Miocene benthic foraminiferal isotope records: A synthesis

${}^{18}\text{O}{}^{16}\text{O}$ and ${}^{13}\text{C}{}^{12}\text{C}$ ratios of Miocene benthic foraminifera from a number of Atlantic, Pacific and Indian Ocean DSDP sites (71, 77B, 206, 208, 238, 279, 289, 296, 329, 357 and 366A) have been compiled. These provide a rather detailed history of Miocene deep water especially in the Pacific Ocean. Bottom-water temperatures rose during the early Miocene and then declined rapidly during the middle Miocene. This decline was accompanied by an increase in Antarctic glaciation. Late Miocene bottom temperatures and Antarctic ice volumes are inferred to be similar to today's, but exhibited some fluctuation. The early Miocene ocean was less thermally stratified at intermediate and abyssal depths while the late Miocene deep ocean had a thermal structure generally similar to the modern ocean.Foraminiferal carbon isotope ratios at most of the sites varied quasi-sympathetically throughout the Miocene. These variations must reflect comparable variations in the mean ${}^{13}\text{C}{}^{12}\text{C}$ of marine HCO₃^?. However, the causes of such variations are not yet clear.     

Laboratory observations of benthic foraminiferal cysts

Long-term observations on living benthic foraminifera in the laboratory were performed to investigate their behaviour and life style. We noticed that the formation of cysts or sedimentary envelopes is a common feature within many groups of foraminifera (organic walled, agglutinated and calcareous) in the laboratory. Several kinds of cyst were observed. In most cases, the entire foraminiferal shell was canopied with detritus or particles, but some specimens covered only parts of their body, such as the aperture region or pseudopods. Cysts were found attached to the glass walls of culture vessels or free in and on the sediment. Foraminifera stayed within cysts for hours to weeks. After leaving sedimentary envelopes, some specimens immediately started to build new ones, others not. The function of cyst formation observed in the present study was not clear, except one case where reproduction took place. Some monitored structures seemed not to be sedimentary cysts but unilocular agglutinated foraminifera with probably allogromiid-like organization. The formation of sedimentary envelopes seems to be a very basic character of foraminifera, suggesting a mechanism for the evolution of shells.     

The benthic foraminiferal communities of the northern Adriatic continental shelf

Off the Po Delta in the northern Adriatic, three different foraminiferal associations were identified in the fraction >0.125 mm of 25 sediment samples. The first association concerns sample stations between depths of 5 and 13.5 m in the shallow, nutrient-rich belt and is characterised by dominant Ammonia beccarii. The second occupies the 29.5–38.5-m-deep, nutrient-rich zone of clay-rich sediment, where inputs from the Po are concentrated, and is dominated by Nonionella opima. The third association, in the 21–46-m-deep and nutrient-poor zone, is dominated by Textularia agglutinans. The same distribution of communities was obtained using total associations (living and dead specimens) and biocenosis (stained individuals). The good correspondence with the foraminiferal distributions described in the past indicates that, in spite of the occurrence of anoxic and dysoxic events, the environmental state in the northern Adriatic may be considered stable.     

Atlantic paleobathymetry, paleoproductivity and paleocirculation in the late Albian: the benthic foraminiferal record

Spatial distribution patterns of benthic foraminifers in upper Albian sediments from 25 DSDP/ODP sites and 31 onshore sections of the North and South Atlantic Ocean are used to generate paleobathymetric reconstructions and to identify areas of high primary production such as coastal and equatorial upwelling zones. New paleobathymetric estimates are provided for DSDP/ODP sites and onshore locations that are not situated on oceanic crust. Paleobathymetric reconstructions indicate shallow water exchange between the North and South Atlantic but show the existence of a deep-water connection between the western and eastern Tethys (>2500 m) through the Gibraltar Gateway. Strikingly, there is no evidence for a strong latitudinal gradient in deep-water benthic foraminiferal distribution during the late Albian: South Atlantic assemblages show close affinity to North Atlantic and Tethyan assemblages, exhibiting only a minor degree of provincialism. Biogeographic patterns reveal a distinct asymmetry in late Albian paleoproductivity for the North Atlantic. As for the present day, the eastern margins of the Atlantic were generally more productive than the western margins, and a belt of enhanced carbon flux export to the seafloor can be traced around the north African coast, which probably corresponded to a zone of vigorous coastal upwelling. By contrast, assemblage composition in the South Atlantic generally reflects mesotrophic to oligotrophic conditions. Benthic foraminiferal distribution patterns, thus, provide robust proxy data to test predictions from paleocirculation and paleobathymetric models for the mid-Cretaceous Atlantic Ocean and adjacent margins.     

