Comparison of radiolarian and sedimentologic paleoproductivity proxies in the latest Miocene–Recent Benguela Upwelling System

Estimating past ocean productivity from ocean sediments often gives different results depending on the measurement used. We have examined a suite of paleoproductivity proxies in latest Miocene–Recent sediments from DSDP Site 532 and ODP Site 1084, two deep-sea sections underlying the Benguela Upwelling System off the Atlantic coast of southern Africa. The productivity history of this system has been previously established via organic carbon concentration, diatom floras and alkenone based estimates of surface water temperature, and shows a change from low productivity in the early Pliocene to sustain high productivity in the late Pliocene–Recent. Each of our samples was split and simultaneously analysed for several proxies of ocean productivity, including organic carbon (TOC%), carbonate, abundance of opaline radiolarians, accumulation rate of benthic foraminifera (BFAR); the radiolarian faunal composition indices Upwelling Radiolarian Index (URI) and the Water Depth Ecology index (WADE); other proxies for opal and carbonate dissolution, plus stable isotopes of benthic foraminifera. Comparisons between proxies in the same measured samples, between sites in downcore plots and to the published productivity record for this region suggest that TOC and radiolarian faunal composition, particularly the WADE index, are good indicators of past productivity, albeit with different sensitivities (log–linear correlation WADE–TOC% r = 0.78, n = 65, p < 0.01). In contrast, carbonate, and carbonate-based proxies such as BFAR primarily reflect changes in dissolution. Radiolarian faunal composition indices do not appear to be affected by bulk opal accumulation or changes in opal preservation. WADE analysis of radiolarian faunas and TOC% measurements appear to be useful proxies for productivity in late Neogene sediments, particularly for sections where opal or carbonate dissolution is significant.     

Latest Cretaceous to Late Paleocene radiolarian biostratigraphy: A new zonation from the New Zealand region

The scarcity of records of Early Paleocene radiolarians has meant that while radiolarian biostratigraphy is firmly established as an important tool for correlation, there has been a long-standing gap between established zonations for the Cretaceous and from latest Paleocene to Recent. It has also led to considerable speculation over the level of faunal change across the Cretaceous/Tertiary (K/T) boundary. Consequently, the discovery of rich and diverse radiolarian assemblages in well-delineated K/T boundary sections within siliceous limestones of the Amuri Limestone Group in eastern Marlborough, New Zealand, is of great significance for biostratigraphy and K/T boundary research.This initial report is restricted to introducing a new latest Cretaceous to mid Late Paleocene zonation based on the radiolarian succession at four of these sections and a re-examination of faunas from coeval sediments at DSDP Site 208 (Lord Howe Rise). Three new Paleocene species are described:Amphisphaera aotea, Amphisphaera kina andStichomitra wero. Six new interval zones are defined by the first appearances of the nominate species. In ascending order these are:Lithomelissa? hoplites Foreman (Zone RK9, Cretaceous),Amphisphaera aotea n. sp. (Zone RP1, Paleocene),Amphisphaera kina n. sp. (RP2),Stichomitra granulata Petrushevskaya (RP3),Buryella foremanae Petrushevskaya (RP4) andBuryella tetradica (RP5). Good age control from foraminifera and calcareous nannofossils permits close correlation with established microfossil zonations. Where age control is less reliable, radiolarian events are used to substantially improve correlation between the sections.No evidence is found for mass extinction of radiolarians at the end of the Cretaceous. However, the K/T boundary does mark a change from nassellarian to spumellarian dominance, due to a sudden influx of actinommids, which effectively reduces the relative abundance of many Cretaceous survivors. An accompanying influx of diatoms in the basal Paleocene of Marlborough, together with evidence for an increase of total radiolarian abundance, suggests siliceous plankton productivity increased across the K/T boundary. Possible causes for this apparently localised phenomenon are briefly discussed.     

