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.     

Microtaphonomy of bioclasts and paleoecology of microencrusters from Upper Jurassic spongiolithic limestones (External Prebetic,Southern Spain)

Microencrusters and microtaphonomic features of the Oxfordian spongiolithic limestones of the External Prebetic were studied using thin-section analysis. The spongiolithic limestone is a bioclastic-rich packstone with common echinoderm, mollusc and brachiopod remains. The bioclasts show a high fragmentation index and frequent microborings. The encrustation index (E _i) is higher for fragments of serpulids, ammonoids and bivalves, and increases with the initial grain-size of bioclasts. The main microencrusters consist of benthic microbial communities (BMC) and nubeculariids, as well as subordinate calcareous and siliceous agglutinated foraminifera, serpulids and bryozoans. BMC are usually the first colonizers, and encrusting foraminifera mainly appear on bioclasts larger than 2 mm. BMC dominate in well-developed encrustations on upward facing surfaces of larger bioclasts that are also colonized by foraminifera (nubeculariids and Subdelloidina). Bullopora, serpulids and bryozoans are more common on lower surfaces. The fact that the values of encrustation index, encrustation thickness and diversity of the microencrusters increase with the size of bioclasts is related to a higher stability and exposure time of the available bioclastic substrate. The microencruster distribution on upper and lower surfaces of large bioclasts may be related to photic control, space competition and/or predation avoidance.     

Larger foraminifera as a substratum for encrusting bryozoans (Late Oligocene,Tethyan Seaway,Iran)

Considering the diversity and abundance of larger foraminifera examined from a wide range of Late Oligocene to Early Miocene palaeoenvironments in the Tethyan Seaway, encrusting bryozoans make extremely little use of their tests as substratum. Significant encrustations by bryozoans were exclusively found on large (ø c. 6 cm), undulating tests of Lepidocyclina spp., on which, however, a remarkable 34 taxa of encrusting bryozoans were recorded. This shallow-water fauna of Chattian age was analyzed in respect of the bryozoan taxa present, colony growth type, and mode of budding, colony size, as well as onset of reproduction. Taxic and morphological similarities between the fossil assemblage and modern faunas encrusting mobile substrata indicate a long history of bryozoans as part of the interstitial habitat, while the tests of certain larger foraminifera may have played a significant role in the evolution of shallow-water bryozoans by providing substrata for encrusting species in otherwise unfavorable environments.     

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.     

Biodiversity and community structure of Late Pleistocene foraminifera from Kish Island, Persian Gulf (Iran)

The raised coral reef sequences at Kish Island provide a rare window into the depositional setting and paleoenvironment of a high-latitude, shallow-water coral reef that developed under turbid conditions in the Persian Gulf during Marine Isotope Stage 7 (~200 to 250?ka). Six sedimentary facies and eight foraminiferal assemblages can be identified throughout the sequence. A ninth assemblage can be defined for the modern subtidal realm. At the base of the sequence is a marl rich in hyaline foraminifera (Elphidium, Ammonia, Asterorotalia, Bulimina, Nonion, and Quinqueloculina) and ostracods, which was deposited in about 30–40?m water depth in a turbid deltaic setting. Shallowing resulted in the marl becoming sandy, and changing to a mollusc-rich facies with rare foraminifera (mostly smaller miliolid taxa) that formed the substrate for coral recruitment. The coral marl layer contains many large corals embedded in situ in an aggregate and coralline algae-rich marl. Two abundance peaks in the foraminifera occur at the base and mid-way through this layer, which also correspond to a change from Murrayinella-dominated to Placopsilina-dominated assemblages, indicating deepening and more open-marine conditions, but elevated turbidity. Towards the top of the layer, abundance of foraminifera decreases and miliolid foraminifera become dominant. The top-most layer is dominated by coral and mollusc fragments and has an Amphistegina-rich reef-related assemblage. Of the Late Pleistocene foraminiferal assemblages, the Murrayinella-, Pararotalia-, and Placopsilina-dominated assemblages are no longer present in the modern gulf for unknown reasons. Of the other five assemblages, only the Amphistegina assemblage is found within proximity to the modern Kish Island. The Elphidium and Asterorotalia-Bulimina assemblages are from deeper areas of the gulf. The Ammonia and Quinqueloculina assemblages occur in lagoonal sediments on the Arabian side of the gulf. Like the modern Persian Gulf, the diversity of foraminifera was low (~80 common species) during the Pleistocene and does not correlate with foraminiferal abundance.     

