Assessment of anthropogenic pollution in Guanabara Bay (SE Brazil) through biogeochemical data and stable isotope mixing models.

This work intends to identify pollution sources along the margins of Guanabara Bay (GB; SE Brazil) through a multiproxy approach and Bayesian stable isotopic mixture model (BSIMM). For this purpose, 33 surface sediment samples were collected and analyzed for granulometry, geochemistry (heavy metals, total organic carbon-TOC, stable isotopes of carbon and nitrogen-δ13C and δ15N, Rock-Eval pyrolysis parameters-REPP), and physicochemical parameters. Metal concentrations (E) dissolved in water (EW), adsorbed by organic matter (EOM) and by Mn hydroxides (EMn), and total extracted concentrations (ET) were analyzed. Sampling was conducted in 2018 after an oil spill from Reduc Oil Refinery. Potential Ecological risk index (PERI), based on metals, classified 85% of the analyzed stations as having moderate to considerable ecological risk. The metals with the potential to cause the highest ecological risk were CdW, CdOM, PbOM, and HgOM. The combination of BSIMM and REPP data was an effective proxy for oil spill detection by indicating the presence of polycyclic aromatic hydrocarbons (PAHs). Relatively high TOC contents suggested that the analyzed stations are eutrophicated environments. BSIMM discriminated three groups of stations with different sources of organic matter (OM), endorsing the result previously shown by the cluster analysis: (A) Niterói region, Botafogo marina, Glória marina, Fiscal and Fundão islands with diffuse sources of OM, including marine phytoplankton and material of continental origin from highly polluted rivers and domestic sewage; (B) region near Fundão and Governador islands and Mangue Channel outlet with OM (≃70%) supplied by highly polluted streams and a small contribution of PAHs; (C) Duque de Caxias and Botafogo-Urca inlet with significant contributions of PAHs, materials from C-3 plants and rivers polluted by urban sewage. Results of linear regressions in conjunction with BSIMM indicate that HgMn and PbOM mainly affect Group A's stations. Although the eastern margin of GB (Niterói; Group A) showed greater oceanic interaction than the other groups, it presented substantial concentrations of metals, potentially harmful (i.e., Hg and Pb) to marine biota and human health.

Inferring the ecological quality status based on living benthic foraminiferal indices in transitional areas of the Guanabara bay (SE Brazil).

Using benthic foraminifera, we evaluate the ecological quality status (EcoQS) of transitional waters of the Guanabara Bay (SE Brazil) by applying the diversity-based index exp (H'bc) and the sensitivity-based Foram-AMBI for the first time in South America. The Guanabara Bay was selected for this study as it is one of the largest transitional ecosystems in the State of Rio de Janeiro and has been severely impacted by anthropogenic activities. Concentrations of potentially toxic elements (PTEs) were assessed by sequential chemical extraction in three phases (i.e., dissolved in water, adsorbed on organic matter, and Mn oxy-hydroxides). Total organic carbon, total nitrogen, and stable isotope (δ13C and δ15N) signatures of organic matter were analyzed to trace environmental stress. The Ammonia/Elphidium ratio suggests hypoxic conditions at most of the sampled sites. Principal component analysis identifies the first component as environmental stress underlying organic matter and PTE enrichment (in all three phases), which is positively related to Foram-AMBI and negatively to exp (H'bc). The exp (H'bc) and Foram-AMBI indices reveal that stations near the Governador Island and Niterói margin have the worst EcoQS, showing medium to extreme pollution. Additionally, Foram-AMBI and exp (H'bc) provide a congruent EcoQS classification for ∼64% of the sites. Although these results are promising, they suggest that a significant effort should be made to obtain better knowledge of foraminiferal ecological requirements to employ benthic foraminifera as a biomonitoring and management method.

Ecological quality status of the NE sector of the Guanabara Bay (Brazil): A case of living benthic foraminiferal resilience.

