Diversity and geographic distribution of benthic foraminifera: a molecular perspective

The diversity and distribution of modern benthic foraminifera has been extensively studied in order to aid the paleoecological interpretation of their fossil record. Traditionally, foraminiferal species are identified based on morphological characters of their organic, agglutinated or calcareous tests. Recently, however, new molecular techniques based on analysis of DNA sequences have been introduced to study the genetic variation in foraminifera. Although the number of species for which DNA sequence data exist is still very limited, it appears that morphology-based studies largely underestimated foraminiferal diversity. Here, we present two examples of the use of DNA sequences to examine the diversity of benthic foraminifera. The first case deals with molecular and morphological variations in the well-known and common calcareous genus Ammonia. The second case presents molecular diversity in the poorly documented group of monothalamous (single-chambered) foraminifera. Both examples perfectly illustrate high cryptic diversity revealed in almost all molecular studies. Molecular results also confirm that the majority of foraminiferal species have a restricted geographic distribution and that globally distributed species are rare. This is in opposition to the theory that biogeography has no impact on the diversity of small-sized eukaryotes. At least in the case of foraminifera, size does not seem to have a main impact on dispersal capacities. However, the factors responsible for the dispersal of foraminiferal species and the extension of their geographic ranges remain largely unknown. Special Issue: Protist diversity and geographic distribution. Guest editor: W. Foissner.     

Grazing of intertidal benthic foraminifera on bacteria: Assessment using pulse-chase radiotracing

Although foraminifera are a dominant component of many marine benthic communities, quantification of their predation on prokaryotes remains an experimental challenge. We have developed an approach that allows us to study grazing by adult specimens of the calcareous species Haynesina germanica and Ammonia beccarii, and the single-chambered agglutinated species Psammophaga sp., on bacteria (Halomonas sp.), pulse-chase-labelled with ³H- and ¹⁴C-Leucine. The bacterivorous ciliate Uronema sp. and flagellate Pteridomonas sp. were used as positive controls. The rate of release of ³H when protozoa were incubated with the labelled bacteria indicated the predator's grazing rate; the proportion of ¹⁴C found in the foraminiferal biomass and shell indicated the prey assimilation rate. All three foraminiferal species grazed bacteria at a rate of 3.2-5.7 ng C ind⁻¹ h⁻¹ depending on bacterial concentrations. About 23% of the biomass of the ¹⁴C-labelled prey was most likely assimilated into foraminiferal pseudopodia, 12% was expelled in dissolved waste material, about 62% was respired and only 0.1% was incorporated into the carbonate shell. Extracellular digestion associated with pseudopodia could explain the very low proportion of the labelled food assimilated in the cell body and the significant proportion located in pseudopodial networks. These experiments also suggest that very little of the carbon ingested by adult calcareous foraminifera is incorporated into the shell. However, we cannot conclude that diet has no influence on the stable isotope composition of the shell since none of our calcareous specimens grew new chambers during the experiments.     

Oxygen respiration rates of benthic foraminifera as measured with oxygen microsensors

Oxygen respiration rates of benthic foraminifera are still badly known, mainly because they are difficult to measure. Oxygen respiration rates of seventeen species of benthic foraminifera were measured using microelectrodes and calculated on the basis of the oxygen fluxes measured in the vicinity of the foraminiferal specimens. The results show a wide range of oxygen respiration rates for the different species (from 0.09 to 5.27 nl cell⁻¹ h⁻¹) and a clear correlation with foraminiferal biovolume showed by the power law relationship: R = 3.98 10⁻³ BioVol^0.88 where the oxygen respiration rate (R) is expressed in nl O₂ h⁻¹ and in μm³ biovolume (BioVol) (n = 44, R² = 0.72, F = 114, p < 0.0001). The results expressed per biovolume unit (1.82 to 15.7 nl O₂ 10⁻⁸ μm⁻³ h⁻¹) allow us to compare our data with the previous published data showing similar ranges. A comparison with available data for other microbenthos groups (nematodes, copepods, ostracods, ciliates and flagellates) suggests that benthic foraminifera have a lower oxygen respiration rates per unit biovolume. The total contribution of benthic foraminifera to the aerobic mineralisation of organic matter is estimated for the studied areas. The results suggest that benthic foraminifera play only a minor role (0.5 to 2.5%) in continental shelf environments, which strongly contrasts with their strong contribution to anaerobic organic matter mineralisation, by denitrification, in the same areas.     

