Molecular evidence for an independent origin of modern triserial planktonic foraminifera from benthic ancestors

Gallitellia vivans is the only Recent representative of the triserial planktonic foraminiferal family Guembelitriidae. The origin and evolution of this interesting albeit poorly known family are enigmatic. To elucidate the phylogenetic relationships between G. vivans and other planktonic foraminifera, we sequenced the small subunit ribosomal DNA (SSU rDNA) for comparison to our extensive database of planktonic and benthic species. Our analyses suggest that G. vivans represents a separate lineage of planktonic foraminifera, which branches close to the benthic rotaliids Stainforthia and Virgulinella. Both genera resemble Gallitellia in general morphological appearance, having elongate triserial tests at least in their early ontogenic stages. The divergence time of G. vivans is estimated at ca. 18 Ma (early Miocene), suggesting an origin independent from the Cretaceous and Paleogene triserial planktonic foraminifera. Our study thus indicates that modern triserial planktonic foraminifera are not related to the Cretaceous–Paleogene triserial species, and that the sporadic occurrences in the fossil record are not the result of poor preservation, but reflect multiple transitions from benthic to planktonic mode of life.     

Benthic origin and earliest evolution of the first planktonic foraminifera after the Cretaceous/Palaeogene boundary mass extinction

There are several hypotheses on the origin and evolution of the earliest Danian planktonic foraminifera. Most experts suggest that they descended from a few opportunist planktonic foraminifera species, mainly of the genera Guembelitria and Hedbergella, which are usually considered to be survivors of the Cretaceous/Paleogene (K/Pg) boundary mass extinction. Nevertheless, early Danian specimens of Guembelitria and Hedbergella remained morphologically well separated from the associated parvularugoglobigerinids (i.e. Parvularugoglobigerina and Palaeoglobigerina), the first trochospiral planktonic foraminifera appearing after the K/Pg boundary event. The most likely alternative is a benthic origin for the parvularugoglobigerinids, which would be consistent with molecular phylogenetic studies that have suggested several episodes of benthic-planktonic transitions in the evolutionary history of planktonic foraminifera. A review of material from the El Kef section and other Tunisian sections supports the previous hypothesis that the buliminid genus Caucasina is the ancestor of the first parvularugoglobigerinids (i.e. Parvularugoglobigerina longiapertura and Palaeoglobigerina alticonusa), on the basis of similarities in test and apertural morphologies and wall texture. The intermediate morphotypes between caucasinids and parvularugoglobigerinids, which appeared approximately 3–5 kyr after the K/Pg boundary, are assigned to Pseudocaucasina antecessor gen. nov. sp. nov.     

Experimental dissolution of a fossil foraminiferal assemblage (Paleocene–Eocene Thermal Maximum,Dababiya, Egypt): Implications for paleoenvironmental reconstructions

Dissolution experiments were carried out on a foraminiferal assemblage from the Paleocene–Eocene Thermal Maximum (PETM) at Dababiya, Egypt, in order to: 1) reveal the effects of differential dissolution on the composition of the foraminiferal assemblage and 2) develop objective criteria for the evaluation of dissolution in foraminiferal assemblages used in early Paleogene paleoenvironmental reconstructions, particularly with respect to neritic Midway-type assemblages from the Paleocene/Eocene transition. Our results confirm two general observations on modern foraminifera: 1) planktic foraminifera are much more vulnerable to dissolution than benthic foraminifera, leading to depressed P/B ratios and 2) dissolution susceptibility differs between size fractions, with the smaller specimens dissolving more rapidly than the bigger ones, leading to a larger average size of the remaining assemblage. Within a size fraction, wall structure and thickness are considered to be the main factors controlling differential dissolution susceptibility. We propose a ranking scheme for taxa with respect to dissolution resistance. Among the benthic taxa, Lenticulina is most resistant, followed by the agglutinated Gaudryina cf. ellisorae and Alabamina midwayensis. Biserial and triserial hyaline taxa and the porcelaneous Spiroloculina sp. are most susceptible to dissolution, whereas rotaliines, such as Cibicidoides and Anomalinoides have an intermediate susceptibility. This implies that mild dissolution of a Midway-type benthic assemblage leads to a relative enrichment in Lenticulina, Gaudryina and rotaliines. Amongst planktic foraminifera, the muricate taxa Acarinina and Morozovella are most resistant, followed by the cancellate Subbotina. The smooth and generally small Globanomalina and Zeauvigerina are least resistant to dissolution. Our data enable to objectively evaluate various degrees of dissolution in benthic and planktic foraminiferal assemblages retrieved from the lower Paleogene Tethyan outcrops. In this way taphonomic artifacts can be readily distinguished from paleoenvironmental signals affecting the primary composition of the assemblages. More generally, we propose that the combined use of foraminiferal numbers, P/B ratio and relative abundances of non-calcareous agglutinated taxa and Lenticulina may provide a powerful proxy for assessing dissolution in hemipelagic assemblages from Cenozoic and upper Cretaceous continental margins. In order to achieve more robust pre-Quaternary paleoenvironmental reconstructions based on quantitative foraminiferal data, application of dissolution proxies, like proposed here, or in slightly modified form, should become a more widely used micropaleontologic procedure. Particularly continental margin studies dealing with major biotic events (e.g. PETM) or employing P/B ratios for sea-level reconstructions should benefit from such an approach.     

