Exceptional record of submarine cave communities from the Pleistocene of Sicily (Italy)

An Early Pleistocene benthic community, discovered inside the Rumena Cave in NW Sicily, Italy, was studied. Analysis of the community led to the recognition of several encrusting species – notably scleractinians, bryozoans, serpuloideans, cirripeds, foraminifera and brachiopods – and borings mostly referable to the ichnogenus Gastrochaenolites. All fossils detected are typical of the present‐day hard‐surface submarine cave biota, at both high taxonomic rank and species level. The biogenic crust, restricted to a few sectors of the cave but locally up to few centimetres thick, largely consist of scleractinians, mainly represented by dendrophylliids. Bryozoans and serpuloideans are also present with Hippaliosina depressa and Spiraserpula massiliensis, locally forming multi‐layered sheets and dense specimen aggregates, respectively. Basing mostly on the composition of the encrusting community and on morphological/morphometric features of some species, it has been hypothesized that at least part of the cave was blind when the crust formed, possibly at relatively shallow depth in a sheltered setting or, more probably, at higher depths, below the fair weather swell zone. Encrustations and borings on the rocky cave ceiling and on occasional speleothems broken surfaces document subsequent phases of cave submersion/colonization separated by emersion/erosion phases. The importance of the Early Pleistocene fossils of the Rumena Cave for the knowledge of submarine cave communities through time and for the understanding of sea‐level variations and the uplift of the area has been remarked.     

Changes in quaternary oceanographic bottom conditions as documented by foraminifera in sediment cores from the slope of the exmouth plateau (off Western Australia)

Benthonic foraminifera from 15 surface- and 70 core-sediment samples (three cores) from the Western Australian continental margin (Exmouth Plateau) were quantitatively investigated. As far as possible, the occurrences of the species were correlated with the recent oceanographic conditions especially as related to water depths. The sampled sites are separated from the Australian Shelf by the Montebello Trough so that the taphocoenoses cannot become adulterated by foraminifera from the shelf. Only 1% or less of the foraminifera are benthonic species. The benthonic foraminiferal faunas of the surface sediment samples from 3800 to 1700 m are composed of about the same deep-water species as described from southeastern Indian Ocean basins by Corliss (1979a). They differ from one another only because different species dominate the faunas dependent on different water depth and/or water mass. Above 1700 m the character of the benthonic foraminiferal fauna is slowly changing from abyssal to bathyal. Downcore the benthonic foraminifera show marked changes in abundance patterns, which partly parallel the changes which can be observed with the “cool” and “warm” indicating planktonic foraminifera, partly show regular phase displacement with these changes. The different abundances of the same species in a core from, for instance, 3000 m or from 2000 m water depth, respectively, prevent a direct correlation between the cores. It also proved impossible to correlate the changes in the frequency patterns of certain species to certain paleo-oceanographic conditions. Examples for this are given.     

Fossil planktic foraminifera (an overview)

Planktic foraminifera are unicellular marine organisms able to produce calcareous tests, which can be fossilized and, therefore, can give important information about the geologic record. In this overview a summary of the present knowledge of fossil planktic foraminifera together with the gradual steps from the earliest and pioneering to the more recent studies on this microfossil group, is given. In particular, the criteria at the base of the classification of these organisms from the earlier studies until the present, are described and summarized. An overview of the biostratigraphic schemes based on species first and last occurrences and assemblages from different latitudes and different regions is also given. The evolution and the response of planktic foraminifera to changing environmental condition are summarized from the more primitiveGlobigerina-like Jurassic forms, to the specialized and diversified Cretaceous species, until their dramatic crisis across the Cretaceous/Tertiary boundary, their successive recovery in the Paleocene and their evolution toward modern organisms. Finally, an overview is given of the use of planktic foraminifera to reconstruct paleoenvironments from the more simple methods to those implying more sophisticated and recent techniques.      

