Benthic Foraminiferal Communities Research Articles

Benthic foraminiferal community structure and its response to environmental factors revealed using high-throughput sequencing in the Zhoushan Fishing Ground, East China Sea

Benthic foraminifera are excellent tools for monitoring marine environments and reconstructing paleoenvironments. This study investigated the structure and diversity of benthic foraminiferal communities in 20 superficial sediment samples obtained from the Zhoushan Fishing Ground (ZFG) using high-throughput sequencing based on small subunit ribosomal DNA and RNA amplification. The results revealed Rotaliida as the most dominant group, with spatial heterogeneity in foraminiferal distribution. Total benthic foraminiferal communities exhibited higher species richness and diversity compared to active communities. While heavy metal pollution in the ZFG was moderate, areas with elevated concentrations of heavy metals exhibited low diversity and richness in foraminiferal communities. Total foraminiferal community structure was primarily influenced by factors such as water depth and Hg, Pb, Cd, and Zn levels. Notably, Hg levels emerged as a critical factor impacting the structure and diversity of the active foraminiferal community. The dominant species, Operculina, exhibited tolerance toward heavy metal pollution.

Distinct oxygenation modes of the Gulf of Oman over the past 43 000 years – a multi-proxy approach

Abstract. Changing climatic conditions can shape the strength and extent of the oxygen minimum zone (OMZ). The presence and variability of the OMZ in the Arabian Sea is of importance to the latter's ecosystem. The state of oxygenation has, for instance, an impact on the pelagic and benthic faunal community or the nitrogen and carbon cycles. It is important to understand the dynamics of the OMZ and related marine environmental conditions because of their climate feedbacks. In this study, we combined three independent proxies to reconstruct the oxygenation state of the water column and bottom water in the Gulf of Oman for the past 43 kyr approximately. This multi-proxy approach is done for the first time at the northeastern Oman margin located in the Gulf of Oman. We used bulk sedimentary nitrogen isotopes (δ15N) and the alkane ratio (lycopane +n-C35)/n-C31 and benthic foraminiferal faunal analysis to reconstruct the strength of the OMZ in the water column and bottom water oxygenation, respectively. Our results show that the Gulf of Oman experienced strong pronounced OMZ and bottom water deoxygenation during the Holocene. In contrast, during Marine Isotope Stage 2 (MIS 2), including the Last Glacial Maximum (LGM), the Gulf of Oman was very well ventilated, with a highly diverse benthic foraminiferal community. This may have been caused by stronger wind-induced mixing and better ventilation by oxygen-rich water masses. Our results also show moderate oxygenation during MIS 3, with deoxygenation events during most of the warmer Dansgaard–Oeschger (D–O) events. We propose two distinct oxygenation modes for the Gulf of Oman: (1) a stable period of either strongly pronounced water column OMZ and bottom water deoxygenation or well-oxygenated water column and bottom water conditions and (2) an unstable period of oscillating oxygenation states between moderately oxygenated (stadials) and deoxygenated (interstadials, D–O events) conditions. The unstable period may be triggered by an interstadial Atlantic meridional overturning circulation (AMOC) mode, which is required to initiate D–O events.

Biogeochemical impact of historical submarine mine tailings on benthic ecosystems in the Repparfjord (Northern Norway)

Historical copper mine tailings deposited in the Repparfjord, Northern Norway, provided new insight into the biogeochemical impact of submarine tailings disposals on high-latitude coastal ecosystems. The submarine tailings disposal in the Repparfjord represents a product of mining activities between 1972 and 1979. Their environmental impact has been extensively studied during the last decade, but geochemistry of the sediment pore water, which is crucial to assess and monitor the in-situ metal leaching and bioavailability, has never been analysed. The actual impact on the benthic fauna remains poorly known. Therefore, this study couples the pore water chemistry and the foraminiferal analysis obtained from selected sediment cores (gravity core, multicore, box cores) to examine metal stability and the past and current status of the foraminifera community. We measured down-core sulfate and trace metal concentrations and Eh-Ph and applied the Shannon index, the AZTI's Marine Biotic Index (F-AMBI) index and the foraminiferal abnormality index. This study confirms the ongoing leaching of Cu from the underlying mine tailings and release across the sediment-water interface. Leaching of Ni, Zn and Pb have been attributed to weathering of natural bedrock lithologies. The original benthic foraminiferal community disappeared almost entirely during the disposal period, and now it is dominated by stress-tolerant and opportunistic species like Bulimina marginata and Spiroplectammina biformis. Anyhow, against previous assumptions, the community composition changed, while the overall diversity and abnormalities (FAI) shell formation is unaffected by elevated Cu concentrations.