Phylogeny and Rates of Molecular Evolution of Planktonic Foraminifera: SSU rDNA Sequences Compared to the Fossil Record

Planktonic foraminifera are marine protists, whose calcareous shells form oceanic sediments and are widely used for stratigraphic and paleoenvironmental analyses. The fossil record of planktonic foraminifera is compared here to their molecular phylogeny inferred from ribosomal DNA sequences. Eighteen partial SSU rDNA sequences from species representing all modern planktonic families (Globigerinidae, Hastigerinidae, Globorotaliidae, Candeinidae) were obtained and compared to seven sequences representing the major groups of benthic foraminifera. The phylogenetic analyses indicate a polyphyletic origin for the planktonic foraminifera. The Candeinidae, the Globorotaliidae, and the clade Globigerinidae + Hastigerinidae seem to have originated independently, at different epochs in the evolution of foraminifera. Inference of their relationships, however, is limited by substitution rates of heterogeneity. Rates of SSU rDNA evolution vary from 4.0 × 10⁻⁹ substitutions/site/year in the Globigerinidae to less than 1.0 × 10⁻⁹ substitutions/site/year in the Globorotaliidae. These variations may be related to different levels of adaptation to the planktonic mode of life. A clock-like evolution is observed among the Globigerinidae, for which molecular and paleontological data are congruent. Phylogeny of the Globorotaliidae is clearly biased by rapid rates of substitution in two species (G. truncatulinoides and G. menardii). Our study reveals differences in absolute rates of evolution at all taxonomic levels in planktonic foraminifera and demonstrates their effect on phylogenetic reconstructions. Received: 21 January 1997 / Accepted: 17 April 1997     

Integrated biostratigraphy based on planktonic foraminifera and dinoflagellates across the Cretaceous/Paleogene (K/Pg) transition at the Izeh section (SW Iran)

The present work is based on semi-quantitative study carried on detailed sampling (samples are spaced by 5, 10 and 15 cm close to the boundary) of an essentially continuous and expanded section crossing the Cretaceous–Paleogene (K/Pg) boundary in Iran. By this work, we attempt to detail biostratigraphy based on planktonic foraminifera biozones and correlate biozones and subzones with dinocyst events. The entire Cretaceous–Paleogene interval contains rich, diversified and well-preserved planktonic foraminifera and dinoflagellate cyst assemblages. Four planktonic foraminiferal biozones have been recognized across the Cretaceous–Paleogene transition (K/Pg): Abathomphalus mayaroensis Biozone including Plummerita hantkeninoides Subzone from the Late Maastrichtian and Guembelitria cretacea (including Hedbergella holmdelensis and Parvularugoglobigerina longiapertura subzones), Parvularugoglobigerina eugubina Biozone and Parasubbotina pseudobulloides Biozone belonging to the Early Danian. These biozones have been correlated with four dinocyst biozones: the Manumiella seelandica Biozone belonging to the Late Maastrichtian and the Alisocysta reticulata, Senoniasphaera inornata and Damassadinium californicum biozones from the Early Danian. At this section, like at the El Kef section (GSSP for the K/Pg) and the auxiliary sections, an Ir anomaly is detected indicating the K/Pg boundary. This geochemical anomaly coincides also with mass extinctions of planktonic foraminifera species. The extinct species are in particular the large, complex tropical and subtropical taxa dwelling in subsurface and lower photic water. The mass extinctions at the Izeh section occurred over a succinct period of time similar to the K/Pg type section at El Kef (Tunisia). These sudden mass extinctions indicate a catastrophic pattern event occurring at the Maastrichtian/Danian boundary. In contrast the organic-walled dinocysts were less affected by the mass extinction and most species crossed the K/Pg boundary without showing mass and sudden extinctions. Nevertheless, they showed changes in their assemblages’ structure beyond the K/Pg boundary. Especially, Manumiella seelandica and M. druggii, typical species of Antarctic Maastrichtian dinocysts assemblages, occur in coeval deposits at the Izeh section; they persist through the Lower Danian and, like in Tunisia (e.g., El Kef section, Ellès section) show an obvious increase in relative abundance.     

