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Thapsus was one of the Roman Empire's largest harbors and is situated next to an easily defended promontory on Tunisia's coast in northern Africa. It was provided with a huge stone and cement breakwater mole that extended almost 1 km into the sea. We examined sedimentological and micropaleontological proxies from 14C-dated core material and shifts in microfauna and macrofauna community structure to infer patterns of sediment dynamics and the chronology of events that shaped the coastal evolution in the Dzira Lagoon at Thapsus over the past 4000 years. The sedimentological and faunal record of environmental changes reflect a sequence of events that display a transition from an open to a semi-closed lagoon environment. At around 4070 cal yr BP and again between 2079 and 1280 cal yr BP, the data reveal two transgressive events and a deposition of sandy sediments in a largely open marine lagoon environment. The transgressive sands overlay marine carbonate sandstones that are upper Pleistocene in age. A gradual closure of the lagoon from 1280 cal yr BP until today is indicated by decreasing species richness values, lower abundances of typical marine taxa, and increasing percent abundances of fine-grained sediments. The environmental transition from an open to a closed lagoon setting was also favored by the construction of an extensive harbor breakwater mole, changes in longshore current drift patterns, and the formation of an extensive sand spit.

International audience; The Lower Devonian Klerf Formation is an exceptional Konservat-Lagerst€ atte, exposed at multiple sites in the Waxweiler region in the Eifel area, western Germany. It has been studied for its various fossils, mainly arthropods, fishes, plants, molluscs, brachiopods and crinoids. At Waxweiler, the strata are palaeoecologically interpreted as a prograding deltaic depositional system elongate from NW to SE in the Ardenno-Rhenish area. The Klerf Formation has, however, not been studied in full in terms of its microflora and microfauna. Our study of the formation from two different quarries in the Waxweiler area yielded fairly diverse miospore assemblages dominated by abundant organic matter in varying degrees of coalification. The miospore assemblages are mainly composed of classic Lower Devonian taxa of the Old Red Continent (Laurussia). These belong, among others, to the genera Ambitisporites, Apiculiretusispora and Retusotriletes. Biostratigraphically more important species recovered include Acinosporites lindlarensis, Apiculiretusispora brandtii, Cymbosporites asymetricus, Diatomozonotriletes franklinii, Emphanisporites annulatus, Verruciretusispora dubia and Verrucosisporites polygonalis. In addition, Emphanisporites foveolatus, which is known only from a limited area in the Ardenno-Rhenish region, is also identified, indicating an earliest Pragian to middle Emsian age for the composite section. These assemblages are found to be accompanied by reworked phytoplankton to a much lesser extent. Our results reveal a much larger palaeobotanical diversity from the Rhineland outcrops than previously known, indicating a well-developed Psilophyton-type vegetation with related plants. The results further suggest a likely presence of plants such as Leclercqia and Pertica.

Mangrove forests are extremely productive ecosystems, are sources and sinks of organic carbon, and provide essential services both to the marine environment and people. We have studied the composition and species richness of modern benthic foraminiferal assemblages from mangrove swamps along the Mamanguape River in Paraíba, northern Brazil. Sampling points for foraminifera were selected to acquire information on the composition of foraminiferal assemblages from dense mangrove stands collected along a river transect. Almost 100 species of benthic foraminifera were identified within the shallow mangrove habitats. The large number of identified mangrove taxa is the highest recorded so far for true mangrove habitats. The high species richness rivals shallow-water assemblages recorded from nearby offshore and reef environments and indicates that a particularly large number of species is capable of growing and flourishing under multiple stressor conditions. Numerical analysis of the faunal assemblages shows that specific taxa, which were previously known to be uncommon in mangrove environments, are abundant in the Mamanguape River estuary. The atypical foraminiferal fauna found in the Mamanguape River estuary resembles shallow-water offshore assemblages, is characterized by high percent abundances of perforate and miliolid taxa, and contains only very few of the otherwise typical and numerically abundant agglutinated mangrove taxa. The unusual structure of the assemblages recorded provides insight into what combination of environmental variables controls their composition and novel perspectives to reconstruct past mangrove environments. Distribution, diversity, and species-specific analysis will provide guidance on the use of Brazilian mangrove foraminifera as indicators for the strength of tidal activity, pollution, and anoxia in coastal waters and sea-level reconstructions.

Global warming threatens the viability of tropical coral reefs and associated marine calcifiers, including symbiont-bearing larger benthic foraminifera (LBF). The impacts of current climate change on LBF are debated because they were particularly diverse and abundant during past warm periods. Studies on the responses of selected LBF species to changing environmental conditions reveal varying results. Based on a comprehensive review of the scientific literature on LBF species occurrences, we applied species distribution modeling using Maxent to estimate present-day and future species richness patterns on a global scale for the time periods 2040-2050 and 2090-2100. For our future projections, we focus on Representative Concentration Pathway 6.0 from the Intergovernmental Panel on Climate Change, which projects mean surface temperature changes of +2.2°C by the year 2100. Our results suggest that species richness in the Central Indo-Pacific is two to three times higher than in the Bahamian ecoregion, which we have identified as the present-day center of LBF diversity in the Atlantic. Our future predictions project a dramatic temperature-driven decline in low-latitude species richness and an increasing widening bimodal latitudinal pattern of species diversity. While the central Indo-Pacific, now the stronghold of LBF diversity, is expected to be most pushed outside of the currently realized niches of most species, refugia may be largely preserved in the Atlantic. LBF species will face large-scale non-analogous climatic conditions compared to currently realized climate space in the near future, as reflected in the extensive areas of extrapolation, particularly in the Indo-Pacific. Our study supports hypotheses that species richness and biogeographic patterns of LBF will fundamentally change under future climate conditions, possibly initiating a faunal turnover by the late 21st century.