Benthic foraminiferal assemblages in the coastal zone of the yellow river delta: Environmental significance and characteristics of microbial diversity

Unicellular eukaryotes known as benthic foraminifera have sophisticated survival mechanisms and ecological environmental indicators. Ten surface samples from the Yellow River Delta's coastal region were taken for this study to analyze the distribution of foraminifera in relation to environmental factors and to examine their microbial diversity. We looked at the physicochemical aspects of the environment at each sampling site (such as pH, TN, TOC, EC, δ13C, δ15N, PAHs, etc.), and we employed 16S rRNA high-throughput sequencing analysis to look into the interactions between microbial communities and foraminiferal species. The findings demonstrated that the Yellow River Delta's advantage benthic foraminiferal species were Quinqueloculina complanata, Ammonia beccarii, and Ammonia aomoriensis, and that TOC and TN were significant determinants of the distribution of benthic foraminiferal communities. PAH enrichment in the coastal zone effects on microbial communities and benthic foraminifera are not yet readily apparent. Sediment organic carbon and nitrogen isotope analysis revealed that marine plankton may be the dominant source of organic matter in the coastal zone sediments in the research area, which was made up of both land-based and marine organic matter. It implies that foraminifera have some environmental indicators, especially when combined with the distributional traits of the benthic foraminiferal assemblages and their considerable association with environmental parameters. The findings revealed that Q. complanata was found in the Yellow River water-affected estuarine coastal zone, A. aomoriensis was significantly influenced by sediment organic matter content and reflected the estuarine and nearshore environments and the combination of Ammonia beccarii-Ammonia aomoriensis-Elphidium excavatum assemblage indicated a semi-open intertidal shallow marine environment. Additionally, benthic foraminifera showed a responsive association with microorganisms, indicating that microbial diversity may be one of the driving forces behind benthic foraminifera's ability to adjust to environmental changes. The findings of this study will open up new avenues for research into the environmental importance of coastal zone ecosystems and for understanding how benthic foraminiferal communities survive.

Deciphering the impact of decabromodiphenyl ether (BDE-209) on benthic foraminiferal communities: Insights from Cell-Tracker Green staining and eDNA metabarcoding

This study investigates the ecotoxicological effects of BDE-209, a persistent organic pollutant (POP) prevalent in Kuwait's coastal-industrial areas, on benthic foraminiferal communities. We conducted a mesocosm experiment in which we exposed benthic foraminiferal communities sampled from the coastal-industrial areas of Kuwait to a gradient of BDE-209 concentrations (0.01 to 20 mg/kg). The impact of exposure was assessed using live-staining and metabarcoding techniques. Despite the significantly different taxonomic compositions detected by the two techniques, our results show that BDE-209 significantly affects foraminiferal communities, with moderately high concentrations leading to reduced α-diversity and considerable taxonomic shifts in both molecular and morphological assemblages. At concentrations of 10 and 20 mg/kg, no living foraminifera were detected after 8 weeks, suggesting a threshold for their survival under BDE-209 exposure. The parallel responses of molecular and morphological communities confirm the reliability of both assessment methods. This study is the first to investigate the reaction of eukaryotic communities, specifically foraminifera, to POPs such as BDE-209, generating valuable insights that have the potential to enhance field studies and aid the refinement of sediment quality guidelines.