Benthic foraminiferal turnover across the Cretaceous/Paleogene boundary at Agost (southeastern Spain): paleoenvironmental inferences

We studied Upper Cretaceous and Lower Paleogene benthic foraminifera from the Agost section (southeastern Spain) to infer paleobathymetrical changes and paleoenvironmental turnover across the Cretaceous/Paleogene (K/P) transition. Benthic foraminifera indicate uppermost bathyal depths at Agost during the Abathomphalus mayaroensis Biochron (from about 400 kyr before the K/P boundary) through the early Plummerita hantkeninoides Biochron (about 120–150 kyr before that boundary). The depth increased to middle bathyal for the remainder of the Cretaceous, and remained so for the Danian part of the studied section (Parasubbotina pseudobulloides Biochron, at least 200 kyr after the K/P boundary). There were no perceivable bathymetrical changes at the K/P boundary, where 5% of the species became extinct, and the species composition of the benthic foraminiferal fauna changed considerably. Below the boundary, infaunal morphogroups constitute up to 65–73% of the faunas. Directly above the boundary, in the black clays of the lower Guembelitria cretacea Biozone, benthic foraminifera are rare. Several opportunistic taxa (e.g. the agglutinant Haplophragmoides sp.) have short peaks in relative abundance, possibly reflecting low-oxygen conditions as well as environmental instability, with benthos receiving food from short-lived, local blooms of primary producers. Above the clays through the end of the studied interval, epifaunal morphogroups dominate (up to 70% of the assemblages) or there is an even mixture or epifaunal and infaunal morphogroups. Infaunal groups do not recover to pre-extinction relative abundances, indicating that the food supply to the benthos did not recover fully over the studied interval (about 200 kyr after the K/P boundary). The benthic foraminiferal faunal changes are compatible with the direct and indirect effects of an asteroid impact, which severely destabilized primary producers and the oceanic food web that was dependent upon them.     

First radiation of Cretaceous planktonic foraminifera with radially elongate chambers at Angles (Southeastern France) and biostratigraphic implications

The study of planktonic foraminifera from the Lower Cretaceous succession at Angles (Southeastern France) directly correlated with ammonites, confirms that the origin of morphotypes with radially elongated chambers occur earlier than previously recorded. In particular, the species Hedbergella semielongata and Hedbergella roblesae bearing subclavate to clavate chambers first appear in the upper Hauterivian, just predating the onset of the oceanic anoxic Faraoni event. Based on these observations, a new zonation is proposed.     

Paléoécologie des foraminifères benthiques de nouvelles coupes de basse latitude du passage Crétacé-Paléogène : Coupes de l’Oued Es Smara et de l’Oued Abiod (région de Téjerouine,NW Tunisie)

Due to an impact of a bolide at the K/Pg boundary, the planktonic foraminifera have suffered sever mass extinction. However, no small Benthic Foraminifera species have documented mass extinction at the K/Pg boundary. Nevertheless, many species showed disturbance. The Maastrichtian assemblages may be different from those of the lower Paleogene by their species content, diversity and frequencies. At Oued Es Smara and Oued Abiod sections, the small benthic foraminifera indicate lower bathyal environment, and manifest significant faunal turnover. Until the uppermost Maastrichtian, their assemblages are highly diversified, with 77 species and 76 species respectively at Oued Es Smara and Oued Abiod sections. These are dominated by endobenthic morphotypes. At the K/Pg boundary, although 33 species (42,85%) (Oued Es Smara section) and 27 species (35,52%) (Oued Abiod section) of them seem to disappear, but only few species have really extinct such as Arenobulimina obesa. Nevertheless, the majority of species persist elsewhere at the Danian (e.g., Pseudoglandulina manifesta, Cibicioides proprius, Clavulinoides amorpha, Coryphostoma plaitum, Pullenia coryelli). At the lower Danian, the survivor Maastrichtian species are of 58% (Oued Es Smara) and 65% (Oued Abiod). Throughout the Parasubbotina pseudobulloides subzone, 4 others species were progressively disappeared. They are oligotrophic and low oxygen tolerant. About the Masstrichtian species, at the two studied sections (e.g. Gaudryina inflata and Tritaxia midwayensis) they seem to be more trophic exigent. Consequently, the benthic Foraminifera did not suffer massive extinction at the K/Pg boundary, but their assemblages underwent a significant faunal turnover which reflects important environmental changes. These changes are compatible with the catastrophic scenario induced by the large asteroid impact.     