Recent abyssal benthic foraminiferal biofacies of the eastern equatorial Indian Ocean

Study of Recent abyssal benthic foraminifera from core-top samples in the eastern equatorial Indian Ocean has identified distinctive faunas whose distribution patterns reflect the major hydrographic features of the region. Above 3800 m, Indian Deep Water (IDW) is characterized by a diverse and evenly-distributed biofacies to whichGlobocassidulina subglobosa, Pyrgo spp.,Uvigerina peregrina, andEggerella bradyi are the major contributors.Nuttalides umbonifera andEpistominella exigua are associated with Indian Bottom Water (IBW) below 3800 m. Within the IBW fauna,N. umbonifera andE. exigua are characteristic of two biofacies with independent distribution patterns.Nuttalides umbonifera systematically increases in abundance with increasing water depth. TheE. exigua biofacies reaches its greatest abundance in sediments on the eastern flank of the Ninetyeast Ridge and in the Wharton-Cocos Basin. The hydrographic transition between IDW and IBW coincides with the level of transition from waters supersaturated to waters undersaturated with respect to calcite and with the depth of the lysocline. Carbonate saturation levels, possibly combined with the effects of selective dissolution on the benthic foraminiferal populations, best explain the change in faunas across the IDW/IBW boundary and the bathymetric distribution pattern ofN. umbonifera. The distribution of theE. exigua fauna cannot be explained with this model.Epistominella exigua is associated with the colder, more oxygenated IBW of the Wharton-Cocos Basin. The distribution of this biofacies on the eastern flank of the Ninetyeast Ridge agrees well with the calculated bathymetric position of the northward flowing deep boundary current which aerates the eastern basins of the Indian Ocean.     

Measuring and controlling data quality in biological assemblage surveys with special reference to stream benthic macroinvertebrates

1. Biological assemblage surveys primarily aim to characterise species composition and relative abundance at one or more spatial or temporal scales. Data interpretation and conclusions are dependent on how well samples characterise the assemblage of interest. 2. Conventional measures of data quality, e.g. standard deviations or coefficients of variation, were designed for single variable estimation, and they are either insufficient or invalid for assessing the quality of data describing entire assemblages. Similarity indices take species composition and relative abundance into account and may be used to effectively measure and control the quality of data used to characterise assemblage structure. 3. The average Jaccard coefficient (JC) calculated across multiple pairs of replicate samples, i.e. autosimilarity JC (AJC), is conceptually and numerically related to the average coefficient of variation in the densities of all species recorded, a measure of sampling precision, and to the proportion of total species richness sampled, a measure of sampling accuracy. 4. We explored how AJC can be used to assess the effect of different potential sources of error on the quality of assemblage survey data, including the sampling effort used both within regions and at individual sites, the individuals collecting samples, sub‐sampling procedures, and consistency of taxon identification. 5. We found that the autosimilarity‐based approach overcomes most weaknesses associated with conventional measures of data quality and can be used to effectively measure and control the quality of assemblage survey data.     

The integral structure of a benthic infaunal assemblage

It has been suggested that the scaling relationships of many features of the physical environment and biological traits are fractal-like, but for the marine benthic infauna certain aspects of the environment clearly are not. These include temporal features such as the cycles of annual climate, primary production and tides, and also some spatial features such as sediment granulometry and the size of the primary producers that constitute the food supply. We explicitly addressed these issues by determining the degree to which infaunal assemblage structure (diversity, species composition, spatial pattern) varies with mesh size, sample size and sample dispersion within an apparently homogeneous area of coarse intertidal sand in the Isles of Scilly, UK. Samples were extracted using a standard range of 5 mesh sizes (63, 125, 250, 500, 1000 μm), with the sample areas and distances between samples scaled to the mesh size. All metazoans were identified to species level. Diversity and species composition did not show a gradual and even degree of change over the size range. Instead, they showed a dramatic stepwise change between the 250 and 500 μm mesh size samples, being relatively constant in the < 500 and > 500 μm categories, with diversity higher in the former. Higher proportions of species in the < 500 μm categories had values of an index of dispersion significantly different to 1 than among species in the > 500 μm categories. This suggests a fractal structure within but not between the < 500 and > 500 μm body size categories. The implications of this for rapid diversity assessment by extrapolation between size classes are discussed. Although the interplay between 3 and 2 dimensional processes in what is a essentially a 2-D study may account for some of the observations, comparative studies suggest that these patterns do not simply correspond to the physical scaling of habitat complexity, and they must therefore relate to some more universal scaling relationships that are not fractal-like. We suggest that the important relationships are those between body size and various biological characteristics such as feeding behaviour, reproductive mode and life history as they are affected by the spatial and temporal structure of the environment.     