Calcifying organisms such as benthic foraminifera are susceptible to changes in ocean pH and alkalinity. Responses to these changes include variations in mortality, calcification rates or assemblage composition, which have been observed in field and experimental studies. Here we applied a growth experiment with benthic foraminiferal propagules under different pH conditions to gather insights into the effect of pH on the composition of grown assemblages. A homogeneous propagule assemblage from a local mudflat in Corfu Island (Greece) was exposed to a range of pH conditions (6.5, 7.2, 7.8 and 8.5) for 5 weeks. In a second experiment, the assemblages were first exposed to low and subsequently to high conditions for a total of 8 weeks. After termination of the experiments, we recorded high survivability and growth throughout the treatments. Analysis of the assemblage composition of the first experiments revealed a shift from porcelaneous dominated taxa in the higher pH treatments to an assemblage with higher numbers of agglutinated taxa in the lower pH treatments. Soft-shelled monothalamous species were common throughout. The second experiment revealed assemblages that were significantly dominated by porcelaneous taxa with monothalamous taxa being almost absent. The results of this study are congruent with other observations on changing assemblage compositions with changing pH from both laboratory and field studies. The fast response of the assemblages through activation of potentially dormant propagules adds insights into the mechanisms behind seasonal composition changes in naturally variable environments such as river estuaries. They also shed new light on possible effects of continuous decreases in ocean pH on shallow-water foraminiferal assemblages in future.

Sediments across Lagos Lagoon have been sampled and analyzed to investigate the extent and distribution of Potentially Toxic Elements (PTEs). The bioavailable fraction of PTEs have been evaluated to explore the relationship between PTE concentration and the spatial distribution, composition, abundance, and species richness of benthic foraminifera biotas. The sediments have been found to show a wide range reflecting a diffuse contamination, where Contamination and Enrichment Factor suggest low to extremely polluted sediments. Our survey of the benthic foraminifera inhabiting Lagos Lagoon revealed diverse assemblages of benthic taxa, species-specific distribution patterns, gradients of species richness and abundance, and a disjunct distribution of agglutinated and hyaline-perforate/porcelaneous taxa along a pronounced salinity gradient. All PTE total concentrations have been shown to positively correlate with mud and Total Organic Carbon (TOC) and two of the most abundant agglutinated taxa, Ammotium salsum, and Trochammina sp. 1, according to our correlation matrix analysis. Moreover, both species display significant positive correlations with CrF4-CoF2-F3-F4-total-CuF4-total-NiF3-F4-total-AlF4-total-FeF3-F4-total-ZnF3-F4-total. On the other hand, both foraminifers correlate negatively with PbF4-SeF3-Setotal. The overall significant positive correlation of these PTEs suggests that they behave as micronutrients when complexed with organic matter. No significant positive correlation with none of the PTEs in any fraction was found for neither species richness nor for the most abundant hyaline perforate species (Ammonia aoteana). Some PTE fractions were found to correlate either positively or negatively with individual foraminifera species, suggesting that they function as either micronutrients and/or stressors. The resulting Contamination Factor of the PTE total concentrations shows that only a few sample sites can be classified as “moderately” polluted for Cr, Zn, and Cu, and that all sampled sites are classified as “highly polluted” for Se. The highest concentrations for Cr, Cu, Ni, and Zn were found towards the industrialized western part, an area that is characterized by moderate to high diversity but low abundances of benthic foraminifera.

Jullienella foetida is probably the largest agglutinated foraminifer in modern oceans and can reach a length of up to ~14 cm. Because of its large size, the species was initially considered to be a bryozoan, but later correctly described as a single-chambered (monothalamous) foraminifer with a large, flat or slightly undulating plate-like test, leaf-like, or fan-like in overall shape and with the chamber interior subdivided by longitudinal partitions. It occupies a restricted geographical range around part of the NW African margin where it is found in eutrophic settings with a preference for energetic environments.We have applied a suite of non-destructive methods, namely light microscopy, SEM, X-ray and high-resolution micro-computed tomography (micro-CT) to 1) explore its external and internal test characteristics and 2) to provide a first-order estimate of its possible contribution to sea floor biomass. High-resolution SEM images show the test wall to comprises a smooth, outer veneer of small mineral grains that overlies the much thicker inner layer, which has a porous structure and is composed of grains measuring several hundreds of microns in size. X-ray images of the test reveal an elaborate system of radial partitions that subdivides the test interior into channels that may serve to direct the flow of the cytoplasm, and perhaps increase its surface to volume ratio. Micro-CT scans suggest that much of the test interior is filled with cytoplasm with a biomass comparable to that of slightly larger xenophyophores. This remarkable species appears to play an important, perhaps keystone, role in benthic ecosystems where it is abundant, providing the only common hard substrate on which sessile organisms can settle.

Shallow-water banks of foraminiferal propagules (tiny juveniles To date, propagule assemblages have only been analyzed “passively” through growth experiments with propagule banks under laboratory conditions. Here, we apply for the first time a combination of a multi-week cultivation experiment and eDNA metabarcoding of the different size fractions.Sediment samples were taken from a shallow lagoon in Corfu (Greece) and sieved over 63 µm to separate the propagule bank from the coarser fraction. The in-situ material of the sampling site was used as a baseline for the subsequent experiment (T0). The finer fraction (63 µm) every 5 weeks (T1–T3). At the same intervals, samples were taken and re-sieved over 63 µm for eDNA metabarcoding of both size fractions.The morphology-based count data of the foraminiferal specimens revealed significantly different assemblage compositions after each harvest (T0–T3). The differences between the in-situ (T0) and the experimental samples (T1–T3) were most distinct, indicating a specific composition of the propagule bank. The differences between in-situ and experimental assemblages as well as the shifting assemblage compositions over time were mirrored by the metabarcoding data from the respective intervals. Our results highlight the potential for metabarcoding to complement and expand insights gained from morphology-based approaches in foraminiferal studies.

