Biotic crises and stages of radiolarian evolution in the Phanerozoic

The questions of ecology and taphonomy of radiolarians are reviewed and reexamined. In Recent oceans and seas, the major part of the “radiolarian rain” (dead individuals) comes from the layer of the water column which is not deeper than 500 m; therefore, the bottom radiolarian thanatocenoses and taphocenoses are formed in all oceanic zones, including the coastal and central oligotrophic regions of oceans. However, radiolarians should not be regarded as indicators of exclusively deepwater oceanic conditions. The crucial moments in the evolution of radiolarians at the major Phanerozoic boundaries are recognized. A dynamic model of cyclic development of radiolarians in the Phanerozoic is proposed and four phases and nine stages in their evolution are recognized. The absence of an outburst of radiolarian biodiversity in the Holocene is shown. Many great extinctions of radiolarians occurred at the boundaries between seasons of galactic years.     

Lower Darriwilian radiolarians from the Argentine Precordillera

The first radiolarians discovered in the Ordovician of the Argentinian Precordillera are reported. The material is associated with conodonts indicating the upper part of the Lenodus variabilis Biozone (Lower Darriwilian, Da1-2). The low diversity radiolarian fauna shows similarities with Middle Ordovician radiolarians from the Akzhal Mountains of Kazakhstan. However, a close comparison is difficult due to the poor preservation of the material, in which internal structures of the radiolarian tests are not preserved.     

Radiolarian flux in Antarctic waters (Drake Passage,Powell Basin,Bransfield Strait)

Summary The study of radiolarians collected during sediment trap experiments in the Drake Passage, the northern Powell Basin, and the King George Basin of the Bransfield Strait provides new information on the fluxes of radiolarian shells in Antarctic waters, on the annual flux pattern, the species distribution and its ecological significance, and on alteration processes of the radiolarian shells in the water column and at the sediment/water interface. A 28-month monitoring with time-series sediment traps in the Bransfield Strait indicates an annual flux pattern characterized by short-term flux pulses during austral summer, which reach daily fluxes of up to 5 × 10³ shells m^–2 and which account for more than 90% of the total annual flux. The distinct seasonal variations are linked to variations in the sea ice coverage. Other controlling factors are the production of phytoplankton and the impact by zooplankton grazers, e.g., krill. During the summer flux pulses the vertical fluxes of radiolarians range between ca. 3 and 21 × 10⁴ shells m^–2, values that are one or more orders of magnitudes lower than fluxes observed at sites in the tropical and northern high-latitude ocean. Significant lateral transport of radiolarians was documented during the austral summer in the Bransfield Strait by a factor of 10 increase of the radiolarian flux in the lower portion of the water column and the species composition trapped in deeper waters. Radiolarian assemblages associated with pelagic and neritic environments characterized by typical Antarctic taxa (Antarctissa spp.) and a group of species with bipolar distribution (e.g. Plectacantha oikiskos, Phormacantha hystrix), respectively, are distinguished. While the signal of polycystine radiolarians is relatively well recorded in the sediments, the shells of phaeodarians, which were observed at fluxes of up to 1 × 10³ shells m^–2day^–1 in the upper portion of the water column, are almost completely dissolved during settling through the water column.     

Mesozoic Radiolaria of Bosnia and Serbia: New data

Jurassic and Cretaceous radiolarian faunas were discovered in bedded chert of the Dinaric and Vardar tectonic zones of Bosnia and Herzegovina. Only Triassic radiolarians have previously been described in Bosnia and Herzegovina, while the finds of Bajocian, Bathonian-Callovian, Oxfordian-Kimmeridgian, Tithonian-Berriasian, and Campanian are new. Additional localities of Triassic and Jurassic radiolarians were investigated in Serbia. By correlation with radiolarians from the previous studies within Serbia, the Late Aalenian-Bajocian and Bathonian-Callovian Serbian radiolarian beds are newly dated. The first find of Cretaceous radiolarians in Serbia is reported. The oldest Mesozoic Radiolaria-bearing formations outcropping in the Western Belt of the Vardar Zone are dated Mid-Upper Triassic. The youngest radiolarians come from the Upper Cretaceous (Campanian) of the northern part of this belt of the Vardar Zone, where they co-occur with planktonic foraminifers. The distribution of 70 radiolarian samples within sections is shown. The taxonomic composition of 39 samples is analyzed. Radiolarian species extracted from 13 samples are described and figured. The list of 72 taxa and 3 plates of Jurassic radiolarians of Bosnia and Herzegovina, 3 plates of Triassic, 1 plate of Middle Jurassic, and 1 plate of Upper Cretaceous radiolarians of Serbia are presented.     