The ecological quality status of the NE region of the Guanabara Bay (SE Brazil), one of the most important Brazilian embayments, is evaluated. For this purpose, sediment samples from in the inner of the Guanabara Bay (GB) were collected and analyzed (grain-size, mineralogy, geochemistry and living foraminifera). In this study, it is hypothesized that the potentially toxic elements (PTEs) concentrations, in solution and associated with organic matter (OMPTEs, potential nutrient source), may represent two potential pathways to impact benthic foraminifers. A multiproxy approach applied to complex statistical analyses and ecological indexes shows that the study area is, in general, eutrophic (with high organic matter and low oxygen content), polluted by PTEs and oil. As a consequence, foraminifera are not abundant and their assemblages are poorly diversified and dominated by some stress-tolerant species (i.e., Ammonia tepida, Quinqueloculina seminula, Cribroelphidium excavatum). The results allow us to identify a set of species sensitive to eutrophication and OMPTEs. Factors such as the increase of organic matter contents and OMPTEs and, in particular of Zn, Cd and Pb, the oxygen depletion and the presence of oil, altogether contribute to a marked reduction in the abundance and diversity of foraminifera. Ammonia-Elphidium Index and the Foram Stress Index confirm that the NE zone of GB is, in general, "heavily polluted", with "poor ecological quality status" and experiences suboxic to anoxic conditions. In light of it, special attention from public authorities and policymakers is required in order to take immediate actions to enable its environmental recovery.

Lunar cycles and rainy seasons drive growth and reproduction in nummulitid foraminifera, important producers of carbonate buildups.

Representatives of the foraminifer Nummulites are important in Earth history for timing Cenozoic shallow-water carbonates. Taphonomic complexity explains the construction of carbonate buildups, but reproduction and life span of the constructing individuals are unknown. During the 15-month investigation period, asexually reproduced schizonts and gamonts showed equal proportions in the first half of this period, whereas gamonts predominated in the second half. Oscillations in cell growth are mainly caused by light intensities during chamber construction when minor differences in water depth increase the photosynthetic rate of endosymbiotic diatoms during neap tides. The continuous reproduction rate of N. venosus throughout the year is increased in subtropical calms by higher summer temperatures and the marginal input of inorganic nutrients during rainy seasons. The expected life span of both gamonts and schizonts are 18 months.

Morphometric analysis of Eocene nummulitids in western and central Cuba: taxonomy, biostratigraphy and evolutionary trends.

Megalospheric specimens of Nummulitidae from eight localities in western and central Cuba were morphometrically investigated using test characters described by 11 growth-independent and growth-invariant attributes that provide a complete geometric reconstruction of nummulitid equatorial morphology. The species Nummulites striatoreticulatus, Palaeonummulites trinitatensis, Operculinoides floridensis and O. soldadensis were classified by an agglomerative cluster analysis. Discriminant analysis yielded significant morphological separators between the species such as the backbend angle, marginal radius increase, perimeter ratio and first chamber length. The transition of tightness to laxity of the spiral was an important morphological separator at the generic level, representing a clear general trend coupled with the change in palaeodepth. Based on further discriminant analysis, an increase in proloculus size was detected in Nummulites striatoreticulatus from the middle Eocene to early late Eocene, supporting this important evolutionary pattern in many lineages of Nummulites. Operculinid forms showed an opposite and more weakly pronounced time-dependent trend in the size decrease of the proloculus. In the Cuban localities, Nummulites striatoreticulatus occurs from the Lutetian to Priabonian, while Palaeonummulites trinitatensis is restricted to the Bartonian to Priabonian. The moderately to loosely coiled operculinid taxa O. floridensis and O. soldadensis have longer stratigraphical ranges from the middle Eocene to probably the early Oligocene. Operculinoides floridensis and O. soldadensis show a broader variability in marginal radius increase, and thus probably occupied wider niches than N. striatoreticulatus. The latter seems to be restricted to the shelf edge and to the shallowest parts of the upper slope. A possible phylogenetic connection between Heterostegina and Operculinoides is suggested by the closest equatorial morphology of Heterostegina sp. indet. to tightly coiled forms of Operculinoides floridensis. Discriminant analysis documents the strongest similarities in perimeter ratio, backbend angle, initial marginal radius and proloculus mean diameter.

Dissimilarity between living and dead benthic foraminiferal assemblages in the Aveiro Continental Shelf (Portugal).