The ecological response of benthic foraminifera to bioavailable Potentially Toxic Elements (PTEs) was evaluated in Lagos Lagoon (Nigeria). We sampled and analyzed PTEs across Lagos Lagoon with the aim to investigate the extent of contaminated sediments, to document their distribution, and to explore the relationship between PTE concentration and the spatial distribution, composition, abundance, and species richness of benthic foraminifera biotas. PTE’s recordings showed a wide range reflecting a diffuse contamination, where Contamination and Enrichment Factor suggest low to extremely polluted sediments. Findings of a previous survey of the benthic foraminifera inhabiting Lagos Lagoon revealed diverse assemblages of benthic taxa, species-specific distribution patterns, gradients of species richness and abundance, and a disjunct distribution of agglutinated and hyaline-perforate/porcelaneous taxa along a pronounced salinity gradient. Correlation matrix analysis shows that except for Selenium, all PTE total concentrations positively correlate with mud and Total Organic Carbon (TOC) and two of the most abundant agglutinated taxa, Ammotium salsum, and Trochammina sp. 1. Moreover, both species display significant positive correlations with CrF4-CoF2-F3-F4-total-CuF4-total-NiF3-F4-total-AlF4-total-FeF3-F4-total-ZnF3-F4-total. On the other hand, both foraminifers correlate negatively with PbF4-SeF3-Setotal. The overall significant positive correlation of these PTEs suggests that they behave as micronutrients when complexed with organic matter. No significant positive correlation with none of the PTEs in any fraction was found for neither species richness nor for the most abundant hyaline perforate species (Ammonia aoteana). Some PTE fractions were found to correlate either positively or negatively with individual species, suggesting that they function as either micronutrients and/or stressors. The resulting Contamination Factor of the PTE total concentrations shows that only a few sample sites can be classified as “moderately” polluted for chromium, zinc, and copper and that all sampled sites are classified as “highly polluted” for selenium. The highest concentrations for Cr, Cu, Ni, and Zn were found towards the industrialized western part, an area that is characterized by moderate to high diversity but low abundances.

We present new observations on

This study explores the paleoenvironmental changes of the Acholla coast over the last 2000 years by means of subsurface sediment analyses of lagoonal deposits via a multiproxy approach. A sequence of 5 steps is shown to have shaped the coastal development at Acholla and includes (i) the settlement of an open marine embayment during the Upper Pleistocene (substage 5e) in response to a transgressive event as evidenced by the deposition of carbonate sandstones rich in mollusc associations and marine elemental ratios Sr/Al, Ca/Al, Cl/Al, and S/Al; (ii) a sea-level rise towards183 AD, after a long period of emergence of the coast, leading to transgressive deposits of poorly sorted fine sands that are rich in diversified mollusc and ostracod taxa; (iii) a high-energy event (towards 417AD) with a peculiar biofacies characterized by coarse bioclastic mollusc-rich sands, charcoal particles, Tyrrhenian lithoclasts, and high values of marine elemental ratios Cl/Al, S/Al, and Ca/Al; (iv) a progradation of coastal habitats (between 417 and 1577 AD), favored by a period of flooding/detrital input and longshore current drifts, resulting in the development of an extensive sandspit and the formation of the Acholla lagoon. The construction of a Roman pier, originally constructed to protect the Acholla harbor from sedimentation, was counterproductive and favored the expansion of coastal swash bars and extension of the sandspit, ultimately resulting in the abandonment of the Acholla harbor; (v) the closure of the lagoon (from 1577 AD onward) and the built-up of well-sorted silt deposits are marked by the decrease of detrital elemental ratios (Ti/Al, Si/Al, and K/Al) and by reduced ostracod species richness values. The inferred scenario for the evolution at Acholla coast matches those for Sfax and Skhira coast in the Gulf of Gabes (Tunisia). Unlike Sfax and Skhira, however, the Acholla coast has been subjected to uplift by active faults, resulting in a time shift of depositional patterns.

Mangrove forests are extremely productive ecosystems, sources and sinks of organic carbon, and provide essential services both to the marine environment and people. We have studied the composition and species richness of modern benthic foraminiferal assemblages from mangrove swamps along the Mamanguape River in Paraíba, northern Brazil. Sampling points for foraminifera were selected to acquire information on the composition of foraminiferal assemblages from dense mangrove stands collected along a river transect. Almost 100 species of benthic foraminifera were identified within the shallow mangrove habitats. The large number of identified mangrove taxa is the highest recorded so far for true mangrove habitats. The high species richness rivals shallow-water assemblages recorded from nearby offshore and reef environments and indicates that a particularly large number of species is capable to grow and flourish under conditions of multiple stressors. Numerical analysis of the faunal assemblages shows that specific taxa, which were previously known to be uncommon in mangrove environments, are abundant in the Mamanguape River estuary. The atypical foraminiferal fauna found in the Mamanguape River Estuary resembles shallow-water offshore assemblages, is characterized by high percent abundances of perforate and miliolid taxa and contains only a very few of the otherwise typical and numerically abundant agglutinated mangrove taxa. The unusual structure of the assemblages recorded provide insight of what combination of environmental variables controls their composition and novel perspectives to reconstruct past mangrove environments. Distribution, diversity and species-specific analysis will provide guidance on the use of Brazilian mangrove foraminifera as indicators for the strength of tidal activity, pollution and anoxia in coastal waters and sea-level reconstructions.

Benthic foraminifera are important indicators for ecological studies. The assemblage composition of local communities can be used to analyze influences of environmental variables such as temperature, salinity, pH, and others. In recent years, the experimental propagule method has emerged as an effective tool to evaluate the influence of these variables on assemblage dynamics of benthic foraminifera. Propagules (tiny juveniles) of benthic foraminifera are widespread and can survive outside of a species’ natural distribution range. Their ability to become dormant and be re-activated once local conditions become suitable, is an important driver behind the capacity of foraminiferal assemblages to react quickly to environmental changes. In the laboratory, the propagules are first separated from the coarser fractions by sieving and then cultured under different conditions.In the present study, we analyzed the effect of ocean pH on the composition of shallow-water assemblages from Corfu Island (Greece). Like other calcifying organisms, assemblages of foraminifera are susceptible to pH variations and have revealed compositional shifts along natural or experimental pH gradients. Our experimental set-up included four pH treatments between 6.5 and 8.5 at constant temperature and salinity (22°C and 38 ppt) for 5 weeks.At the conclusion of the cultivation experiment, we found high numbers of grown specimens (825–1564 per replicate) and a high survivability rate throughout all treatments (78–87%). Higher pH (7.8 and 8.5) resulted in assemblages that were dominated by monothalamous and porcelaneous species, whereas lower pH (6.5 and 7.2) lead to a reduction in porcelaneous and an increase in agglutinated species. Several taxa showed significant positive or negative correlations with decreasing pH values.Our results are congruent with previous findings that reported compositional shifts from calcareous to agglutinated taxa with decreasing pH (both from culture and field observations). Our study also indicates that the activation of propagules is an important mechanism behind assemblage dynamics in shallow-water foraminifera. As such, it offers an improved insight into potential resilience and recovery mechanisms of foraminiferal assemblages with regard to local or seasonal pH variations as well as ongoing ocean acidification.