Cluster analysis of recent benthic foraminifera from the northwestern Mediterranean coast of Egypt

A suite of 74 surface sediment samples, collected from two areas along the Egyptian Mediterranean coast (the Western Harbor of Alexandria and its environs, and the area comprising the Gulf of Kanayis and the Abu Hashafa Bay), have been examined for their benthic foraminiferal faunas. A total of 82 species were identified. Census data were obtained for different species in each sample and the statistically significant fractional abundances values (≥5%) were analyzed using a Q-mode cluster analysis. Samples were segregated into four clusters, each having its peculiar benthic faunal assemblage (biotope), reflecting particular environmental conditions. These clusters are: (1) the Ammonia beccarii forma tepida Biotope, found in samples located in areas with waters of low energy, characterizing semi-closed basin conditions, with muddy or sandy mud bottom sediments (Harbor Proper); (2) the Quinqueloculina spp. Biotope, found in samples collected from depths bathed by turbid inner shelf conditions with some fresh water inflow and sandy bottom sediments (Harbor’s environs); (3) the Peneroplis-Amphistegina Biotope, found in samples collected from depths characterizing marine shelf environments with calcareous algae, and medium to very coarse calcareous sands (the Gulf of Kanayis and the Abu Hashafa Bay); (4) the Triloculina trigonula-Adelosina laevigata Biotope was represented by only one sample (at the far eastern part of the Gulf of Kanayis), collected from a site exhibiting very restricted environmental conditions. The study suggests that nutrients, turbidity, light intensity, type of substrate, and salinity are the main ecological factors controllingthe distribution of benthic foraminifera.     

Abyssal NE Atlantic benthic foraminifera during the last 15 kyr: Relation to variations in seasonality of productivity

Benthic foraminiferal faunas (> 63 μm) and stable isotopes from the last 15 kyr were studied in BENGAL programme (high-resolution temporal and spatial study of the BENthic biology and Geochemistry of a north-eastern Atlantic abyssal Locality) kasten core 13078#16 from the Porcupine Abyssal Plain, NE Atlantic (48°49.91 N, 16°29.94 W, water depth 4844 m). Changes occurred in the accumulation rates, species composition, diversity, and stable isotopes during the last 15 kyr. Today, the area is strongly influenced by seasonal inputs of phytodetritus following the spring blooms in surface water primary productivity. Variations in the relative abundance of the two most abundant species, Epistominella exigua and Alabaminella weddellensis, which today show significant increases in abundance with the presence of phytodetritus on the sea-floor, are interpreted as resulting from changes in the seasonality of productivity. Seasonal productivity was higher during the Holocene than during the last deglaciation and Younger Dryas, probably coinciding with the retreat of the polar front to higher latitudes. This hypothesis is consistent with simultaneous decreases in the percentage of the polar planktic foraminifera Neogloboquadrina pachyderma (s), and increases in the percentage of Globigerina bulloides, a warmer water planktic foraminifera indicative of phytoplankton blooms and enhanced productivity. The relative abundance of the ‘phytodetritus species’ (E. exigua and A. weddellensis) covary between 14.7 and 8.1 kyr, but not between 7.8 and 1.2 kyr. Major decreases in the numbers per gram and accumulation rates of planktic and benthic foraminifera occurred at ∼ 12–8.5 kyr and at ∼ 4 kyr which correspond to decreases in the % sediment coarse fraction and published data on inorganic carbon contents suggesting that dissolution may have increased at these times. Relationships between benthic foraminiferal faunas and benthic stable isotope records suggest no simple relationship between faunal abundances and test isotope chemistry. For example, the abundances of phytodetritus species do not show strong correlations with either the δ¹³C values of E. exigua or the Δδ¹³C _{E. exigua − P. wuellerstorfi} record, which have previously been suggested as indicative of seasonality of productivity.     