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.     

Age of the latest marine sedimentation in the western Kunlun area constrained by planktic foraminifera

Surface uplift of the Tibetan Plateau in the Cenozoic has dramatically affected evolutions of climate, environment, and biota in East Asia. The onset time of plateau uplift can be roughly constrained by the termination of the latest marine sedimentation, which shows large spatial differences in the Tibetan Plateau. In the northwest Tibetan Plateau of the western Kunlun area, the Tielongtan Group represents the latest marine sedimentation. However, the age of its depositional cessation is still unknown. For the first time, we conduct taxonomic, paleoecologic and paleogeographic studies of planktic foraminifera in the limestones of the Tielongtan Group. Despite poor preservation and low abundance, ten planktic foraminiferal species are identified, including Globotruncana falsostuarti, G. cf. linneiana, G. cf. ventricosa, Planohedbergella prairiehillensis, P. cf. yaucoensis, Pseudotextularia nuttalli, Planoheterohelix globulosa, P. cf. praenuttalli, Muricohedbergella holmdelensis and Whiteinella brittnensis. The foraminiferal assemblages suggest an age of late Campanian to Maastrichtian, indicating a shallow, neritic marine environment. Combining available research data from other regions, we show that these planktic foraminifera were widely distributed in the Tarim basin, southern Tibet, and the Mediterranean area during the latest Cretaceous. Our result of planktic foraminifera in the Tielongtan Group implies that the surface uplift of the western Kunlun area did not begin until after ∼76–66 Ma.     

Planktonic versus benthic foraminifera response to Milankovitch forcing (Late Jurassic,Betic Cordillera): testing methods for cyclostratigraphic analysis

Foraminiferal assemblages from the Bifurcatus Zone (Oxfordian, Upper Jurassic) are studied in the Navalperal section (Betic Cordillera, southern Spain). A total of 24 sampling stations and around 5,700 specimens of foraminifera were recognized on thin-section analysis and classified into two major categories: planktonic and benthic. The abundance of foraminifera (number of specimens/cm²) was calculated for both categories and the cyclic pattern was studied by spectral analysis, considering the autochthonous and para-autochthonous character of the studied assemblages. The Lomb-Scargle periodogram together with a permutation test were tested for performing the high-resolution spectral analysis, being particularly well suited for working with short time series and uneven sampling. Spectral analysis reveals the influence of orbital-scale Milankovitch cyclicity at the eccentricity, obliquity, and precession bands. Moreover, this incidence is significantly different depending on the analyzed group (benthic versus planktonic). Whereas the long-range eccentricity band is not distinguishable from a trend and the short-range eccentricity band is not statistically significant (at 90% confidence level), the obliquity band is better represented in the planktonic component and the precession band is better developed in the benthic group. Variations in temperature affecting upper waters, determined by obliquity-scale fluctuations, could be responsible for changes in the planktonic foraminiferal assemblage, while changes in nutrient availability and substrate oxygenation, as a consequence of input variations from source areas at the precession-scale cycles, could affect the benthic foraminiferal assemblage.     