Assessing the effect of Amphistegina lobifera invasion on infralittoral benthic foraminiferal assemblages in the Sicily Channel (Central Mediterranean)

This article documents, through a quantitative approach, the negative effect of the highly invasive species Amphistegina lobifera Larsen, 1976 on native benthic foraminiferal assemblages of coastal areas in the Sicily Channel (Central Mediterranean). A nested sampling design was applied through the comparison of benthic foraminiferal community structure across three areas that are known to be at different stages of invasion (i.e. Maltese Islands - advanced, southern Sicily – medium, and eastern Sicily - early). Results suggested that both diversity and richness of benthic foraminiferal community from the Maltese Islands were strongly modified by increased abundances of A. lobifera. In contrast, this phenomenon is less evident in southern and eastern Sicily, where the invader displayed lower abundances and the community structure was more diversified. Collected data also allowed for predicting what could happen in the near future in the whole Sicily Channel, as well as in the rest of the Mediterranean Sea.

Deep-sea response to interglacial-glacial variability on the South Australian margin over the last 94 ka

The continuous record offered by deep-sea sediments has been extensively used to constrain shifting continental and oceanographic conditions. Yet, past fluctuations in deep-sea benthic conditions and bottom-currents are in numerous parts of the globe scarcely documented, one such example being the South Australian margin. Indeed, though variations in surface water masses and continental aeolian dust and river outflow are well documented in the area, little is known about benthic environments and their dynamics during the last interglacial-glacial cycle. We focus here on benthic foraminiferal assemblages sampled from a sediment core recovered at 2420 m depth from a small plateau south of Kangaroo Island within the underwater Murray Canyons Group (South Australian margin). Benthic foraminiferal assemblages show a distinct separation between interglacial and glacial periods over the last 94 ka, and indicate that the benthic environment was well-ventilated and oligotrophic during glacial periods, whilst being rather marked by reduced oxygenation associated to higher food input during the Holocene and Marine Isotope Stage 5a-c. We demonstrate that autochtonous deep-sea benthic foraminiferal communities neither respond to changes in the Murray River's discharges, nor do they follow variations in aeolian dust input from South Australia. Instead, the deep-sea and the terrestrial realm appear decoupled. Moreover, our observations suggest that bottom-water slope currents were stronger during the Holocene and Marine Isotope Stage 5a-c. We propose that this strengthening was triggered by an intensification of the poleward-circulating deep eastern boundary current transporting carbon-rich Indian Deep Water. In contrast, glacial seafloor conditions, especially during the Last Glacial Maximum, may reflect a greater influence and a shoaling of oxygen-rich Antarctic Bottom Water of South Australia. This bottom-water shift would follow the northward displacement of the Subtropical and Subantarctic Fronts and coincide with a withering influence of the Leeuwin Current within surface waters.

Benthic foraminifera as bio-indicator of marine pollution in the southwestern Bay of Bengal, India.

The benthic foraminiferal community is an excellent bio-indicator of pollution in coastal and marginal marine settings. Their abundance, diversity and its relationship with environmental parameters, viz. grain size, organic carbon (Corg) and trace metals concentration, from the surface sediment layer of the southwestern shelf of the Bay of Bengal, have been used to understand the marine pollution in this study. A total of 32 surface sediment samples were collected at various water depths.The samples were analysed for grain size distribution, organic matter (OM), foraminiferal and trace metals studies. Statistical analyses such as correlation matrix, PCA (Principal Component Analysis) and cluster analysis show that the foraminiferal species composition is significantly influenced by the organic carbon (Corg), Co, Pb, Zn, Cr, and Ni concentrations. The dominance of stress-tolerant species, viz. Ammonia beccarii, Ammonia tepida, Nonion faba, Bulimina marginata, Bolivina robusta, Elphidium craticulatum and Elphidium advenum, as well as less species diversity, demonstrates the elevated Corg and trace metal pollution in the marine environment.

The Middle-Upper Eocene benthic foraminifera from north-west Fayoum area, Egypt: Paleoecology and their similarity to the Tethyan provinces