Dynamics of spicule production in the marine sponge <Emphasis Type="Italic">Hymeniacidon perlevis</Emphasis> during in vitro cell culture and seasonal development in the field

To characterize the formation of silica spicules, the dynamics of spiculogenesis of an intertidal marine sponge Hymeniacidon perlevis (Montagu 1818) (Porifera: Demospongiae) were investigated by measuring the gene expression of silicatein (the enzyme responsible for spicule silicification) and the dimensional changes of spicules during the developmental process of individual sponges and in cell cultures of primmorphs of archaeocyte-dominant cell populations. The different developmental stages of spicules were documented by time-lapse microscopy and observed by transmission electron microscopy during a 1-month culture period. During its annual life cycle, H. perlevis has four different developmental stages: dormancy, resuscitation, bloom, and decline. Field-grown individual sponge samples at different stages were collected over 7 months (March to September 2005). The dimensions of the silica spicules from these samples were microscopically measured and statistically analyzed. This analysis and the material properties of the spicules allowed them to be classified into four groups representing the different developmental stages of spiculogenesis. Silicatein expression in the bloom stage was more than 100 times higher than that in the other stages and was correlated with the spicule developmental stage. The trend of spicule formation in field-grown sponges was consistent with the trend in cell culture. A new parameter, the maturation degree (MD) of spicules (defined as the ratio of actual to theoretical silica deposition of mature spicules), was introduced to quantify spicule development. Silica spiculogenesis during H. perlevis development was delineated by comparing MD and silicatein expression.     

Spicule records of <Emphasis Type="Italic">Ephydatia fluviatilis</Emphasis> as a proxy for hydrological and environmental changes in the shallow Lake Trasimeno (Umbria,Italy)

This research shows the results of an analysis of siliceous spicules found in sediment cores collected in Lake Trasimeno (Umbria, Italy), a shallow lake that experienced an important water level lowering during the last century. A morphological analysis of sedimentary sponge records revealed that the spicules accumulated in the lake sediments over the last 150 years are attributable to Ephydatia fluviatilis, the only sponge species found in the lake in recent years. The stratigraphic analysis of the cores showed that the abundance and size of the sponge spicules (megascleres) have remarkably decreased, suggesting that a significant depletion of the sponge fauna occurred, particularly during the first half of the twentieth century. A correlation analysis has identified morpho-hydrological and related variables (the theoretical fraction of lake surface subjected to resuspension and the amount of total suspended solids) as the most significant factors explaining the change in density of sponge spicules. Two ecological explanations of the sponge decline are proposed, based on the sensitivity of the sponge both to the availability of suitable hard substrata for colonization, and to the amount of wind-resuspended solids. One-ended sigmoid response curves were obtained by regression and corresponding transfer functions were derived, which allow the mean water depth of the lake and total suspended solids to be inferred from spicule density records. The results support the use of sponge spicules as a paleohydrological and paleoecological proxy, application of which appears particularly promising for shallow-water systems.     

The pre-cephalic development of the skeleton of Amphimelissa setosa (Actinopoda: Nassellarida)

The ontogeny of the skeleton of the nassellarian radiolarian Amphimelissa setosa begins with the formation of three radial bars of silica; these correspond to the medial bar and spines L₁ and L₂ in the adult. By a precise sequence of the addition of various siliceous arcs and spines, this basic structure develops into the eucephalus. A new growth mechanism, in which existing skeletal elements serve as templates for the formation of new skeletal members, is hypothesized to explain the observed developmental patterns. The early ontogeny of nassellarian skeletons can be used to determine which skeletal features develop earliest. Among the structural characteristics used by radiolarian systematists, these conservative features are most suitable for use in interpretation of the taxonomic relationships and phylogeny of the group.     

Radiolarian and sedimentologic paleoproductivity proxies in late Pleistocene sediments of the Benguela Upwelling System,ODP Site 1084

