the composition of the assemblages. We compare diversity fluctuations to the stable isotope record of Pseudohastigerina naguewichiensis and use multispecies stable isotope analyses to determine the life habitats of the most important species. A major shift in the evenness occurs at ~ 33.8 Ma associated with the extinction of the T cerroazulensis group suggesting acute ecological disturbance. We propose that the extinction of the T. cerroazulensis group at ~ 33.8 Ma was directly related to cooling of sea surface temperatures, while the extinction of Hantkeninidae was due to modifications in the thermal structure of the oceans and associated productivity changes. After the extinctions, renewed origination and diversification occurred, leading to a characteristic Oligocene planktonic foraminifer assemblage.     

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”).     

Recent benthic foraminiferal assemblages in deep high-energy environments from the Gulf of Cadiz (Spain)

The benthic environment in the Gulf of Cadiz, north-eastern Atlantic, is strongly affected by the Mediterranean outflow water undercurrent (MOW) which flows northwards along the western Iberian Margin at 500–1500 m water depth. Foraminiferal census counts of living and dead assemblages from 27 surface samples ranging from 103 to 1917 m water depth, and the examination of hard substrates reveal a close correlation of the fauna with the local hydrography and sediment facies. Four different faunal groups are separated by factor analysis of the living fauna. Assemblage 1 contains typical lower slope species and dominates samples from the lower MOW core layer and in the North Atlantic deep water below. Shelf edge foraminifera are common in assemblage 2a which shows the highest proportions in samples from 103 to 272 m. Assemblage 2b is dominated by upper slope species and suspension-feeders that are frequent in the upper MOW core layer and in distal settings between 396 and 901 m. Species from assemblage 3 prefer epibenthic habitats and are recorded with high proportions exclusively in the immediate flow paths of the upper MOW between 496 and 881 m. Colonisation structures and species composition of epibenthic assemblages from the proximal facies largely differ from those in distal settings. In general, epibenthic foraminifers only use elevated substrates under the influence of near-bottom flow. Under high current velocities, epibenthic foraminifers prefer large and heavy objects. They colonise high attachment levels where a maximum yield of advected food particles can be achieved. In distal settings at lower flow velocities, the elevation height does not exceed 20 mm above the surrounding sediment surface. This level is related to a hydrologic transition layer with high concentrations of suspended particles. The comparison of microhabitat preferences and faunal structure under high and low current velocities reveal that substrate stability may be a confining environmental variable for endobenthic and shallow epibenthic foraminifers. The observations also indicate that the preferential settling height of epibenthic foraminifera is related to the highest lateral flux rates of food particles within reach from the sea floor. A dynamic selection of elevated microhabitats is only used by 7.8% of all species recognised in the Gulf of Cadiz area.     

The Madenli (Central Taurides) Upper Cretaceous platform carbonate succession: Benthic foraminiferal biostratigraphy and platform evolution

The Upper Cretaceous succession in the Madenli area (western Central Taurides, Southern Turkey) consists of platform carbonate rocks deposited in entirely peritidal environments, which are sensitive to sea level changes driven by global eustasy, but also strongly affected by local and regional tectonics. It includes economically important bauxite deposits. Previous works suggest different ages for bauxite formation ranging from the Albian to the Santonian. Benthic foraminiferal biostratigraphy and facies analysis of the Madenli and Doğankuzu outcrop sections allow for a more precise dating of the platform emersion periods. The footwall limestones of the bauxite deposits consist of well-bedded limestones (Unit-1), which contain a benthic foraminiferal assemblage (BFA) including mainly Biconcava bentori and Pastrikella biplana, Chrysalidina gradata (BFA I), assigned to the middle-upper Cenomanian. In the Madenli section, the first bauxite deposit occurs in the upper part of Unit-1 as a layer interbedded with pinkish sparitic and dolomitic beds (subunit-1a) deposited in supratidal environment. Subunit-1a is stratigraphically equivalent to the Doğankuzu and Mortaş bauxite deposits considered as karst-related, unconformity-type deposits. The hanging-wall limestones of the bauxite are represented by the massive limestones (Unit-2) starting locally with either the upper Cenomanian characterized mainly by the presence of Pseudolituonella reicheli or upper Campanian comprising mainly Murciella cuvillieri and Moncharmontia apenninica (BFA II). There is no field evidence of a discontinuity surface at the contact between the lower part of Unit-2, including BFA I, and the upper part of Unit-2, including BFA II. This contact is defined as a paraconformity indicating a stratigraphic gap from the Turonian to the early Campanian. The top of Unit-2 is truncated by another discontinuity surface associated with a minor bauxite deposit. The overlying Unit-3 is characterized by well-bedded, rudist-bearing limestones topped by laminated and dolomitized limestones organized in shallowing upward cycles. It is assigned to the upper Maastrichtian based on the presence of Rhapydionina liburnica (BFA III) and rudist assemblage. A third emersion period of the platform corresponds to the early Maastrichtian.     