Benthic foraminiferal assemblages of the Cretaceous platform carbonate succession in the Yavca area (Bolkar Mountains, S Turkey): biostratigraphy and paleoenvironments

The microbiostratigraphic analysis of the three outcrop sections from the Cretaceous inner platform carbonate succession in the Yavca area (Bolkar Mountains) allows to recognize the four local benthic foraminiferal zones. These are: (1) Voloshinoides murgensis and Praechrysalidina infracretacea Cenozone in the Lower Aptian; (2) Pseudorhapydionina dubia and Biconcava bentori Cenozone in the Middle-Upper Cenomanian; (3) Ostracoda and Miliolidae Interval Zone in the probable Turonian, represented by dolomitized limestones without any significant markers; (4) Moncharmontia compressa and Dicyclina schlumbergeri Cenozone in the Coniacian-Santonian. The benthic foraminiferal assemblages correspond to those in other areas of the Mediterranean realm, with the exception of a lack of alveolinids and orbitolinids due to unfavorable environmental conditions (inner platform, restricted shelf). After the regionally well-known emergence during the late Aptian, Albian and early Cenomanian, very shallow subtidal to intertidal conditions were re-established during the middle-late Cenomanian time. The Coniacian-Santonian benthic foraminiferal assemblage shows an increase in diversity and abundance as a result of open marine influence, confirmed by the presence of larger foraminifera (Dicyclina), Rotaliidae and radiolitid fragments. Thaumatoporella and Aeolisaccus-bearing wackestone intercalations still indicate the existence of sporadic restricted environment conditions. The Cretaceous shallow-water platform carbonate succession of the Yavca area is conformably overlain by gray pelagic limestones with calcispheres and planktonic foraminifera. The Campanian flooding of the Bolkar Da? carbonate platform resulted in drowning of the pre-existing biota and facies.     

Neogene stratigraphy,foraminifera, diatoms,and depositional history of Maria Madre Island,Mexico: Evidence of early Neogene marine conditions in the southern Gulf of California

Foraminifera and diatoms have been analyzed from an upper Miocene through Pleistocene(?) sequence of marine sediments exposed on Maria Madre Island, largest of the Trés Marias Islands off the Pacific coast of Mexico. The Neogene stratigraphic sequence exposed on Maria Madre Island includes a mid-Miocene(?) non-marine and/or shallow marine sandstone unconformably overlain by a lower upper Miocene to uppermost Miocene upper to middle bathyal laminated and massive diatomite, mudstone, and siltstone unit. This unit is unconformably overlain by lower Pliocene middle to lower bathyal sandstones and siltstones which, in turn, are unconformably overlain by upper Pliocene through Pleistocene(?) upper bathyal to upper middle bathyal foraminiferal limestones and siltstones. These beds are unconformably capped by Pleistocene terrace deposits. Basement rocks on the island include Cretaceous granite and granodiorite, and Tertiary(?) andesites and rhyolites. The upper Miocene diatomaceous unit contains a low diversity foraminiferal fauna dominated by species of Bolivina indicating low oxygen conditions in the proto-Gulf Maria Madre basin. The diatomaceous unit grades into a mudstone that contains a latest Miocene upper to middle bathyal biofacies characterized by Baggina californica and Uvigerina hootsi along with displaced neritic taxa. An angular unconformity separates the upper Miocene middle bathyal sediments from overlying lower Pliocene siltstones and mudstones that contain a middle to lower bathyal biofacies and abundant planktonic species including Neogloboquadrina acostaensis and Pulleniatina primalis indicating an early Pliocene age. Significantly, this Pliocene unit contains common occurrences of benthic species restricted to Miocene sediments in California including Bulimina uvigerinaformis. Pliocene to Pleistocene(?) foraminiferal limestones and siltstones characterize submarine bank accumulations formed during uplift of the Trés Marias Island area, and include abundant planktonic foraminifera such as Pulleniatina obliquiloculata and Neogloboquadrina duterteri. Common benthic foraminifera in this unit are indicative of upper bathyal water depths. The Neogene depositional history recorded on Maria Madre Island involves an early late Miocene subsidence event marking formation of the Trés Marias Basin with relatively undiluted diatomaceous sediment deposited in a low oxygen setting. Subsidence and deepening of the basin continued into the early Pliocene along with rapid deposition of terrigenous clastics. Uplift of the basinal sequence began in late Pliocene time accompanied by deposition of upper Pliocene-Pleistocene foraminiferal limestones on a rising submarine bank. Continued episodic uplift of the Neogene deposits brought the island above sea level by late Pleistocene time.     