This study deals with the Middle–Late Eocene benthic foraminiferal assemblage from the Um Rigl succession that is exposed in the northwest of Fayoum area, Egypt. Stratigraphically, two rock units are recognized from base to top as the Gehannam Formation and Birket Qarun Formation. The examination of benthic foraminiferal community identifies 63 species belonging to 36 genera, 22 families and four suborders. Their distribution aids to recognize one benthic foraminiferal biozone, Bulimina jacksonensis-Uvegerina jacksonensis Zone. To detect the paleocological conditions that prevailed during the deposition of the studied succession, a quantitative analysis is done on the foraminiferal assemblage, including the number of taxa, the relative abundances of the recorded suborders, calcareous species, agglutinated species and common species. The morphology of benthic foraminiferal tests is strongly associated with their microhabitat preferences. Therefore, the percentages of epifaunal and infaunal species are also calculated to detect the level of oxygenation. Accordingly, two ecozones are established, ecozone 1 represents the Gehannam Formation, whereas ecozone 2 represents the Birket Qarun Formation (excluding the barren zone in the upper part). Ecozone 1 is dominated by calcareous species (90–100%) and infaunal species (38–99%), while epifaunal species range from 2 to 62%. The most abundant taxa are Bolivina, Bulimina, Uvigerina, Nonion and Lenticulina. Therefore, the ecozone 1 was deposited under an inner-outer neritic environment, with normal salinity and eutrophic to mesotrophic conditions. On the other side, ecozone 2 is characterized by changes in the composition of foraminiferal assemblage, where infaunal species dominate the base of this ecozone, followed by a strong decline in the middle part and then increase again in the upper part. Also, a notable increase in epifaunal species is documented in this ecozone. This variation in foraminiferal assemblage reflects changes in water depth and oxygen content. Consequently, ecozone 2 was deposited under an inner to middle neritic environment, with normal salinity and mesotrophic to oligotrophic conditions. Regarding the paleobiogeographic outline, the results of multivariate analyses indicate that the Egyptian species show a strong similarity to those from the Southern Tethyan Province (Tunisia and Libya), whereas they show a reasonable similarity to the Western Tethyan Province (Spain, Italy, France). This deduction reflects a probable marine connection between both provinces during Eocene.

Foraminifera associated with cold seeps in marine sediments

Cold seep foraminifera have attracted considerable attention as they provide valuable insights into the study of cold seeps. This study provides a comprehensive overview of the manifestations of foraminifera in cold seep environments and methane seepage activities. Certain taxa of benthic foraminifera, such as Uvigerina, Bolivina, Bulimina, Chilostomella, Globobulimina, Nonionella, Melonis, Epistominella, Cibicidoides, and Globocassidulina, are known to inhabit geochemical conditions induced by methane-rich environments and may feed on associated methanotrophic microbial communities. Secondary mineralization on foraminifera shells is a widespread manifestation in seep sediments, and alters the microstructure, elementary composition, and isotopic signatures of foraminifera. On one hand, the precipitation of secondary authigenic Mg-rich, Mn-rich, Sr-rich, and Ba-rich calcite coatings have been observed on microfossils. On the other hand, micron-sized crystal pyrite and gypsum aggregates can also grow on the foraminifera walls. The negative δ13C and positive δ18O anomalies in both planktonic and benthic foraminifera from seep-associated sediments can serve as proxies for tracing past seepage activities, either in their live form or being adulterated by methane-derived authigenic carbonate after deposition. Seeping activities are recognized with a significant impact on benthic foraminifera, and the presence of cold seep-related species and significant isotopic anomalies in shells can be used to reconstruct past methane seepage events. Intensive methane seepage tends to suppress benthic foraminifera populations, while moderate intensity seepage may lead to a thriving benthic foraminiferal community, with hypoxia-enduring taxa such as Uvigerina, Melonis, and Bulimina being predominant. In contrast, oxygen-loving epibenthic taxa such as Cibicidoides often occur in areas of low methane flux. Compared to planktonic foraminifera, the single species of benthic foraminifera can provide a more comprehensive record of seepage evolution. Live benthic specimens are preferred for in-situ seep studies, while the superimposition of secondary minerals on the original shells should not be ignored when observing dead individuals. The significance of the evolution of methane seepage, changes in environmental parameters of the living habitat, and species sensitivity in cold seeps are emphasized in explaining the variation in foraminiferal assemblages and fluctuations in stable isotopes.

Through the eDNA looking glass: Responses of fjord benthic foraminiferal communities to contrasting environmental conditions.