Radiolaria associated with large diversity of marine alveolates

We have isolated cells of unculturable radiolarians from marine coastal waters. Individual cells were subjected to single cell whole genome amplification (SCWGA) and gene-targeted PCR. Using this approach we recover a surprisingly large diversity of sequences related to the enigmatic marine alveolate groups 1 and 2 (MALV I and MALV II) that most likely represent intracellular symbionts or parasites of the radiolarian cells. 18S rDNA phylogeny of the MALV sequences reveals 4 distinct clades of radiolarian associates here named Radiolarian Associated Sequences (RAS) 1-4. One clade of both phaeodarian and radiolarian associates and one clade of only phaeodarian associates are also identified. The MALV sequences cluster according to host type, i.e. sequences from associates identified in radiolarians, fish, copepods, ciliates or dinoflagellates are not intermixed but separated into distinct clades. This implies several independent colonizations of host lineages and links a large diversity of MALV to radiolarian-associated species. This demonstrates that radiolarians may be an important reservoir for MALV, making them a key group for understanding the impact of intracellular symbionts on the marine ecosystem. This study shows that applying SCWGA on unculturable cells is a promising approach to study the vast diversity and interactions of intracellular eukaryote organisms.     

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。     

Deviations in skeletons of radiolarians

Deviations in the skeleton structure of radiolarians are extremely rare in the fossil record. They are known from the Devonian to Recent. This is a very ancient and periodically repeated phenomenon, which should be regarded as a regularity or “lost chance” rather than exotic or accident. Emergence of deviant “mutants” in radiolarians is probably connected with two main causes, i.e., disturbance during reproduction and mutations in particular modules. From the point of view of the morphogenesis of skeletal structures, deviancy facilitates the recognition of relatively “weak” modules–blocks in the general skeleton pattern, which are subject to structural changes. Nonheritable deviant variants have been revealed in skeletons of 61 radiolarian in all of three subphyla of the phylum Radiolaria: Polycystina, Phaeodaria, and Collodaria. Three deviant types are recognized: multiplicative, posterioric and supplemental.     

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.     

Radiolarian-based sea surface temperatures and paleoceanographic changes during the Late Pleistocene–Holocene in the subantarctic southwest Pacific

A high-resolution record of radiolarian faunal changes from Site Y8 south of the Subtropical Front (STF), offshore eastern New Zealand, provides insight into the paleoceanographic history of the last 265 kyrs. Quantitative analysis of radiolarian paleotemperature indicators and radiolarian-based sea surface temperature (SST) estimates reveal distinct shifts during glacial–interglacial (G-I) climate cycles encompassing marine isotope stages (MIS) 8–1. Faunas at Site Y8 are abundant and diverse and consist of a mixture of species typical of the subantarctic, transitional and subtropical zones which is characteristic of subantarctic waters just south of the STF. During interglacials, diverse radiolarian faunas have increased numbers of warm-water taxa (~ 15%) while cool-water taxa decrease to ~ 11% of the assemblage. Warmest climate conditions occurred during MIS 5.5 and the early Holocene Climatic Optimum (HCO) at the onset of MIS 1 where SSTs reach maxima of 12.8 and 12.9 °C, respectively. This suggests that temperatures during the HCO were comparable to the Eemian, one of the warmest interglacial intervals of the Late Quaternary. Glacials are characterized by less diverse radiolarian faunas with cool-water taxa increasing to 49% of the assemblage. Coolest climate conditions occurred in MIS 4 and 2 where SSTs are reduced to 5.4 °C and 4.3 °C, respectively. Radiolarian faunal changes and SST estimates clearly identify major water masses and oceanic fronts in the offshore eastern New Zealand area. During warmest MIS 5.5 and early MIS 1 substantial influence of northern-sourced Subtropical Surface Water (STW) is evident at Site Y8. This implies southward incursions of STW around the eastern crest of Chatham Rise with the STF displaced towards higher latitudes and spinning off eddies as far south as Campbell Plateau. Additionally, increased flow of the Southland Current (SC) might have enhanced the local occurrence of warm-water radiolarians derived from the subtropical Tasman Sea. Coolest glacials are marked by a strong inflow of cool, southern-sourced waters at Site Y8 indicating a more vigorous flow along the Subantarctic Front (SAF).     