Modern foraminiferal distribution and diversity in two atolls from the Maldives,Indian Ocean

Foraminifera from the coral-reef lagoons of two atolls are used to investigate the distribution and diversity of assemblages in the central Indian Ocean. Eight assemblages and 270 species of foraminifera are identified. Three assemblages are reefal and dominated by Amphistegina and Calcarina. Only one lagoon assemblage is present in both atolls, which is characterized by abundant Ammonia sp. 1 and smaller miliolid foraminifera. In Ari Atoll, abundant Amphistegina sp. 1, Operculina ammonoides, Amphistegina radiata and Nummulites venosa characterize one lagoon assemblage and abundant Neoeponides bradyi and Textularia cf. T. cushmani characterize the other. In Rasdhoo Atoll, abundant Textularia gr. foliacea and Spiroloculina nummiformis characterize one lagoon assemblage and abundant Textularia cf. T. cushmani and Textularia sp. 5 characterize the other. The assemblages compare well with sedimentological characteristics, which is also reflected in their distributions. Species diversity in the Maldives appears to be higher than the western Indian Ocean region due to the stronger influence of central Indo-Pacific faunas. It is not however, as high as the central Indo-Pacific region, which may be due to lower habitat diversity.     

Spatial distribution of Late Pliensbachian (Early Jurassic) calcareous nannofossils within the Lusitanian basin (Portugal)

Quantitative analyses of Pliensbachian calcareous nannofossils have been carried out on a proximal-distal transect in the Lusitanian Basin (Portugal). The studied sections Vale Venteiro near Tomar and Peniche represent proximal and distal environments with respect to the emerged land of the Iberian Meseta (to the East). The upper portion of the Vale das Fontes Fm (Davoei and Margaritatus ammonite Zones) is studied in both sections and correlated by means of ammonite and nannofossil integrated biostratigraphy. A careful analysis of the preservation state of nannofossils is performed. Preservation state is moderate to good in the two settings; changes in nannofossil assemblages are therefore considered as primary. Samples were analysed for nannofossil absolute and relative abundances, species diversity and wt%CaCO₃. The analysis of calcareous nannofossils (absolute abundance, percentage, average abundance) shows changes in the community structure in space (i.e., from proximal to distal), and vertically within the water column. This pattern suggests a partitioning of habitats within the photic zone, and with respect to emerged lands. Proximal environments within the Lusitanian Basin were probably more eutrophic, because of the proximity of emerged lands to the East (Iberian Meseta). This was the source area for nutrients delivered into the basin by river transport. Distal environments were likely characterized by a greater water depth and by a deeper light penetration leading to a relatively more expanded photic zone. The assemblages recorded in the proximal setting exhibit high mean relative abundance of placolith-bearing coccolithophorids (Lotharingius spp., Biscutum spp. and Similiscutum spp.) and of Schizosphaerella spp., while distal assemblages are dominated by Crepidolithus crassus and Schizospharella spp. Crepidolithus crassus is interpreted as a deep-dweller coccolithophorid, inhabiting preferentially distal and deeper settings in the Lusitanian Basin. Placolith-bearing coccolithophorids were more abundant in proximal settings with respect to emerged lands, under relatively elevated trophic conditions. The probable calcareous dinocyst Schizospharella spp. proliferated in surface waters of both proximal and distal environments.     

Cyclic changes in Turonian to Coniacian planktic foraminiferal assemblages from the tropical Atlantic Ocean

Abundance patterns of planktic and benthic foraminifera from a tropical Atlantic drill site (Ocean Drilling Program Site 1259, Demerara Rise, Suriname margin) display a pronounced 400 kyr cyclicity, uninterrupted throughout our  87.8–92 Ma record, between two clearly distinguishable assemblages: (1) a pelagic foraminifer fauna, which represents a deep oxygen minimum zone, and (2) another assemblage representing a shallow oxygen minimum zone where the foraminifer fauna is dominated by a higher diversity population of mostly small clavate and biserial species common in epicontinental seas. The cyclic changes in the long eccentricity band (400 kyr) between these two assemblages are proposed to reflect changes in the mean latitudinal position of the Intertropical Convergence Zone (ITCZ). Associated fluctuations in precipitation and trade wind strength may have influenced the upwelling regime at Demerara Rise leading to the observed cyclicity of planktic foraminiferal assemblages. The severe Turonian to Coniacian paleoclimatic and paleoceanographic changes in the Atlantic Ocean (e.g., gateway opening, cooling, and glaciation), however, seem to have no influence on the composition of tropical planktic foraminiferal faunas. There is no apparent relationship between foraminifer abundances and a major deflection in the stable isotope record interpreted elsewhere as a sign of the growth and decay of a large polar ice sheet.     