Recent benthic foraminiferal assemblages from cold-water coral mounds in the Porcupine Seabight

Cold-water coral ecosystems are characterised by a high diversity and population density. Living and dead foraminiferal assemblages from 20 surface sediment samples from Galway and Propeller Mounds were analysed to describe the distribution patterns of benthic foraminifera on coral mounds in relation to different sedimentary facies. Hard substrates were examined to assess the foraminiferal microhabitats and diversities in the coral framework. We recognised 131 different species, of which 27 prefer an attached lifestyle. Epibenthic species are the main constituents of the living and dead foraminiferal assemblages. The frequent species Discanomalina coronata was associated with coral rubble, Cibicides refulgens showed preference to the off-mound sand veneer, and Uvigerina mediterranea displayed abundance maxima in the main depositional area on the southern flank of Galway Mound, and in the muds around Propeller Mound. The distribution of these species is rather governed by their specific ecological demands and microhabitat availability than by the sedimentary facies. Benthic foraminiferal assemblages from coral mounds fit well into basin-wide-scale distribution patterns of species along the western European continental margin. The diversity of the foraminiferal faunas is not higher on the carbonate mounds as in their vicinity. The living assemblages show a broad mid-slope diversity maximum between 500 and 1,300 m water depth, which is the depth interval of coral mound formation at the Celtic and Amorican Margin. The foraminiferal diversity maximum is about 700 m shallower than comparable maxima of nematodes and bivalves. This suggests that different processes are driving the foraminiferal and metazoan diversity patterns.     

Planktic foraminiferal molecular evolution and their polyphyletic origins from benthic taxa

Phylogenetic analyses based on partial sequences of the small subunit (SSU) ribosomal (r) RNA gene have shown that the planktic and benthic foraminifera form a distinct monophyletic group within the eukaryotes. In order to determine the evolutionary relationships between benthic and planktic foraminifers, representatives of spinose and non-spinose planktic genera have been placed within a molecular SSU rDNA phylogeny containing sequences of the benthic suborders available to date. Our phylogenetic analysis shows that the planktic foraminifers are polyphyletic in origin, not evolving solely from a single ‘globigerinid-like’ lineage in the Mid-Jurassic, but derived from at least two ancestral benthic lines. The benthic ancestor of Neogloboquadrina dutertrei may have entered the plankton later than the Mid-Jurassic, and further investigation of related extant species should provide an indication of the timing of this event. The evolutionary origin of the non-spinose species Globorotalia menardii remains unclear. The divergences of the planktic spinose species generally support recent phylogenies based on the fossil record, which infer a radiation from a globigerinid common ancestor in the Mid- to Late Oligocene. The branching pattern indicates that there are possibly four distinct groups within the main spinose clade, with large evolutionary distances being observed between them. Globigerinoides conglobatus clusters strongly with Globigerinoides ruber and are divergent from Globigerinella siphonifera, Orbulina universa and Globigerinoides sacculifer.Conserved regions of the SSU rRNA gene show sufficient variation to discriminate foraminifers at the species level. Large genetic differences have been observed between the pink and white forms of Gs. ruber and between Ge. siphonifera Type I and II. The two types of Ge. siphonifera cannot be discriminated by traditional palaeontological methods, which has considerable implications for tracing fossil lineages and for the estimation of molecular evolutionary rates based upon the fossil record. The conserved regions show a high degree of sequence identity within a species, providing signature sequences for species identification. The variable regions of the gene may prove informative for population level studies in some species although complete sequence identity was observed in G. sacculifer and O. universa between specimens collected from the Caribbean and Western Pacific.     