New radiolarians from the Devonian of the southern Ural Mountains: 1. Early-Middle Devonian (Late Emsian-Early Eifelian)

Late Emsian-Early Eifelian radiolarians from the southern Ural Mountains are recorded and described for the first time. They come from the Kamennaya Gora section on the Tanalyk River near the town of Buribay, Bashkiria. The new Late Emsian-Early Eifelian radiolarian assemblage consists of 20 species of 11 genera of spherical, discoidal, spindle-shaped, pylomate, and spiny radiolarians. Four new species, i.e., Entactinia rostriformis sp. nov., Palaeodiscaleksus tumefactus sp. nov., Primaritripus buribayensis sp. nov., and P. chuvashovi sp. nov., are described. The diagnosis of the genus Primaritripus Afanasieva, 2000 is amended. The fact that the oryctocenosis is dominated by discoidal radiolarians suggests that, in the Late Emsian-Early Eifelian, the area under study was occupied by a relatively shallow marine basin. Based on radiolarians, a new Upper Emsian-Lower Eifelian biostraton, beds with Primaritripus buribayensis-Entactinia rostriformis, is established. The history of the study of Early Devonian radiolarians is considered.     

A 350-ky radiolarian record off Lüderitz,Namibia—evidence for changes in the upwelling regime

The study of radiolarian assemblages from Core MD 962086 provides new information on the variability in the upwelling intensity and origin of upwelled water masses over the past 350 ky in one of the major filamentous regions of the Benguela Upwelling System (BUS), located off Lüderitz, Namibia. The use of key radiolarian species to trace the source of upwelled waters, and the use of a radiolarian-based upwelling index (URI) to reconstruct the upwelling intensity represent the first use of radiolarians for paleoceanographic reconstructions in the BUS. These radiolarian-based proxies indicate strongest upwelling during Marine Isotope Stages (MIS) 3, 5, and 8, which compares well with other studies. While during MIS 3 and 8, the radiolarian-based proxies indicate the influx of waters of Southern Ocean origin, they also point to the increased influence of tropical waters during the lower portion of MIS 5. During MIS 2, 4 and 6 the radiolarian assemblages indicate generally lower upwelling intensities, although this signal is complicated by the increased occurrence of organic carbon in the sediments during these intervals. During MIS 2 there appears to be less of an input of Southern Ocean waters to the BUS, although during the also glacial MIS 4 and 6, there is evidence for an increased influence of cold Antarctic waters. The comparison of the results from Core MD 962086 with other studies in the BUS area indicates a non-uniform pattern of upwelling intensity and advection of cold, southern waters into this system during MIS 2. Weaker upwelling signaled by the radiolarian-based proxy in MIS 4 is in contrast to other studies that indicate higher productivity during this time period. In general, the data show that there is a strong spatiotemporal complexity in upwelling intensity in the BUS and that the advection of water into it is not strongly tied to glacial–interglacial variations in climate.     