The human-mediated translocation of marine alien species beyond their natural ranges started as early as people began navigating the sea and is of growing concern to nature conservation. The Mediterranean Sea is among the most severely affected areas by biological invasions, a phenomenon that has been fostered by the opening and recent extension of the Suez Canal, the transport and release of ballast water, aquaculture and aquarium trade, ichthyochory and other active or passive dispersal mechanisms. The increase of marine invasions has stimulated considerable research, but for some important groups, in particular microorganisms, data are still limited. In this paper we have reviewed the current status of marine alien foraminifera in the Mediterranean Sea. Our survey includes a comrephensive taxonomic revision of previously recognized alien taxa, and new information obtained from the fossil record and from molecular studies. Our survey and reexamination of alien benthic foraminifera yielded a total of 43 validly recognized species and two species of cryptogenic taxa and reduces the number of previous recordings. The revised list includes both larger symbiont-bearing and smaller benthic foraminifera, including 16 hyaline-perforate, 3 agglutinated and 24 porcelaneous taxa. The vast majority of alien foraminifera recorded so far have become established in the Eastern and Central Mediterranean Sea, indicative for translocation and introduction via the Suez Canal pathway. Only one species, Amphistegina lobifera, causes significant ecological impacts and fulfills the criteria to be considered as an invasive alien. This species is a prolific carbonate producer, and displays extreme forms of ecosystem invasibility with capabilities to reduce native diversity and species richness. The proliferation and rates of recently observed range extensions, track contemporary sea surface temperature increases, provide strong support for previous species distribution models, and corroborate findings that rising water temperatures, global climate change and the extension of climate belts are major drivers fueling the latitudinal range expansion of larger symbiont-bearing and smaller epiphytic foraminifera. Intensified efforts to study alien foraminifera on a molecular level, in dated cores and in ballast water are required to trace their source of origin, to identify vectors of introduction and to verify their status as true aliens.

The Lagos Lagoon is among Africa’s largest estuarine ecosystems. It is bordered by one of the fastest growing megacities in the world (Lagos) and the ultimate repository of contaminants carried in industrial, municipal and agricultural wastes. The high levels of pollutants have progressively deteriorated the water quality, adversely affected marine ecosystems, impacted the livelihood of the coastal population and pose serious risks to human health. Benthic foraminifera are excellent proxies and sensitive bioindicators of environmental disturbances but comprehensive studies on the structure, distribution, diversity and impact of pollution upon foraminiferal communities have not yet been conducted. In order to demonstrate the potential of foraminifera as proxies of environmental perturbations, benthic foraminifera were investigated on a lagoon-wide basis.Lagos Lagoon has areas that range from low levels of direct impact to severely affected by various forms of anthropogenic disturbance. The goal of this study was to elucidate foraminiferal community structure, to analyze patterns of distribution and species richness and to identify taxa that track documented records of pollution in Lagos Lagoon sediments. The sediments were analyzed for a range of physicochemical properties via a multi-parameter sensor probe-device including temperature, pH, depth and total dissolved solids (TDS) measurements. Quantitative analysis of 24 sediment samples yielded a total 3872 individuals of benthic foraminifera that belong to 42 species and 25 genera. They comprise 10 porcelaneous, 22 hyaline perforate and 10 agglutinated species. Ammobaculites aff. A. exiguus, Ammotium salsum, Ammonia tepida, Ammonia parkinsoniana and Trochammina sp. have been found to be the most abundant species.Heat maps were generated from abundance records for selected species to illustrate environmental preferences and relative resistance levels to individual forms of anthropogenic disturbance. The features recorded allow to delineate the spatial effects of hydrocarbon and heavy metal pollutants, urban sewages, and sand dredging activities. The data generated may ultimately form the basis to assess the progressive deterioration of Lagos Lagoon environments from cores by using benthic foraminifera as bioindicators of environmental perturbation.

Raja Ampat (Indonesia) is a remote archipelago west of Papua mainland and situated at the eastern edge of the Coral Triangle. The archipelago is considered one of the species-richest and most pristine localities and home to the world’s most diverse coral reefs. For centuries, reefs of Raja Ampat have enjoyed natural protection and remained largely untouched due to their isolated location. The region represents a key area for evaluating richness and biogeographic patterns of tropical shallow-water organisms, yet the foraminiferal fauna of Raja Ampat is virtually unexplored. Benthic foraminifera are an integral part of the reef fauna, prolific carbonate producers and essential for reefal accretion and substrate stability. Previous studies on Indo-Pacific benthic reef foraminifera indicate that diversity is highest in the Central Indo-Pacific and broadly correlates with diversity patterns of other tropical marine taxa. This report presents the first illustrated catalog and comprehensive analysis of the structure, composition, and diversity of the species-rich benthic foraminiferal biotas of the Raja Ampat Archipelago including an assessment of local reef vitality with the Foraminifera in Reef Assessment and Monitoring (FoRAM) Index. We examined shallow-water sediment samples from fringing reefs, reef platforms, sheltered bays and reef-associated environments to cover the full range of existing micro- and macrohabitats. Atotal number of 421 species were recovered, among them five new species and one newly described genus of the porcelaneous Miliolida. The fauna is dominated by hyaline taxa but the Miliolida represent the species-richest order and are especially diverse in fine-grained sediments of deep fore-reef slopes. Extraordinary high diversity of the benthic foraminiferal communities indicates that the waters of Raja Ampat probably represent one of the world’s biologically richest locations in reefs.We identified a total of 35 species of larger symbiont-bearing benthic foraminifera, a number that is among the highest reported so far for reefs in modern oceans. The analysis of the FoRAM Index revealed that water quality at all sites examined is suitable for reef growth and recovery. The results of our study show that the Raja Ampat Archipelago harbors unique and particularly diverse assemblages of modern benthic foraminifera and provides a protistan perspective for the hotspot of diversity in the Central Indo-Pacific.

