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

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

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”).     

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

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

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

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

Relationship of Foraminifera and thecamoebian distributions to sediments contaminated by pulp mill effluent: Saguenay Fiord, Quebec, Canada

Gravity cores and grab samples collected in the Saguenay Fiord between 1976 and 1988 contain the record of a 20th century benthic marine environment contaminated primarily by organic matter discharges from several local pulp and paper mills.Spiroplectammina biformis is the dominant arenaceous species. Its living percent abundance decreases between 1982 and 1988 as a consequence of the apparent recolonization of the upper reaches of the fiord by several arenaceous taxa, the most important beingTextularia earlandi. Cassidulina reniforme, the dominant living calcareous species, shows about a three fold increase in relative abundance over the six year interval. Grab sample observations also suggest a recolonization of some formerly barren benthic environments near the head of the fiord by foraminiferal species between 1982 and 1988. This recolonization may reflect the combined effect of government regulations imposed on local industrial polluters in the early 1970's and the capping of a large area of contaminated sediment by a layer of clay that was transported to the basin at the head of the fiord as the result of a catastrophic landslide in 1971.X-radiographs of core sections from a basin at the head of the fiord show sharp laminations that reflect both the absence of bioturbators and the seasonal variation of fluvial depositional processes. Organic waste deposition from pulp and paper mill outfalls first increased around 1910 and shows a first order inverse relationship to benthonic foraminiferal assemblage diversity. The presence of only allochthonous thecamoebian and reworked planktonic Foraminifera tests in many benthonic Foraminifera-barren core intervals supports the idea that common indigenous estuarine species such asSpiroplectammina biformis were excluded from these areas at certain times as a consequence of pollution “stress” rather than having been removed from the sediment record as a result of post-depositional diagenetic effects. However, future studies of estuarine pollution history based on proxy foraminiferal data from cores must give greater emphasis to distinguishing ecological stress responses from diagenetic process (i.e. test dissolution effects) since both mechanisms can be reflected in the proxy pollution “signal” in a similar way.     

Benthische Foraminiferen auf dem Island-Schottland Rücken: Umwelt-Anzeiger an der Grenze zweier ozeanischer Räume

Surface sediment samples taken by box corer from 32 stations on the Iceland-Scotland Ridge have been investigated for their benthic foraminiferal content. The live (Rose Bengal stained) benthic foraminiferal fauna was differentiated from empty tests comprising the foraminiferal death assemblage. Principal component analysis of both the live and dead faunal data from the Iceland-Scotland Ridge reveals eight live species assemblages and six corresponding dead assemblages. Bottom water current conditions, surface sediment characteristics, particulate organic matter supply, and to some extent also the bottom water temperatures are the main factors limiting and governing the composition and distribution of live benthic foraminiferal species assemblages on the Iceland-Scotland Ridge. On the Atlantic slope of the Iceland-Scotland Ridge the dead species assemblages differ greatly from the foraminiferal fauna living there today due to winnowing processes and redeposition of Pleistocene sediments. In this area an investigation of distribution patterns of the empty tests only would lead to wrong results concerning ecologic interrelations between benthic foraminiferal species assemblages and their environment.     

Dasycladaleans from the Upper Turonian to Santonian of Austria (Gosau Group pro parte) and paleobiogeographic considerations

Previously unreported dasycladaleans and one morpho-taxon of assumed algal origin are described from Upper Turonian to Santonian rocks of the Lower Gosau Subgroup (LGS) of the Northern Calcareous Alps. A taxonomic inventory of green-algal/benthic foraminiferal assemblages shows that assemblages of “pure” carbonate environments are more diverse than those of siliciclastic and mixed siliciclastic-carbonate settings. A comparison of the taxonomic inventory of the LGS with assemblages in similar sedimentary successions of the Alpine-Mediterranean realm shows the highest similarity with the “Mirdita Zone” of the internal Dinarids. Comparability of assemblages, however, is limited due to narrow chronostratigraphic overlap and/or because of scarcity of data from areas outside the Alps. Although higher than previously known, the total diversity of the green-algal/benthic foraminiferal assemblage of the LGS is clearly inferior to that of the peri-Adriatic carbonate platforms.     