江苏仑山地区上奥陶统五峰组放射虫动物群及其地质意义

五峰组以盛产笔石动物群著称,时代为凯迪晚期(相当于英国的阿什极尔期),其中也发现了放射虫.该时期的放射虫在全球并不多见.本文首次描述江苏句容仑山地区五峰组的放射虫动物群,该动物群以不具刺的球形放射虫最为发育,并出现少量Inaniguttidae科的分子,化石保存较差,属种单调,分异度很低,这些特点与大西洋沿岸地区陆表海...     

Host-symbiont associations of polycystine Radiolaria: epifluorescence microscopic observation of living Radiolaria

In all 29 polycystine radiolarian species were obtained from surface seawater on May 28, 1999, using a plankton-net at one station (Site 990528; 26°37′18″N, 127°47′35″E) approximately 5 km northwest of Okinawa Island, Japan. In most polycystine radiolarians of the orders Nassellarida and Spumellarida symbiotic algae were observed under light microscopy. The light microscopic (LM) images of the symbionts, however, varied in clarity among individuals because of the variations in microanatomy of the host radiolarian cells. On the other hand, epifluorescence microscopic (EFM) observation easily detected and confirmed the existence of the algal symbionts within the host cytoplasm even in radiolarians such as Dictyocoryne truncatum (Ehrenberg) that include algal symbionts in the depth of the cytoplasm. The chloroplasts of the algal symbionts emitted autofluorescence in ultraviolet irradiation and they appeared red. That is, the autofluorescence images of the chloroplasts can be used to recognize the existence of the algal symbionts within the host radiolarians. Moreover, staining of the symbiont cells with 4′,6-diamido-2-phenylindle permitted visualization of the nucleus in the center of the symbiont cell, confirming the existence of living endosymbiotic algae within the polycystine radiolarians. Both the LM and EFM observations of eight polycystine radiolarian species revealed the specific patterns of various host-symbiont associations. (1) The investigated polycystine radiolarians all possess algal symbionts, except for one species, i.e. Dictyocoryne profunda Ehrenberg. (2) The size of the algal symbionts depends on the radiolarian species. The symbionts are largely classified into two types based on the size of their diameters, i.e. about 8–10 μm for the larger group and about 5 μm for the smaller one. (3) The algal symbionts show a variety of locations within the host radiolarian cytoplasm. The types of distribution of algal symbionts may be a useful characteristic for radiolarian taxonomy.     

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.     

Early Cambrian radiolarians and sponge spicules from the Niujiaohe Formation in South China

Most of the reported Cambrian radiolarians are from middle Cambrian (Miaolingian Series, Wuliuan Stage) and onward, the radiolarians from lower Cambrian (Terreneuvian and Cambrian Series 2), on the other hand, are poorly documented, thus their morphological characteristics have not been well understood. In this study, we extracted spherical radiolarians and sponge spicules from the chert of the Cambrian Series 2 Niujiaohe Formation in southern Jiangxi Province, China. The well-preserved radiolarian fossils, identified as Paraantygopora porosa, consist of perforate plate shells pierced by dense pores with elevated rims, and display similarities to those from Lower Ordovician. Seven forms of sponge spicules are recognized, including monaxons, diaxons, tetractines, hexactines, triaxon tetractines, triaxon pentactines and irregular tetraxon tetractines, which are similar to those from the Qiongzhusian Stage in South China. Our results, combined with previously reported early Cambrian radiolarians from South China, indicate that the early Cambrian radiolarians developed advanced spherical skeletons.     

Techniques for determining opal in deep-sea sediments: A comparison of radiolarian counts and x-ray diffraction data

Detailed records of two estimators of the biogenic silica content of deep-sea sediments, x-ray diffraction analysis opal concentrations and quantitative radiolarians per gram counts from microscope slides were compared in downcore sediments from northwest Pacific core RC14-105 to evaluate and to intercalibrate the two techniques. Both the degree to which the data were similar and the degree to which inferences from the data were similar were considered. Although both data sets are internally consistent and reflect surface water processes, they are not equivalent and there is no way to convert one linearly into the other. The numbers of radiolarians per gram of sediment appear to respond to the same factors which control the fluctuations in the radiolarian assemblage distrbutions in the area. The opal contents respond to the factors which control the total siliceous productivity of the area.     

