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.     

Live benthic foraminiferal faunas along a bathymetrical transect (140-4800 m) in the Bay of Biscay (NE Atlantic)

In a 10-stations bathymetrical transect in the Bay of Biscay, we observed important changes in the density, composition and microhabitats of live foraminiferal faunas from the outer continental shelf to the abyssal plain. Four zones are recognised: (1) at the upper continental shelf (140 m water depth), foraminiferal densities are very high and the superficial sediment is occupied by Bolivina subaenariensis and Valvulineria bradyana. Globobulimina spp., Chilostomella oolina and Nonion fabum dominate the infaunal niches, which are positioned close to the sediment-water interface due to a strong compaction of the vertical succession of redox zones. (2) At the upper continental slope stations (300-1000 m), foraminiferal densities are high and the superficial sediments are dominated by Uvigerina mediterranea/peregrina. Deeper in the sediment, intermediate infaunal niches are occupied by Melonis barleeanus. Due to a deeper oxygen penetration, the deep infaunal taxa Globobulimina spp. and C. oolina live at a considerable depth in the sediment. (3) At the mid and lower slope stations (1000-2000 m) in the superficial sediment Cibicidoides kullenbergi and Hoeglundina elegans progressively replace U. mediterranea. U. peregrina is still a dominant taxon, reflecting its preference for a somewhat intermediate organic flux level. Deep infaunal taxa become increasingly rare. (4) At the lower slope and abyssal plane stations (deeper than 2000 m), faunal densities are very low and the fauna is composed exclusively by shallow infaunal species, such as Nuttallides umboniferus and Melonis pompilioides. The foraminiferal data together with the pore water data in the sediment give evidence of the presence of a trophic gradient from very eutrophic settings at the upper continental shelf towards oligotrophic settings at the abyssal area.     

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.     

Taphonomy of fossil macro-invertebrate assemblages as a tool for ecostratigraphic interpretation in Upper Jurassic shelf deposits (Prebetic Zone, southern Spain)

Composition and taphonomy of macro-invertebrate fossil assemblages, together with facies analysis, have been approached in order to interpret shifting paleoenvironmental conditions in the External Prebetic (S-SE Spain) during the early Late Jurassic (Middle Oxfordian). In oolitic and spongiolitic limestones, the size of fossil remains, mode of preservation, within-bed position, corrasion, fragmentation, epibiont and biogenic encrustation, disarticulation and uncoupling, allow recognition of two taphofacies, respectively. Identified ecostratigraphic events and trends accord with rapid flooding under high-energy conditions related to ecospace enlargement for cephalopods and then the persistence of lower energy, long-lasting exposure of skeletals and higher sedimentary rates. The paleoenvironmental interpretation is consistent with neritic environments shifting from shallow carbonate to hemipelagic sedimentation and enlarging of shelf ecospace for marine invertebrates.     

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.     

Stable isotope composition of Late Cretaceous benthic foraminifera from the southern South Atlantic: Biological and environmental effects

The stable carbon and oxygen isotope composition of different benthic foraminiferal species of the latest Campanian and earliest Maastrichtian from Ocean Drilling Project Hole 690C (Weddell Sea, southern South Atlantic, ∼1800 m paleowater depth) have been investigated. The total range of measured isotope values of all samples exceeds ∼4‰ for δ¹³C and 1.1‰ for δ¹⁸O. Carbon isotope values of proposed deep infaunal species are generally similar or only slightly lower when compared to proposed epifaunal to shallow infaunal species. Interspecific differences vary between samples probably reflecting temporal changes in organic carbon fluxes to the sea floor. Constantly lower δ¹³C values for Pullenia marssoni and Pullenia reussi suggest the deepest habitat for these species. The strong depletion of δ¹³C values by up to 3‰ within lenticulinids may be attributed to a deep infaunal microhabitat, strong vital effects, or different feeding strategy when compared to other species or modern lenticulinids. The mean δ¹⁸O values reveal a strong separation of epifaunal to shallow infaunal and deep infaunal species. Epifaunal to shallow infaunal species are characterized by low δ¹⁸O values, deep infaunal species by higher values. This result possibly reflects lower metabolic rates and longer life cycles of deep infaunal species or the operating of a pore water [CO₃²⁻] effect on the benthic foraminiferal stable isotopes.Pyramidina szajnochae shows an enrichment of oxygen isotopes with test size comprising a total of 0.6‰ between 250 and 1250 μm shell size. Although δ¹³C lacks a corresponding trend these data likely represent the presence of changes in metabolic rates during ontogenesis. These results demonstrate the general applicability of multi-species stable isotope measurements of pristine Cretaceous benthic foraminifera to reconstruct past microhabitats and to evaluate biological and environmental effects on the stable isotope composition.     

