Middle–Late Ordovician chitinozoans from Songliang of Qiaojia,western South China,and their biostratigraphic implications

Ordovician chitinozoans are as yet inadequately documented from the western Yangtze Platform, South China. Here we present a systematic study on chitinozoans from a Middle–Upper Ordovician succession at Songliang of Qiaojia, northestern Yunnan, southwestern China. Altogether 34 species of 12 genera are identified from the upper Hungshihyen and the Huadan formations. The top of the Hungshihyen Formation yields typical Early and Middle Ordovician forms, for instance, Lagenochitina obeligis and Belonechitina chenjiawuensis, with the latter only known from the lower Darriwilian in South China, thus suggesting an early Darriwilian age for this interval. The chitinozoans recovered from the Huadan Formation include Lagenochitina prussica and Spinachitina fossensis, both are mainly confined in Katian and sometimes ranging into younger strata. This indicates an age younger than the middle to late Darriwilian previously determined for the Huadan Formation. However, conflicts exist in age assignment of the Huadan Formation based on chitinozoan and other evidence, and further work is required.     

Conodont biostratigraphy and carbon isotope chemostratigraphy of the Middle to Upper Ordovician on the western Yangtze Platform,South China

Middle and Late Ordovician strata in the nearshore area of the western Yangtze Platform are investigated. In total, 241 δ¹³C_carb samples and 108 conodont samples from the Songliang and Tuanjie sections in the northeastern Yunnan Province were collected for chemostratigraphic and biostratigraphic studies. Altogether, 21 species belonging to 16 conodont genera were recovered from the two sections, among which Ansella jemtlandica, Drepanoistodus basiovalis, Hamarodus brevirameus, and Microzarkodina hagetiana are important age indicators within the Dapingian to early Katian interval. Four carbon isotope shifts are recognized in the studied sections: a negative shift in the early Darriwilian, the middle Darriwilian carbon isotope excursion (MDICE), the early Katian Guttenberg carbon isotope excursion (GICE), and a further positive carbon isotope shift later in the early Katian which is a new record of this event in South China. Integrated carbon chemostratigraphy and conodont biostratigraphy indicate early Darriwilian and early Katian ages for the base and top, respectively, of the Huadan Formation at the Songliang section, and a Darriwilian to early Katian age for the Shihtzupu Formation at the Tuanjie section.     

The biological affinity of Amsassia: new evidence from the Ordovician of North China

Amsassia shaanxiensis sp. nov. occurs in the Middle Ordovician part of the Jinghe Formation in Yongshou and the lower part of the Upper Ordovician Beiguoshan Formation in Longxian, Shaanxi Province, north‐central China. In addition to module increase by bipartite longitudinal fission, which is also known in other species of Amsassia, tripartite and rare quadripartite fission are recognized in A. shaanxiensis. All species previously assigned to Lichenaria from the Middle to Upper Ordovician of Shaanxi probably belong to Amsassia. Therefore, Amsassia, rather than the tabulate coral Lichenaria, should be credited as an important contributor to reef‐building in this area. Reports of Lichenaria from elsewhere in the North China Platform require confirmation in the light of the present study. Some morphological characteristics of Amsassia are comparable to those of tabulate corals, tetradiids and chaetetid sponges. Consequently, various authors have assigned Amsassia to the Lichenariida, Tetradiida (now Prismostylales; florideophycean rhodophyte algae) and Chaetetida. Other important characters, however, seem to exclude Amsassia from those taxonomic groups. The phacelocerioid organization of modules having separate walls would not be expected in sponges. The basic symmetry of individuals may have been radial, unlike the tetramerous symmetry of tetradiids. Module increase by longitudinal fission, involving infoldings of the wall, is fundamentally different from modes of increase in corals, tetradiids and chaetetids. The skeleton was probably aragonitic, whereas that of tabulates was calcitic. The affinity of Amsassia remains unresolved, but it is unlikely to have been a coral, tetradiid or sponge. Perhaps, like the tetradiids, Amsassia was an alga.     