Paleoenvironmental controls on the morphology and abundance of the coccolith Watznaueria britannica (Late Jurassic,southern Germany)

The coccolithophore species Watznaueria britannica is dominant in Middle-Upper Jurassic calcareous nannofossil assemblages and presents morphological variation, including different coccolith size, shape and length of the central area and of the bridge. Six morphotypes can be recognized in the polarizing light microscope. The aim of this work is to better understand the morphological variability of W. britannica and determine if this variability is controlled by paleoecological factors. In order to investigate the potential paleoecological controls on W. britannica morphology and abundance, we carried out a biometric study on a restricted temporal interval: the Late Oxfordian, in the Swabian Alb (southern Germany), characterized by increasing carbonate production linked to climatic changes. The Balingen–Tieringen section, where previous works on sedimentology, nannofossil assemblage composition, and δ¹⁸O and δ¹³C analyses were performed, was selected for this study. The variations in morphology and abundances of W. britannica were studied on 40 samples of the Balingen–Tieringen section, presenting variable lithologies and calcium carbonate contents. For each level, seven biometric parameters (coccolith length, width and ellipticity, central area length, width and ellipticity and central area proportion with respect to the coccolith) were measured or calculated on digitally captured images of the first 100 W. britannica coccoliths observed in the light microscope. The relationships between the different biometric variables were described using bivariate and Principal Component Analyses. Biometric parameters and Principal Component factors extracted from nannofossil assemblages as well as other paleoenvironmental proxies, were investigated using regression, and their stratigraphic trends were compared. Principal component analysis of the six biometric variables (3938 measurements) on W. britannica coccoliths shows a reduced morphological variability compared to a significant size gradient. An allometric trend recognized on the total placolith and on the central area within the W. britannica assemblages suggests that the different morphotypes may represent intra-specific variability rather than different species. The general trend through Late Oxfordian shows an increase in size of W. britannica coccoliths, mainly driven by an increase in the contribution of the large morphotypes. Increasing placolith size is associated with drier and warmer climatic conditions during the latest Oxfordian.     

Gregoryceras in the Oxfordian of Kachchh (India): Diverse eventful implications

The Kachchh Oxfordian ammonoid stratigraphic record for over 150 years remained restricted largely to Early Oxfordian condensed oolitic facies in the relatively distal Mainland Kachchh. Recently, it has been enlarged with the discovery in 1994 of over 200 m-thick uncondensed Middle to Late Oxfordian succession at Kantkote in the proximal part of the basin. Apart from abundant Perisphinctinae and Mayaitinae, a 10 m-thick sediment interval in the lower half of the above succession yielded a few examples of Gregoryceras. The specimens are here identified as G. gr. devauxi Bert and Enay and in view of their association with Larcheria subschilli (Lee) are assigned to the Middle Oxfordian Subschilli Horizon of the Schilli Subzone. Gregoryceras distribution south of the equator in Kachchh, Chile, Mexico and Madagascar, all within 35-40° latitude marks the southern limit of its latitudinal expansion during the first order maximum flooding surface (MFS) of the Schilli Subzone of the Kachchh Toarcian-Hauterivian Sequence. The maximum ammonoid diversity, density and frequency of the Kachchh first order sequence coupled with dominance of mayaitins and perisphinctins suggest over 20 m depth in the Gregoryceras interval. Distinctly greater bathymetry in the 200 km distally away basinal part causes sediment starved situation, and explains the Middle Oxfordian to early Early Kimmeridgian submarine nondepositional hiatus in the Mainland Kachchh.     