The changing composition of radiolarian faunas from late Neogene deep-sea sediments has been used in recent years as a proxy for changes in marine paleoproductivity. We examine radiolarian faunas, organic carbon content (TOC), opal and coarse-fraction components over the last 270,000 years in sediments from ODP Hole 1084A, drilled in a high productivity upwelling region within the Benguela Upwelling System off the west coast of Africa. Age control is provided by stable oxygen isotope measurements of benthic foraminifera. Prior research has established that late Pleistocene glacial intervals in this upwelling system generally had higher productivity than interglacials. The radiolarian WADE (water-depth ecology) paleoproductivity index correlates well with TOC and opal in these samples, and all three parameters change in synchrony with the benthic isotope curve over all but the MIS 5e–6 time interval. WADE inferred productivity is significantly higher in glacials than interglacials. We conclude that the WADE index is a useful proxy for paleoproductivity at this location, as are also opal and organic carbon accumulation rates. Carbonate and carbonate based indices such as the accumulation rate of benthic foraminifera (BFAR) by contrast do not correlate well either to productivity indices or to the glacial–interglacial cycle, and are interpreted to primarily reflect carbonate dissolution.     

Hexapylocapsa anachoreta n.gen., n.sp., type of a new monocyrtid nassellarian family (Radiolaria) with double-shelled cephalis

The Lower Tithonian Mühlheim Member of the Mörnsheim Formation in the Solnhofen area (Franconian Alb, Southern Germany) contains a very well preserved and rich radiolarian fauna with many new species of Spumellaria, Entactinaria and Nassellaria.Hexapylocapsa anachoreta n. gen., n. sp. described herein is one of such nassellarian taxa. Having a skeleton formed of a small cephalis enclosed within a spongy, bell-shaped cortical test, this new radiolarian represents the type of the new family Hexapylocapsidae which presents structural affinities with the Late Paleozoic family Pylentonemidae.     

南海断面春季放射虫残骸的空间分布及其与表层沉积记录对比

放射虫是揭示古海洋环境信息的重要载体。本文采用虎红染色方法,首次揭示了南海典型断面现代放射虫残骸群的空间分布特征,并比较分析现代水体中放射虫残骸群与海底表层沉积物中放射虫化石群的相互关系。结果表明:南海春季放射虫残骸群比较丰富,其丰度一般呈现出随深度增加而增高的趋势,最高值主要出现在75—300m深即位于叶绿素最大值层和活体高峰深度之下;分析发现研究区放射虫残骸丰度的深度分布受到海流的横向输运影响;春季水体和表层沉积物中放射虫属种组成和丰度分布的差异表明,除了现代放射虫具有季节性变化之外,海流的横向传输也是一个很重要的原因。在150—300m水体中除了Cornutella profunda和Cyrtopera laguncula,还发现了Cycladophora davisiana的残骸,推测典型冷水种C.davisiana由于受到径向翻转流的作用,从中深层水被带到了中上层水体中。     

Distribution of living polycystine radiolarians in the Gulf of Mexico and Caribbean Sea,and comparison with the sedimentary record

Plankton tows and bottom samples from the Gulf of Mexico and Caribbean Sea were examined to study the distribution, ecology, and preservation of modern radiolarians. Abundances of radiolarians from this region, an area of low primary productivity, are low. Diversities are variable in the surface waters; diversity indices decrease with increasing water depth, indicating that fewer niches are occupied by radiolarians in deep water. Living radiolarians are confined to specific water masses, some species ranging between two masses, and can be used as indicators of these water masses in ecologic or paleo-oceanographic studies.The record of radiolarian tests present in the sediment is dissimilar to the record of living radiolarians in the water column, but is similar to the record of empty tests falling through the water column. This indicates that a considerable length of time is involved in the settling of radiolarian tests. It also confirms the validity of ecologic interpretations based on a study of sediments only.     

Provenance analysis of Cenozoic conglomerates: Insights from Lower Cretaceous radiolarians in chert clasts from the Xigaze area,southern Tibet