Late Darriwilian–early Katian (Ordovician) ostracods of northeastern Yunnan,Southwest China

Ordovician silicified ostracods of the western Yangtze Platform, South China, are documented for the first time from the Huadan Formation of Qiaojia County, northeastern Yunnan Province, Southwest China. Twenty-one species of 17 genera are identified with new taxa including Vendona spinifera n. sp. and Nikitinella songliangensis n. sp. These two species, together with Medianella gigantea are the most common constituents of the fauna. Faunal comparison shows close connections between South China, Baltica, the Argentine Precordillera, Kazakhstan, Tarim, and Sibumasu during this time interval. This fauna is considered to be related to a nearshore carbonate environment, and biostratigraphical analysis of it supports a late Darriwilian to early Katian age for the Huadan Formation.     

Spicules and microspheres in the heliolitid tabulates from the Silurian strata of Gorny Altai,Siberia

Calcareous spicules closely similar to those previously reported in several species of Heliolites, have been found in another heliolitid species, Squameolites junggarensis Lin et Wang, from a clayey limestone of the lower part of Kuimov Formation, of Homerian (Wenlock) age, in the western part of the Gorny Altai, Siberia. The sclerites are associated with ‘microspheres’, identified also in colonies of an unidentified species of Propora. It is suggested that the microspheres associated with the tabulate corals were their photoautotrophic symbionts.     

Late Moscovian (mid-Pennsylvanian) rugose corals from Wadi Araba (Egypt,Eastern Desert): Taxonomy,palaeoecology and palaeobiogeography

An important late Moscovian rugose coral association is described from the Rod El Hamal Formation of the Wadi Araba area (northern Eastern Desert, Egypt). The upper part of the formation yielded a moderately abundant but poorly diversified coral fauna composed of large dissepimented solitary rugose corals. In total, six species belong to the Bothrophyllidae and the Geyerophyllidae, including three new species: Bothrophyllum suezensis, Bothrophyllum cylindricum, and Amygdalophylloides omarai. Three undeterminable Bothrophyllidae taxa are left in open nomenclature. The corals were attached to the soft substrate by talons and rootlets, either rooted and growing upward as mud-stickers or as secondary recliners. They show frequent rejuvenations and constrictions. Encrustation and/or bioerosion are scarce. Carbonate microfacies indicate a habitat in littoral and inner neritic zones adjacent to a low terrigenous hinterland. Colonial taxa are missing and tabulate corals are rare. The corals from the Rod El Hamal Formation are the only example of a late Moscovian coral fauna on the southern margin of the Palaeotethys, that is to say along the northern margin of Gondwana. The association shows similarities with northwestern Spain (Cantabrian Mountains), the Donets and Moscow basins, and thus a general attribution to the Palaeotethys realm.     

Halysitid tabulates: sponges in corals' clothing

Abundant pyritic pseudomorphs of monaxonic siliceous spicules (ophirhabds and ?heloclones) have been found entrapped in the calcareous skeleton of the halysitid tabulate Quepora ?agglomeratiformis (Whitfield) from late Ordovician limestones of Frobisher Bay, Baffin Island, Canada. The finding indicates a poriferan (choristid or sublithistid) affinity of halysitids, early Palaeozoic marine fossils related so far to corals. They probably derived from a monaxonic group of early demosponges that adapted during the Ordovician to Ca²⁺ stress conditions in epicontinental seas by excreting the excessive Ca²⁺ influx to their tissues as variously designed chains of basally secreted calcareous tubes.     

A New Endosymbiont in Late Ordovician Tabulate Corals from Anticosti Island,Eastern Canada

Cylindrical tubes of the trace fossil Chaetosalpinx occur within the skeletal walls of Late Ordovician sarcinulid tabulate corals preserved on Anticosti Island. A large host-specific embedment structure, Chaetosalpinx rex isp. nov., is described from the reefal sarcinulid genus Columnopora that occurs in (Rawtheyan, Ashgill) coral-stromatoporoid patch reefs of the upper Vauréal Formation (Mill Bay Member). While locally abundant in Columnopora, this newly described embedment structure is absent in other frame-builders in the reefs.     