Benthic foraminifera of a Miocene canyon and fan

We present benthic foraminiferal assemblage data from an exhumed Miocene canyon and fan system from the Tabernas Basin (SE Spain). The presence of good indicator taxa and unique assemblages occupying specific environments allows the distinction of slope, canyon and fan environments within the Tabernas Basin by foraminiferal assemblages alone. Five assemblages are defined on the basis of the occurrence of the indicator taxa. Primary control on the distribution of these assemblages is consistent with trends of physical disturbance and consequent defaunation. Barren samples, which are predominantly found in high-energy parts of the proximal canyon, are recognized as representing recently defaunated substrates (i.e. early successional assemblages). High diversity assemblages containing a high abundance of agglutinated taxa are recognised in the undisturbed slope sequences as being the regional equilibrium (“climax”) fauna. Intermediate between these end-members are assemblages with low diversity, dominated by calcareous taxa typically found in the relatively low-energy canyon and fan environments, which are recognized as representing the middle phases of the ecological succession. Two further assemblages, a low diversity assemblage typified by Cassidulina laevigata and Bulimina costata and a very low diversity assemblage dominated (> 10% of all benthic tests) by Globobulimina spp., are restricted to low-energy parts of the canyon and fan, and are absent from the proximal canyon and slope. The composition of these assemblages indicate that nutrient supply/oxygenation is a secondary control on the palaeoecology of the canyon system. A conceptual model for the recolonisation of defaunated substrates in El Buho Canyon is proposed, in which either an oligotrophic climax assemblage or a eutrophic climax assemblage can be achieved at the completion of recolonisation of defaunated substrates, depending on environmental conditions.     

Ammonite-benthic Foraminifera turnovers across the Lower-Middle Jurassic transition in the Lusitanian Basin (Portugal)

This paper describes and characterises the co-occurrence of ammonite and benthic foraminiferal assemblages across the São Gião outcrop (Central Portugal), a reference section for the Lower-Middle Jurassic boundary in the Lusitanian Basin. The upper Toarcian-lower Aalenian marls and marly-limestones in this section provide a precise and detailed ammonite-based biostratigraphic zonation, with a mixed assemblage of northwest European and Mediterranean faunal elements, associated with benthic foraminifera assemblages with northern hemisphere affinities, both correlatable with the Aalenian GSSP at the Fuentelsaz section (Iberian Cordillera, Spain). A total of 447 well-preserved ammonite specimens and 13.116 foraminifera have been studied; no evidence was detected of any taphonomic processes that could have changed the original assemblages. From a biostratigraphic point of view, the ammonite record has enabled four biostratigraphic units to be recognised (the Mactra and Aalensis subzones of the Aalensis Biozone in the upper Toarcian, and the Opalinum and Comptum subzones of the Opalinum Biozone in the lower Aalenian). With regard to the benthic foraminifera, the taxa identified have enabled the Astacolus dorbignyi Zone and 11 bioevents to be identified, most of which representing local biostratigraphic proxies. However, the increase in the relative abundance of Lenticulina exgaleata Dieni from the upper part of the Opalinum Subzone to the lower part of the Comptum Subzone has a regional value. The constant and continuous ammonite record of northwest European taxa, together with typical Mediterranean taxa – namely Grammoceratinae – throughout the section, the high relative abundance of Miliolina representatives – generally interpreted as foraminifers typical of shallow waters – and the absence of foraminiferal forms typical of cool waters, do not support the inference of cool seawater temperatures attributed to the Early Aalenian, or the global character of the “Comptum cooling event”, at least with reference to the Lusitanian Basin.     