The health of coastal marine environments is severely declining with global changes. Proxies, such as those based on microeukaryote communities, can record biodiversity and ecosystem responses. However, conventional studies rely on microscopic observations of limited taxonomic range and size fraction, missing putatively ecologically informative community components. Here, we tested molecular tools to survey foraminiferal biodiversity in a fjord system (Sweden) on spatial and temporal scales: Alpha and beta diversity responses to natural and anthropogenic environmental trends were assessed and variability of foraminiferal environmental DNA (eDNA) compared to morphology-based data. The identification of eDNA-obtained taxonomic units was aided by single-cell barcoding. Our study revealed wide diversity, including typical morphospecies recognized in the fjords, and so-far unrecognized taxa. DNA extraction method impacted community composition outputs significantly. DNA extractions of 10 g sediment more reliably represented present diversity than of 0.5-g samples and, thus, are preferred for environmental assessments in this region. Alpha- and beta diversity of 10-g extracts correlated with bottom-water salinity similar to morpho-assemblage diversity changes. Sub-annual environmental variability resolved only partially, indicating damped sensitivity of foraminiferal communities on short timescales using established metabarcoding techniques. Systematically addressing the current limitations of morphology-based and metabarcoding studies may strongly improve future biodiversity and environmental assessments.

Unique benthic foraminiferal communities (stained) in diverse environments of sub-Antarctic fjords, South Georgia

Abstract. Sub-Antarctic fjords are among the environments most affected by the recent climate change. In our dynamically changing world, it is essential to monitor changes in these vulnerable settings. Here, we present a baseline study of “living” (rose-bengal-stained) benthic foraminifera from fjords of South Georgia, including fjords with and without tidewater glaciers. Their distribution is analyzed in the light of new fjord water and sediment property data, including grain size and sorting, total organic carbon, total sulfur, and δ13C of bulk organic matter. Four well-defined foraminiferal assemblages are recognized. Miliammina earlandi dominates in the most restricted, near-shore and glacier-proximal habitats, Cassidulinoides aff. parkerianus in mid-fjord areas, and Globocassidulina aff. rossensis and an assemblage dominated by Ammobaculites rostratus, Reophax subfusiformis, and Astrononion echolsi are in the outer parts of the fjords. Miliammina earlandi can tolerate strong glacial influence, including high sedimentation rates in fjord heads and sediment anoxia, as inferred from sediment color and total organic carbon / sulfur ratios. This versatile species thrives both in the food-poor inner reaches of fjords that receive mainly refractory petrogenic organic matter from glacial meltwater and in shallow-water coves, where it benefits from an abundant supply of fresh, terrestrial, and marine organic matter. A smooth-walled variant of C. aff. parkerianus, apparently endemic to South Georgia, is the calcareous rotaliid best adapted to inner-fjord conditions characterized by moderate glacial influence and sedimentation rates and showing no preference for particular sedimentary redox conditions. The outer parts of fjords with clear, well-oxygenated bottom water are inhabited by G. aff. rossensis. Ammobaculites rostratus, R. subfusiformis, and A. echolsi dominate in the deepest-water settings, with water salinities ≥ 33.9 PSU and temperatures 0.2–1.4 ∘C, characteristic of winter water and Upper Circumpolar Deep Water. The inner- and mid-fjord foraminiferal assemblages seem specific to South Georgia, although with continued warming and deglaciation, they may become more widespread in the Southern Ocean.

Next-generation sequencing and metabarcoding to understand the ecology of benthic foraminiferal community in the Bering Sea

The sedimentary archives from the Bering Sea can provide excellent records of the global climate shift. However, paleoclimatic studies from the Bering Sea are limited due to the lack of well calibrated proxies. Various characteristics of foraminifera are frequently used as paleoclimatic and paleoceanographic proxies. A precise understanding of the ecology of foraminiferal species is required to use it as proxy. Here, we use next-generation sequencing and metabarcoding to understand the ecology of benthic foraminiferal community in the Bering Sea. The total genomic deoxyribonucleic acid (DNA) was extracted from 11 surface sediment samples collected from the Bering Sea. The foraminiferal small sub unit (SSU) rDNA was amplified with foraminiferal-specific primers. A total of 2,170,762 effective tags and 1046 operational taxonomic unit (OTUs) were obtained after next-generation sequencing, strict quality control, and data processing. The results showed that the station BR07 had the highest number of OTUs. The highest foraminiferal Margalef index and Shannon-Wiener index were obtained at BR06. Rotaliida accounted for the highest proportion, followed by Textulariida and Monothalamida, with the lowest proportion of Miliolida. From the principal component analysis (PCA), Leptohalysis scottii, Stainforthiidae_X sp., and Rectuvigerina phlegeri were found to be the more important species in the Bering Sea. Based on the redundancy analysis (RDA) and Spearman correlation analysis, we found that the grain size and water depth exhibited significant control on foraminiferal community. The foraminiferal reads were positively correlated with the grain size, whereas the Shannon-Wiener index was negatively correlated with it. The water depth was positively correlated with OTUs, Leptohalysis scottii, and Rectuvigerina phlegeri. The clustering analysis showed that Bering Sea stations cluster together with Yellow Sea stations of similar water depth. The water depth was a bigger factor for foraminiferal community than the geographical distance.

