Graptolites from glacial erratics of the Laerheide area,northern Germany

Ordovician and Silurian glacial erratics of the Laerheide area (Lower Saxony, north-western Germany) bear well-preserved graptolites. The faunas provide important information on the origin and transport direction of the sediments preserved in a kame, representing the Drenthe stadial of the Saalian glaciation. The faunas even include species not commonly encountered in the successions of mainland Sweden, from where the erratics presumably originated. The most common graptolites are from Upper Ordovician (Sandbian to Katian) limestones and from Katian black shales. More common, however, are greenish limestones, sand- and siltstones, often combined in the term ‘Grünlich-Graues Graptolithengestein’, in which upper Wenlock to Ludlow (upper Silurian) graptolites are common.     

The position of graptolites within Lower Palaeozoic planktic ecosystems

An integrated approach has been used to assess the palaeoecology of graptolites, both as a discrete group and as a part of the biota present within Ordovician and Silurian planktic realms. Study of the functional morphology of graptolites and comparisons with Recent ecological analogues demonstrates that graptolites most probably filled a variety of niches as primary consumers, with modes of life related to the colony morphotype. Graptolite coloniality was extremely ordered, lacking any close morphological analogues in Recent faunas. To obtain maximum functional efficiency, graptolites would have needed varying degrees of coordinated automobility. A change in lifestyle related to ontogenetic changes was prevalent within many graptolite groups. Differing lifestyle was reflected by differing reproductive strategies, with synrhabdosomes most likely being a method for rapid asexual reproduction. Direct evidence in the form of graptolithophage 'coprolitic' bodies, as well as indirect evidence in the form of probable defensive adaptations, indicate that graptolites comprised a food item for a variety of predators. Graptolites were also hosts to a variety of parasitic organisms and provided an important nutrient source for scavenging organisms. □ Feeding, graptolites, hydrodynamics, ontogeny, parasitism, palaeoecology, plankton, predation.     

Facies distribution patterns of some marine benthic faunas in early paleozoic platform environments

Worldwide Late Cambrian—Silurian lithofacies patterns indicate that the platforms of that time were sites of accumulation of two essentially different rocks suites: the platform carbonate rocks and the platform terrigenous rocks. Most of the platform rocks accumulated as sediments in shallow marine environments similar to those of the present but far more widely spread.Present-day marine benthic faunas are distributed in depth zones which are primarily controlled by temperature. Faunas tend to occur in substrate-related discrete clusters (communities) within each life zone; similar substrates in different depth zones commonly have different faunal associations. Individual phyletic stocks may encounter environmental optimum or near-optimum conditions in certain areas, that commonly are revealed by an abundance of species and individuals within species in each stock. Environmental optimum conditions depend upon availability of food that may be utilized, modes of feeding of the animals present, water motion, and substrate, among other factors. Organisms in past seas were distributed in patterns similar to those of the present.Carbonate platforms were particularly widespread during the latest Cambrian—Early Ordovician. Intertidal environments spread widely across those platforms during that time and characteristic faunal associations developed in them. Saukiid and related tribolites dominated latest Cambrian carbonate platform intertidal faunas. The Early Ordovician carbonate platform intertidal was dominated by archeogastropod-nautiloid cephalopod faunas. These animals were joined by tabulate corals and certain brachiopods during the latter part of the Ordovician and Silurian as prominent faunal elements in the carbonate platform intertidal—shallow subtidal. Cruziana and related trace fossils, bivalves, and certain tribolites (notably homalonotids and dalmanitids) dominated most terrigenous platform intertidal—shallow subtidal faunas of the Ordovician and Silurian.Articulate brachiopods (primarily orthoids, strophomenoids, and rhynchonelloids) appear to have been relatively prominent during the Early Ordovician in shallow subtidal environments on both carbonate and terrigenous platforms and to have spread down the bathymetric gradient into increasingly deeper subtidal areas of both platforms during the latter part of the Ordovician. Tribolites dominated faunas in relatively moderate to deep subtidal environments on both platforms during the early part of the Ordovician. They were gradually replaced by brachiopods in first the shallower, and later the deeper subtidal as dominant members of the faunas. Brachiopods (primarily pentameroids and spiriferoids) dominated nearly all Silurian warm-water subtidal environments from the shallow subtidal to the edges of the platforms.Platform uplifts in the Middle Ordovician and glacio-eustatic sea-level fluctuations in the Late Ordovician caused environmental changes across the platforms that were accompanied by marked replacements among marine benthic faunas in all environments. The distribution of Ordovician carbonate platforms and glacial deposits suggests that an Ordovician polar region may have been close to present-day equatorial Africa and that Ordovician warm temperate-tropical regions lay close to the present-day North Pole.     