Integrated biostratigraphy based on planktonic foraminifera and dinoflagellates across the Cretaceous/Paleogene (K/Pg) transition at the Izeh section (SW Iran)

The present work is based on semi-quantitative study carried on detailed sampling (samples are spaced by 5, 10 and 15 cm close to the boundary) of an essentially continuous and expanded section crossing the Cretaceous–Paleogene (K/Pg) boundary in Iran. By this work, we attempt to detail biostratigraphy based on planktonic foraminifera biozones and correlate biozones and subzones with dinocyst events. The entire Cretaceous–Paleogene interval contains rich, diversified and well-preserved planktonic foraminifera and dinoflagellate cyst assemblages. Four planktonic foraminiferal biozones have been recognized across the Cretaceous–Paleogene transition (K/Pg): Abathomphalus mayaroensis Biozone including Plummerita hantkeninoides Subzone from the Late Maastrichtian and Guembelitria cretacea (including Hedbergella holmdelensis and Parvularugoglobigerina longiapertura subzones), Parvularugoglobigerina eugubina Biozone and Parasubbotina pseudobulloides Biozone belonging to the Early Danian. These biozones have been correlated with four dinocyst biozones: the Manumiella seelandica Biozone belonging to the Late Maastrichtian and the Alisocysta reticulata, Senoniasphaera inornata and Damassadinium californicum biozones from the Early Danian. At this section, like at the El Kef section (GSSP for the K/Pg) and the auxiliary sections, an Ir anomaly is detected indicating the K/Pg boundary. This geochemical anomaly coincides also with mass extinctions of planktonic foraminifera species. The extinct species are in particular the large, complex tropical and subtropical taxa dwelling in subsurface and lower photic water. The mass extinctions at the Izeh section occurred over a succinct period of time similar to the K/Pg type section at El Kef (Tunisia). These sudden mass extinctions indicate a catastrophic pattern event occurring at the Maastrichtian/Danian boundary. In contrast the organic-walled dinocysts were less affected by the mass extinction and most species crossed the K/Pg boundary without showing mass and sudden extinctions. Nevertheless, they showed changes in their assemblages’ structure beyond the K/Pg boundary. Especially, Manumiella seelandica and M. druggii, typical species of Antarctic Maastrichtian dinocysts assemblages, occur in coeval deposits at the Izeh section; they persist through the Lower Danian and, like in Tunisia (e.g., El Kef section, Ellès section) show an obvious increase in relative abundance.     

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.     

Radiolarian faunal turnover across the Oligocene/Miocene boundary in the equatorial Pacific Ocean