Conglomerates, exposed on either side of the Yarlung Tsangpo suture zone (YTSZ) in southern Tibet, have attracted wide attention in elucidating uplift and erosion histories of the Himalayan-Tibetan orogen. However, the provenance of these conglomerates remains controversial. Although radiolarian-bearing chert clasts within these conglomerates have received little focus, identification of the radiolarian assemblages they contain could shed light on the provenance of these sedimentary units. We present the first report of Upper Jurassic to Lower Cretaceous (upper Kimmeridgian–lower Barremian) radiolarian assemblages recovered from chert clasts within the Liuqu and Gangrinboche conglomerates in the Xigaze area. To extract radiolarian fossils from independent clasts in the conglomerates, a detailed and efficient experimental process is illustrated. The assemblages are well correlated to those in the YTSZ and Tethyan Himalaya, showing typical Tethyan characteristics. The lithology of chert clasts and ages of constituent radiolarian faunas suggest that the chert clasts in Liuqu and Gangrinboche conglomerates were derived from the Bainang terrane. This interpretation implies that Early Cretaceous accretionary complexes in the YTSZ had been exposed and eroded before deposited as clasts in the Cenozoic Liuqu and Gangrinboche conglomerates.     

Molecular phylogeny of acantharian and polycystine radiolarians based on ribosomal DNA sequences, and some comparisons with data from the fossil record

Polycystines (spumellarians, nassellarians, and collodarians), phaeodarians, and acantharians are marine planktonic protists that have been conventionally and collectively called "radiolaria". Recent molecular phylogenetic studies revealed radiolarian polyphyly with phaeodarians being a separate offshoot. Collodarians and nassellarians are also shown to form a monophyletic group, but other aspects of radiolarian phylogeny, such as interrelations among polycystines and acantharians, remained uncertain. Here, we present molecular phylogenetic analyses including new ribosomal RNA sequences from ten spumellarians and nine nassellarians, based on Bayesian and maximum-likelihood methods. Results indicate that the Polycystinea is a paraphyletic group, with Bayesian analysis suggesting that spumellarians form a clade with acantharians. The heliozoan-like protist Sticholonche appears as a sister to the spumellarian clade. The nassellarian Eucyrtidium is located outside the clade including the other nassellarians and collodarians. The mineralogy of the test of extant radiolarians and the tree topology obtained in this work suggest that acantharians and spumellarians evolved from an ancestor with a siliceous skeleton. Collodarians and nassellarians form a well-supported clade and one might infer from the fossil record that they may have diverged between the Jurassic and the Eocene.     

Silica deposition in Demosponges: spiculogenesis in Crambe crambe

Transmission electron-microscopy images coupled with dispersive X-ray analysis of the species Crambe crambe have provided information on the process of silica deposition in Demosponges. Sclerocytes (megasclerocytes) lie close to spicules or surround them at different stages of growth by means of long thin enveloping pseudopodia. Axial filaments occur free in the mesohyl, in close contact with sclerocytes, and are triangular in cross section, with an internal silicified core. The unit-type membrane surrounding the growing spicule coalesces with the plasmalemma. The axial filament of a growing spicule and that of a mature spicule contain 50%-70% Si and 30%-40% Si relative to that contained in the spicule wall, respectively. The extracellular space between the sclerocyte and the growing spicule contains 50%-65%. Mitochondria, vesicles and dense inclusions of sclerocytes exhibit less than 10%. The cytoplasm close to the growing spicule and that far from the growing spicule contain up to 50% and less than 10%, respectively. No Si has been detected in other parts of the sponge. The megascleres are formed extracellularly. Once the axial filament is extruded to the mesohyl, silicification is accomplished in an extracellular space formed by the enveloping pseudopodia of the sclerocyte. Si deposition starts at regularly distributed sites along the axial filament; this may be related to the highly hydroxylated zones of the silicatein-alpha protein. Si is concentrated in the cytoplasm of the sclerocyte close to the plasmalemma that surrounds the growing spicules. Orthosilicic acid seems to be pumped, both from the mesohyl to the sclerocyte and from the sclerocyte to the extracellular pocket containing the growing spicule, via the plasmalemma.     