Distribution of some planktonic foraminifera species in deep-sea cores from the red sea and their relation to eustatic sea-level changes

Distribution patterns of planktonic foraminifera in four sediment cores from the Red Sea are studied. The most common species are Globigerinoides ruber, G. sacculifer, Globigerinella siphonifera and Orbulina universa. G. ruber and G. sacculifer show opposite trends of distribution in the sediment cores. Abundance of the foraminifera during the glacial periods suggests that the connection of the Red Sea to the Indian ocean was not completely interrupted and the salinity conditions were not extreme.However, higher salinities appear to have existed in the northern Red Sea, where most of the planktonic foraminifera that occur in the southern Red Sea are absent. It is inferred that the salinity in the southern Red Sea during the glacial period was less than 50%, whereas higher salinity might have existed in the north where the influence of the Indian Ocean was minimal.     

Improving detection of foraminifera by cathodoluminescence

Cathodoluminescence (CL) studies of Lower–Middle Oxfordian marls and limestones, as well as clasts from the uppermost Turonian–?Early Coniacian conglomerates of the Cracow Upland (southern Poland), reveal that the CL view of foraminifers from some lithologies differs from that in transmitted light. In particular, the CL technique revealed abundant tests of planktonic species Globuligerina oxfordiana in the Middle Oxfordian glauconitic marls, which under transmitted light are either poorly visible or remain completely undetected. Bright red–orange luminescence characterizes originally hyaline aragonitic tests of G. oxfordiana, but also several calcitic benthic species, in spite of their different taxonomic position and original test structure and mineralogy. In sponge microbial boundstones, foraminifers generally do not show the CL emission, or show a weak luminescence. Similarly, Late Cretaceous foraminifera represented mostly by planktonic taxa were detected or their view was clearly improved under CL only in some clasts from the uppermost Turonian–?Early Coniacian conglomerates filling karstic cavities. In other clasts, foraminifera are clearly visible only under normal transmitted light, therefore the luminescence signature is highly spatially variable. These results indicate a strong influence of lithology and diagenesis and rather minor effects of shell structure on luminescence of microfossils. The CL technique can be a useful tool in the detection and documentation of abundance patterns of foraminifers that are poorly preserved under transmitted light.     

Biostratigraphy and geodynamic impact in the uppermost part of the northeastern coastal basin of Togo

Lithological and micropaleontological studies of core-samples from five boreholes drilled in the northeastern part of the coastal basin of Togo allow for a specification of the stratigraphy and the paleogeography of this area during the Maastrichtian and Paleogene. A lithological analysis reveals a marine series consisting of a Lower Maastrichtian unit, a Middle Paleocene to Eocene unit, and an upper unit attributed to the Continental terminal sensu lato. The biostratigraphical study, based on planktonic foraminifera, has led to a characterization of the basin in terms of biozones ranging from the Globotruncana aegyptiaca to the Abathomphalus mayaroensis biozones and biozones P5 to P11, thus specifying a Middle to Upper Maastrichtian, an Upper Paleocene and a Lower to Middle Eocene units. The paleogeographical evolution of the area shows that the series recorded two sedimentary cycles: the first one stops at the end of the Cretaceous and the second one in the Paleogene.     