Total and active benthic foraminiferal community and their response to heavy metals revealed by high throughput DNA and RNA sequencing in the Zhejiang coastal waters, East China Sea.

Benthic foraminifera, large protists abundant in marine environments, have been widely used as bioindicators of environmental conditions. In this study, high-throughput sequencing based on small subunit rDNA and rRNA amplifications was used to investigate total and active benthic foraminifera community composition and diversity from nineteen and twelve superficial marine sediment samples in the Zhejiang coastal waters, respectively. The results showed that the dominant taxa of total foraminifera changed from Buliminellidae (hyaline) to Saccamminidae (agglutinated) from north to south along the coastal waters of Zhejiang Province. According to our survey, heavy metal contamination was moderate in Zhejiang coastal waters, and the potential ecological risks posed by Cd and Hg were higher. The contamination level of heavy metals at Yueqing Bay was the highest, followed by those at Sanmen Bay and Hangzhou Bay. Cd, Cu and grain size may be key factors affecting the distribution and composition of active foraminiferal communities.

COI metabarcoding of large benthic Foraminifera: Method validation for application in ecological studies.

Monitoring community composition of Foraminifera (single-celled marine protists) provides valuable insights into environmental conditions in marine ecosystems. Despite the efficiency of environmental DNA (eDNA) and bulk-sample DNA (bulk-DNA) metabarcoding to assess the presence of multiple taxa, this has not been straightforward for Foraminifera partially due to the high genetic variability in widely used ribosomal markers. Here, we test the correctness in retrieving foraminiferal communities by metabarcoding of mock communities, bulk-DNA from coral reef sediment samples, and eDNA from their associated ethanol preservative using the recently sequenced cytochrome c oxidase subunit 1 (COI) marker. To assess the detection success, we compared our results with large benthic foraminiferal communities previously reported from the same sampling sites. Results from our mock communities demonstrate that all species were detected in two mock communities and all but one in the remaining four. Technical replicates were highly similar in number of reads for each assigned ASV in both the mock communities and bulk-DNA samples. Bulk-DNA showed a significantly higher species richness than their associated eDNA samples, and also detected additional species to what was already reported at the specific sites. Our study confirms that metabarcoding using the foraminiferal COI marker adequately retrieves the diversity and community composition of both the mock communities and the bulk-DNA samples. With its decreased variability compared with the commonly used nuclear 18 S rRNA, the COI marker renders bulk-DNA metabarcoding a powerful tool to assess foraminiferal community composition under the condition that the reference database is adequate to the target taxa.

Co-occurrence patterns and community assembly mechanisms of benthic foraminiferal communities in South Chinese bays

