Cytological Observations of the Large Symbiotic Foraminifer Amphisorus kudakajimensis Using Calcein Acetoxymethyl Ester

Large benthic foraminifera are unicellular calcifying reef organisms that can form symbiotic relationships with a range of different microalgae. However, the cellular functions, such as symbiosis and calcification, and other aspects of cellular physiology in large benthic foraminifera are not fully understood. Amphisorus kudakajimensis was used as a model to determine the detailed cellular characteristics of large benthic foraminifera. We used calcein acetoxymethyl ester (calcein AM) as a fluorescent indicator for live confocal imaging. We demonstrated that calcein AM is a useful fluorescent indicator to stain the fine network of reticulopodia and the cytoplasm in living A. kudakajimensis. We showed that at least two types of reticulopodia exist in A. kudakajimensis: the straight bundle of reticulopodia that spreads from the aperture and the fine reticulopodia along the surface of the aperture and chamber walls. The cytoplasm in outer chambers was highly branched and contained a few dinoflagellates. In contrast, the inner chamberlets contained condensed cytoplasm and many dinoflagellates, suggesting that the cytoplasm of A. kudakajimensis performs different functions based on its location within the large test. Our confocal detailed image analysis provides real-time cellular morphology and cell physiology of living foraminifera.     

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

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

Assessing the ecological health status using macrobenthic communities of tropical coastal water

The relative contributions of natural and anthropogenic fluctuations are different in shaping habitat health status and natural benthic communities in tropical coastal water. Understanding responses of coastal benthic communities to these fluctuations are still equivocal and thus available data are inadequate. Here, multiple analytical approaches were used to address the significant risk factors and their impacts on coastal benthic habitat health through space and time. A total abundance of 1436 ± 612 individuals of 33 benthic species were counted and identified from 22 sampling stations across eight sampling periods over the two years of study. Bioassay results showed that the benthic community is moderately exposed to anthropogenic pollutants in Klang Strait coastal water. The results showed that there were no significant temporal changes of habitat health status and macrobenthic community structure; however, spatial changes were significant and synchronized with anthropogenic and natural fluctuations. This study demonstrates that Cd and Hg levels and sediment characteristics played the primary role in shaping the habitat health and macrobenthic assemblages, whereas the influence of other factors were insignificant.     

Biostratigraphy and paleoecology of the Oligo-Miocene succession in Fars and Khuzestan areas (Zagros Basin,SW Iran)

Paleontological and biostratigraphical studies on carbonate platform succession from southwest Iran documented a great diversity of shallow-water benthic foraminifera during the Oligocene–Miocene. Larger foraminifera are the main means for the stratigraphic zonation of carbonate sediments. The distributions of larger benthic foraminifera in two outcrop sections (Abolhayat and Lali) in the Zagros Basin, Iran, are used to determine the age of the Asmari Formation. Four assemblage zones have been recognized by distribution of the larger benthic foraminifera in the study areas. Assemblage 3 (Aquitanian age) and 4 (Burdigalian age) have not been recognized in the Abolhayat section (Fars area), due to sea-level fall. The end Chattian sea-level fall restricted marine deposition in the Abolhayat section and Asmari Formation replaced laterally by the Gachsaran Formation. This suggests that the Miocene part of the formation as recognized in the Lali section (Khuzestan area) of the Zagros foreland basin is not present in the Abolhayat outcrop. The distribution of the Oligocene larger benthic foraminifera indicates that shallow marine carbonate sediments of the Asmari Formation at the study areas have been deposited in the photic zone of tropical to subtropical oceans. Based on analysis of larger benthic foraminiferal assemblages and microfacies features, three major depositional environments are identified. These include inner shelf, middle shelf and outer shelf. The inner shelf facies is characterized by wackestone–packstone, dominated by various taxa of imperforate foraminifera. The middle shelf is represented by packstone–grainstone to floatstone with a diverse assemblage of larger foraminifera with perforate wall. Basinwards is dominated by argillaceous wackestone characterized by planktonic foraminifera and large and flat nummulitidae and lepidocyclinidae. Planktonic foraminifera wackestone is the dominant facies in the outer shelf.     

Is increased calcarinid (foraminifera) abundance indicating a larger role for macro-algae in Indonesian Plio-Pleistocene coral reefs?