Modern benthic foraminifera on Banco Chinchorro,Quintana Roo,Mexico

Benthic foraminiferal assemblages may be used as environmental indicators on Banco Chinchorro, an isolated carbonate platform off the eastern coast of the Yucatan Peninsula. Seventy-seven species from 44 genera were identified in 14 surface sediment samples, which were collected along an E–W windward-to-leeward transect across the platform. A total of 15,493 foraminiferal tests (max. 1,200 tests per sample) were investigated and served as the basis of this study. As many taxa range throughout several platform zones, assemblages are better environmental indicators as compared to individual species. Four foraminiferal assemblages were identified using statistical methods including (1) a Homotrema assemblage, which occurs at the windward platform margin, (2) an Archaias-Homotrema assemblage which is found on the leeward margin and on platform interior coral patch reefs, (3) a Quinqueloculininae-Archaias-Rosalina assemblage of the western platform, and (4) an Archaias-Quinqueloculininae assemblage characteristic of the eastern platform interior. Environmental factors which influence variation in foraminiferal distribution and diversity on Banco Chinchorro include exposure to waves and currents and substrate (plant and algal growth). Sediment transport does not play a major role in Banco Chinchorro based on the observation that there are only limited amounts of taxa found outside their typical habitats, and, that mean grain-size and sorting of foraminiferal tests do not exhibit clear patterns. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Benthic foraminiferal distribution and abundance changes in Skagerrak surface sediments: 1937 (Höglund) and 1992/1993 data compared

Both living (stained) and dead (unstained) foraminiferal assemblages from surface sediments (0–2 cm) in the northwestern part of the Skagerrak have been studied in order to (1) define and characterize the distribution of various modern benthic environments and (2) by comparing these findings with surface samples collected 40–60 years ago, to document possible faunal changes that might have occurred. The investigated area is physiographically divided into the Norwegian slope, the Skagerrak Basin, and the Danish slope. The latter is under the influence of the Jutland Current, while the basin and the investigated parts of the Norwegian slope are bathed in Atlantic water. All areas have bottom waters with a high oxygen concentration. Three living (stained) and three dead (unstained) assemblages occupy the three physiographic areas. Only one assemblage (on the Norwegian slope) is common to both the living and dead assemblages but the boundaries between them lie at comparable depths. The higher standing crops are found on the fertile Danish slope while the lower ones are in the deep basin where the diversity is at a maximum. In the dead assemblages, the relative abundance of agglutinated tests increases with depth. Comparison with data collected 40 to 60 years ago shows increases in absolute numbers of tests, especially in the deep basin. There are changes in assemblage compositions in all areas. The dominant species found in 1937 are different from those of 1992/1993. There is a major change in the basin where one agglutinated species has changed its depth distribution downslope and two present day abundant species are new arrivals. These faunal events are probably linked to environmental changes.     

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.     

Benthic foraminifera off West Africa (1°N to 32°S): Do live assemblages from the topmost sediment reliably record environmental variability?

Recent benthic foraminifera (> 125 μm) were investigated from multicorer samples on a latitudinal transect of 20 stations between 1°N and 32°S along the upper slope off West Africa. Samples were selected from a narrow water depth interval, between 1200 and 1500 m, so that changes in water masses are minimized, but changes in surface productivity are important and the only significant environmental variable. Live (Rose Bengal stained) benthic foraminifera were counted from the surface sediment down to a maximum of 12 cm. Dead foraminifera were investigated in the top 5 cm of the sediment only. Five live and five dead benthic foraminiferal assemblages were identified using Q-mode principal component analysis, matching distinct primary productivity provinces, characterized by different systems of seasonal and permanent upwelling. Differences in seasonality, quantity, and quality of food supply are the main controlling parameters on species composition and distribution of the benthic foraminiferal faunas.To test the sensitivity of foraminiferal studies based on the uppermost centimeter of sediment only, a comparative Q-mode principal component analysis was conducted on live and dead foraminiferal data from the top 1 cm of sediment. It has been demonstrated that, on the upper slope off West Africa, most of the environmental signals as recorded by species composition and distribution of the “total” live and dead assemblages, i.e., including live and dead foraminifera from the surface sediment down to 12 cm and 5 cm, respectively, can be extracted from the assemblages in the top centimeter of sediment only. On the contrary, subsurface abundance maxima of live foraminifera and dissolution of empty tests strongly bias quantitative approaches based on the calculation of standing stocks and foraminiferal numbers in the topmost centimeter.     