Discovery of Anticostia uniformis from the Xiazhen Formation at Zhuzhai,South China and its stratigraphic implication

The Xiazhen Formation is an Upper Ordovician lithostratigraphic unit in the Jiangshan-Changshan-Yushan (JCY) area, which contains series of Late Ordovician reef successions. The reef successions of the Xiazhen Formation at Zhuzhai are critical for evaluation of the Late Ordovician marine diversity and palaeoecology. However, their age has long been uncertain and generally is regarded as of upper Katian, based on the occurrences of shelly fossils and correlation with the stratigraphic equivalent Changwu Formation. The newly discovered graptolite species Anticostia uniformis, in the Xiazhen Formation, together with the combined evidence of brachiopods and sedimentology, indicates an age range for the graptolite locality from the Dicellograptus complanatus Biozone to the Diceratograptus mirus Subzone of late Katian, but the graptolites do not rule out the possibility that it is Hirnantian.     

从底栖生物化石再释贵州独山泥盆系龙洞水段的时代

贵州独山的猴儿山组龙洞水段是一套浅海相碳酸盐沉积,盛产底栖牛物化石.通过对该段地层中所含四射珊瑚、床板珊瑚和腕足类等化石属种的详尽分析,再次证实龙洞水段的地质时代仍应属中泥盆世早期(Eifelian).而不是像某些根据痕迹化石或层序地层学的研究那样将它置于下泥盆统.     

Plicathyridine brachiopods (Athyridida) from the Frasnian (Late Devonian) of Western Europe and Middle East

Plicathyridine brachiopods (Athyridida) from the early–middle Frasnian of southern Belgium and northern France (Dinant Synclinorium) are systematically described for the first time. They include two species: Anathyris (Anathyris) calestiennensis nov. sp., and A. (A.) sp. indet. 1. They are uncommon in the mainly shally La Prée (Nismes Formation) and Ermitage (Moulin Liénaux Formation) members and are absent from the mixed argillaceous–carbonate late Frasnian succession in this area contrary to what is observed in Russia, notably in the East-European Platform and Siberia. In the Middle East, two species of Anathyris are recognized on the basis of a limited material from the Dascht-e-Nawar area in Afghanistan (A. (A.) sp. indet. 2) and the Kuh-e Kaftar mountains in Central Iran (A. (A.) sp. indet. 3). Anathyris (A.) calestiennensis nov. sp. is sometimes encrusted by epizoans (tabulate and rugose corals, and bryozoans) and rarely displays single, small circular drill holes. The past and current epizoan–brachiopod interactions are also discussed (Anathyris vs. Lingula, respectively).     

Middle–Late Ordovician brachiopods from Ningnan,southern Sichuan Province,Southwest China: Implications for macroevolution and palaeogeography

Middle to Late Ordovician brachiopods from the Huadan Formation (upper Darriwilian–Sandbian) of Ningnan County, southern Sichuan Province, are systematically documented here for the first time. The locality belongs to the western margin of the Upper Yangtze Platform, South China palaeoplate, and the brachiopod fauna includes one new genus and three new species as well as five other constituents: Hingganoleptaena sp., Acculina zhongliangziensis n. sp., Ningnanmena longisepta n. gen. n. sp., Kassinella (Trimurellina) minuta n. sp., Lepidorthis typicalis Wang, 1955, Protoskenidioides weixinensis Zhan and Jin, 2005, Porambonites transversus Xu, Rong and Liu, 1974, and Psilocamerella sp. Taxonomically it is a typical representative of a Middle to early Late Ordovician brachiopod fauna, and, together with some other evidence from other fossil groups like trilobites, conodonts, chitinozoans, a late Darriwilian–Sandbian age could be inferred for the horizon yielding this fauna. According to the richness of each constituent, this fauna is suggested to be called the Acculina-Ningnanmena fauna (ANF). Numerical palaeogeographical analysis shows that two broad palaeobiogeographic provinces could be recognized during this particular time interval, and, although the ANF is grouped into the South China cluster, it shares very little similarity with other representatives of that group except for two cosmopolitans. It further confirms that the Great Ordovician Biodiversification Event (GOBE), in other words the Ordovician radiation, was actually manifested by the strong localization of major marine organisms such as brachiopods, trilobites, graptolites, etc.     