Sediment (grain size and clay mineralogy) and organic matter quality control on living benthic foraminifera

The paleoecological interpretation of fossil foraminiferal assemblages depends on an understanding of the ecological processes operating at the present. This study investigates both the quality of organic matter (OM) by elemental analysis as well as the sediment grain size and clay mineralogy to understand their relative influence on distribution and abundance of benthic foraminifera. This study is carried out on 15 samples regularly spaced from the mudflat to the tidal marsh. The results indicate that grain size is the most limiting parameter. Living (stained) benthic foraminiferal density and species richness are both very low within coarser sediments. OM is the second limiting factor. The density of foraminifera is the lowest and the species richness is the highest with the lowest organic carbon (C_org) contents and C/N < 12. Conversely, when the C_org is very high and C/N > 12, the density is high and the species richness medium. A high smectite proportion within the clay-size fraction seems to favor the development of Miliammina fusca. Trochammina inflata and Jadammina macrescens are both favored by an increase of organic carbon proportion but Trochammina inflata preferentially feeds on algal-derived OM when compared with Jadammina macrescens.     

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 foraminifera and trace metal distribution: a case study from the Burullus Lagoon,Egypt

Benthic foraminifera are widely used to detect the health of their habitat, where they are very sensitive to even slight variations in the ecosystem. Therefore, the main objectives of this study are to examine the benthic foraminiferal assemblages in the sediments of Burullus Lagoon, evaluate the pollution levels and deduce the impact of trace metals on foraminifera. The continuous discharge of trace metals from agricultural, industrial and domestic sources into the lagoon may lead to a severe environmental problem. The concentrations of Mn, Cu, Cd, Zn and Pb within the sediments were measured. Recently, the assessment of contamination is principally based on the contamination indices which provide fast and simple quantitative values on the degree of pollution in a given aquatic environment. Thus, some indices, including the contamination factor, the degree of contamination, pollution load index, geoaccumulation index, ecological risk factor and potential ecological risk index are applied in this investigation. Based on the contamination factors, the sediments are very highly contaminated with Cd, considerably to very highly contaminated with Cu and Zn, moderately contaminated with Mn, low to moderately contaminated with Pb. All sites display very high values for the degree of contamination. Moreover, the values of the pollution load index are higher than 1, indicating that the lagoon is polluted. Depending on the geoaccumulation index, the contaminants are arranged as follows Cd > Zn > Cu > Mn > Pb. It is clear that Cd is the main contributor to the ecological risk factor in Burullus Lagoon. Concerning the richness of the foraminiferal assemblages, it fluctuates between 1–5 species per sample. Because of its higher tolerance to extreme conditions (changes in salinity and pollution), Ammonia tepida is the most abundant species. The occurrence of rare living individuals (25) is restricted only to sites close to El-Boughaz Inlet where higher salinity and lower levels of pollution are recorded. The same trend of distribution is shown by Cribroelphidium excavatum and miliolids, where they occur at sites with higher salinities. The occurrence of test deformities in all the studied sites may be related to the response of benthic foraminifera to trace metal. The forms of deformation include spiroconvex, reduced chambers, twisted tests, twinning, additional chamber and complex forms. The deformation depends on the nature of pollutants. Twinning and reduced chambers are the most dominant forms in areas close to the agricultural drainage (southern drains), while complex forms are abundant in areas close to industrial drainage (El-Gharbia drain). Thus, salinity and pollution may be the most regulatory factors controlling the distribution of foraminifera. This investigation confirmed the role of benthic foraminifera as a good ecological indicator in Burullus Lagoon.     