Microbial encrustations from the Middle Oxfordian-earliest Kimmeridgian lithofacies in the Prebetic Zone (Betic Cordillera, southern Spain): characterization, distribution and controlling factors

Two main types of microbial encrustation were identified in Middle Oxfordian to lowermost Kimmeridgian deposits in the Prebetic Zone (southern Spain), showing existing relationships between skeletal content, fabric and morphology of these organosedimentary structures. Laminated planar and concentric encrustations relate to peloidal fabrics (mainly constituted of microbes = microbial laminated fabrics s. str. and microbial oncoids s. str.), as well as to dense microbial fabrics periodically colonized by encrusting foraminifera (microbial laminated fabrics with nubeculariids and microbial oncoids with nubeculariids). Sedimentation rates, substrate stability and grain size, as well as illumination, influenced microbial growth pattern as major controlling factors in low-energy conditions, and forced palaeogeographic and stratigraphic patterns of distribution. Significant encrustation was identified in terrigenous-poor lithofacies from the middle (Transversarium-Bifurcatus zones) to the outer (Transversarium-Bimammatum zones) shelf in the Prebetic Zone. Rare-to-absent encrustation characterized terrigenous-rich deposits (Bimammatum and Planula zones) in the area.     

Palaeogeographic and stratigraphic distribution of mid-late Oxfordian foraminiferal assemblages in the Prebetic Zone (Betic Cordillera, Southern Spain)

Middle-Upper Oxfordian assemblages of foraminifera in the Prebetic Zone (Betic Cordillera, SE Spain) were analysed at the genus level to determine their composition, relative abundance, diversity, and dominance, as well as the size of the specimens. A relationship has been established between lithofacies, palaeogeography and composition of foraminiferal assemblages, the former two also determining the stratigraphic record of these microfossil assemblages. Two assemblages of foraminifera serve to identify relatively distal and proximal areas in the Prebetic shelf. The distal assemblage is characterized by higher diversity, specimens of greater size, and more abundant planktic and agglutinated forms. Benthic forms include Ophthalmidium, Epistomina and colonies of encrusting foraminifera. The proximal assemblage shows lower diversity, lower abundance of planktic forms, Epistomina and encrusting nubeculariids, and a greater abundance of spirillinids and Reofax. On the whole, planktic foraminifera decrease upwards in the studied succession, which, together with decreasing nodularity, could be related to system tract conditions previously proposed for Oxfordian deposits in the southern palaeomargin of Iberia.     

A conceptual model explaining benthic foraminiferal microhabitats

We present a conceptual model which explains benthic foraminiferal microhabitat preferences in terms of differences in the downward organic flux. We argue that under oligotrophic conditions the microhabitat depth is controlled by the availability of metabolizable food particles in the sediment. Under more eutrophic conditions, the ecosystem is no longer food-controlled, but instead, a critical oxygen level determines down to what depth we find a living fauna. Under food-limited conditions, anaerobic degradation of organic matter may provide an additional food source around the redox front, which could explain deep infaunal maxima reported in the literature. In a sample transect through the Adriatic Sea, both microhabitat controls (food-limited and oxygen-limited) are present. On the shelf and the upper part of the slope, the rather shallow depth of the microhabitat is controlled by a critical oxygen level. In the 1250 m deep southern basin and on the lower part of the slope, on the contrary, the availability of metabolizable organic matter, and not a critical oxygen level, determines down to what depth living foraminifera are found.     

Stable oxygen and carbon isotopes of live benthic foraminifera from the Bay of Biscay: Microhabitat impact and seasonal variability