Tropical tide and rock pools are extreme environments with regard to temperature. They undergo diurnal heating and constitute natural laboratories to assess the fate of marine calcifiers under scenarios of global change. This study focuses on benthic foraminiferal assemblages from tide and rock pool sites from eastern Africa as “natural” laboratories to document the diverse spectrum of foraminiferal biotas potentially capable to succeed under conditions of future global warming. To date, the majority of foraminiferal thermal tolerance data have been compiled from laboratory experiments, anthropogenically heat-polluted or extreme hydrothermal vent areas. A total of 111 species of benthic foraminifera were recorded within the tide pool samples. Perforate species were most abundant (up to 84%), followed by porcellaneous taxa (up to 45%), while agglutinated species were only rare (up to 5%). Larger benthic foraminifera represented between 30 and 75% of the total assemblages with 8 observed genera. Species richness varied between 23 and 63 species per sample. The foraminiferal communities of most sites were dominated by larger perforate taxa such as Neorotalia calcar and Amphistegina spp., followed by a variety of small miliolid taxa (54 species) and small rotaliids (27 species). The surprisingly high foraminiferal species richness suggests that a large range of shallow-water benthic taxa appear to be capable to tolerate exposure to diurnally occurring temperature extremes (35°–> 40 °C). The large spectrum of benthic taxa recorded provides insight into potential acclimatization capacities of foraminiferal assemblages with regard to ongoing ocean warming and projected temperature changes. We infer that calcification in heat-tolerant benthic foraminifera, a group of productive carbonate producers, is more widespread than previously thought and will continue in future decades when water temperatures are significantly elevated. Tolerance to thermal stress is imperative for intertidal foraminifera and assemblages from tropical rock and tide pools appear to be less susceptible to future change. This capacity is central to the resilience to rising ocean temperatures and dictates how warming affects these delicate environments. Intertidal foraminiferal assemblages could potentially act as source populations for restocking and to mitigate detrimental effects of global change.

The sequence stratigraphic architecture of the Neogene section from the subsurface Nile Delta is resolved based on exceptionally well-preserved microfossils from the offshore NDOB-1 borehole. The architecture is configured on bathymetry variations and statistical parameters as deduced from the relative abundance of foraminiferal associations. The statistical parameters include ratios of planktic versus benthic foraminifera, and cluster and correspondence analysis of the twelve most commonly used benthic foraminiferal genera as proxies of water depths. The combination of cluster and correspondence analysis was employed to decipher the evolution and dynamics of the basin and the mechanisms that controlled the deposition of Neogene sequences in the Nile Delta region. Four basic environmental settings were recognized: 1.) a middle to outer neritic eutrophic setting, 2.) a middle to outer neritic mesotrophic setting, 3.) an outer neritic to upper bathyal mesotrophic setting and 4.) an upper bathyal mesotrophic setting. Eight stratigraphic sequences were identified in the Neogene Nile Delta section. Three sequence in the Miocene (MSeq1, MSeq2 and MSeq3), three in the Pliocene (PSeq1, PSeq2 and PSeq3) and two in the Pleistocene (PtSeq1 and PtSeq2). These sequences are systematically measured and described in terms of time, space and water bathymetry. The sequence boundaries and flooding surfaces were dated using high-resolution microfossil biochronology and stratigraphic index markers. Individual sequences and boundaries were correlated with international and local sequence stratigraphic models. The new sequence stratigraphic model established here provides age calibrated surfaces for inter-basinal correlations and opens new avenues for hydrocarbon reservoir exploration.

The disproportionately low presence of marine species in the list of invasive alien species (IAS) of Union concern of the European Union (EU) Regulation 1143/2014 does not fully acknowledge the threat they pose to the EU marine environment. In this study, the first EU-scale Horizon Scanning (HS) focusing on marine alien species was performed, aiming to deliver a ranked list of species that should be of high priority for risk assessment (Article 5 of the EU IAS Regulation). Species absent from or with a limited distribution in EU marine waters were targeted. In total, 363 alien species were initially screened for HS by a panel of experts, including a broad range of taxonomic groups. Species were scored for their likelihood of arrival, establishment, spread, and impact in EU waters. A consensus workshop ranked 267 species, including a subset of 26 prioritized species. These species are considered to be mainly introduced by shipping (fouling and ballast water), via the Suez Canal, and aquaculture activities. The 26 priority species were also scrutinized in terms of feasibility of their management; 18 of them were suggested for performing risk assessments on the basis of the EU IAS Regulation. Since biological invasions are dynamic and connected with accelerated globalization and diversified human activities, we recommend HS to be repeated periodically to review the species already listed and assess new ones.