Calcite crystal orientation patterns in the bilayers of laminated shells of benthic rotaliid foraminifera

Shells of calcifying foraminifera play a major role in marine biogeochemical cycles; fossil shells form important archives for paleoenvironment reconstruction. Despite their importance in many Earth science disciplines, there is still little consensus on foraminiferal shell mineralization. Geochemical, biochemical, and physiological studies showed that foraminiferal shell formation might take place through various and diverse mineralization mechanisms.In this study, we contribute to benthic foraminiferal shell calcification through deciphering crystallite organization within the shells. We base our conclusions on results gained from electron backscattered diffraction (EBSD) measurements and describe microstructure/texture characteristics within the laminated shell walls of the benthic, symbiontic foraminifera: Ammonia tepida, Amphistegina lobifera, Amphistegina lessonii. We highlight crystallite assembly patterns obtained on differently oriented cuts and discuss crystallite sizes, morphologies, interlinkages, orientations, and co-orientation strengths.We show that: (i) crystals within benthic foraminiferal shells are mesocrystals, (ii) have dendritic-fractal morphologies and (iii) interdigitate strongly. Based on crystal size, we (iv) differentiate between the two layers that comprise the shells and demonstrate that (v) crystals in the septa have different assemblies relative to those in the shell walls. We highlight that (vi) at junctions of different shell elements the axis of crystal orientation jumps abruptly such that their assembly in EBSD maps has a bimodal distribution. We prove (vii) extensive twin-formation within foraminiferal calcite; we demonstrate (viii) the presence of two twin modes: 60°/[0 0 1] and 77°/~[6 –6 1] and visualize their distributions within the shells.In a broader perspective, we draw conclusions on processes that lead to the observed microstructure/texture patterns.     

Thanetian–Ilerdian coralline algae and benthic foraminifera from northeast India: microfacies analysis and new insights into the Tethyan perspective

Thanetian–Ilerdian carbonate deposits from the Lakadong Limestone in Assam Shelf, Meghalaya, northeast India, are studied with respect to microfacies distributions and controlling ecological factors on dominant biogenic components. Palaeoenvironmental implications are inferred following the detailed analysis of microfacies characterized by rich assemblages of coralline red algae and benthic foraminifera. The carbonate sediments have been interpreted as lagoonal to outer shelf facies. It is envisaged that the analysed benthic communities thrived in a meso‐oligotrophic regime above the fair‐weather wave base. The Lakadong Limestone constitutes a well‐preserved record of Late Palaeocene–earliest Eocene shallow marine carbonate ecosystem and has high potential to decipher its response to an interval of distinct changes in climate and tectonic settings. The abundance of oligotrophic larger benthic foraminifera in the Lakadong Limestone is comparable to the foraminiferal assemblages of west Tethys. The phylogenetic changes (‘Larger Foraminiferal Turnover’, LFT) and subsequent rapid radiation of typical Eocene larger benthic foraminifera (Alveolina, Nummulites) usually observed in the west Tethys have also been observed in the upper part of the Lakadong Limestone. The eastward migration of Eocene foraminifera from the west coincided with the India‐Asia collision and global warming events at the Palaeocene–Eocene boundary that may have produced a wide array of modifications in biogeography, seasonal run‐offs and ocean circulation pathways. The data indicate that rapid rate of migration from west before the onset of geographic barriers and/or timely restoration of pan‐Tethyan environmental conditions ensured the incidence of these forms in the earliest Eocene sediments.     

Benthic foraminifera and their role to decipher paleoenvironment during mid-Cretaceous Oceanic Anoxic Events - the “anoxic benthic foraminifera” paradox

Oceanic Anoxic Events (OAEs) are a widespread, recurring and well-studied feature of the mid-Cretaceous greenhouse climate, reflecting severe perturbations in Earth's climate and oceanography. To investigate bottom-water environments before, during and after these OAEs and to understand the dynamic fluctuations of seafloor environments, benthic foraminifera are a commonly employed tool. Here, a review of the response of benthic foraminiferal assemblages to OAEs (i.e., OAEs 1a, 1b, 1d, 2, and 3) will be given. I will discuss in detail how benthic foraminifera can be used to identify the paleoclimatic and paleoceanographic mechanisms responsible for OAE formation. The main feature of benthic foraminiferal assemblages during oceanic anoxic events is a dramatic decrease in both diversities and abundances or their total absence. However, some studies record the permanent occurrence of benthic foraminifera throughout OAEs or short-term repopulation events. These patterns clearly indicate that mid-Cretaceous OAEs are complex events. The paradox of anoxic (and even euxinic) conditions during OAEs coincident with the occurrence of benthic foraminifera (here called the “anoxic benthic foraminifera” paradox) is most readily explained by the sampling (and therefore temporal) resolution of typical foraminiferal studies that have homogenized hundreds to thousands of years in a single sample.     