This study compares living (LA) and dead (DA) benthic foraminiferal assemblages and identifies different factors that possibly cause differences in the distribution of both assemblages in the Aveiro Continental Shelf (Portugal). A total of 44 sediment samples was collected during summers of 1994 and 1995 along transects (east-west direction) and between 10 and 200 m water depth. Complex statistical analyses allow us to compare the abundance and composition of the LAs and DAs in function of depth, grain-size and total organic matter in all studied stations even in those where the numbers of individuals were rare in one or both assemblages. The highest densities and diversities of the LAs are found in the middle continental shelf on gravel deposits (coarse and very coarse sands) mostly due to the substrate stability, reduced deposition of fine sedimentary particles, availability of organic matter with high quality related to oceanic primary productivity likely induced by upwelling events, and oxygenated porewaters conditions. The DAs have, in general, higher densities and diversities than the LAs. In the outer continental shelf, the dissimilarity between both assemblages is higher due to the accumulation of tests, low dilution by sedimentary particles and scarcity of living foraminifera. Based on the comparison of LAs and DAs and considering the characteristics of the study area and the species ecology, it has been possible to understand the cause of temporal deviation between the LAs and DAs of benthic foraminifera. This deviation is much more pronounced in the inner shelf where the energy of the waves and the currents induce very dynamic sedimentary processes preventing the development of large LAs and the preservation of DAs. Some deviation also occurs in the middle shelf due to the seasonal loss of empty tests. The most well-preserved time-averaged DAs were found in the outer continental shelf.

Growth estimation of the larger foraminifer Heterostegina depressa by means of population dynamics.

In Heterostegina depressa, the flagship species of laboratory investigations of larger benthic foraminifera (LBF) since the 70's, the timing of reproduction, longevity and natural chamber building rates are still understudied. A recently developed method, the natural laboratory (sensu Hohenegger), has been applied on H. depressa populations from Sesoko Jima, NW Okinawa, Japan. An averaged chamber building rate and longevity of H. depressa were calculated based on 17 monthly samplings at fixed stations. All samples were collected at 20 and 50 m water depths using SCUBA. Live populations were dried and investigated by microCT. The monthly frequency distributions of chamber numbers and test diameters have been decomposed in normally distributed components. For each month, mean and standard deviations of the components were used to calculate the maximum chamber number and maximum test diameter. Based on these values, the natural chamber building rate (CBR) or diameter increase rate (DIR) could be estimated using the Michaelis-Menten function. CBR and DIR were inverted to estimate the 'birthdate' of all investigated individuals. Based on frequencies of these 'birthdates', main reproduction events could be detected and compared to the reproduction timing of other subtropical and tropical LBF taxa. Furthermore, peaks in reproduction could be linked to monsoon wet seasons (="rainy seasons") and winter rains.

Growth, chamber building rate and reproduction time of Palaeonummulites venosus (Foraminifera) under natural conditions.

We investigated the symbiont-bearing benthic foraminifer Palaeonummulites venosus to determine the chamber building rate (CBR), test diameter increase rate (DIR), reproduction time and longevity using the 'natural laboratory' approach. This is based on the decomposition of monthly obtained frequency distributions of chamber number and test diameter into normally distributed components. Test measurements were taken using MicroCT. The shift of the mean and standard deviation of component parameters during the 15-month investigation period was used to calculate Michaelis-Menten functions applied to estimate the averaged CBR and DIR under natural conditions. The individual dates of birth were estimated using the inverse averaged CBR and the inverse DIR fitted by the individual chamber number or the individual test diameter at the sampling date. Distributions of frequencies and densities (i.e., frequency divided by sediment weight) based on both CBR and DIR revealed continuous reproduction throughout the year with two peaks, a stronger one in June determined as the onset of the summer generation (generation 1) and a weaker one in November determined as the onset of the winter generation (generation 2). This reproduction scheme explains the presence of small and large specimens in the same sample. Longevity, calculated as the maximum difference in days between the individual's birth date and the sampling date, is approximately 1.5 yr, an estimation obtained by using both CBR and DIR.

Growth of Heterostegina depressa under natural and laboratory conditions.

The use of micro-computed tomography (µCT) provides a unique opportunity to look inside the shells of larger benthic foraminifera to investigate their structure by measuring linear and volumetric parameters. For this study, gamonts/schizonts and agamonts of the species Heterostegina depressa d'Orbigny were examined by µCT; each single chamber's volume was digitally measured. This approach enables cell growth to be recognised in terms of chamber volume sequence, which progressively increases until reproduction occurs. This sequence represents the ontogeny of the foraminiferal cell and has been used here to investigate controlling factors potentially affecting the process of chamber formation. This is manifested as instantaneous or periodic deviations of the realised chamber volumes derived from modelled growth functions. The results obtained on naturally grown specimens show oscillations in chamber volumes which can be modelled by sums of sinusoidal functions. A set of functions with similar periods in all investigated specimens points to lunar and tidal cycles. To determine whether such cyclic signals are genuine and not the effects of a theoretical model, the same analysis was conducted on specimens held in a closed laboratory facility, as they should not be affected by natural environmental effects. Surprisingly, similar cyclicities were observed in such samples. However, a solely genetic origin of these cycles couldn't be verified either. Therefore, detailed analysis on the phase equality of these growth oscillations have been done. This approach is pivotal for proving that the oscillatory patterns discovered in LBF are indeed genuine signals, and on how chamber growth might be influenced by tidal currents or lunar months.