Evidence for spicule homology in calcareous and siliceous sponges: biminerallic spicules in Lenica sp. from the Early Cambrian of South China

Botting, J.P., Muir, L.A., Xiao, S., Li, X. & Lin, J.‐P. 2012: Evidence for spicule homology in calcareous and siliceous sponges: biminerallic spicules in Lenica sp. from the Early Cambrian of South China. Lethaia, Vol. 45, pp. 463–475. The relationships of the extant sponge classes, and the nature of the last common ancestor of all sponges, are currently unclear. Early sponges preserved in the fossil record differ greatly from extant taxa, and therefore information from the fossil record is critical for testing hypotheses of sponge phylogenetic relationships that are based on modern taxa. New specimens of the enigmatic sponge Lenica sp., from the Early Cambrian Hetang Biota of South China, exhibit an unusual spicule structure. Each spicule consists of a siliceous core with an axial canal, an organic outer layer and a middle layer interpreted to have been originally calcium carbonate. This finding confirms previous work suggesting the existence of biminerallic spicules in early sponges. Combined with data from other early sponges, the new findings imply that the two fundamental spicule structures of modern sponges were derived from a compound, biminerallic precursor. Spicules are therefore homologous structures in Calcarea and Silicea, and if sponges are paraphyletic with respect to Eumetazoa, then spicules may also have been a primitive feature of Metazoa. □Calcarea, Early Cambrian, Hetang Biota, phylogeny, Silicea, taphonomy.     

Radiolarian skeleton: Morphology of spines,internal framework,and primary sphere

The morphology and evolution of the internal framework, primary inner sphere, and various spines of radiolarian skeletons are considered. A new scheme of successive stages of spine formation is offered. The convergent similarity of radiolarian spines and sponge spicules are discussed.     

A functional evenness index for microbial ecology

Microbial ecologists attempting to describe community structures through the use of synthetic parameters face enormous difficulties. These stem in part from the necessity of using standard taxonomic reference levels in a field where the species level is poorly defined. This paper presents an attempt to obviate this problem. A “functional evenness” index (E) is defined using information measures; it is based directly on the characteristics of the bacteria, as determined, for example, with the API 20B method. Comparisons of this index with classic structure indices, such as taxonomic evenness (Pielou) or systematic dominance (Hulburt), show that it behaves like an evenness index, while bypassing the taxonomic study required before computation of the classic indices. Its use is illustrated with samples of aerobic heterotrophic bacteria obtained from brackish lagoon sediments. We are grateful to our statistician colleagues, Dr. Yves Lepage from Université de Montréal and Dr. André Plante from Université du Québec à Montréal. Both have contributed important elements to the discussion presented in this Appendix.     

Freshwater sponge silicateins: comparison of sequences and exon-intron structure of genes

Siliceous sponge spicules contain silicateins--proteins taking part in biogenic silica precipitation and determination of the spicule morphological features. The exon-intron structure of four silicatein-alpha isoforms: -alpha1,-alpha2, -alpha3 and -alpha4 from endemic baikalian sponge Lubomirskia baicalensis was studied. For eight sponge species, including both cosmopolitan (Spongilla lacustris, Ephydatia muelleri, E. fluviatilis) and Baikal endemic (L. baicalensis, L. incrustans, Baikalospongia intermedia, B. fungiformis, Sw. papyracea) species, seventeen gene fragment sequences of different silicatein isoforms were determined. It was shown that cosmopolitan and endemic Baikalian sponges differ from each other by gene structure (have different length ofintrons). Among Baikalian sponges silicatein-alpha1 has the most variable intron length, and silicatein-alpha4 is the most conservative. Phylogenetic analysis of amino-acid silicatein sequences allow identify different silicatein isoforms, which authentically differ form four clusters on phylogenetic tree. Phylogenetic analysis of exon-intron sequences gives the possibility to separate different sponge species in the clusters.     

DOES COSMOPOLITANISM RESULT FROM OVERCONSERVATIVE SYSTEMATICS? A CASE STUDY USING THE MARINE SPONGE CHONDRILLA NUCULA

The sponge species Chondrilla nucula has a simple morphology and a very wide geographical distribution. To verify whether the latter might be an artifact of the former, samples of this species were collected across 10,000 km of its range, in the Mediterranean, the Caribbean, and the southwestern atlantic. The classical (spicule morphology) and molecular (allozymes) systematic approaches were compared, to try to define the geographic limits between populations and detect possible cryptic species. We found five distinct genetic forms within C. nucula that sometimes showed morphological homogeneity and other times plasticity. The difference in size of spicules could not be related to the clear-cut genetic differences, suggesting that the use of spicule sizes for sponge systematics should be reappraised. The population of one of the genetic forms along 3000 km of the Brazilian coast was highly structured (F_ST = 0.21; N_em = 0.96). Our results reject the null hypothesis of cosmopolitanism of C. nucula and indicate that the putative worldwide distribution of some marine sponges, and possibly many other benthic invertebrates, may be the result of overly conservative systematics. Cryptic species appear to be particularly prevalent when genera are well defined but species are characterized by only a few morphological characters.     