Climate-Driven Range Extension of Amphistegina (Protista,Foraminiferida): Models of Current and Predicted Future Ranges

Species-range expansions are a predicted and realized consequence of global climate change. Climate warming and the poleward widening of the tropical belt have induced range shifts in a variety of marine and terrestrial species. Range expansions may have broad implications on native biota and ecosystem functioning as shifting species may perturb recipient communities. Larger symbiont-bearing foraminifera constitute ubiquitous and prominent components of shallow water ecosystems, and range shifts of these important protists are likely to trigger changes in ecosystem functioning. We have used historical and newly acquired occurrence records to compute current range shifts of Amphistegina spp., a larger symbiont-bearing foraminifera, along the eastern coastline of Africa and compare them to analogous range shifts currently observed in the Mediterranean Sea. The study provides new evidence that amphisteginid foraminifera are rapidly progressing southwestward, closely approaching Port Edward (South Africa) at 31°S. To project future species distributions, we applied a species distribution model (SDM) based on ecological niche constraints of current distribution ranges. Our model indicates that further warming is likely to cause a continued range extension, and predicts dispersal along nearly the entire southeastern coast of Africa. The average rates of amphisteginid range shift were computed between 8 and 2.7 km year⁻¹, and are projected to lead to a total southward range expansion of 267 km, or 2.4° latitude, in the year 2100. Our results corroborate findings from the fossil record that some larger symbiont-bearing foraminifera cope well with rising water temperatures and are beneficiaries of global climate change.     

川西,藏东晚三叠世有孔虫及海参骨片

金沙江上游两岸为川西、藏东地区。此区的四川省白玉县、西藏贡觉县及江达县的晚三叠世诺利期沉积中发现了有孔虫１８属２１种,海参骨片５属８种。三叠纪海参在中国为首次报道：白玉县及贡觉县的三叠世有孔虫也为首次发现。这一动物群在区域地质史及古地理研究上具有一定意义。     

The meaning of birth and death (in macroevolutionary birth-death models)

Birth–death models are central to much macroevolutionary theory. The fundamental parameters of these models concern durations. Different species concepts realize different species durations because they represent different ideas of what birth (speciation) and death (extinction) mean. Here, we use Cenozoic macroperforate planktonic foraminifera as a case study to ask: what are the dynamical consequences of changing the definition of birth and death? We show strong evidence for biotic constraints on diversification using evolutionary species, but less with morphospecies. Discussing reasons for this discrepancy, we emphasize that clarity of species concept leads to clarity of meaning when interpreting macroevolutionary birth–death models.     

Foraminifères benthiques complexes du Bathonien supérieur au Kimméridgien inférieur de la marge atlantique marocaine (« Bassin » d’Agadir) : stratigraphie et paléobiogéographie

The carbonate sediments of the Agadir Basin (Morocco) deposited during the Upper Bathonian-Lower Kimmeridgian p.p. interval are rich in agglutinated complex benthic foraminifera. The detailed inventory of several field-sections allows to identify and to describe these benthonic foraminifera, some of them being recorded for the first time in Morocco. They belong to the main genera Archaeosepta, Andersenolina, Praekurnubia, Pseudocyclammina, Everticyclammina, Alveosepta, Kurnubia, Parurgonina, Neokilianina, Labyrinthina and Rectocyclammina. The stratigraphic and paleobiogeographic distribution of these foraminifera seem to be related to the geodynamic events occurring on the Moroccan Atlantic margin during this time-interval. The stratigraphic distribution of the principal species is generally controlled by the occurrence of ammonites, brachiopods and of some foraminifera which are considered as good stratigraphic markers.     

Distribution of living benthic Foraminifera on the back-reef and outer slopes of a high island (Moorea French Polynesia)

Living foraminifera were examined on both sides of the barrier reef across a section situated on the north-west part of the high island of Moorea (Society Archipelago, French Polynesia). In all, 87 species were found: 62 in the back-reef area (fringing reef, channel and barrier reef) and 72 on the outer slope; 47 were common to both zones. This study points out the importance of substrates in the distribution pattern of foraminifera. The sediments are essentially inhabited by epipsammitic species, whereas algae are colonized mainly by free-living foraminifera: miliolids in the back-reef area; rotaliids in the outer slope. Free-living individuals are much less abundant in the fringing reef, channel and inner part of the barrier reef than in the other parts of the study area. The two major groups on the two sides of the barrier reef are very different in size, test building and probably nutritional modes. In the back-reef area, epipsammitic taxa, mainly composed of small agglutinated species, dominate the fauna; in the outer slope large calcareous symbiotic Amphistegina are the major components of the living association (50 to 70%).     