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

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

Impact of intentionally injected carbon dioxide hydrate on deep-sea benthic foraminiferal survival

Sequestration of carbon dioxide (CO₂) in the ocean is being considered as a feasible mechanism to mitigate the alarming rate in its atmospheric rise. Little is known, however, about how the resulting hypercapnia and ocean acidification may affect marine fauna. In an effort to understand better the protistan reaction to such an environmental perturbation, the survivorship of benthic foraminifera, which is a prevalent group of protists, was studied in response to deep-sea CO₂ release. The survival response of calcareous, agglutinated, and thecate foraminifera was determined in two experiments at ∼3.1 and 3.3 km water depth in Monterey Bay (California, USA). Approximately 5 weeks after initial seafloor CO₂ release, in situ incubations of the live–dead indicator CellTracker Green were executed within seafloor-emplaced pushcores. Experimental treatments included direct exposure to CO₂ hydrate, two levels of lesser exposure adjacent to CO₂ hydrate, and controls, which were far removed from the CO₂ hydrate release. Results indicate that survivorship rates of agglutinated and thecate foraminifera were not significantly impacted by direct exposure but the survivorship of calcareous foraminifera was significantly lower in direct exposure treatments compared with controls. Observations suggest that, if large scale CO₂ sequestration is enacted on the deep-sea floor, survival of two major groups of this prevalent protistan taxon will likely not be severely impacted, while calcareous foraminifera will face considerable challenges to maintain their benthic populations in areas directly exposed to CO₂ hydrate.     

Rafting of benthic macrofauna: important factors determining the temporal succession of the assemblage on detached macroalgae

Rafting on biotic and abiotic substrata has been reported for many benthic marine invertebrates. Here, I describe important characteristics of common floating substrata and review published studies examining the succession of the assemblage on detached macroalgae to identify the most important factors determining this succession. Floating substrata differ in survival time (i.e. before they disintegrate) and in food value, with abiotic materials having high survival times but low food value and some biotic substrata (seagrasses, fresh wood) having high food values but short survival times. Large macroalgae with pneumatocysts may combine high survival times and high food values. Substratum survival and food value have consequences for the species composition of the rafting macrofauna. In general, suspension feeders dominate rafting assemblages on abiotic substrata, while grazing and boring species abound on macroalgae and wood. The succession of the rafting assemblage can be subdivided into three phases during which different processes predominate. During the initial phase some of the original colonizers (if present) disappear from the substratum, either due to active emigration or predation. This short, initial phase is followed immediately by the colonization phase, during which new organisms colonize the rafting substratum. Colonization may be rapid and intense in the beginning but then slows down. Towards the end of the colonization phase, some rafting organisms may reproduce and their offspring recruit within the parental raft. Results from two long-term studies confirmed that the proportion of species with direct development increased with duration of rafting. These successional changes will be most pronounced for large biotic substrata, rendering these particularly suitable for long-distance dispersal of organisms with direct development.     

High assemblage persistence in heterogeneous habitats: an experimental test with stream benthic algae

1. The persistence of biological assemblages is positively affected by spatial heterogeneity. This influence may be indirect, through increased species richness. Another possibility is the increased availability of refuges from disturbances, which would prevent local loss. 2. We conducted a field experiment to test the hypothesis that greater roughness (a form of spatial heterogeneity) on the surface of substrata allows higher persistence of assemblages of stream benthic algae and that this relationship does not depend on species richness. Samples were taken on six occasions from smooth and rough artificial substrata used for algal colonisation. We calculated the persistence of assemblages using two analytical approaches: the mean distance to group centroid and the sum of the Euclidean distances between consecutive sampling occasions, both in a multivariate space. We also subsampled the data to take into account differences in species richness between treatments and thus to evaluate the effect of species richness on persistence. 3. Assemblages on rough substrata were more persistent than assemblages on smooth substrata. The effects detected were not due to the greater species richness on rough substrata, since a higher persistence of the assemblages on rough substrata remained after the subsampling procedures. 4. Our results indicate a strong positive relationship between substratum roughness and the persistence of stream benthic algal assemblages. We suggest that this is due to the presence of physical refuges in heterogeneous habitats.     

Stable-isotope analysis of a deep-sea benthic-fish assemblage: evidence of an enriched benthic food web

In this study, fishes and invertebrates collected from the continental slope (1000 m) of the eastern North Pacific Ocean were analysed using stable-isotope analysis (SIA). Resulting trophic positions (T(P) ) were compared to known diets and habitats from the literature. Dual isotope plots indicated that most species groups (invertebrates and fishes) sorted as expected along the carbon and nitrogen axes, with less intraspecific variability than interspecific variability. Results also indicated an isotopically distinct benthic and pelagic food web, as the benthic food web was more enriched in both nitrogen and carbon isotopes. Trophic positions from SIA supported this finding, resulting in the assignment of fishes to different trophic positions from those expected based on published dietary information. These differences can be explained largely by the habitat of the prey and the percentage of the diet that was scavenged. A mixing model estimated dietary contributions of prey similar to those of the known diet of Bathyraja trachura from stomach-content analysis (SCA). Linear regressions indicated that trophic positions calculated from SIA and SCA, when plotted against B. trachura total length for 32 individuals, exhibited similar variation and patterns. Only the T(P) from SCA yielded significant results (stomach content: P < 0·05, stable isotope: P > 0·05).     