Benthic foraminifera are among the most important components of marine benthic communities and can provide valuable information about the structure and activity of those communities. The distribution and diversity of benthic foraminifera have been well documented, but their co-occurrence patterns and community assembly processes have rarely been elucidated. To fill this gap, we used network analysis and null/neutral model approaches to benthic foraminiferal communities in five bays in South China to evaluate the influences of environmental factors and interspecies interactions on species co-occurrence and the relative importance of deterministic and stochastic processes in shaping community structure, respectively. The spatial heterogeneity of foraminiferal communities across bays was characterized by nonmetric multidimensional scaling analysis. Variation partitioning analysis showed that heavy metals (Cr, Co, Cu, As, Sb, and Hg) were the most important factor influencing community structure, while less importance was attributed to organic matter and grain sizes, which was likely due to the low TOC contents and small variations in TOC and grain size across sites. Network analysis showed that closely related species tended to co-occur with each other, which may be mediated by environmental filtering. The network was divided into three distinct clusters: one cluster consisted of miliolid, and two were dominated by rotaliids, each of which represented a special niche according to the habitat preference of the dominant taxa. The structure of the network was highly informative, as illustrated by the fact that the observed structure was denser than that of randomly constructed networks with the same size or the same degree sequence. The frequency of occurrence of most species (70%) met neutral model expectations, indicating that neutral processes were important for the assembly of benthic foraminiferal communities. However, PER-SIMPER analysis revealed that niche-assembly processes were more important than dispersal-assembly processes in shaping species abundance. Overall, this study provides the first insight into the mechanisms maintaining the community structure and diversity of benthic foraminifera.

Santonian deep sea benthic foraminifera from IODP Site U1513, Mentelle Basin (SW Australia): Reactions of benthic foraminiferal assemblages to surface water cooling at southern high latitudes

International Ocean Discovery Program (IODP) Expedition 369 recovered a stratigraphically complete, Santonian-aged, pelagic sedimentary sequence at Site U1513 on the Naturaliste Plateau (33°47.6084′S, 112°29.1338′E) at ~2800 m water depth. The site was located at ~60°S paleolatitude in Cretaceous times. A total of 59 samples studied from Holes 1513A and 1513B yielded 140 calcareous and nine agglutinated benthic foraminiferal taxa. Gavelinellids (Notoplanulina, Gavelinella, Anomalinoides) and Gyroidinoides spp., are the most abundant taxa overall and also dominate most samples. We also report 382 foraminiferal stable isotopic measurements and infer cooling [of ~5 °C in surface waters and 2 °C at the seafloor] based on changes in δ18O values for benthic and planktonic taxa through the Santonian at Site U1513. Inferred cooling was accompanied by changes in planktonic and benthic foraminiferal assemblages. At Site U1513, benthic foraminiferal taxonomic richness declines by 15 species (from 20– 25 to 5–10 taxa) towards the upper Santonian. Within this interval, epifaunal, oxic foraminifera diminish in absolute and relative abundance, and there is a parallel relative increase in opportunistic taxa and infaunal foraminifera. Changes in benthic foraminiferal communities indicate a shift from oligotrophic to eutrophic conditions in the bottom water despite the general cooling trend. This paper explores the causes and effects of the paleoenvironmental changes in the bottom waters.

Testing an indirect palaeo-seagrass indicator: Benthic foraminifera from the Lower Pleistocene Posidonia meadow of Fauglia (Tuscany, Italy)

Well-preserved remains of a Posidonia oceanica meadow have been found in a Lower Pleistocene (Calabrian) succession cropping out near Fauglia (Tuscany, Italy). This paper analyses and describes the benthic foraminiferal community associated with this well-preserved Pleistocene P. oceanica meadow, with the purpose of testing the usefulness of foraminifera as an Indirect Palaeo-Seagrass Indicator (IPSI), providing both qualitative and quantitative parameters to recognize other meadow-related environments where fossil remains of seagrasses are not preserved. Despite being influenced by some diagenetic processes that might have affected the foraminiferal assemblage, the Fauglia succession represents a suitable setting for testing benthic foraminifera as IPSI in a temperate environment. Considering this limitation, together with other ecological constraints such as seawater temperature, this study provides evidence of the high potential of benthic foraminifera as IPSI. Several parameters such as the modified Foraminifera Index (FI’), the “long vs. short life-span index” (ILS), and the presence of permanently attached, encrusting foraminifera were tested. New indexes were also developed, namely the K/R ratio, consisting of the ratio between keeled Elphidium and the sum of rounded elphidiids (e.g., Elphidium translucens, Cribroelphidium, Porosononion) and Haynesina, and the K/REXT ratio, consisting of the ratio between keeled Elphidium and the sum of rounded elphidiids, Haynesina and other related genera that display a rounded periphery (i.e., Astrononion, Melonis, Nonion, Nonionella, Valvulineria). All these indexes were examined, suggesting their potential use for recognizing seagrass-related palaeoenvironments. The K/REXT ratio (and K/R) in association with the presence of permanently attached foraminifera revealed to be the most reliable palaeo-seagrass indicators, suggesting that this combination could be very useful also in other case studies where diagenesis altered the foraminiferal association.