The Mediterranean Nile Delta is a major target for hydrocarbon reservoirs. In the search for hydrocarbons, high-resolution biostratigraphy has long been established as high-impact and low-cost solution to face the challenges in the move towards structurally more complex and unexplored areas. To date, the biozonation of the Nile Delta is largely provided by benthic and planktonic index foraminifera. We have examined the foraminifera assemblages from NDOB-1, a borehole in the northern part of the Nile Delta. The material recovered provides unique and exceptionally well-preserved microfossil biotas that completely cover the Neogene succession (Middle Miocene to Pleistocene). This allows the establishment of an intact biozonation scheme for the Nile Delta region with reliable chronostratigraphic correlation and suitable industrial applications. The stratigraphic succession of the Nile Delta region consists of seven formations: Sidi Salim Formation, Qawasim Formation, Rosetta Formations, Abu Madi Formation, Kafr El Sheikh Formation, El Wastani Formation and Mit Ghamr Formation. Examination of the planktonic foraminifera assemblages of the NDOB-1 borehole from the deep part of the Nile Delta lead to the identification of twelve biozones: Globorotalia inflata Interval Zone, Globorotalia aemiliana Interval Zone, Globorotalia puncticulata Interval Zone, Globorotalia puncticulata- Globorotalia margaritae Concurrent Range Zone, Globorotalia margaritae Interval Zone, Sphaeroidinellopsis seminulina Acme Zone, Non-distinctive Zone, Globorotalia suterae Interval Zone, Globigerina woodi Interval Zone, Globorotalia mayeri Interval Zone, Globorotalia fohsi peripheroronda Interval Zone and Praeorbulina glomerosa Interval Zone. The biozones fully cover the time span from the Middle Miocene to the Pleistocene. The benthic foraminifera recovered allow the identification of five benthic biozones for the Pliocene and Pleistocene: Hyalinea balthica Interval Zone, Bulimina marginata Interval Zone, Bulimina aculeata Interval Zone, Gyroidinoides laevigata Interval Zone and Uvigerina rutila Interval Zone. The identified planktonic bioevents were used to calibrate the studied Neogene section with the Astronomically Tuned Neogene Time Scale (ATNTS, 2012).

The Ologe Lagoon in southwestern Nigeria is a natural habitat located closely to both residential and industrial estates. The lagoon is heavily affected by effluents of agricultural and industrial runoff including organochlorine compounds and heavy metals (chloropesticides, DDT and polychlorinated biphenyls, cadmium, manganese, zinc, copper, lead, iron and nickel). Ologe Lagoon is situated in a semi-isolated position and its brackish waters traverse several creeks and lagoons before it reaches the Atlantic Ocean. The lagoon experiences minor tidal but noticeable salinity and pH variations during the rainy season. During the dry season, however, the waters have a comparatively long resident time where sediments act as sink and aggregate contaminants over time. Investigations on benthic foraminifera, thecamoebian and diatom species were carried out to document the current status, composition, spatial distribution, and response of microfaunal assemblages to environmental stressors within the lagoon. The benthic foraminifer assemblage is composed of four highly specialized agglutinated species of Ammotium salsum, Miliammina fusca, Ammobaculites dilatatus and Ammobaculites aff. A. exiguus. They are present at all sample sites with varying abundances. The high abundance of Ammotium salsum is considered an indication of environmental stress. Thecamoebian species of Difflugia oblonga, Difflugia proteiformis and Pontigulasia compressa were also recorded in varying proportions and generally represent less than one percent of the total foraminifera/thecamoebian assemblage. The presence of marine brackish water larger siliceous diatoms of Actinoptychus and Campylodiscus species indicates an interaction between the lagoon and the marine waters although on the minimal scale with regards to actual salinity ranges. The faunal assemblages recovered are characterized by low species richness and alpha diversity indices and are typical of shallow, marginal-marine hyposaline environments. Numerical abundances and spatial distribution patterns suggest taxon-specific habitat preferences and highlight their potential for environmental monitoring studies.

New data evaluating the relative sea level changes and sedimentary dynamics along the coast of Tunisia are presented. Data are inferred from archaeological sites of Punic-Roman age located between Ras Boutria (Jebeniena) and El Awebed (Sfax). New results are based on high-resolution micropalaeontological, sedimentological and geochemical analyses of several sediments cores coupled to new 14C datings. The coast of Jebeniana (Ras Boutria historic site) recorded the first marine transgression, overlying the quartz sand Tyrrhenian deposits, 2210 a BP in age. This transgression, also recognized in Sfax coast, revealed by the dominance of lagoonal ostracod associated to coastal and marine taxa, allows the settlement of an open lagoon. It reveals also, a high value of species number and a reduced density. The pelagic influences are augmented by the high marine geochemical elements concentrations such as the chloride (Cl) and the sulfur (S). The increase of the brackish ostracod mark the settlement of a closed lagoon toward 768 a BP. The enrichment of the sediment in quartz grains coupled with the action of the current drift authorized the build-up of sandy spits and the genesis of the lagoon that preserve a rich assemblage of euryhaline ostracods and mollusks species. This process was probably responsible of the obstruction of the historic Boutria harbor and the abandon of the historic city.

The quantitative study of foraminiferal associations coupled with geochemical and sedimentological proxies, of a 94-cm sediment core from Korba lagoon (Cap Bon, Tunisia), enabled us to better understand the dynamics of depositional environments and identify different stages of evolution of the lagoon. Three major periods were identified. The first revealed high abundance of foraminifera dominated by lagoonal taxa associated with high TOC, TIC and Ca contents. These parameters marked the settlement of a closed lagoon separated from the open ocean by a sandy bar with high foraminiferal productivity in fine sediments. The second period showed the development of estuarine foraminifera coupled with the appearance of charophyte oogonia in addition to planktonic foraminifera, high Ti and Fe concentrations and dominant coarse fraction. These parameters are indicative of an open lagoon environment with the fluvial inputs. The disappearance of charophyte oogonia and planktonic foraminifera coupled with the enrichment of marine taxa marked the opening of our lagoon to the sea. The progressive modification of foraminiferal association revealed by the increase of lagoonal species and the decrease of estuarine taxa prove the settlement of the present-day lagoon conditions.