An important question in coral reef ecology is whether algal abundance in coral reef eco-systems is a natural phenomenon, or has increased as a result of coral reef degradation ultimately resulting in coral–algal regime shifts. Regime shifts, from coral to macro-algae dominated, alter the three-dimensional habitat structure in coral reef ecosystems. Surprisingly, few studies have looked at the effects for species that inhabit the reefs without being the architects of the three-dimensional structure. In this study the effects of a change in habitat characteristics on the community structure of large benthic foraminifera (LBF) is compared between an area with high (Kepulauan Seribu) and lower (Spermonde Archipelago) anthropogenic influence. The results indicate a general relationship between habitat and LBF assemblage structure. The largest difference was observed in shallow habitats. Habitats dominated by algae are inhabited by a specific group of LBF, the Calcarinidae, and domination of this group increases with higher algal prevalence. The fossil record of this group indicates that they evolved following a major change in settings of the central Indo-West-Pacific coral reefs from land detached platforms to fringing reefs, about 5 million years ago. Understanding the biotic response to this transition in reef morphology and the associated increase in terrestrially derived nutrients forms an excellent challenge to gain insights in present-day threats to coral reef ecosystems.     

Benthic foraminifera in deep trenches of the Kara Sea and the relation of <Emphasis Type="Italic">Saccorhiza ramosa</Emphasis> (Brady) to the distribution of natural radionuclides

The abundance and structure of the foraminifera community have been investigated. The role of agglutinating and secreting species in bottom sediments of the deep trenches of the Kara Sea has been studied. It has been found that the abundance and dominance of the agglutinated foraminifera Saccorhiza ramosa depend on the depth of the sea and are related to the genetic type of origin of such sediments. A correlation has been revealed between the distribution of S. ramosa and the concentrations of natural radionuclides (²²⁶Ra, ²³²Th, and ²¹⁰Pb) in bottom sediments. The degree of dominance of S. ramosa in the community of benthic foraminifera increases along with natural radioactivity.     

Benthic foraminifera off West Africa (1°N to 32°S): Do live assemblages from the topmost sediment reliably record environmental variability?

Recent benthic foraminifera (> 125 μm) were investigated from multicorer samples on a latitudinal transect of 20 stations between 1°N and 32°S along the upper slope off West Africa. Samples were selected from a narrow water depth interval, between 1200 and 1500 m, so that changes in water masses are minimized, but changes in surface productivity are important and the only significant environmental variable. Live (Rose Bengal stained) benthic foraminifera were counted from the surface sediment down to a maximum of 12 cm. Dead foraminifera were investigated in the top 5 cm of the sediment only. Five live and five dead benthic foraminiferal assemblages were identified using Q-mode principal component analysis, matching distinct primary productivity provinces, characterized by different systems of seasonal and permanent upwelling. Differences in seasonality, quantity, and quality of food supply are the main controlling parameters on species composition and distribution of the benthic foraminiferal faunas.To test the sensitivity of foraminiferal studies based on the uppermost centimeter of sediment only, a comparative Q-mode principal component analysis was conducted on live and dead foraminiferal data from the top 1 cm of sediment. It has been demonstrated that, on the upper slope off West Africa, most of the environmental signals as recorded by species composition and distribution of the “total” live and dead assemblages, i.e., including live and dead foraminifera from the surface sediment down to 12 cm and 5 cm, respectively, can be extracted from the assemblages in the top centimeter of sediment only. On the contrary, subsurface abundance maxima of live foraminifera and dissolution of empty tests strongly bias quantitative approaches based on the calculation of standing stocks and foraminiferal numbers in the topmost centimeter.     

Deep-sea benthic foraminiferal recolonization of the 1991 Mt. Pinatubo ash layer in the South China Sea

A census count of Rose Bengal stained benthic foraminifera from the surface area on top of a 2 to 6 cm thick ashfall layer at three deep water stations along the western margin of the Philippines exhibits a unique assemblage composition of benthic foraminifera. The total number of benthic foraminifera is low and the ratio of living individuals to empty tests is high. Specific diversity is low, with a significant dominance of infaunal morphotypes including species of the genus Reophax (R. scorpiurus, R. bilocularis and R. dentaliniformis), which are regarded as successful recolonizers. Assemblages below the ash layers are diverse and contain many epifaunal suspension-feeding agglutinated and calcareous foraminifera. The 1991 Mt. Pinatubo eruption caused mass mortality of benthic foraminifera in a vast area of the eastern South China Sea followed by step-wise recolonization of the ash substrate. Three years after the eruption, the benthic foraminiferal community structure is still far from recovery to background levels.     