Environmental RNA outperforms eDNA metabarcoding in assessing impact of marine pollution: A chromium-spiked mesocosm test

Environmental (e)DNA metabarcoding holds great promise for biomonitoring and ecotoxicological applications. However, few studies have compared the performance of eDNA versus eRNA metabarcoding in assessing organismal response to marine pollution, in experimental conditions. Here, we performed a chromium (Cr)-spiked mesocosm experimental test on benthic foraminiferal community to investigate the effects on species diversity by analysing both eDNA and eRNA metabarcoding data across different Cr concentrations in the sediment. Foraminiferal diversity in the eRNA data showed a significant negative correlation with the Cr concentration in the sediment, while a positive response was observed in the eDNA data. The foraminiferal OTUs exhibited a higher turnover rate in eRNA than in the eDNA-derived community. Furthermore, in the eRNA samples, OTUs abundance was significantly affected by the Cr gradient in the sediment (Pseudo-R2 = 0.28, p = 0.05), while no significant trend was observed in the eDNA samples. The correlation between Cr concentration and foraminiferal diversity in eRNA datasets was stronger when the less abundant OTUs (<100 reads) were removed and the analyses were conducted exclusively on OTUs shared between eRNA and eDNA datasets. This indicates the importance of metabarcoding data filtering to capture ecological impacts, in addition to using the putatively active organisms in the eRNA dataset. The comparative analyses on foraminiferal diversity revealed that eRNA-based metabarcoding can better assess the response to heavy metal exposure in presence of subtle concentrations of the pollutant. Furthermore, our results suggest that to unlock the full potential for ecosystem assessment, eDNA and eRNA should be studied in parallel to control for potential sequence artifacts in routine ecosystem surveys.

Paleoceanographic inferences from benthic foraminifera across the early Aptian Ocean Anoxic Event 1a in the western Tethys

The paleoenvironmental impact of the early Aptian Ocean Anoxic Event 1a (OAE 1a, ca. 121 Ma) has been investigated in detail in the Cismon Core (Lombardy Basin, western Tethys) by using different geochemical and micropaleontological proxies. We provide the first high-resolution data of benthic foraminiferal assemblages through the upper Barremian - lower upper Aptian stratigraphic interval. Benthic foraminifera data are integrated with calcareous nannofossil and planktonic foraminifera records to create a comprehensive characterization of bottom and surface waters across OAE 1a. Benthic foraminiferal communities are indicative of a marked change in bottom-waters around the “nannoconid decline” (latest Barremian) due to increased flux of organic matter to the seafloor and intermittent dysoxic conditions probably promoted by pulses of higher productivity during the initial Greater Ontong Java Event (GOJE). Benthic foraminifera experienced a marked crisis in abundance (“benthic foraminiferal crisis” BFC) ca. 35 kyr before the OAE 1a, in correspondence with the “nannoconid crisis” and the onset of the most intense GOJE phase. The literature survey shows that the BFC is commonly recorded before the OAE 1a onset in several stratigraphic sections worldwide, and therefore, it is here proposed as a global event. At Cismon, deep-water anoxia was reached at the OAE 1a onset and lasted for ca. 300 kyr, promoted by higher productivity and eventually enhanced water stratification during the super-greenhouse climate. The continuation of OAE 1a was marked by a benthic foraminiferal repopulation event, probably resulting from the influx of relatively cooler and oxygenated waters. In turn, the OAE 1a was marked by intermittent anoxic to dysoxic conditions, likely in response to primary productivity sustained by N-fixing bacteria. The distribution and abundance of benthic foraminifera documented in other sections across the Selli Level equivalent show different features that point to local factors such as paleodepth and increased runoff. The post-OAE 1a was characterized by dysoxic conditions coupled with moderate organic matter flux to the seafloor. At the Cismon site and worldwide, the termination of OAE 1a coincided with the return of relatively more abundant benthic taxa in response to the restoration of favorable conditions allowing the development of diversified benthic foraminiferal communities.