Evidence of photosymbiosis in Palaeozoic tabulate corals

Coral reefs form the most diverse of all marine ecosystems on the Earth. Corals are among their main components and owe their bioconstructing abilities to a symbiosis with algae (Symbiodinium). The coral–algae symbiosis had been traced back to the Triassic (ca 240 Ma). Modern reef-building corals (Scleractinia) appeared after the Permian–Triassic crisis; in the Palaeozoic, some of the main reef constructors were extinct tabulate corals. The calcium carbonate secreted by extant photosymbiotic corals bears characteristic isotope (C and O) signatures. The analysis of tabulate corals belonging to four orders (Favositida, Heliolitida, Syringoporida and Auloporida) from Silurian to Permian strata of Europe and Africa shows these characteristic carbon and oxygen stable isotope signatures. The δ¹⁸O to δ¹³C ratios in recent photosymbiotic scleractinians are very similar to those of Palaeozoic tabulates, thus providing strong evidence of such symbioses as early as the Middle Silurian (ca 430 Ma). Corals in Palaeozoic reefs used the same cellular mechanisms for carbonate secretion as recent reefs, and thus contributed to reef formation.     

Origin and early diversification of the phylum Cnidaria Verrill: major developments in the analysis of the taxon's Proterozoic–Cambrian history

Diploblastic eumetazoans of the phylum Cnidaria originated during the Neoproterozoic Era, possibly during the Cryogenian Period. The oldest known fossil cnidarians occur in strata of Ediacaran age and consist of polypoid forms that were either nonbiomineralizing or weakly so. The oldest possible anthozoans, including the genus Ramitubus, may be related to tabulate corals and occur in the Doushantuo Lagerstätte (upper Doushantuo Formation, South China), the age of which is poorly constrained (approximately 585 Ma?). Conulariid scyphozoans may first appear as early as 635–577 Ma (Lantian Formation, South China). A definite conulariid, most similar to Palaeozoic species assigned to the genus Paraconularia, occurs in association with the possible scyphozoan, Corumbella werneri, in the latest Ediacaran (c. 543 Ma) Tamengo Formation of Brazil. Basal Cambrian (c. 540 Ma) phosphorites in the upper Kuanchuanpu Formation (South China) yield solitary polyps of the oldest probable anthozoan (Eolympia pediculata), which appears to have been a stem hexacorallian. This same formation contains fossils interpreted by some authors as pentaradial cubozoan polyps; however, both the oldest known cubozoans and the oldest hydrozoans, all medusae, may actually occur in the Cambrian (Series 3, c. 505 Ma) Marjum Formation (Utah, USA). Although these recently published palaeontological data tend to corroborate the hypothesis that Cnidaria has a relatively deep Neoproterozoic history, the timing of major internal branching events remains poorly constrained, with, for example, the results of some molecular clock analyses indicating that the two cnidarian subphyla (Anthozoaria and Medusozoa) may have originated as many as one billion years ago. Further progress towards elucidating the evolution and early fossil record of cnidarians may accrue from: (1) an intensive search for phosphatized soft parts in possible anthozoans from the Ediacaran Doushantuo Formation; (2) an expanded search for Ediacaran conulariids; and (3) additional detailed analyses of the taphonomy and preservation of Ediacaran and Cambrian cnidarians, including possible pentaradial cubozoan polyps from the Fortunian upper Kuanchuanpu Formation.     