Oxygen isotopic evidence for the depth stratification of tertiary and cretaceous planktic foraminifera

Oxygen isotope analyses of Tertiary and Cretaceous planktic foraminifera indicate that species have been stratified with respect to depth in the water column at least since Albian time. There is a relationship between morphology and depth habitat. Species with globigerine morphology have consistently occupied shallower depths than have species with globorotalid morphology. Biserially arranged species occupied both shallow and deep levels in the water column. On the average, it appears that ancient species with shallow habitats have been more susceptible to dissolution and have been preserved less well than species dwelling in deeper habitats. This relationship is similar to that observed for Recent planktic foraminifera. Comparison of carbon isotope ratios of adult and juvenile forms indicates that either the source of the carbon found in the shell or the carbon isotopic fractionations which occur during calcite secretion change during the development of individual foraminifera. The carbon isotopic ratios do not provide a reliable means for reconstructing the depth habitats of ancient species. Temperature-depth profiles for tropical Tertiary oceans have been reconstructed from the isotopic temperatures of planktic and benthic foraminifera. The vertical thermal structure of Oligocene oceans resembled that of modern oceans most closely. Those of Paleocene and Maastrichtian times differed most from that of modern oceans.     

渤海莱州湾表层沉积物中底栖有孔虫分布特征及其环境意义

对渤海莱州湾海域240个站位表层沉积物中底栖有孔虫群落进行了分析,共鉴定常见的底栖有孔虫42种。结果表明,莱州湾表层沉积物中底栖有孔虫主要以玻璃质壳为主(平均丰度达70.9%),瓷质壳含量次之,胶结壳含量最低;玻璃质壳占有孔虫全群的百分含量,随水深的增加而增加;从黄河口向外海方向,有孔虫分异度和丰度都逐渐增大。该海域底栖有孔虫平面分布的主要控制因素为盐度和底质沉积物类型,大体可分为两个组合分区,I区为Ammonia beccarii-Quinqueloculina spp.组合,代表盐度较低的近岸海陆过渡浅水环境;II区为Cribrononionsub-incertum-Protelphidium tuberculatum组合,代表盐度较高的远岸内陆架环境。     

Compound‐specific isotope analysis of benthic foraminifer amino acids suggests microhabitat variability in rocky‐shore environments

The abundance and biomass of benthic foraminifera are high in intertidal rocky‐shore habitats. However, the availability of food to support their high biomass has been poorly studied in these habitats compared to those at seafloor covered by sediments. Previous field and laboratory observations have suggested that there is diversity in the food preferences and modes of life among rocky‐shore benthic foraminifera. In this study, we used the stable nitrogen isotopic composition of amino acids to estimate the trophic position, trophic niche, and feeding strategy of individual foraminifera species. We also characterized the configuration and structure of the endobiotic microalgae in foraminifera using transmission electron microscopy, and we identified the origin of endobionts based on nucleotide sequences. Our results demonstrated a large variation in the trophic positions of different foraminifera from the same habitat, a reflection of endobiotic features and the different modes of life and food preferences of the foraminifera. Foraminifera did not rely solely on exogenous food sources. Some species effectively used organic matter derived from endobionts in the cell cytoplasm. The high biomass and species density of benthic foraminifera found in intertidal rocky‐shore habitats are thus probably maintained by the use of multiple nitrogen resources and by microhabitat segregation among species as a consequence.     

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

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

Predation contributes to invasion resistance of benthic communities against the non-indigenous tunicate <Emphasis Type="Italic">Ciona intestinalis</Emphasis>

Marine anthropogenic structures offer novel niches for introduced species but their role in the subsequent invasion to natural habitats remains unknown. Upon arrival in new environments, invaders must overcome biotic resistance from native competitors and predators if they are to establish successfully in natural habitats. We tested the hypotheses that (1) artificial structures (e.g., suspended aquaculture installations) present a niche opportunity for invasive species by providing a refuge from native benthic predators, and (2) native predators in natural benthic habitats suppress successful colonization by invaders. A recruitment experiment showed that the ascidians Pyura chilensis (native) and Ciona intestinalis (invasive) could recruit to both suspended artificial structures and natural benthic habitats. Ciona, however, was only able to establish adult populations on artificial structures. In natural benthic habitats Ciona only recruited and grew in predator-exclusion cages, because without this protection predation prevented its establishment. In predation experiments, native invertebrate and fish predators removed all invasive ascidians (recruits and adults) in benthic habitats, which contrasted with the high adult survival of the native ascidian P. chilensis. The refuge from a number of benthic predators facilitates the establishment of large populations of invasive species on suspended structures. We present a conceptual model of the invasion processes that includes the anthropogenic structures as a transitional stepping-stone that facilitates invasion by enhancing and prolonging propagule supply to surrounding natural communities. Those established invaders might then overcome biotic resistance during time periods when populations of consumers or competitors are weakened by natural or anthropogenic disturbances. Our results suggest that the conservation of natural habitats with a high diversity of native predators can be an effective means to prevent the spread of invasive species growing on suspended structures.     