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.     

Habitat selective factors influencing the distribution of larger benthic foraminiferal assemblages over the Kepulauan Seribu

Symbiont-bearing foraminifera are used to study the effects of habitat deterioration on benthic communities in coral reefs dominated by macroalgae. It is shown that, despite their preference for nutrient deprived conditions, some symbiont-bearing foraminifera occur on reefs heavily affected by nutrient stress and macro-algal dominance, thus highlighting the need for a better understanding of the autecology of species and assemblages in these conditions. Both diversity and habitat fractionation increases as terrestrial and nutrient influence decline. The assemblage structure in the most nearshore reefs are dominated by generalist species, while, additionally, more specialistic species occur at the more offshore reefs. Apart from larger scale gradients in ambient seawater quality, local scale variation in physical environmental conditions, such as habitat structure, are important for the assemblage structure as well.     

Spatial distribution of live benthic foraminifera in the Rhône prodelta: Faunal response to a continental–marine organic matter gradient

Benthic foraminifera were collected in the Rhône prodelta (Gulf of Lions, Mediterranean Sea), an enriched zone with high organic matter content. In June 2005, sediment cores were sampled at depths ranging from 20 to 100 m. Four distinct foraminiferal assemblages were determined in the study area, reflecting the geographical distribution of the impact of river supply. The living foraminiferal faunas present a typical picture, with strongly impoverished faunas composed exclusively of stress-tolerant taxa (Fursenkoina fusiformis, Bulimina aculeata, Leptohyalis scottii, and Adelosina longirostra) in the immediate vicinity of the river mouth. This assemblage is well adapted to a high input of continental organic matter and a minimum oxygen penetration depth into the sediment. To the southwest, under the main corridor followed by the river plume, high organic input with a dominantly terrestrial signature (more refractory) may be stressful for many taxa which need organic matter of a more labile quality. In this area, Nonion scaphum, Nonionella turgida and Rectuvigerina phlegeri are present in low densities. On the edge of this area, these taxa show much higher densities. A greater proportion of marine organic carbon could explain their increasing abundances in this area. Towards the east and towards the deepest stations, in the outer part of the enriched zone, biodiversity increases. Faunas at these stations have intermediate densities and contain a number of taxa (Cassidulina carinata, Epistominella vitrea, Valvulineria bradyana, Nonionella iridea/bradyi) at the deepest stations; Bolivina dilatata/spathulata and Textularia porrecta at the eastern stations) that seem to benefit from more marine organic matter. The comparison of geochemical measurements and foraminiferal data strongly suggests that the spatial distribution of foraminifera in the Rhône prodelta is mainly governed by the quality and the quantity of organic matter reaching the sediment–water interface. Since bottom waters are well oxygenated (215–260 µmol/L), and oxygen penetration into the sediment is less than 1 cm at all stations, benthic ecosystem oxygenation appears to have only a minor impact on regional differences in faunal distribution.     

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

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

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

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

Suitable sediment fraction for paleoenvironmental reconstruction and assessment of contaminated coastal areas based on benthic foraminifera: A case study from Augusta Harbour (Eastern Sicily,Italy)

Since the 1990s several studies noticed that, along coastal marine areas, the mean size of benthic foraminifera may be reduced due to heavy metal pollution, even if no biometric studies were carried out to quantify this aspect. The Augusta harbour (Sicily, Italy), is characterized by a strong contamination due to several anthropogenic activities, the most important of which are a petrochemical pole and an important industrial harbour. Taking into account the previous studies carried out in the area, which recorded small-sized foraminifera, the present study compared assemblage composition and faunal parameters in the >125 μm and >63 μm fractions of a sediment core collected in the most polluted sector of Augusta harbour. The aim was to understand if the two fractions have comparable environmental significance providing reliable information on the environmental status. In order to quantify the amount of smaller foraminifera in a community and to determine species loss between size fractions, two new indices are used: the Foraminiferal Size Index (FSI) and the Lost Species Index (LSI). Species richness, diversity and composition of the two assemblages were determined to characterize their structure. The results highlighted great depletion and different composition of the >125 μm assemblage with respect to the >63 μm one, showing a selective loss of particular ecological groups (stress-tolerant infaunal taxa). Also the better correlation of Foraminiferal Number (FN) and H’ index of >63 μm fraction with Polychlorobiphenyls (PCBs), Polycyclic Aromatic Hydrocarbons (PAHs), Barium (Ba) and Mercury (Hg), demonstrated the higher reliability of this size fraction for environmental assessment purposes.     