We determined the stable oxygen and carbon isotopic composition of live (Rose Bengal stained) benthic foraminifera (> 150 μm size fraction) of seven taxa sampled along a downslope transect between 140 to 2000 m water depth in the Bay of Biscay. At the five stations, Hoeglundina elegans, Cibicidoides pachydermus, Uvigerina peregrina, Uvigerina mediterranea preferentially occupy shallow infaunal niches, whereas Melonis barleeanus and Uvigerina elongatastriata occupy an intermediate infaunal microhabitat, and Globobulimina spp. live in a deep infaunal niche close to the zero oxygen boundary.When compared with δ¹⁸O values of calcite formed in equilibrium with bottom waters, U. peregrina forms its test in close equilibrium with bottom water δ¹⁸O. All other foraminiferal taxa calcify with a constant offset to calculated equilibrium calcite. There is no systematic relationship between the foraminiferal microhabitat depth and the Δδ¹⁸O between foraminiferal and equilibrium calcite. We calculated correcting factors for the various taxa, which are needed for constructing multispecies-based oxygen isotope records in paleoceanographic studies of the study area.The δ¹³C values of foraminiferal taxa investigated in this study do neither record bottom water δ¹³C_DIC in a 1 : 1 relationship nor with a constant offset, but appear to be mainly controlled by microhabitat effects. The increase of δ¹³C values of shallow infaunal taxa with increasing water depth reflects the decrease of the exported flux of organic carbon along the bathymetric transect and early diagenetic processes in the surface sediment. This is particularly the case for the shallow infaunal U. peregrina. The δ¹³C values of deep infaunal Globobulimina spp. are much less dependent on the exported organic matter flux. We suggest that the Δδ¹³C between U. peregrina and Globobulimina spp. can shed light on the various pathways of past degradation of organic detritus in the benthic environments.At a station in 550 m water depth, where periodic eutrophication of sediment surface niches was demonstrated previously, we performed a two-year seasonal survey of the isotopic composition of foraminiferal faunas. No marked seasonal changes of the stable carbon isotopic composition of shallow, intermediate and deep infaunal foraminiferal taxa were observed. Thus, the δ¹³C values of foraminiferal individuals belonging to the > 150 μm fraction may result from rather long-term calcification processes lasting for several weeks or months, which limit the impact of ephemeral ¹²C enrichment of shallow infaunal niches on the isotope chemistry of adult individuals during eutrophic periods. Only highly opportunistic taxa reproducing or calcifying during phytoplankton bloom periods and the subsequent deposits of phytoplankton remains in the benthic environment may exhibit a particularly low δ¹³C, indicative of such short productive periods.     

Regional and interannual productivity of biogenic components and planktonic foraminiferal fluxes in the northwestern Pacific Basin

Time-series sediment trap experiments at subtropical (WCT-1) and subarctic (WCT-2) stations in the northwestern Pacific indicate seasonal, latitudinal and depth variations in total particulate, biogenic and foraminiferal fluxes. At the subtropical station, the average total mass flux was 19.4 mg m⁻² day⁻¹ in the shallow trap (1060 m) and 21.5–26.1 mg m⁻² day⁻¹ in the deep trap (3930 m) during the sampling period. At subarctic station, these values were 91.5–176.9 mg m⁻² day⁻¹ in the shallow and 68.6–112.3 mg m⁻² day⁻¹ in the deep trap. We recognized 12 and 15 planktonic foraminiferal species at Station WCT-1 and Station WCT-2, respectively. The planktonic foraminiferal flux and species turnover are related to seasonal and interannual changes in source water and water column conditions at both stations. At Station WCT-1, the highest flux was recorded during the summer, with a peak in mid to late June associated with similar flux patterns of the dominant species, Globigerinoides ruber and Globigerinita glutinata. The total flux of foraminiferal tests at the shallow and deep traps is similar in numbers and magnitude. At Station WCT-2, the peaks of total flux of foraminiferal tests at the two trap depths differ in number, and their magnitude in the deep trap is almost half of that in the shallow trap. A distinctive seasonal pattern occurred in the shallow and the deep trap, with a peak in total foraminiferal flux in mid June to mid July. Globigerina quinqueloba, Neogloboquadrina pachyderma and Neogloboquadrina dutertrei dominate the planktonic population throughout the year.Subtropical Station WCT-1 was characterized by low total foraminiferal fluxes and low total mass flux, which is dominated by calcium carbonate and depleted in opal, whereas high foraminiferal fluxes and a high total mass flux dominated by high biogenic opal, and less calcium carbonate and organic matter characterize subarctic Station WCT-2. The foraminiferal carbonate that reaches the seafloor accounts for an average 20–27% and 22–23% of the total calcium carbonate at Station WCT-1 and Station WCT-2, respectively. The primary reason for the difference in flux at both stations thus lies in the different contributions of siliceous and calcareous planktonic assemblages. The seasonal variation in biogenic particulate flux at both stations implies that temporal changes in biological productivity are governed by large-scale seasonal climatic variability and local hydrography.     

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.     