A re-examination of marine alien species or Non Indigenous Species (NIS) reported in Italian Seas, until December 2018, is provided, focusing on establishment success, year of first record, origin, potential invasiveness, and likely pathways, in particular. Furthermore, their distribution is assessed according to the marine subregions outlined by the European Union (EU) Marine Strategy Framework Directive: Adriatic Sea (ADRIA), Ionian Sea and Central Mediterranean Sea (CMED), and Western Mediterranean Sea (WMED). In Italy, 265 NIS have been detected with the highest number of species being recorded in the CMED (154 species) and the WMED (151 species) subregions, followed by the ADRIA (143) subregion. Most of these species were recorded in more than one subregion. One hundred and eighty (180 or 68%) NIS have established stable populations in Italian Seas among which 26 have exhibited invasive traits. As regards the taxa involved, Macrophyta rank first with 65 taxa. Fifty-five of them are established in at least one subregion, mostly in the ADRIA and the CMED. Crustacea rank second with 48 taxa, followed by Polychaeta with 43 taxa, Mollusca with 29 taxa, and Fishes with 28 taxa, which were mainly reported from the CMED. In the period 2012-2017, 44 new alien species were recorded, resulting in approximately one new entry every two months. Approximately half of the NIS (~52%) recorded in Italy have most likely arrived through the transport-stowaway pathway related to shipping traffic (~28% as biofoulers, ~22% in ballast waters, and ~2% as hitchhikers). The second most common pathway is the unaided movement with currents (~19%), followed by the transport-contaminant on farmed shellfishes pathway (~18%). "Unaided" is the most common pathway for alien Fishes, especially in the CMED; escapes from confinement account for ~3% and release in nature for ~2%. The present NIS distribution hotspots for new introductions were defined at the first recipient area/location in Italy. In the ADRIA, the hotspot, Venice, accounts for the highest number of alien taxa introduced in Italy, with 50 newly recorded taxa. In the CMED subregion, the hotspots of introduction are the Taranto and Catania Gulfs, hosting 21 first records each. The Strait of Sicily represents a crossroad between alien taxa from the Atlantic Ocean and the Indo-Pacific area. In the WMED, bioinvasion hotspots include the Gulfs of Naples, Genoa and Livorno. This review can serve as an updated baseline for future coordination and harmonization of monitoring initiatives under international, EU and regional policies, for the compilation of new data from established monitoring programs, and for rapid assessment surveys.

Foraminiferal propagule banks occur in fine sediment fractions that contain small individuals of benthic foraminifera. These sediments include locally sourced juveniles and propagules, as well as allochthonous propagules that have dispersed from surrounding areas. Such propagules can remain viable even under unfavorable local conditions. When exposed to more favorable conditions, they may grow to adult stages. Accordingly, during environmental changes, propagule banks have the potential to function as species pools and allow quick assemblage reactions. The propagule method was designed to study responses of foraminiferal assemblages by exposing propagule banks to controlled conditions in the laboratory, an approach that is applicable to a variety of ecological questions. Therefore it is important to understand the nature and dynamics of propagule banks, including local and seasonal influences. To obtain insights into the composition of local propagule banks, we studied experimentally grown assemblages from two shallow-water lagoons on Corfu Island in western Greece, and compared the results with in situ assemblages. We sampled in spring and autumn of 2017 and experimental treatments included the use of different substrates in our experiments to account for potential effects on assemblage compositions. Results revealed that sediments from each lagoon contained a distinct propagule bank. We found abundant allochthonous taxa among specimens grown in all experimental treatments, indicating dispersal of propagules, and possibly also juveniles, from adjacent regions into both lagoons. The time of sampling had a significant effect on experimental assemblages, indicating that the composition of propagule banks can vary throughout the year. However, no significant differences were found in assemblages grown in different substrata, suggesting a stronger influence of water variables (e.g., temperature or salinity) on assemblage compositions. Moreover, the experimental set-ups favored small, fast-growing, sediment-dwelling species tolerant of relatively high organic content. Our findings highlight the potential of propagule banks as species pools and will help to refine and improve future applications of the method.

This study updates the current distribution, range expansion and establishment status of the non-indigenous species Amphistegina lobifera Larsen, 1976 and other foraminifera that are cryptogenic in the Sicily Channel. Prior to this study, amphisteginids were reported from the Levantine Basin, the Central Mediterranean (Tunisia, Malta, Pelagian islands) and the southern Adriatic Sea. Here, we provide new records documenting a north-western expansion in the Central Mediterranean. In summer-autumn 2017 and spring-summer 2018, we collected algae and sediment samples from shallow coastal habitats along the shores of the Maltese archipelago, southern and north-western Sicily, Pantelleria and the Aegadian islands. Analysis of the foraminiferal assemblages showed that A. lobifera is effectively established around Malta and in southern/south-eastern Sicily, and has reached the oceanographic boundary between the Central and Western Mediterranean. Our results also show that the thermotolerant A. lobifera is at an advanced stage of invasion in the Sicily Channel, probably favoured by a recent rise in Mediterranean sea surface temperatures. New species distribution models are provided for the years 2040–2050 and 2090–2100, indicating that the predicted warming trend will facilitate north-westward migration of Mediterranean amphisteginids along the coast of northern Africa into the Alboran Sea, and deep into the Adriatic Sea.

Two new genera and eight new species of benthic foraminifera are described from the shallow water, tropical lagoon of Madang, Papua New Guinea. The new hauerinid genus Pseudolachlanella is characterized by juvenile cryptoquinqueloculine, adult almost massiline arranged chambers, and a slitlike, curved aperture with parallel sides and a long, slender, curved miliolid tooth. Pitella haigi n. gen., n. sp. is a new foraminifera with cryptoquinqueloculine arranged chambers, an almost entirely pitted shell surface (pseudopores) and a rounded aperture with a short simple tooth. Among the other species described as new are four hauerinids and two agglutinated foraminifera All new species described here occur sporadically in the shallow water back- and forereef environments of the lagoon (0–55m), and live infaunally and epifaunally in well-oxygenated, fine and coarse grained biogenic sediments. They are absent in muddy, organic-rich, low-oxygen sedimentary environments within bay inlets where variations of salinity are considerable.