Bacterial Diversity and Distribution in the Holocene Sediments of a Northern Temperate Lake

Sediments contain an abundance of microorganisms. However, the diversity and distribution of microorganisms associated with sediments are poorly understood, particularly in lacustrine environments. We used banding patterns from denaturing gradient gel electrophoresis (DGGE) and 16S rDNA sequences to assess the structure of bacterial communities in the Holocene sediments of a meromictic lake in Minnesota. Cluster analysis of the DGGE banding patterns indicates that the early- and middle-Holocene samples group separately from the late-Holocene samples. About 79% of the recovered bacterial sequences cluster with the α-, β-, δ-, ɛ-, and γ- Proteobacteriaceae and Firmicutes. The remaining ∼21% lack cultured representatives. The taxonomic lineages of bacteria differ statistically among the early-, middle-, and late-Holocene samples, although the difference is smallest between early- and middle-Holocene samples. Early- and middle-Holocene samples are dominated by ɛ-Proteobacteriaceae, and late-Holocene samples are dominated by sequences from uncultured subphyla. We only recovered δ-Proteobacteriaceae in late-Holocene sediments and α- and γ- Proteobacteriaceae in late- and middle-Holocene sediments. Diversity estimates derived from early-, middle-, and late-Holocene clone libraries indicate that the youngest (late-Holocene) samples had significantly greater bacterial diversity than the oldest (early-Holocene) samples, and the middle-Holocene samples contained intermediate levels of diversity. The observed patterns of diversity may be caused by increased bacterial niche-partitioning in younger sediments that contain a greater abundance of labile organic matter than older sediments. D. M. Nelson and S. Ohene-Adjei contributed equally to this work     

Molecular diversity of alveolates associated with neritic north atlantic radiolarians

Seven species of polycystine radiolarians and one phaeodarian species were investigated in order to determine the diversity of their associate organisms and their species specificity. Twelve partial 18S ribosomal DNA (rDNA) sequences were obtained showing a high diversity of associates, both within spumellarian and nassellarian radiolarians and among species. Two of the sequences obtained are highly similar to Scrippsiella, a dinoflagellate genus already reported as a symbiont of polycystine radiolarians. Nine of the new 18S rDNA sequences group with various alveolates. Some of these groups include parasites, such as the lethal endoparasite Amoebophrya, while others consist of non-annotated novel organisms found worldwide in various types of marine environments. We also obtained a sequence from a bacillariophytan highly similar to the 18S rDNA of the diatom species Diatoma tenue, which may derive from radiolarian food. Additionally, this is the first study to report on a phaeodarian associate.     

The biology of the marine sponge Microciona prolifera (Ellis and Solander). III. Spicule secretion and the effect of temperature on spicule size

The secretion of siliceous spicules in the marine demosponge Microciona prolifera (Ellis and Solander) is by three different means. Styles are secreted by sclerocytes with archeocyte characteristics (nucleolate nucleus, phagosomes). chelas are formed by small sclerocytes with anucleolate nuclei, and toxas are apparently formed extracellularly within membranous material. Genetically and physiologically equivalent explants of this sponge were grown at 15, 20, and 25 C for four weeks. Analyses of spicule dimensions show little correlation of temperature with spicule length, except in the case of toxas. but a clear inverse relationship of spicule width with temperature. It is suggested that thicker spicules are formed at lower temperatures due to the more efficient entrapment of silicon rather than to effects upon silicon transport. Chela dimensions are very uniform implying an all or none process in their secretion. Differences in spicule dimensions between individual sponges grown at these temperatures may be due to the highly complex pathways of silicon transport and/or to genetic differences.