Biostratigraphy and paleoecology of the Oligo-Miocene succession in Fars and Khuzestan areas (Zagros Basin,SW Iran)

Paleontological and biostratigraphical studies on carbonate platform succession from southwest Iran documented a great diversity of shallow-water benthic foraminifera during the Oligocene–Miocene. Larger foraminifera are the main means for the stratigraphic zonation of carbonate sediments. The distributions of larger benthic foraminifera in two outcrop sections (Abolhayat and Lali) in the Zagros Basin, Iran, are used to determine the age of the Asmari Formation. Four assemblage zones have been recognized by distribution of the larger benthic foraminifera in the study areas. Assemblage 3 (Aquitanian age) and 4 (Burdigalian age) have not been recognized in the Abolhayat section (Fars area), due to sea-level fall. The end Chattian sea-level fall restricted marine deposition in the Abolhayat section and Asmari Formation replaced laterally by the Gachsaran Formation. This suggests that the Miocene part of the formation as recognized in the Lali section (Khuzestan area) of the Zagros foreland basin is not present in the Abolhayat outcrop. The distribution of the Oligocene larger benthic foraminifera indicates that shallow marine carbonate sediments of the Asmari Formation at the study areas have been deposited in the photic zone of tropical to subtropical oceans. Based on analysis of larger benthic foraminiferal assemblages and microfacies features, three major depositional environments are identified. These include inner shelf, middle shelf and outer shelf. The inner shelf facies is characterized by wackestone–packstone, dominated by various taxa of imperforate foraminifera. The middle shelf is represented by packstone–grainstone to floatstone with a diverse assemblage of larger foraminifera with perforate wall. Basinwards is dominated by argillaceous wackestone characterized by planktonic foraminifera and large and flat nummulitidae and lepidocyclinidae. Planktonic foraminifera wackestone is the dominant facies in the outer shelf.     

Morphology and microhabitat preferences of benthic foraminifera from the northwest Atlantic Ocean

The distribution of Rose Bengal stained calcareous benthic foraminifera was determined in six ☐ cores raised from water depths between 200 and 3000 m on the Nova Scotian continental margin and Gulf of Maine. The taxa can be separated into four microhabitats within the surficial sediments. Epifaunal taxa are generally found in the top cm, intermediate infaunal taxa are found from about 1 to 4 cm and deep infaunal taxa are found at > 4 cm sediment depth in at least one ☐ core. A fourth group, shallow infaunal taxa, is found in the top 2 cm and is inferred to be infaunal based on wall porosity characteristics and test shapes similar to infaunal taxa. The epifaunal, shallow infaunal and intermediate infaunal taxa maintain their positions within the sediments from core to core, whereas the deep infaunal taxa are found at progressively shallower sediment depths in cores within increasing organic carbon contents from shallower water depths.Each microhabitat category has distinct morphological characteristics. Epifaunal taxa have plano-covex or biconvex cross sections, trochospiral coiling and large pores absent or found on only one side. Shallow infaunal taxa have uniserial, triserial, or planispiral coiling, with surface ornamentation present on a number of taxa. The intermediate infaunal taxa have rounded peripheries, pores over the entire test and planispiral coiling, with the exception ofCibicidoides bradyi which has trochospiral coiling. The deep infaunal taxa have, in general, planispiral or triserial coiling with cylindrical or ovate shaped tests.     

Factors influencing the distribution of Subantarctic deep-sea benthic foraminifera,Campbell and Bounty Plateaux,New Zealand

We investigate the combination of environmental factors that influence the distribution patterns of benthic foraminiferal tests (> 63 μm) in a topographically varied region crossed by both the Subtropical and Subantarctic Fronts, south-east of New Zealand. Seafloor sample sites, extending from outer shelf (50 m) to abyssal (5000 m) depths, are bathed by five different water masses, and receive phytodetritus from Subtropical, Subantarctic and Circumpolar surface water masses. Eight mappable associations are recognised by Q-mode cluster analysis of the benthic foraminiferal census data. Similar associations are identified using cluster analysis based solely on the presence or absence of species. Canonical correspondence analysis and a correlation coefficient matrix were used to relate the faunal data to a set of environmental proxies. These show that factors related to water depth (especially decreasing food supply with increasing depth) are the most significant in determining the overall foraminiferal distribution. Other contributing factors include surface water productivity and its seasonality; bottom water ventilation; energetic state of the benthic boundary layer and resulting substrate texture; and bottom water carbonate corrosiveness. Three shallow-water associations (50–700 m), dominated by Cassidulina carinata, Trifarina angulosa, Globocassidulina canalisuturata, Gavelinopsis praegeri, and Bolivina robusta, occur in coarse substrates on the continental shelf, and on the crests and upper slopes of four seamounts under well-oxygenated, high energy regimes, and high food input. Three mid bathyal to upper abyssal associations (500–3300 m), dominated by Alabaminella weddellensis, C. carinata, and Epistominella exigua, occur in biopelagic sandy mud, beneath a region of strongly seasonal food supply, with their composition influenced by total food flux, ventilation (Oxygen Minimum Zone), and bottom current strength. An unusual lower bathyal association (1200–2100 m), dominated by T. angulosa and Ehrenbergina glabra, occurs in a belt of coarser sandy substrate that runs along the crest of the submarine plateaux slopes beneath the strongly-flowing Subantarctic Front-related currents. A deep abyssal association (3500–5000 m), dominated by Nuttallides umbonifer and Globocassidulina subglobosa, occurs on the abyssal plain beneath oligotrophic lower Circumpolar Water south-east of the Subantarctic Front and is strongly influenced by the cold, carbonate-corrosive conditions.     