Tabulate Corals after the Frasnian/Famennian Crisis: A Unique Fauna from the Holy Cross Mountains,Poland

Famennian tabulate corals were very rare worldwide, and their biodiversity was relatively low. Here we report a unique tabulate fauna from the mid- and late Famennian of the western part of the Holy Cross Mountains (Kowala and Ostrówka), Poland. We describe eight species (four of them new, namely ?Michelinia vinni sp. nov., Thamnoptychia mistiaeni sp. nov., Syringopora kowalensis sp. nov. and Syringopora hilarowiczi sp. nov.); the whole fauna consists of ten species (two others described in previous papers). These corals form two assemblages—the lower, mid-Famennian with Thamnoptychia and the upper, late Famennian with representatives of genera ?Michelinia, Favosites, Syringopora and ?Yavorskia. The Famennian tabulates from Kowala represent the richest Famennian assemblage appearing after the F/F crisis (these faunas appear some 10 Ma after the extinction event). Corals described here most probably inhabited deeper water settings, near the limit between euphotic and disphotic zones or slightly above. At generic level, these faunas show similarities to other Devonian and Carboniferous faunas, which might suggest their ancestry to at least several Carboniferous lineages. Tabulate faunas described here represent new recruits (the basin of the Holy Cross mountains was not a refuge during the F/F crisis) and have no direct evolutionary linkage to Frasnian faunas from Kowala. The colonization of the seafloor took place in two separate steps: first was monospecific assemblage of Thamnoptychia, and later came the diversified Favosites-Syringopora-Michelinia fauna.     

Stratigraphical distribution of the Ordovician graptolite Azygograptus Nicholson & Lapworth in the Central Andean Basin (northwestern Argentina and southern Bolivia)

We analyzed new occurrences of Azygograptus lapworthi from the Cordillera Oriental, Argentina. The bearer sandstones levels, corresponding to the Acoite Formation, are overlying the deposits, in which the Didymograptellus bifidus Biozone (Lower Ordovician, upper Floian, Fl3) was previously recognized, and are overlain by younger pelitic levels yielding Xiphograptus lofuensis (Middle Ordovician, early Dapingian, Dp2). Previous records from the Central Andean Basin are also reviewed in detail and accurately correlated, allowing us to conclude that the Azygograptus lapworthi Biozone corresponds to the Middle Ordovician (lower Dapingian, Dp1). This biostratigraphic framework documents that the transition between the Lower and Middle Ordovician deposits occurs in the uppermost levels of the Acoite Formation in the Argentine Cordillera Oriental. It is additionally integrated with up to date conodont records to establish a high-resolution regional correlation, with equivalent deposits from the Puna of northwestern Argentina and Cordillera Oriental of Bolivia, and to discuss new insights for global correlation.     

Cyclocrinitids from the Lower Palaeozoic Tethyan sequence of Spiti,India

Well preserved cyclocrinitids (calcareous green algae) are reported from early Palaeozoic deposits of Spiti, Himachal Pradesh, India. Casts of the algal fossils are preserved in calcareous grey siltstone unit of the Takche Formation (Late Ordovician–Early Silurian) which appears rusty/earthy brown due to weathering. The assemblage includes Cyclocrinites favus, Cyclocrinites pyriformis, Cyclocrinites cf. welleri, Cyclocrinites sp. and Cyclocrinites globosus. Both Cyclocrinites cf. welleri and Cyclocrinites globosus are reported for the first time from the entire Tethyan Himalaya, India. The described algal flora basically denotes relatively shallow marine depositional setting and low to moderate hydrodynamic conditions.     