The global warming trend of the latest Oligocene was interrupted by several cooling events associated with Antarctic glaciations. These cooling events affected surface water productivity and plankton assemblages. Well-preserved radiolarians were obtained from upper Oligocene to lower Miocene sediments at Ocean Drilling Program (ODP) Leg 199 Sites 1218 and 1219 in the equatorial Pacific, and 110 radiolarian species were identified.Four episodes of significant radiolarian faunal changes were identified: middle late Oligocene (27.5 to 27.3 Ma), latest Oligocene (24.4 Ma), earliest Miocene (23.3 Ma), and middle early Miocene (21.6 Ma). These four episodes approximately coincide with increases and decreases of biogenic silica accumulation rates and increases in δ¹⁸O values coded as “Oi” and “Mi” events. These data indicate that Antarctic glaciations were associated with change of siliceous sedimentation patterns and faunal changes in the equatorial Pacific.Radiolarian fauna was divided into three assemblages based on variations in radiolarian productivity, species richness and the composition of dominant species: a late Oligocene assemblage (27.6 to 24.4 Ma), a transitional assemblage (24.4 to 23.3 Ma) and an early Miocene assemblage (23.3 to 21.2 Ma). The late Oligocene assemblage is characterized by relatively high productivity, low species richness and four dominant species of Tholospyris anthophora, Stichocorys subligata, Lophocyrtis nomas and Lithelius spp. The transitional assemblage represents relatively low values of productivity and species richness, and consists of three dominant species of T. anthophora, S. subligata and L. nomas. The characteristics of the early Miocene assemblage are relatively low productivity, but high species richness. The two dominant species present in this assemblage are T. anthophora and Cyrtocapsella tetrapera. The most significant faunal turnover of radiolarians is marked at the boundary between the transitional/early Miocene assemblages.We also reviewed changes in other microfossil assemblages in the low latitudes during the late Oligocene through early Miocene. The microfossil assemblages of major groups show sequential changes near the Oligocene/Miocene (O/M) boundary (23.8 Ma). Many extinction events and some first occurrences of calcareous nannofossils and many occurrences of radiolarians are found from about 24.8 to 23.3 Ma, and first occurrences of planktic foraminifers and diatoms followed from 23.2 through 22 Ma. Hence, the O/M boundary is identified as a significant level for microfossil evolutions.     

Ammonia beccarii and Ammonia tepida (Foraminifera): morphofunctional arguments for their distinction

New observations from Yeu Island and the Bay of Bourgneuf on the Atlantic coast of France show that Ammonia beccarii s.s. and Ammonia tepida have different morphofunctional adaptations to their habitats and environments. Adult A. beccarii s.s. may be epiphytic, living on the calcareous alga Corallina officinalis or on the red alga Gigartina acicularis along rocky shores, while A. tepida is endopelic, living in sediments of brackish environments. Rapid 3-D emission of pseudopodia from interlocular spaces on both umbilical and spiral sides of adult A. beccarii s.s. can support its test within the seaweed framework. This behaviour is characteristic of phytal species. Thus, it appears that the major difference between A. beccarii s.s. and A. tepida is the presence of large interlocular spaces on spiral side of the former, corresponding to a potential morphofunctional adaptation to the epiphytic mode of life. We believe that these observations could help to solve the long-lasting A. beccarii problems of specific distinction.     

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.     

Cyclic environmental changes during the Early Toarcian at the Mochras Farm Borehole (Wales): a variable response of the foraminiferal community

Cyclostratigraphical analysis of the foraminiferal assemblages from the Early Toarcian at the Mochras Farm Borehole (Wales) was conducted in order to evaluate the incidence of cyclic palaeoenvironmental changes on the foraminiferal community. Different variables such as type of morphogroup, evolutionary strategy, habitat, particular taxa, diversity and abundance were studied using the Lomb–Scargle periodogram implemented in the computer program SLOMBS. A well‐developed cyclostratigraphical pattern is recognized, with the presence of several cycles (in metres) at 3.4–4/7.2–7.5/10.1–10.6/32.1–33.3/104.2–111.2/128.2/166.7, belonging to the high‐, middle‐, middle‐/low‐, and low‐frequency bands. The incidence and relevance of the cycles is found to be related to particular variables marking the global and local character of the involved processes. Cyclic changes in the organic matter input are found to be the most relevant palaeoenvironmental factor, oxygenation being secondary. A correspondence with specific Milankovitch cycles is, at present, difficult to determine.     

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.     

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.     

New occurrences of the wood Protocupressinoxylon purbeckensis Francis: implications for terrestrial biomes in southwestern Europe at the Jurassic/Cretaceous boundary

Abstract:  Previously known from the Kimmeridgian–Portlandian of Dorset (UK) only, Protocupressinoxylon purbeckensis wood is reported here from the Kimmeridgian of Asturias (Spain) and Ajoie (Switzerland). The morphospecies taxonomy and nomenclature are discussed, and new supplementary illustrations are given. The P. purbeckensis tree was growing in dry strongly seasonal (tropophilous) environments, and the new occurrences suggest that such a climate prevailed on land all over southwestern Europe at the end of the Jurassic (Kimmeridgian sensu anglico – Portlandian). The review of fossil wood data indicates that such a stressful environment may have constrained terrestrial biocoenoses and their evolution at the Jurassic/Cretaceous boundary. But wood generic diversity curves are also strikingly similar to that drawn 20 years ago for nonmarine tetrapods, implying a fossil Lagerstätte effect.     