Stable isotopic study of Oligocene-Miocene sediments from DSDP Site 354, Equatorial Atlantic

The oxygen- and carbon-isotope compositions of planktic and benthic foraminifera and calcareous nannofossils from Middle Oligocene-Early Miocene Equatorial Atlantic sediments (DSDP Site 354) indicate two important paleoceanographic changes, in the Late Oligocene (foraminiferal Zone P.21) and in the Early Miocene (foraminiferal Zone N.5). The first change, reflected by a δ¹⁸O increase of 1.45‰ inGlobigerina venezuelana, affected only intermediate pelagic and not surface, deep or bottom waters. The second change affected surface and intermediate waters, whereas deep and bottom waters showed only minor fluctuations. In the case of the former the isotope effect of the moderate ice accumulation on the Antarctic continent is amplified in the Equatorial Atlantic by changes in the circulation pattern. The latter paleoceanographic change, reflected by a significant increase in¹⁸O in both planktic and benthic forms (about 1.0‰ and 0.5‰, respectively), may have been caused by ice volume increase and temperature decrease. Both oxygen- and carbon-isotope compositions indicate a marked depth-habitat stratification for planktic foraminifera and calcareous nannofossils. Three different dwelling groups are recognized: shallowGlobigerinoides, Globoquadrina dehiscens, Globorotalia mayeri and nannofossils; intermediateGlobigerina venezuelana; and deepCatapsydrax dissimilis. The comparison of foraminifera and calcareous nannofossils suggests that the isotopic compositions of nannofossils are generally controlled by the same parameters which control the isotopic composition of shallow-dwelling foraminifera, but the former are more enriched in¹⁸O.     

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

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

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.     

New paleobiogeographical and paleoenvironmental insight through the Tortonian brachiopod and ichnofauna assemblages from the Mediterranean-Atlantic seaway (Guadix Basin,SE Spain)

The paleogeography of the Late Neogene Atlantic-Mediterranean seaway via the Betic-Rifean Domain is quite complex due to the presence of several marine corridors. The study of transitional basins in these seaways is crucial to understand the configuration and evolution of the Mediterranean-Atlantic inter-connection. A mixed skeletal-siliciclastic sandstone succession located in one of these transitional areas (Guadix Basin, Southern Spain) was studied from a comprehensive paleontological standpoint focused on the main benthic assemblages (foraminifera, brachiopods, and trace fossils), integrating the data with the study of planktic foraminifera for an accurate biostratigraphic framework. Brachiopods are mostly represented by the Aphelesia-Gryphus assemblage. Two trace fossil assemblages were observed, dominated by Ophiomorpha with Bichordites (1) and Macaronichnus (2), respectively. The benthic foraminiferal assemblage is mostly represented by Planulina and Cibicides. The data gathered from the benthic communities reveal habitats with high-energy and turbulent conditions in an outer neritic-upper bathyal bathymetric range. Brachiopods from the Alicún section show a Mediterranean paleobiogeographic affinity. They were constrained in the Late Tortonian to the restricted basins of the Betic-Rifean Seaway and after the Messinian Salinity Crisis proliferated in both Mediterranean- and Atlantic-type basins of the Betic-Rifean Domain. The Guadix Basin contributed to the Mediterranean-Atlantic faunal inter-connection through the Betic-Rifean Seaway during the Late Tortonian and facilitated the earliest Pliocene expansion of brachiopods in the Mediterranean.     

Fossil planktic foraminifera (an overview)

Planktic foraminifera are unicellular marine organisms able to produce calcareous tests, which can be fossilized and, therefore, can give important information about the geologic record. In this overview a summary of the present knowledge of fossil planktic foraminifera together with the gradual steps from the earliest and pioneering to the more recent studies on this microfossil group, is given. In particular, the criteria at the base of the classification of these organisms from the earlier studies until the present, are described and summarized. An overview of the biostratigraphic schemes based on species first and last occurrences and assemblages from different latitudes and different regions is also given. The evolution and the response of planktic foraminifera to changing environmental condition are summarized from the more primitiveGlobigerina-like Jurassic forms, to the specialized and diversified Cretaceous species, until their dramatic crisis across the Cretaceous/Tertiary boundary, their successive recovery in the Paleocene and their evolution toward modern organisms. Finally, an overview is given of the use of planktic foraminifera to reconstruct paleoenvironments from the more simple methods to those implying more sophisticated and recent techniques.