Distribution of gravel-sized empty tests of large benthic foraminifers as practical depositional indicators in tropical reef and shelf carbonate environments

To explore the utility of gravel-sized tests of large benthic foraminifers (LBFs) as practical paleoenvironmental indicators of tropical reef and shelf carbonate environments, depth and spatial distributions of gravel-sized empty tests of LBFs were examined using 39 surface sediment samples collected from depths shallower than 200 m off the west coast of Miyako Island (Ryukyu Islands, northwest Pacific). Distributions of the LBF tests were mainly related to water depth, topography, and substrate type. Q-mode cluster analysis based on the binary (presence/absence) data of LBF associations (4–2-mm size fraction) clearly delineates four depositional environments: bay, back reef to fore reef, flat shelf, and shelf slope. Application of this modern dataset to fossil LBF data from larger foraminiferal limestones of the Pleistocene Ryukyu Group indicate that a test section was deposited in an outer flat shelf at depths between 54 and 99 m. Comparisons of these results with previous reports suggest that our foraminiferal analysis using gravel-sized tests is methodologically easier than conventional analyses including smaller sized tests to distinguish similar levels of depositional environments. However, taxonomic and environmental similarities make the applicability of this dataset to fossil LBF data from Quaternary tropical carbonate environments in the northwest Pacific.     

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

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

Middle-Upper Jurassic Foraminifera from Jumara Hills, Kutch, India

A prolific foraminiferal assemblage comprising 51 species is reported from Jurassic sediments of the Chari Formation, Jumara Hills, Kutch, India. The assemblage is dominated by the families Vaginulinidae and Nodosariidae. Sixteen species are reported for the first time from the Indian region including one new species. The foraminiferal assemblage suggests a Callovian to Oxfordian age for the studied sequence. The depositional environment of the studied sequence is interpreted based on foraminiferal evidence combined with lithology and megafauna which indicates that the sediments of the Chari formation exposed at Jumara Hills were deposited in a shallow-water, near shore, environment with fluctuating shoreline in a tectonically unstable shelf zone. The Jumara Hills foraminiferal assemblage exhibits close affinity with certain other Jurassic assemblages of the Tethyan Realm and helped in drawing palaeogeographic conclusions, suggesting that during the Middle and Late Jurassic epochs Kutch had close sea connections with Central Arabia, Afghanistan and Rajasthan on the one hand and Somalia and Malagasy on the other.     

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.     

The life habits of the Ordovician illaenine trilobite Bumastoides

The Ordovician illaenine Bumastoides exhibits a distinctive effaced and strongly convex morphology. Orientation of the visual field, the extreme convexity of the cephalon and the nature of the thoracic articulation support an interpretation of Bumastoides as an infaunal trilobite that was poorly suited to epifaunal crawling. The genus may have been sedentary; spending most of its post-larval life cycle within a burrow. Suspension feeding would be a viable existence for a sedentary trilobite such as Bumastoides. Maintenance of a burrow is essential for respiration and would require a firm, cohesive substrate. The infaunal niche had become occupied by trilobites by at least the Late Cambrian and continued to be exploited through the Ordovician, Silurian and. possibly, into the Devonian. Convergence has led to the appearance of the effaced, strongly convex morphotype in a number of unrelated families, including the Illaenidae, Asaphidac, Aulacopleuridae, Plethopeltidae and Scutelluidae. A high numerical abundance of illaenid trilobites, such as Illaenus and Bumastoides , is characteristic of the illaenid–chcirurid association, which persisted from the early Ordovician until at least the Late Silurian. This association has been recorded from shelf-edge and on-shelf carbonate buildups and shallow subtidal level bottom environments. It appears to be confined to firm substrates.