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

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

深海底栖有孔虫在古海洋学研究中的应用

通过对海洋沉积物的调查分析,探讨地质历史上海洋环流模式和海水物理化学性质的变化是古海洋学研究的重要内容之一。深海底栖有孔虫对环境变化敏感、其壳体稳定同位素和微量元素与海水性质密切相关使得它成为研究古海洋学的有力工具。但同时,底栖有孔虫的生命效应、分布的区域性以及海水自身等因素在一定程度上制约了其应用。本文总结了底栖有孔虫在古海洋学研究中的应用以及限制因素,以利于在具体研究中客观分析,提高古海洋环境重建的可靠性。     

Depth estimation using diameter-thickness ratios in larger benthic foraminifera

The diameter-thickness ratio (D/T) of larger benthic foraminifera varies indirectly with depth. Increased turbulence thickens the test, whilst decreased light intensity causes a flatter test. It is shown that in a nearshore-offshore transect the D/T gradient is similar between reefs, but stretched over a larger depth gradient. In facies analysis an increase in D/T can either be caused by decreased hydrodynamic energy, increased transparency or increased depth (or a combination of these).     

Morphological abnormalities in Late Cretaceous and early Paleocene foraminifer tests (northern Patagonia, Argentina)

Morphological abnormalities are common in Late Cretaceous and early Paleocene foraminifer tests at two localities in northern Patagonia, Argentina. Protelphidium sp. in the Auca Mahuevo section (late Campanian–early Maastrichtian) exhibit abnormal size or shape of the later chambers, with the last chamber commonly larger than normal or inflated and variably extending onto one of the lateral sides of the test; modification of the coiling plane; protuberances near the proloculus or on one or more chambers; a double last chamber, and complex forms. Protelphidium hofkeri Haynes in the Cerro Azul section (Danian) exhibit abnormal size or shape of one or more chambers, producing peripheral irregularities. In addition, there are rare multiple tests in planktic species from the Cerro Azul section, probably teratological specimens. The sedimentology of the sections and the character of the accompanying faunas indicate that the abnormalities in the two benthic foraminiferal taxa were most probably caused by hypersalinity and/or fluctuations in salinity. The fossil occurrence of assemblages with abundant deformed specimens suggests that investigators should carefully look at many aspects of the environment before concluding that anthropogenic pollution is the only cause of deformations of living benthic foraminifera.     

Benthic Protozoa (Foraminifera,Allogromiida) As Potential Indicator Species for the Sedimentation Record of the Azov–Black Sea Basin Bottom Deposits

Allogromiida are soft-shelled benthic foraminifera protected by thin proteinaceous shells only. They lack rigid calcareous or arenaceous structures and do not undergo fossilization. Some species of the genus Psammophaga are known to accumulate crystals of certain minerals in the cytoplasm during the life cycle. Information on benthic foraminifera (allogromiids) that inhabit the Black Sea and the Sea of Azov and accumulate mineral crystals of unknown nature inside the cells is given in the present paper for the first time. Information on these organisms is proposed to be relevant for the reconstruction of sedimentation basins that existed during the past geological epochs.     

Applications of nummulitids and other larger benthic foraminifera in depositional environment and sequence stratigraphy: an example from the Eocene deposits in Zagros Basin,SW Iran

The Tale-Zang Formation in Zagros Mountains (south-west Iran) is a Lower to Middle Eocene carbonate sequence. Carbonate sequences of the Tale-Zang Formation consist mainly of large benthic foraminifera (e.g. Nummulites and Alveolina), along with other skeletal and non-skeletal components. Water depth during deposition of the formation was determined based on the variation and types of benthic foraminifera, and other components in different facies. Microfacies analysis led to the recognition of ten microfacies that are related to four facies belts such as tidal flat, lagoon, shoal and open marine. An absence of turbidite deposits, reefal facies, gradual facies changes and widespread tidal flat deposits indicate that the Tale-Zang Formation was deposited in a carbonate ramp environment. Due to the great diversity and abundance of larger benthic foraminifera, this carbonate ramp is referred to as a “foraminifera-dominated carbonate ramp system”. Based on the field observations, microfacies analysis and sequence stratigraphic studies, three third-order sequences in the Langar type section and one third-order sequence in the Kialo section were identified. These depositional sequences have been separated by both type-1 and type-2 sequence boundaries. The transgressive systems tracts of sequences show a gradual upward increase in perforate foraminifera, whereas the highstand systems tracts of sequences contain predominantly imperforate foraminifera.