Integrated stratigraphy and astronomical calibration of the Serravallian/Tortonian boundary section at Monte Gibliscemi (Sicily, Italy)

Results are presented of an integrated stratigraphic (calcareous plankton biostratigraphy, cyclostratigraphy and magnetostratigraphy) study of the Serravallian/Tortonian (S/T) boundary section of Monte Gibliscemi (Sicily, Italy). Astronomical calibration of the sedimentary cycles provides absolute ages for calcareous plankton bio-events in the interval between 9.8 and 12.1 Ma. The first occurrence (FO) of Neogloboquadrina acostaensis, usually taken to delimit the S/T boundary, is dated astronomically at 11.781 Ma, pre-dating the migratory arrival of the species at low latitudes in the Atlantic by almost 2 million years. In contrast to delayed low-latitude arrival of N. acostaensis, Paragloborotalia mayeri shows a delayed low-latitude extinction of slightly more than 0.7 million years with respect to the Mediterranean (last occurrence (LO) at 10.49 Ma at Ceara Rise; LO at 11.205 Ma in the Mediterranean). The Discoaster hamatus FO, dated at 10.150 Ma, is clearly delayed with respect to the open ocean. The ages of D. kugleri first and last common occurrence (FCO and LCO), Catinaster coalitus FO, Coccolithus miopelagicus last (regular) occurrence (L(R)O) and the D. hamatus/neohamatus cross-over, however, are in good to excellent agreement with astronomically tuned ages for the same events at Ceara Rise (tropical Atlantic), suggesting that both independently established timescales are consistent with one another. The lack of a reliable magnetostratigraphy hampers a direct comparison with the geomagnetic polarity timescale of Cande and Kent (1995; CK95), but ages of calcareous nannofossil events suggests that CK95 is significantly younger over the studied time interval. Approximate astronomical ages for the polarity reversals were obtained by exporting astronomical ages of selected nannofossil events from Ceara Rise (and the Mediterranean) to eastern equatorial Pacific ODP Leg 138 Site 845, which has a reliable magnetostratigraphy.Our data from the Rio Mazzapiedi–Castellania section reveal that the base of the Tortonian stratotype corresponds almost exactly with the first regular occurrence (FRO) of N. acostaensis s.s. as defined in the present study, dated at 10.554 Ma. An extrapolated age of 11.8 Ma calculated for the top of the Serravallian stratotype indicates that there is a gap between the top of the Serravallian and the base of the Tortonian stratotype, potentially rendering all bio-events in the interval between 11.8 and 10.554 Ma suitable for delimiting the S/T boundary. Despite the tectonic deformation and the lack of a magnetostratigraphy, Gibliscemi remains a candidate to define the S/T boundary by means of the Tortonian global boundary stratotype section and point (GSSP).     

Benthic foraminiferal morphogroups of mid to outer shelf environments of the Late Jurassic (Prebetic Zone, southern Spain): Characterization of biofacies and environmental significance

Benthic foraminiferal assemblages belonging to a mixed carbonate–siliciclastic shelf succession of the Oxfordian (Upper Jurassic) in the Prebetic, southern Spain, were analyzed. The faunal data, obtained on thin sections of strongly lithified sediments, allowed the detailed differentiation of a foraminiferal morphogroups system to be applied to the interpretation of paleoenvironmental conditions. Eleven morphogroups were differentiated according to shell composition, general morphology, number of chambers and mode of coiling. Paleoenvironmental analysis combines these features with inferred life-style (epifaunal, shallow infaunal and deep infaunal) and feeding strategy (suspension-feeder, deposit-feeder, herbivores, etc.) of the foraminifera. Analogies and differences regarding previously established morphogroup systems could be outlined accordingly.

In the Prebetic Oxfordian, distribution trends of foraminiferal morphogroups allow for a comparison of mid to outer shelf environments occupying the South-Iberian paleomargin, indicating that nutrient supply was the most important factor controlling morphogroup development. The mid shelf settings are dominated by epifaunal active herbivores–phytodetritivores, in shallow to deep substrate position of the redox boundary. The outer shelf is dominated by agglutinated and infaunal detritivore-bacterial scavenger morphogroups revealing a deep substrate position of the redox boundary. Particularly favorable trophic conditions in mid shelf environments are related with higher phytodetritus input from more proximal and shallower shelf areas.