Changes in paleoecology and climate of northern Tunisia during the last 3000 years were reconstructed based on the flood history interpreted from a 172-cm sediment core of Lake Ichkeul (NW Tunisia). Seven wet/dry episodes were identified based on biological (ostracods, foraminifera and mollusks) and biotic indices (H and E index, species richness and abundance). These proxy-based environmental changes were supported by correspondence analyses (CAs) and ecophenotypic responses of the brackish taxon Cyprideis torosa in addition to a grain-size study. Two dry episodes were identified in the lower (EP1) and upper (EP7) parts of the core. These were marked by the dominance of the brackish ostracod C. torosa. High water salinity was indicated by the presence of the brackish ostracod Loxoconcha elliptica, the foraminifer Ammonia sp. and lagoonal mollusks. The occurrence of the freshwater ostracods Ilyocypris sp., Herpetocypris sp., Dawinula stevensoni and Limnocythere inopinata was associated with high species richness and ecophenotypic changes of C. torosa valves indicating lower salinity during three major wet episodes (EP2, EP4, and EP6). The ecological and environmental changes, occurring between 3050 and 50 cal. year BP, are most likely linked to an increase of fluvial inputs which are also recognized in several other Mediterranean lakes. The humid episodes were interrupted by two saline periods (EP3 and EP5) during which freshwater ostracod assemblages declined, diversity indices dropped to the lowest values, and fine-grained sediments became dominant. The top of the core is characterized by the remarkable dominance of L. elliptica coupled with C. torosa, the absence of freshwater ostracods, and the changes in grain-size sediment parameters which are most likely the result of anthropogenic activities. The changes in hydrochemistry and sedimentology were attributed to the deepening of the Bizerte navigation canal, main wadis damming, and Tinja sluice construction which mostly occurred in the XXth century.

Symbiont-bearing Larger Benthic Foraminifera (LBF) are ubiquitous components of shallow tropical and subtropical environments and contribute substantially to carbonaceous reef and shelf sediments. Climate change is dramatically affecting carbonate producing organisms and threatens the diversity and structural integrity of coral reef ecosystems. Recent invertebrate and vertebrate surveys have identified the Coral Triangle as the planet’s richest center of marine life delineating the region as a top priority for conservation. We compiled and analyzed extensive occurrence records for 68 validly recognized species of LBF from the Indian and Pacific Ocean, established individual range maps and applied Minimum Convex Polygon (MCP) and Species Distribution Model (SDM) methodologies to create the first ocean-wide species richness maps. SDM output was further used for visualizing latitudinal and longitudinal diversity gradients. Our findings provide strong support for assigning the tropical Central Indo-Pacific as the world’s species-richest marine region with the Central Philippines emerging as the bullseye of LBF diversity. Sea surface temperature and nutrient content were identified as the most influential environmental constraints exerting control over the distribution of LBF. Our findings contribute to the completion of worldwide research on tropical marine biodiversity patterns and the identification of targeting centers for conservation efforts.

Amphisteginid foraminifera are larger symbiont-bearing foraminifera that are abundant in tropical and subtropical reef and shelf regions of the world’s oceans. There is now unequivocal evidence that climate change has led to an expansion of tropical belts. As temperatures rise, surface isotherms are shifting poleward and habitat ranges are moving towards higher latitudes. Fueled by substantial warming, the Mediterranean has been identified as one of the most severely affected areas. This study documents for the first time that amphisteginid foraminifera have now crossed the Strait of Otranto and invaded the eastern Adriatic coast along southern Albania. Sampling on the opposite side along the southern part of the Italian boot shows, however, that amphisteginid foraminifera have not yet colonized the western coast of the Adriatic. The asymmetric invasion of the Adriatic displays spatial heterogeneity, where the progressive colonization follows the major surface currents with a northerly flow along the eastern coast and a southerly return along the western coastline. Previous sampling along Ionian Sea sample sites provides a baseline chronology allowing computations of range expansion rates. The observed recent rate of range expansion in Amphistegina is computed between 4.0 and 10 km/year−1 and provides strong support for current species model projections. Based on the new data, the range boundary shift is projected to lead to a total northward range expansion of 5.2° latitude in the year 2100. The latitudinal range extension computed for the Mediterranean is thus almost twice as large as those computed for the southern hemisphere. The ongoing range extension into new areas of the Mediterranean Sea is shown to trigger changes in community structures with potential consequences for ecosystem functioning. Numerical abundances and impacts of invasive amphisteginids are greatest at shallow depths

The description of two subspecies, Miscellanea juliettae pfenderae Leppig and M. juliettae villattea Leppig, is emended and a new holotype is designated. Both subspecies were previously described as subordinate units of Miscellanea juliettae, a species that has never been formally described. Here we provide the formal description of the species Miscellanea juliettae n. sp., and show that both subspecies are intergrading units of the same species. This results in retaining a single taxonomic unit, Miscellanea juliettae n. sp., leading us to abandon previous designations of subspecies rank.

Oral diseases such as dental caries, edentulism (tooth loss), periodontal disease (PD), and oral cancer currently constitute an increased major public health burden across the globe, with significant differences between countries. One of the main drivers of caries, edentulism, and PD is the excessive intake of sugars. Here, we aimed to quantify the global sugar-related dental health and cost burden in the year 2010. This study used a health-econometrical model to calculate the disease burden as well as the direct and indirect costs attributable to the intake of free sugars (mono- and disaccharides [MDS]). To this end, several databases from the Institute for Health Metrics and Evaluation (IHME), Organisation for Economic Co-operation and Development (OECD), Food and Agriculture Organization (FAO), and World Bank were used. In total, the corresponding disease burden in 168 countries and economic burden in 31 OECD countries were quantified. In 2010, the consumption of MDS was associated with a global dental disease burden of 4.1 million disability-adjusted life years (DALYs; 95% uncertainty interval [UI]: 2.1 to 7.4 million DALYs), with 2.7 million DALYs from MDS-related caries and 1.4 million DALYs from PD. In terms of economic costs, MDS-related dental diseases were associated with a global financial burden of 172 billion US dollars (USD; 95% UI: 91 to 295 billion USD), the largest share of which (151 billion USD) was incurred in OECD countries. Overall, 26.3% (95% UI: 13.3% to 47.5%) of the total global oral disease burden was attributed to the consumption of MDS. The present study emphasizes the need to further address the role of free sugars in oral health and nutrition policy. Although the largest share of the economic burden was accounted for by OECD countries, emerging economies should address this challenge early on in national public health policies if they are to avoid disease and the prospect of increased cost burdens.