Benthic foraminifera across the Cretaceous–Tertiary (K–T) boundary: a review

The response of the Earth’s biota to global change is of fundamental interest to paleontologists, but patterns of change in paleontologic data are also of interest to a wider spectrum of Earth scientists in that those patterns are of great significance in constraining hypotheses that attempt to explain physical changes in the Earth’s environment. The Cretaceous–Tertiary (K–T) boundary is a case in point. Some paleontologists have criticized the bolide impact hypothesis, not because they deny the impact but because the proposed effects of that impact do not always conform to the available paleontological data. Benthic foraminifera are of particular interest in this context because it has been suggested for over 20 years that shallow-water benthic foraminifera were affected more severely than deep-water benthic foraminifera by events at the K–T boundary. This observation adds to the fact of planktonic foraminiferal extinction and indicates that K–T boundary environmental effects were largely restricted to shallow waters. In this paper I review all published works on smaller benthic foraminifera at the K–T boundary and conclude the following. (1) Shallow-water benthic foraminifera were not more severely affected than deeper dwelling species. True extinction, as opposed to local extinction and/or mass mortality, is generally quite low no matter what the water depth. (2) The data are not sufficient in quality, quantity and geographic range to conclude that there is a latitudinal pattern of extinction. (3) In general, biotic changes (such as they are) begin before the boundary in shallow and intermediate depth waters and at the boundary in deep water. Disagreements about the placement of the boundary and the presence, absence and duration of hiatuses hinder more precise conclusions. (4) There appears to be preferential survivorship of epifaunal species into the early Danian with a short interval dominated by infaunal taxa in the earliest Danian. This pattern can best be explained by short-lived input of increased amounts of organic matter at the boundary followed by a sudden collapse of primary productivity and, hence, major reduction or cessation of organic flux to the seafloor. In summary, based on the current dataset, smaller benthic foraminifera, no matter whether they lived in shallow or deep waters, high or low latitudes, or infaunal or epifaunal microhabitats, survived the environmental events across the K–T boundary quite well. Mass extinction does not characterize this group of organisms at this time.     

Une espèce nouvelle du Kimméridgien du Portugal (Estremadura): Aulacomyella abadiensis nov. sp. (Bivalvia,Posidoniidae)

The Kimmeridgian Series situated at the North ofLisbon is included in the Portuguese Lusitanian basin. Its sandy clays pelagic facies with planctonic foraminifera contains a new species of bivalvia: Aulacomyella abadiensis related to A. neogeaeImlay from the Middle Kimmeridgian of Mexico and from the Kimmeridgian of Crimea. This species confirms the Middle Kimmeridgian.     

Pleistocene agglutinated foraminifera from the Lomonosov Ridge and Amundsen Basin, Arctic Basin. Initial report on piston cores 2177-5 (KAL) and 2176-3 (KAL)

Noncalcareous Pleistocene sediments of the Central Arctic Ocean contain sparse benthic foraminiferal assemblages consisting entirely of agglutinated taxa. Deep water agglutinated foraminifera are studied from two piston cores collected from the Lomonosov Ridge and Amundsen Basin [Cores PS 2177-5 (KAL) and 2176-3 (KAL)]. Core PS 2177-5 (KAL) contains an assemblage of 10 species, dominated by Cyclammina pusilla Brady, and is interpreted to reflect a bathyal environment with variable organic flux and nutrition levels. Core PS 2176-3 (KAL) in the Amundsen Basin yielded a very depauperate benthic foraminiferal assemblage. It is assumed that the environment was inhospitable for agglutinated foraminifera.     

Larger foraminifera as a substratum for encrusting bryozoans (Late Oligocene,Tethyan Seaway,Iran)

Considering the diversity and abundance of larger foraminifera examined from a wide range of Late Oligocene to Early Miocene palaeoenvironments in the Tethyan Seaway, encrusting bryozoans make extremely little use of their tests as substratum. Significant encrustations by bryozoans were exclusively found on large (ø c. 6 cm), undulating tests of Lepidocyclina spp., on which, however, a remarkable 34 taxa of encrusting bryozoans were recorded. This shallow-water fauna of Chattian age was analyzed in respect of the bryozoan taxa present, colony growth type, and mode of budding, colony size, as well as onset of reproduction. Taxic and morphological similarities between the fossil assemblage and modern faunas encrusting mobile substrata indicate a long history of bryozoans as part of the interstitial habitat, while the tests of certain larger foraminifera may have played a significant role in the evolution of shallow-water bryozoans by providing substrata for encrusting species in otherwise unfavorable environments.     