Upper Devonian Radiolarians from the Zeravshan-Gissar Mountainous Area,Uzbekistan

The first finds of Upper Devonian radiolarians in cherty rocks of the Akbasay Formation, Zeravshan-Gissar mountainous area, are reported. Nineteen species of spherical and spiny forms were identified and assigned to 8 genera. Finds of co-occurring conodonts and radiolarians enabled the refinement of the proposed age of radiolarian associations.     

First evidence of Late Carnian radiolarians from the Izmir-Ankara suture complex, central Sakarya, Turkey: implications for the opening age of the Izmir-Ankara branch of Neo-Tethys

Within the Late Cretaceous mélange complex of the Izmir-Ankara suture zone in central Sakarya area, north-western Turkey, a megablock with radiolarian cherts associated with basaltic pillow lavas has been dated by radiolarians. The studied radiolarian assemblage and conodonts yielded an early Late Carnian age. This age is the oldest obtained from the chert blocks all along the suture belt and has important implications for the rifting/opening age of the Izmir-Ankara branch of the Tethys ocean in north-western Turkey. Based on this new data, it is concluded that during the Late Triassic the Izmir-Ankara seaway was connected to the other Tethyan oceanic branches and deep enough to provide chert sedimentation and exchange radiolarians with the main open oceans. It is further suggested that the long-lasting misinterpretation of a Liassic rifting/opening of the Izmir-Ankara ocean should be revised.     

Radiolarian faunal turnover across the Oligocene/Miocene boundary in the equatorial Pacific Ocean

The global warming trend of the latest Oligocene was interrupted by several cooling events associated with Antarctic glaciations. These cooling events affected surface water productivity and plankton assemblages. Well-preserved radiolarians were obtained from upper Oligocene to lower Miocene sediments at Ocean Drilling Program (ODP) Leg 199 Sites 1218 and 1219 in the equatorial Pacific, and 110 radiolarian species were identified.Four episodes of significant radiolarian faunal changes were identified: middle late Oligocene (27.5 to 27.3 Ma), latest Oligocene (24.4 Ma), earliest Miocene (23.3 Ma), and middle early Miocene (21.6 Ma). These four episodes approximately coincide with increases and decreases of biogenic silica accumulation rates and increases in δ¹⁸O values coded as “Oi” and “Mi” events. These data indicate that Antarctic glaciations were associated with change of siliceous sedimentation patterns and faunal changes in the equatorial Pacific.Radiolarian fauna was divided into three assemblages based on variations in radiolarian productivity, species richness and the composition of dominant species: a late Oligocene assemblage (27.6 to 24.4 Ma), a transitional assemblage (24.4 to 23.3 Ma) and an early Miocene assemblage (23.3 to 21.2 Ma). The late Oligocene assemblage is characterized by relatively high productivity, low species richness and four dominant species of Tholospyris anthophora, Stichocorys subligata, Lophocyrtis nomas and Lithelius spp. The transitional assemblage represents relatively low values of productivity and species richness, and consists of three dominant species of T. anthophora, S. subligata and L. nomas. The characteristics of the early Miocene assemblage are relatively low productivity, but high species richness. The two dominant species present in this assemblage are T. anthophora and Cyrtocapsella tetrapera. The most significant faunal turnover of radiolarians is marked at the boundary between the transitional/early Miocene assemblages.We also reviewed changes in other microfossil assemblages in the low latitudes during the late Oligocene through early Miocene. The microfossil assemblages of major groups show sequential changes near the Oligocene/Miocene (O/M) boundary (23.8 Ma). Many extinction events and some first occurrences of calcareous nannofossils and many occurrences of radiolarians are found from about 24.8 to 23.3 Ma, and first occurrences of planktic foraminifers and diatoms followed from 23.2 through 22 Ma. Hence, the O/M boundary is identified as a significant level for microfossil evolutions.