New results on the hydrodynamic behaviour of fossil Nummulites tests from two nummulite banks from the Bartonian and Priabonian of northern Italy.

Settling velocities of 58 well-preserved tests of fossil Nummulites were experimentally determined using a settling tube. The tests were collected from the nummulite banks of Pederiva di Grancona (A forms of N. lyelli and N. striatus, Middle Eocene) and San Germano dei Berici (A and B forms of N. fabianii, Late Eocene), both in the Berici Mts. (Veneto, northern Italy). The data were compared with estimated settling velocities that the same specimens might have had in life conditions. This was done by reconstructing their densities simulating water-filled condition and, to simulate post-diagenetic effects, under calcite-filled condition. These simulations show that A and B forms, even if they greatly diverge in shape, volume and size, still possess comparable settling velocities, and that each nummulite bank is characterized by specific hydrodynamic parameters. The use of settling velocity as a parameter to quantify the hydrodynamic behaviour of particles in seawater enables estimation of palaeoenvironmental conditions such as depth, substrate and the energy scenario. Such information is useful in obtaining further insights into the genesis of nummulite banks, the autochthony or allochthony of which is still being debated. Our results point to an autochthonous interpretation.

Growth oscillation in larger foraminifera.

This work shows the potential for applying three-dimensional biometry to studying cell growth in larger benthic foraminifera. The volume of each test chamber was measured from the three-dimensional model obtained by means of computed tomography. Analyses of cell growth based on the sequence of chamber volumes revealed constant and significant oscillations for all investigated specimens, characterized by periods of approximately 15, 30, 90, and 360 days. Possible explanations for these periods are connected to tides, lunar cycles, and seasonality. The potential to record environmental oscillations or fluctuations during the lifetime of larger foraminifera is pivotal for reconstructing short-term paleoenvironmental variations or for gaining insight into the influence of tides or tidal current on the shallow-water benthic fauna in both recent and fossil environments.

TEST FUSION IN ADULT FORAMINIFERA: A REVIEW WITH NEW OBSERVATIONS OF AN EARLY EOCENE NUMMULITES SPECIMEN.

In foraminifera, so-called "double tests" usually arise due to abnormal growth originating mainly from twinning, but may also be caused by irregularities in the early chambers and by regeneration after test injury that modifies the direction of growth. A fourth cause of double tests has only rarely been reported: the fusion of the tests of two adult individuals. We studied an early Eocene Nummulites double test consisting of two adult individuals that fused after an extended period of independent growth. The specimen was studied using computed tomography with micrometric resolution (micro-CT) that allowed bi- and three-dimensional visualization of the internal structure. Before fusion each individual test had 30-36 chambers, which, by comparison with growth rates in recent nummulitids, implies at least three months of independent growth. After fusion, the compound test grew in two spirals that fused after about one whorl and then continued in a single spiral. To fuse their tests, either adult individuals have to be forced to do so or the allorecognition (ability to distinguish between self and another individual) mechanisms must fail. A possible explanation for the merged Nummulites tests in this study is forced fusion in attached individuals after surviving ingestion and digestion by a metazoan. Alternatively, environmental stress could lead to a failure of allorecognition mechanisms and/or foraminiferal motility. Once fused, subsequent growth seems to be determined mainly by the relative orientation of individual tests. In any case, the frequency in which adult fusion occurs remains unknown.

PALEOBIOLOGICAL APPLICATIONS OF THREE-DIMENSIONAL BIOMETRY ON LARGER BENTHIC FORAMINIFERA: A NEW ROUTE OF DISCOVERIES.