Ontogeny and phylogeny of Upper Cenomanian rotaliporids (Foraminifera)

The fossil record of planktonic foraminifera is a key source of data on the evolution of marine plankton. One of the most distinctive groups of Cretaceous foraminifera, the rotaliporids, widely used as a stratigraphic index, has always been considered to be a monophyletic clade. New data on the coiling direction and persistent morphological features of the late rotaliporids from the Upper Cenomanian of the Western Interior Seaway, USA, and the Vocontian Basin of southeast France is used as a phylogenetic proxy. Dealing with key morphological features, the coiling pattern of these keeled morphotypes proves that the rotaliporids group is polyphyletic and composed of Thalmanninella, that displays a dextral-coiling preference, and Rotalipora s.s., that have a proportionate-coiling mode. The stratigraphically youngest rotaliporids with keels co-occur with globular forms; and all morphologies transitional between these morphotypes are observed. The ontogenetic relationships between them are investigated, indicating that loss of the keel was a selective advantage that enabled those rotaliporids to remain in the surface water, thereby avoiding the expansion of the oxygen minimum zone. Two species are observed: Thalmanninella multiloculata and Rotalipora planoconvexa. These species are interpreted as having arisen by neoteny from Thalmanninella greenhornensis and Rotalipora cushmani respectively.     

Soft-walled monothalamids (Rhizaria: foraminifera) of the Crimean shelf (Black Sea): taxonomic composition and inter-regional patterns of species diversity and distribution

We present new data on monothalamous (single-chambered) foraminifera from the Black Sea Crimean shelf zone between Karkinitsky Gulf in the west to the area near Kerch in the east. Within this region we recognized a total of 40 morphospecies; 8 are assigned, in some cases tentatively, to known species and another 9 to known genera, again sometimes tentatively. Twelve species (all undescribed) are reported here for the first time. The most abundant species are Psammophaga sp. (sensu Gooday et al. 2011), Goodayia rostellata Sergeeva & Anikeeva 2008 and Vellaria pellucida Gooday & Fernando 1992, each of which is evenly distributed in all studied areas. The highest species richness of monothalamous foraminifera was observed in the Yalta region. Based on a multivariate analysis of monothalamid assemblage structure and diversity indices [Shannon (H’), Margalef (D’), eveness Pielou (J’), Simpson (1-λ’) and rarefaction ES(n)], we recognized three groups of stations, corresponding to the Western, Southern and Eastern coasts of the Crimean peninsula. The monothalamid assemblages found in each of these coastal regions exhibit different structural features and are distinguished by certain characteristic species.     

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.     

Phylogenetic placement of Cibicidoides wuellerstorfi (Schwager, 1866) from methane seeps and non‐seep habitats on the Pacific margin

Benthic foraminifera are among the most abundant groups found in deep‐sea habitats, including methane seep environments. Unlike many groups, no endemic foraminiferal species have been reported from methane seeps, and to our knowledge, genetic data are currently sparse for Pacific deep‐sea foraminifera. In an effort to understand the relationships between seep and non‐seep populations of the deep‐sea foraminifera Cibicidoides wuellerstorfi, a common paleo‐indicator species, specimens from methane seeps in the Pacific were analyzed and compared to one another for genetic similarities of small subunit rDNA (SSU rDNA) sequences. Pacific Ocean C. wuellerstorfi were also compared to those collected from other localities around the world (based on 18S gene available on Genbank, e.g., Schweizer et al., 2009). Results from this study revealed that C. wuellerstorfi living in seeps near Costa Rica and Hydrate Ridge are genetically similar to one another at the species level. Individuals collected from the same location that display opposite coiling directions (dextral and sinstral) had no species level genetic differences. Comparisons of specimens with genetic information available from Genbank (SSU rDNA) showed that Pacific individuals, collected for this study, are genetically similar to those previously analyzed from the North Atlantic and Antarctic. These observations provide strong evidence for the true cosmopolitan nature of C. wuellerstorfi and highlight the importance of understanding how these microscopic organisms are able to maintain sufficient genetic exchange to remain within the same species between seep and non‐seep habitats and over global distances.     

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.     

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.     

Effect of the mariculture of the Japanese scallop on foraminifera assemblages in Alekseev Bight, Sea of Japan

The species composition and population densities of foraminifera were studied in scallop farming grounds in Alekseev Bight (Amurskii Bay, Sea of Japan). Ninety-one species of foraminifera were identified; members of the Elphidiidae, Discorbidae, and Ataxophragmiidae were numerically dominant. When the scallop mariculture farm was in operation, the species composition of foraminifera in farming grounds was impoverished and their population densities were an order of magnitude lower than at more distant locations. Between 1988 and 1995, after the liquidation of the farm, the species diversity and population density of foraminifera in farming grounds increased, while there was a general decline of the foraminifera population in the bay. The highest density and species diversity of foraminifera occurred on coarse silty sand and small silty pebbles, and the lowest species diversity was found on silty sediment.