Paleoceanographic implications of smaller benthic and planktonic foraminifera from the Eocene Pazin Basin (Coastal Dinarides,Croatia)

Summary Smaller benthic and planktonic foraminifera from the clastic sediments of the Pazin Basin (Istria, Croatia) were studied in order to obtain more data about paleoceanographic conditions that existed in the Middle Eocene Dinaric foreland basin. The succession investigated corresponds to the Middle Eocene planktonic foraminiferal zones Globigerapsis kugleri/Morozovella aragonensis (P11), Morozovella lehneri (P12), and Globigerapsis beckmanni (P13). Benthic foraminiferal assemblages from the clastic succession are dominated by epifaunal trochospiral genera suggesting oligotrophic to mesotrophic conditions and moderately oxygenated bottom waters. Planktonic foraminiferal assemblages indicate mesotrophic to eutrophic conditions of the surface waters, with increased eutrophication in the upper part of the section. Water depth, based on the ratio between planktonic and epifaunal benthic foraminifera and on the recognized species of cosmopolitan benthic foraminifera, was estimated to have been between about 900 and 1200 m. The basin was elongated and open to marine currents on both sides allowing good circulation and ventilation of the bottom water.     

Isotopic and distributional evidence of a planktonic habit for the foraminiferal genusStreptochilus Bro¨nnimann and Resig, 1971

Oxygen and carbon isotope analyses show that the biserial foraminiferal genusStreptochilus, which was originally described from pelagic sediments on the Eauripik Rise and Ontong Java Plateau, lived deep in the upper water column within the oxygen minimum layer. The species ofStreptochilus average from 4 to 19% of the foraminiferal assemblages in which benthic forms compose less than 1 or 2%. Specimens ofStreptochilus are selectively dissolved when in contact with the bottom water mass. Their rapid evolutionary turnover of less than a few million years and their wide areal distribution in the equatorial Indo-Pacific are indicative of planktonic foraminifera. Aside from usefulness of the species ofStreptochilus as stratigraphic indices, these Neogene biserial planktonic foraminifera are potential indices of paleoceanographic stratification.     

Benthic foraminifera associated with the invasive ascidian, Didemnum sp. A

The invasive ascidian, Didemnum sp. A, first appeared in New England bays and harbors in the early 1990s, and in the waters around Cape Cod in 1993. While ship traffic was the likely vector introducing the species, its origin and precise date and location of its introduction are presently unknown. Colony surfaces of Didemnum sp. A appear very clean and not favorable substrates for epibiota settlement, but closer inspection revealed the presence of benthic foraminifera. During 2003 and 2004, 52 samples of Didemnum sp. A and other ascidians were collected to determine whether or not the foraminiferal assemblages might also be non-native and thus provide a potential clue to the place of origin of Didemnum sp. A. Sample locations included the New England coast from Connecticut to Maine (with a concentration in the Cape Cod area), northern California, Zeeland, The Netherlands, and Shakespeare Bay, New Zealand. From New England samples, 18 species of benthic foraminifera were identified. The most common species represented were Cornuspira involvens, C. planorbis, Elphidium galvestonense, E. margaritaceum, Glabratellina lauriei, Miliolinella subrotunda, Quinqueloculina bicornis, and Rosalina floridana. Foraminiferal assemblages on Didemnum sp. A from other regions sampled were composed of the same cosmopolitan species found in New England, plus other species which were indigenous to each region. Because no exotic foraminifera species were found it is concluded that Didemnum sp. A likely did not introduce non-native foraminifera originating from their native habitats into the New England region.     

Extreme differences in rates of molecular evolution of foraminifera revealed by comparison of ribosomal DNA sequences and the fossil record

Foraminifera have one of the best known fossil records among the unicellular eukaryotes. However, the origin and phylogenetic relationships of the extant foraminiferal lineages are poorly understood. To test the current paleontological hypotheses on evolution of foraminifera, we sequenced about 1,000 base pairs from the 3' end of the small subunit rRNA gene (SSU rDNA) in 22 species representing all major taxonomic groups. Phylogenies were derived using neighbor- joining, maximum-parsimony, and maximum-likelihood methods. All analyses confirm the monophyletic origin of foraminifera. Evolutionary relationships within foraminifera inferred from rDNA sequences, however, depend on the method of tree building and on the choice of analyzed sites. In particular, the position of planktonic foraminifera shows important variations. We have shown that these changes result from the extremely high rate of rDNA evolution in this group. By comparing the number of substitutions with the divergence times inferred from the fossil record, we have estimated that the rate of rDNA evolution in planktonic foraminifera is 50 to 100 times faster than in some benthic foraminifera. The use of the maximum-likelihood method and limitation of analyzed sites to the most conserved parts of the SSU rRNA molecule render molecular and paleontological data generally congruent.      