Size-related isotopic trends in some Maastrichtian planktic foraminifera: methodological comparisons, intraspecific variability, and evidence for photosymbiosis

The serial test dissection and sieve fraction methods for determining the pattern of size-related change in oxygen and carbon isotopic ratios are compared using four Late Cretaceous planktic foraminifer species (Racemiguembelina fructicosa, Planoglobulina acervulinoides, Planoglobulina multicamerata, and Pseudoguembelina palpebra) from a subtropical site in the North Atlantic (DSDP Hole 390A). Despite the extra labor required, we identify several clear advantages of the dissection method, including: (1) it provides a means of obtaining size-dependent changes in isotopic signatures that are unequivocally ontogenetic, whereas isotopic variation observed from sieve-separated size fractions could be ontogenetic or ecotypic; (2) the taxonomic identity of smaller sized specimens using the dissection method is unequivocal, whereas species identification is increasingly ambiguous in smaller size fractions using the sieve method; (3) it reveals a greater total range and a greater complexity in the pattern of ontogenetic change in stable isotopic values, whereas the sieve method averages the isotopic signal across the entire ontogenetic range preserved within the whole tests that are used. Our results from serial dissections demonstrate that among the species analyzed, R. fructicosa and P. acervulinoides yield relatively negative adult δ¹⁸O values, a large size-related change in δ¹³C values (1.32 and 2.05‰, respectively), and virtually no correlation between size-related δ¹³C and δ¹⁸O values. On this basis we suggest that these were photosymbiotic species that inhabited relatively shallow surface waters. Evidence for photosymbiosis is not as compelling for P. palpebra, as this species yields a 1.06‰ shift in δ¹³C and relatively negative δ¹⁸O values in adult chambers, but much stronger correlation between size-related δ¹³C and δ¹⁸O values (r²=0.40) than in R. fructicosa and P. acervulinoides. Planoglobulina multicamerata yields the most positive adult δ¹⁸O values of the species studied, a strong covariance between size-related δ¹³C and δ¹⁸O values (r²=0.77), and a 0.97‰ shift in δ¹³C composition during ontogeny. We conclude that this species lacked photosymbionts and migrated to a deeper surface water paleohabitat as it increased in size. Single specimen analyses of tightly constrained size fractions reveal a high degree of intraspecific variation. δ¹³C and δ¹⁸O values vary by up to 0.70 and 0.28‰ in R. fructicosa, 1.41 and 0.80‰ in P. acervulinoides, 0.66 and 0.82‰ in P. palpebra, and 0.18 and 0.33‰ in P. multicamerata, respectively. Such a range of isotopic variation has been observed in modern day planktic foraminifer assemblages, and likely results from growth of individuals during different phases of the seasonal cycle and/or the kinetic effect of intraspecific variation in shell calcification rates. As suggested by other investigators, large sample sizes should be analyzed to provide the most reliable correlation of stable isotopic stratigraphic records.     

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.     

Origin of double and multiple tests in benthic foraminifera: observations in laboratory cultures

Many juvenile and adult double and multiple tests of benthic foraminifera were observed on specimens grown in laboratory cultures or collected in various natural environments. Our observations bring to light three possible causes for such abnormalities, each one referring to characteristic morphological features. Double tests may result: (1) from an anomaly in the development of a single juvenile, building two or three second chambers or two third chambers, each one possibly developing in an individual whorl; (2) from the early fusion of two juveniles, which both develop after their fusion; (3) from the attachment of a juvenile on a parental test after the schizogony followed by the young's development.