Four specimens of larger benthic foraminifera (the Recent Palaeonummulites venosus and Operculina ammonoides, and the phylogenetically related Paleogene Nummulites fabianii and N. fichteli) were investigated by X-ray tomography. The resulting three-dimensional measurements enabled a comprehensive, quantitative study of shell morphology to interpret cell growth without specific shell preparation and/or destruction. After segmentation and extraction of all scanned lumina, the following characters were measured on all chambers of each specimen: chamber volume, septal distance, chamber height, and chamber width. The sequence of chamber lumina follows either a logistic function (Palaeonummulites, Operculina), where the deceleration in growth rate of the latest chambers could mark the onset of reproduction, or it can be modeled by a series of stepwise functions with differing constants (Nummulites). Variations around the growth model are either periodic, following external cycles, or random as expressed by abrupt deviations. Therefore, they may reflect the response of the cell to environmental changes in terms of cyclic changes (e.g., seasonality) or single events (e.g., predator attack). Correlations between chamber volume and the other chamber parameters show that septal distance always matches the sequence in chamber volume and can therefore be used as a proxy for environmental analyses in both growth models. Chamber height and width often remain constant around their function and rarely deviate drastically to accommodate the needed lumen for retaining test size and shape. Chamber width may vary according to chamber volume in involute specimens, whereas both chamber height and width correlate with volume in those tests following an Archimedean spiral. X-ray-tomography shows particular promise in determining which parameters that can be assessed routinely in two dimensions primarily reflect environmental conditions vs. parameters best used for taxonomic identification and for systematic lineage reconstruction.

AXIALLY ORIENTED SECTIONS OF NUMMULITIDS: A TOOL TO INTERPRET LARGER BENTHIC FORAMINIFERAL DEPOSITS.

The "critical shear velocity" and "settling velocity" of foraminiferal shells are important parameters for determining hydrodynamic conditions during deposition of Nummulites banks. These can be estimated by determining the size, shape, and density of nummulitid shells examined in axial sections cut perpendicular to the bedding plane. Shell size and shape can be determined directly from the shell diameter and thickness, but density must be calculated indirectly from the thin section. Calculations using the half-tori method approximate shell densities by equalizing the chamber volume of each half whorl, based on the half whorl's lumen area and its center of gravity. Results from this method yield the same lumen volumes produced empirically by micro-computed tomography. The derived hydrodynamic parameters help estimate the minimum flow velocities needed to entrain nummulitid tests and provide a potential tool to account for the nature of their accumulations.

Nannofossil biostratigraphy, strontium and carbon isotope stratigraphy, cyclostratigraphy and an astronomically calibrated duration of the Late Campanian Radotruncana calcarata Zone.

A section from the southern (Austro-Alpine Northern Calcareous Alps) margin of the Penninic Ocean in the NW Tethys realm of Late Campanian age is investigated stratigraphically. Plankton foraminifer and nannofossil biostratigraphy designate the presence of the Globotruncana ventricosa Zone and the Radotruncana (Globotruncanita) calcarata Zone, and standard nannofossil zones CC21-UC15cTP and CC22ab-UC15deTP. The combination of carbon isotope stratigraphy, strontium isotopes, and cyclostratigraphy allows a detailed chronostratigraphic correlation. Periodicity was obtained by power spectral analysis, sinusoidal regression, and Morlet wavelets. The duration of the calcarata Total Range Zone is calculated by orbital cyclicity expressed in thickness data of limestone-marl rhythmites and stable carbon isotope data. Precessional, obliquity, and short and long eccentricity cycles are identified and give an extent of c. 806 kyr for the zone. Mean sediment accumulation rates are as low as 1.99 cm/kyr and correspond well to sediment accumulation rates in similar settings. We further discuss chronostratigraphic implications of our data.

How to react to shallow water hydrodynamics: The larger benthic foraminifera solution.

Symbiont-bearing larger benthic foraminifera inhabit the photic zone to provide their endosymbiotic algae with light. Because of the hydrodynamic conditions of shallow water environments, tests of larger foraminifera can be entrained and transported by water motion. To resist water motion, these foraminifera have to build a test able to avoid transport or have to develop special mechanisms to attach themselves to substrate or to hide their test below sediment grains. For those species which resist transport by the construction of hydrodynamic convenient shapes, the calculation of hydrodynamic parameters of their test defines the energetic input they can resist and therefore the scenario where they can live in. Measuring the density, size and shape of every test, combined with experimental data, helps to define the best mathematical approach for the settling velocity and Reynolds number of every shell. The comparison between water motion at the sediment-water interface and the specimen-specific settling velocity helps to calculate the water depths at which, for a certain test type, transport, deposition and accumulation may occur. The results obtained for the investigated taxa show that the mathematical approach gives reliable results and can discriminate the hydrodynamic behaviour of different shapes. Furthermore, the study of the settling velocities, calculated for all the investigated taxa, shows that several species are capable to resist water motion and therefore they appear to be functionally adapted to the hydrodynamic condition of its specific environment. The same study is not recommended on species which resist water motion by adopting hiding or anchoring strategies to avoid the effect of water motion.