Effect of seasonal hypoxia on the benthic foraminiferal community of the Louisiana inner continental shelf: The 20th century record

A species census in sediment core samples reveals significant changes in the composition of the Louisiana-shelf benthic foraminiferal community in the past century; these changes can be explained by an increase in the severity of seasonal hypoxia in bottom waters. Agglutinated and porcelaneous orders living in water depths less than 60 m suffered a noticeable decline during this time. In particular, the genus Quinqueloculina was severely affected by the progression of hypoxia, and nearly disappeared from parts of the study area. In contrast, several hyaline taxa, especially Nonionella basiloba, Buliminella morgani, and Epistominella vitrea, tolerated the progressive oxygen depletion well. Results of cluster and principal components analyses of the foraminiferal assemblage data match the observed species distribution trends and indicate that seasonal hypoxia on the Louisiana shelf, related to eutrophication and water stratification, worsened in the past century, even near the outer edge of the present-day zone of spring and summer oxygen depletion. The temporal trends in the foraminiferal record correspond to that of fertilizer use in the U.S. and nitrogen loading in the Mississippi River, suggesting that the anthropogenic factor has been particularly strong in the development of coastal hypoxia since the early 1940s.     

Spatial pattern indicates an influence of barnacle and ascidian aggregations on the surrounding benthic assemblage

Joint clusters of solitary ascidians Styela spp., Bolthenia echinata (L.), Molgula spp. and barnacles Balanus crenatus Bruguiere in the White Sea subtidal often develop on shells or stones partially buried into muddy sediment. To assess the structuring role of these suspension-feeders' aggregations, we examined the spatial patterns of the surrounding infaunal assemblage. Pairs of cores obtained close to clusters (Close ones) and 20-25 cm away from them (Distant ones) were compared, the distance corresponding to the average distance observed between aggregations. These pairs were spatially grouped in replicas (blocks) tens of meters from each other. Out of 10 dominating invertebrate species found around the clusters (84 in total), 5 demonstrated significant difference in density (N) close to aggregations and between them. All the five, including polychaets Scoloplos armiger (Muller, Orbiniidae), Aricidea nolani (Eliason, Paraonidae), Heteromastus filiformis Zachs (Capitellidae), Chaetozone setosa Malmgren (Cirratulidae) and undetermined oligochaets, were associated with Close cores. Total abundance of polychaets, oligochaets, crustaceans and bivalves was also different in Close and Distant cores and indicated significant correlations (of either sign) with biomass estimates for barnacles and ascidians in each block. Individual mean body weight (IMW) in pairs of cores was different for 3 of 10 dominants, maldanid polychaets being larger close to clusters, and cirratulids being larger between them. For several taxa the difference, observed in both parameters (N and IMW), significantly varied between the replicas.Filter-feeding activity and faeces production are regarded as the main possible factors explaining the effect of barnacles and ascidians presence, since the spatial pattern observed corresponds with feeding types of the infaunal taxa studied. Average distance between the clusters was short enough, which allows us to conclude that the spatial distribution pattern of benthic infaunal species within the research area strongly depends on barnacle and ascidian aggregations mosaic.     

Recent benthic foraminiferal assemblages in deep high-energy environments from the Gulf of Cadiz (Spain)