Spatial Patterns in the Distribution,Diversity and Abundance of Benthic Foraminifera around Moorea (Society Archipelago,French Polynesia)

Coral reefs are now subject to global threats and influences from numerous anthropogenic sources. Foraminifera, a group of unicellular shelled organisms, are excellent indicators of water quality and reef health. Thus we studied a set of samples taken in 1992 to provide a foraminiferal baseline for future studies of environmental change. Our study provides the first island-wide analysis of shallow benthic foraminifera from around Moorea (Society Archipelago). We analyzed the composition, species richness, patterns of distribution and abundance of unstained foraminiferal assemblages from bays, fringing reefs, nearshore and back- and fore-reef environments. A total of 380 taxa of foraminifera were recorded, a number that almost doubles previous species counts. Spatial patterns of foraminiferal assemblages are characterized by numerical abundances of individual taxa, cluster groups and gradients of species richness, as documented by cluster, Fisher α, ternary plot and Principal Component Analyses (PCA). The inner bay inlets are dominated by stress-tolerant, mostly thin-shelled taxa of Bolivina, Bolivinella, Nonionoides, Elongobula, and Ammonia preferring low-oxygen and/or nutrient-rich habitats influenced by coastal factors such as fresh-water runoff and overhanging mangroves. The larger symbiont-bearing foraminifera (Borelis, Amphistegina, Heterostegina, Peneroplis) generally live in the oligotrophic, well-lit back- and fore-reef environments. Amphisteginids and peneroplids were among the few taxa found in the bay environments, probably due to their preferences for phytal substrates and tolerance to moderate levels of eutrophication. The fringing reef environments along the outer bay are characterized by Borelis schlumbergeri, Heterostegina depressa, Textularia spp. and various miliolids which represent a hotspot of diversity within the complex reef-lagoon system of Moorea. The high foraminiferal Fisher α and species richness diversity in outer bay fringing reefs is consistent with the disturbance-mosaic (microhabitat heterogeneity) hypothesis.Calculations of the FORAM Index (FI), a single metric index to assess reef vitality, indicate that all fore- and most back-reef environments support active carbonate accretion and provide habitat suitability for carbonate producers dependent on algal symbiosis. Lowest suitability values were recorded within the innermost bays, an area where natural and increasing anthropogenic influences continue to impact the reefs. The presence of habitat specific assemblages and numerical abundance values of individual taxa show that benthic foraminifera are excellent recorders of environmental perturbations and good indicators useful in modern and ancient ecological and environmental studies.     

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.     

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.     

Holocene trends in distribution and diversity of benthic foraminifera assemblages in atoll lagoons,Belize, Central America

Seven benthic foraminiferal assemblages were identified in vibracores through Holocene lagoons of three Belize atoll lagoons (Glovers Reef, Lighthouse Reef, Turneffe Islands). These include (1) the low-diversity Cribroelphidium assemblage (2) the Cribroelphidium-Elphidium assemblage (3) the Quinqueloculina-Triloculina-Peneroplis assemblage (4) the high-diversity miliolid assemblage (5) the Archaias-miliolid assemblage (6) the low-diversity miliolid assemblage, and (7) the mixed assemblage. Altogether, 109 species and 56 genera were identified. The highest diversities are observed in the largest lagoon (Turneffe Islands), whereas one of the smaller lagoons (Glovers Reef) exhibits the lowest diversities during the Holocene. No significant changes in diversity over time occur, however, a slight trend to higher diversity may be observed through the Holocene, suggesting that the foraminiferal faunas in the atolls are in a diversification stage. Faunal diversity in atoll lagoons appears to be controlled largely by habitat size, habitat heterogeneity, and water circulation. Habitat age and water depth only play minor roles. Substrate texture, water depth, and turbidity influence the predominant modes of life of benthic foraminifera encountered in the lagoons (epifaunal versus infaunal versus symbiont-bearing). Time-averaging effects were not observed, even though lagoonal sedimentation rates fluctuate in individual cores and the three lagoons, and despite the fact that sediments are modified through bioturbation by callianassid shrimps. This finding underlines the potential of benthic foraminifera for paleoecological studies in the fossil record of reefs and carbonate platforms.     

The Foram Stress Index: A new tool for environmental assessment of soft-bottom environments using benthic foraminifera. A case study from the Saronikos Gulf,Greece, Eastern Mediterranean