Enregistrement des fluctuations du niveau marin dans l’Holocène supérieur du lac Retba (Sénégal) par les foraminifères

A study of sea level fluctuation records based on foraminiferal assemblages was carried out for the first time in a sediment core taken at the Lac Retba edge. A total of 37 foraminifera species were collected along the core, among which most are known in the modern estuaries and lagoons of Senegal. A succession of four assemblages dominated by Ammonia parkinsoniana and A. tepida were identified; they correspond to different stages of evolution of the lake. The first association, at the base of the core, indicates a lagoon slightly opened to the sea and bordered with vegetation, under relatively humid climate. The second has the richest and most diversified microfauna with a high proportion of coastal, marine benthic and planktonic species (about 10%) that indicate a small coastal gulf bordered with mangroves. The third association is oligospecific and typical of a closed and hypersaline lagoon under a dry climate. The last association again contains coastal, marine benthic and scarce planktonic species (3%) indicating a change to a saltier and more open lagoon under dry climate. The disappearance of planktonic foraminifera at the top of the core indicates the closure of the lagoon. The biocenotic indicators are evidence for two marine intrusions that are referred to the Dakarian (3000 years B.P.) and Saint-Louisian (2140-680 years B.P.) stages of the Upper Holocene of the senegalo-mauritanian stratigraphic scale. Evidence of the Lac Retba closure since 680 years B. P. appears in the core by high reduction of foraminifers’ abundance and diversity absence of planktonic species, and salts precipitated at the bottom of the lake.     

Magnesium and strontium compositions of recent symbiont-bearing benthic foraminifera

Minor and trace elements in foraminiferal carbonates are potential paleo-proxies of climate, nutrient and seawater composition. There are very few reports of trace element composition of symbiont-bearing, larger foraminifera that are known to be important constituents of shallow-marine, modern and ancient carbonates. In this paper we examine the range of variation in Mg and Sr content of Recent species of these foraminifera from a lagoon of Lakshadweep Atoll (Indian Ocean) and Akajima Islands, Japan. Two hyaline species, Amphistegina lessonii and Neorotalia calcar,and two porcellaneous species, Amphisorus hemprichii and Marginopora vertebralis were collected live from Lakshadweep islands. Mg / Ca in these foraminifera is of an order of magnitude higher than the values reported for planktonic and symbiont-free benthic foraminifera. The Sr / Ca values are, however, comparable with the reported values in other foraminiferal taxa and they are found to vary within a narrow range. Electron-probe micro-analysis of three symbiont-bearing benthic species indicates spatial heterogeneity of high orders in Mg / Ca composition in all the species. The annual variation in temperature and pH of the lagoon water cannot explain the observed amplitude of the compositional variation. The photosynthesis and respiration of the symbionts and host foraminifera are possibly the major cause of compositional heterogeneity in individual tests, as has also been recently postulated for symbiont-bearing planktonic foraminiferal species. It highlights the need to isolate biological factors and necessitates species-specific paleotemperature scale in paleoclimatic analysis. We also analyzed δ¹⁸O, δ¹³C, Ca, Mg and Sr in carefully dissected chambers of a reef-dwelling, porcellaneous benthic foraminifer, Marginopora kudakajimaensis, collected live in four seasons. A moderate positive correlation is observed between Mg / Ca and temperature. However, large inter- and intra-test variation in Mg limits the precision of Mg / Ca as palaeotemperature proxy. The Sr / Ca of the test calcite is unrelated to temperature of the sea water. The δ¹³C of M. kudakajimaensis does not correlate with δ¹⁸O, Mg / Ca or Sr / Ca.     

Book Reviews/Buchbesprechungen

Book Reviewed in this article:
Fossils & Strata, 44 (1997) . Containing: C hristensen , W. K. & S chulz , M.-G.
Capy, P.; B azin , C.; H iguet , D.; L angin , T.: Dynamics and Evolution of Transposable Elements .
W idmark , J. G. V.: Deep-sea benthic foraminifera from Cretaceous/Paleogene boundary strata in the South Atlantic – taxonomy and paleoecology .