The benthic environment in the Gulf of Cadiz, north-eastern Atlantic, is strongly affected by the Mediterranean outflow water undercurrent (MOW) which flows northwards along the western Iberian Margin at 500–1500 m water depth. Foraminiferal census counts of living and dead assemblages from 27 surface samples ranging from 103 to 1917 m water depth, and the examination of hard substrates reveal a close correlation of the fauna with the local hydrography and sediment facies. Four different faunal groups are separated by factor analysis of the living fauna. Assemblage 1 contains typical lower slope species and dominates samples from the lower MOW core layer and in the North Atlantic deep water below. Shelf edge foraminifera are common in assemblage 2a which shows the highest proportions in samples from 103 to 272 m. Assemblage 2b is dominated by upper slope species and suspension-feeders that are frequent in the upper MOW core layer and in distal settings between 396 and 901 m. Species from assemblage 3 prefer epibenthic habitats and are recorded with high proportions exclusively in the immediate flow paths of the upper MOW between 496 and 881 m. Colonisation structures and species composition of epibenthic assemblages from the proximal facies largely differ from those in distal settings. In general, epibenthic foraminifers only use elevated substrates under the influence of near-bottom flow. Under high current velocities, epibenthic foraminifers prefer large and heavy objects. They colonise high attachment levels where a maximum yield of advected food particles can be achieved. In distal settings at lower flow velocities, the elevation height does not exceed 20 mm above the surrounding sediment surface. This level is related to a hydrologic transition layer with high concentrations of suspended particles. The comparison of microhabitat preferences and faunal structure under high and low current velocities reveal that substrate stability may be a confining environmental variable for endobenthic and shallow epibenthic foraminifers. The observations also indicate that the preferential settling height of epibenthic foraminifera is related to the highest lateral flux rates of food particles within reach from the sea floor. A dynamic selection of elevated microhabitats is only used by 7.8% of all species recognised in the Gulf of Cadiz area.     

Relationship of surface sediments and benthic foraminiferal distribution patterns in the Norwegian Channel (northern North Sea)

Our study of the distribution of benthic foraminifers in surface sediment samples from the Norwegian Channel has shown that eight species and five different assemblages are important. The assemblages are dominated byUvigerina peregrina, Bolivina skagerrakensis, Cassidulina laevigata, Elphidium excavatum andBulimina marginata, respectively.Hyalinea balthica, Melonis barleeanum andTrifarina angulosa are also abundant. The first four assemblages and their dominant species can be related to certain water masses with known characteristics. Two species are related to certain types of substrate;M. barleeanum seems to prefer a fine-grained substrate, whileT. angulosa occurs most abundantly where the content of silt and clay is low.B. skagerrakensis andH. balthica occur most abundantly in sediments with a high content of organic carbon, whileT. angulosa shows inverse correlation to the organic carbon content.     

Late Quaternary paleoenvironmental reconstruction of the Tremiti structural high (Central Adriatic Sea) from benthic foraminiferal assemblages

Foraminiferal analyses carried out on the VTR01/8 core (Tremiti Islands, Adriatic Sea) lead to the reconstruction of the paleoenvironmental evolution of the area. Although the core site is located at about 1000 km from the Po delta, this study shows that the installation of the recent sedimentological and run-off system (with a shore parallel clay-belt) is similar to that described at sites much closer to the Po delta. The statistical analysis singled out four associations that correspond to four distinct environments. The ecological preferences of the most abundant taxa that characterize each environment reveal that from the bottom of the core to − 50 cm, there is a slight increase of the water depth after which the water depth increases until the present level. Association A (that comprises subclusters A1, A2 and A3) is characterized by shallow-water taxa (e.g., Ammonia beccarii, Quinqueloculina spp. and Elphidium granosum) showing that in the past an infralittoral environment typified the Tremiti High. The subclusters of Association A show that during the deposition of the lower part of the core there had been an alternation of phases characterized by different degrees of riverine influence: the site was at first under the influence of river outflow (A2) that later bypassed the core site (A3) and then came back, although located further away than before (A1). With the rising of the sea level, the Tremiti High became only marginally affected by river influence because the core site is located at a considerable distance from the Po outlets and seaward with respect to the center of the modern mud-belt. As a matter of fact, association B (Bulimina marginata, Cassidulina carinata and Textularia spp.), which starts at − 46 cm, shows the typical characteristics of the modern mud-belt outer-shelf assemblage, although the presence of opportunistic species is limited by the distant position with respect to the Po delta.     

Relationship between benthic foraminiferal assemblages and environmental factors in atoll lagoons of the central Tuamotu Archipelago (French Polynesia)

Foraminiferal assemblages were studied in ten atoll lagoons in the central Tuamotu Archipelago, French Polynesia, in order to determine which environmental factors influence their distribution. Among geomorphological conditions, the degree of aperture has a major influence on lagoonal communities. Cluster and factor analyses reveal a clear contrast between the foraminiferal associations located in the peripheral and central areas of the lagoons, showing the effect of the vicinity of coral reef, and a limited post-mortem transport of species from the periphery toward the center of the lagoons. The rarity of planktonic species indicates limited penetration of the lagoon by planktonic forms, including planktonic stages of benthic colonizers. Colonization of these remote lagoons, even those with a high degree of aperture, may thus require long periods. Therefore, the Holocene transgression that flooded the deeper lagoons before the shallower ones, allowing a longer colonization period, may have contributed to the higher species richness of foraminifera within them.