Saronikos Gulf, including the industrial zone of Elefsis Bay and the Port of Piraeus, is one of the most anthropogenically impacted coastal regions of Greece. Distinct assemblages of benthic foraminifers in sediment samples, collected from this gulf in February 2012, defined three zones that reflect abiotic parameters of the sediments (e.g., organic carbon, metal content). A low-diversity assemblage, dominated by stress-tolerant Ammonia tepida and Bulimina spp., was characteristic of samples from Elefsis Bay. Samples from the western and central part of Saronikos Gulf were the most variable with respect to both abiotic parameters and the foraminiferal assemblage, characterized by a mix of stress-tolerant and more sensitive taxa, especially Bolivina spp. and Nonion fabum. Samples from the coast of Salamis and at the eastern sector of the gulf were characterized by a diverse assemblage that included Peneroplis pertusus, miliolids, and a variety of small, epiphytic rotaliid taxa. A new biotic index, the Foram Stress Index (FSI), is based on the relative percentages of two ecological groups of benthic foraminiferal species, grouped according to their tolerance/sensitivity to organic matter enrichment and weighted proportionately to obtain a formula to define five ecological-status classes. The FSI produced three rankings for these samples (Poor, Moderate and Good), that strongly correlate with the macroinvertebrate-classification tool known as the BENTIX Index. The FSI provides a new tool to assess sediment or substrata quality based upon the benthic foraminiferal assemblages, which are a significant component of living meiobenthic communities that are generally not considered in most biotic benthic indices.     

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.     

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.     

Late Pliocene benthic foraminifera and mollusks from the Atsipades Section, Central Crete; Palaeoecological distribution and use in palaeoenvironmental assessment

Changes in benthic foraminiferal and mollusk assemblages from the lower part of the Upper Pliocene of the Atsipades Section (Iraklion Basin, central Crete) were studied. The Atsipades Section represents a shallowing-upward sequence from outer shelf blue-gray clays at the bottom of the sequence, deposited below the storm wave base, to shallow inner shelf deposits affected by storm waves at the top. The foraminiferal assemblage at the bottom of the sequence is dominated by Bolivina spathulata, Bolivina dilatata and Uvigerinidae, a microfossil assemblage corresponding to the deepest deposits formed under dysoxic sea-floor conditions. Foraminiferal assemblages of the middle part of the section are highly diversified, predominantly Haynesina depressula, Cassidulina carinata and Reusella spinulosa. The top of the section is mainly characterised by Asterigerinata planorbis, Bolivina pseudoplicata, Cibicides lobatulus and Elphidium sp., a typically epiphytic foraminiferal assemblage which can be correlated with the presence of an algal covered sea-bottom. Within this general environmental trend, a minor shallowing cycle can be differentiated. The boundaries of this cycle can be inferred, based on a substantial microfossil assemblage change and on the coincidence of species diversity maximum and a planktonic/benthic (P/P + B) ratio peak. Nonetheless, upwelling currents and/or over-abundance of nutrients due to continental outflow could also contribute to increased diversity and P/P + B ratio. The character of the mollusk assemblages is in accordance with these trends. Moreover, the increase in diversity and in sculpture constitutes a clear indication of an increase in hydrodynamic energy related to a shallowing-upward trend.     

The surface ocean productivity response of deeper water benthic foraminifera in the Atlantic Ocean

We test the relationship of deep sea benthic foraminiferal assemblage composition to the surface ocean productivity gradient in the low latitude Atlantic Ocean using 81 surface sediment samples from a water depth range between 2800 and 3500 m. The samples are selected so that the surface ocean productivity gradient, controlling the flux of organic carbon to the seabed, will be the most important environmental variable. The first two principal components of the assemblage data account for 73% of data variance and are clearly linked to the productivity gradient across the Atlantic. These components show that under higher productivity the assemblages contain a higher abundance of Uvigerina peregrina, Melonis barleeanum, Globobulimina spp. and other taxa with probable infaunal microhabitats. Alabaminella weddellensis, a species linked to episodic phytoplankton debris falls, is also important in these assemblages. As productivity decreases there is a regular shift in assemblage composition so that low productivity assemblages are dominated by Globocassidulina subglobosa and several Cassidulina species along with Epistominella exigua. We hypothesize that these taxa are epifaunal to very shallow infaunal since nearly all organic carbon oxidation occurs near the sediment-water interface in low productivity settings. Discriminant function analysis of the foraminiferal assemblages, with groups selected on the basis of surface ocean productivity, shows clear separation among five productivity levels we used. This analysis demonstrates that productivity variations have a strong influence on assemblage composition. Finally, we used two groups of samples from the Rio-Grande Rise representing water depths from 2007 to 2340 m and 2739 to 3454 m to test for effects produced by changing water depth. All these samples are from a low productivity region and represent nearly identical environmental conditions. Although the low productivity nature of all the Rio-Grande Rise samples is obvious, there are assemblage differences between our depth groups. We cannot account for the assemblage differences with changes in organic carbon flux, dissolution effects or other physical/chemical properties of the ocean. Thus there are as yet unidentified factors related to water depth which cause some assemblage variation in the low productivity setting we investigated.