Stratigraphic distribution of selected acritarchs in the Ordovician subsurface,Canning Basin,Western Australia

Palynological investigation of 80 core samples from four boreholes, drilled through the Lower to Middle Ordovician marine sections of the central-northeastern Canning Basin, northwestern Australia, reveals diverse acritarch associations including a number of stratigraphically significant species. The stratigraphic succession studied, embracing the Willara Formation and conformably overlying Goldwyer Formation, has been independently dated as early Arenig to Llanvirn by conodont faunas (Oepikodus communis through Phragmodus-Plectodina zonal interval). Nine morphologically distinctive acritarch species have relatively narrow vertical ranges within the study interval and likely constitute serviceable palynostratigraphic indices for the Ordovician in and possibly beyond the Western Australian study area. Three new species, Comasphaeridium setaricum, Polyancistrodorus kunzeanensis, and Stelomorpha calix, are established; and one new combination, Baltisphaeridium variocavatum (Playford and Martin, 1984), is proposed. Other stratigraphically significant species include Petaloferidium comptum Playford and Martin, 1984 and Peteinosphaeridium sp. cf. P. exornatum Tongiorgi et al., 1995; together with Dorsennidium symmetricum (Lu, 1987) Sarjeant and Stancliffe, 1994, Peteinosphaeridium coronula Yin et al., 1998, and Striatotheca rarirrugulata (Cramer et al., 1974) Eisenack et al., 1976, which are found for the first time in Australia. The vertical distribution, and hence the stratigraphic utility, of each of the above species is documented through the studied sections.     

New observations on the skeletons of the earliest bryozoans from the Fenhsiang Formation (Tremadocian,Lower Ordovician), Yichang,China

The earliest unequivocal bryozoans occurred in the Fenhsiang Formation (upper Tremadocian, Lower Ordovician) of Yichang, South China, providing important clues about the early evolution of primitive bryozoans. In this study, SEM and EDAX were used to analyze the microstructure of the walls of two bryozoan genera, Nekhorosheviella and Orbiramus, from the Fenhsiang Formation. All walls of Nekhorosheviella and endozonal walls of Orbiramus show poor preservation in thin sections, with a granular appearance reflecting extensive neomorphism, whereas exozonal walls of Orbiramus are distinctly laminated. These preservational differences may reflect skeletal chemistry, particularly the magnesium content of the calcite. Phosphatic skeletal linings were found to be distributed unevenly in the autozooecia of Nekhorosheviella, but were absent in Orbiramus.     

Darriwilian–Sandbian (Middle–Late Ordovician) shallow warm-water deposits in the Upper Yangtze region,South China: A case study from the Qiaojia area,northeastern Yunnan Province

The Middle–Late Ordovician Nuoduo and Huadan formations are well exposed at Songliang of northeastern Yunnan Province, representing rare neritic carbonate sediments deposited in the Yangtze region, South China during that time. Detailed sedimentological analyses support the recognition of ten microfacies from the succession, including shale (MF 1), argillaceous mudstone/dolostone (MF 2), mollusk-ostracod wacke-packstone (MF 3), peloidal wacke- to pack-grainstone/dolostone (MF 4), ooid-cortoid-peloid pack-grainstone/dolostone (MF 5), peloidal bindstone (MF 6), bioclastic wacke-packstone (MF 7), green algae mud-wackestone to floatstone (MF 8), bioclast-lithoclast float-rudstone (MF 9) and green algae bindstone (MF 10). Types A, B and C of shallowing-upward sequences are constructed according to vertical associations of MFs. The Nuoduo Formation mainly comprises Type A and Type B, both being characterized by abundant peloids, ooids, cortoids and aggregate grains. MF 5 and MF 6 cap the Type A and Type B sequences, respectively, suggesting an overall restricted marine environment shifting cyclically from lagoon to shoal, or lagoon to tidal flat, respectively. Type C sequences, occurring in the topmost Nuoduo to Huadan formations, contain various green algae indicative of open-sea, euphotic subtidal zones. The transition from A and B to C sequences is marked by ferruginous ooidal intervals, possibly correlated to relative sea-level rise and/or tectonic activities. A continuous neritic, warm-water environment is proposed for this area from late Middle to Late Ordovician, with the Nuoduo Formation filling the absence of warm-water deposits in the tropical or subtropical Yangtze region during Middle Ordovician. The increasing diversity and abundance of green algae in the Huadan Formation might represent a global radiation event of marine flora during the studied interval. Unique fossils and environments distinct from those of the interior Yangtze region are documented here, leading to a better understanding of the Ordovician System regionally and globally.