Sheinwoodian (Silurian) conodonts and graptolites from NE Anti‐Atlas,Morocco

Männik, P., Loydell, D.K. & Lubeseder, S. 2010: Sheinwoodian (Silurian) conodonts and graptolites from NE Anti‐Atlas, Morocco. Lethaia, Vol. 44, pp. 410–416. Conodonts and graptolites from a limestone at the base of the Tamaghrout Formation indicate that this is of early Wenlock age. This is the first unequivocal dating of a Sheinwoodian limestone from Morocco. All of the conodonts are known also from several other regions and suggest that there was no major difference in conodont faunas in northern Gondwana, Baltica and Laurentia at this time. Biostratigraphy, conodonts, graptolites, Morocco, Silurian, Sheinwoodian.     

Dendroid graptolites from the Lower Ordovician (Tremadocian) of the Yichang area, Hubei, China

The Yichang area of Hubei Province, China provides important information on Lower to Middle Ordovician fossil faunas and is a center for research on Ordovician graptolites due to the sometimes excellent preservation of the copiously available material. New material collected during a research project on Lower to Middle Ordovician biota in the Yichang Region provided among others specimens a number of dendroid (benthic) graptolites of the genera Acanthograptus, Aspidograptus, Dendrograptus, and Koremagraptus showing unexpected and unknown structural features, useful for structural and evolutionary investigations.     

UPPER ORDOVICIAN BRYOZOANS OF THE PIN FORMATION (SPITI VALLEY, NORTHERN INDIA)

Abstract: Twenty‐nine species of bryozoans from the Upper Ordovician–Lower Silurian Pin Formation (Spiti, India) have been identified. Eight of these are new: Trematopora minima, Ulrichostylus bhargavai, Ptilodictya exiliformis, Phaenopora ordinarius, Oanduellina himalayaica, Pesnastylus? vesiculosum, Ralfina? originalis and Pinocladia triangulata. The fossil record and facies analyses of the area investigated indicate shallow‐water conditions within the subtropical–tropical realm. The distribution pattern of fossils among the Ordovician/Silurian succession on the Northern Gondwana shelf and the influence of the Late Ordovician cooling phases on marine organisms are distinctive owing to a dramatic reduction in diversity globally. As far as the bryozoan taxa of Spiti are concerned, only one (Helopora fragilis) of the 29 species was recorded above the Ordovician/Silurian boundary. Observed bryozoan communities are very similar to faunas of Laurentia, the Baltic, Siberia and southern China of early–late Ordovician age.     

An Unusually Diverse Llandovery (Silurian) Diplograptid Fauna from the Southern Urals of Russia and Its Evolutionary Significance

This vesiculosus Biozone assemblage of chemically-isolated graptolites from the Orenburg District of Russia was obtained from limestones associated with pillow lavas of the Sakmara Formation of Silurian–Early Devonian age. It comprises 30 species, of which 24 are described. Nine are new, one is a new subspecies, and eight are left under open nomenclature. The assemblage is unique in its diversity of minute biserial graptolites referable to the genera ' Glyptograptus ', Normalograptus , Sudburigraptus , and Metaclimacograptus ; and is unusual in that well-preserved graptolite faunas are rare at this stratigraphical level, which is fixed by the occurrence of Cystograptus vesiculosus (Nicholson) s.l. and Paraclimacograptus innotatus (Nicholson). The following new forms are described: ' Glyptograptus ' tamariscus praecursor subsp. nov., G. mazovoensis sp. nov., G. nikitini sp. nov., G. pavlinovi sp. nov., G. telarius sp. nov., G. zimai sp. nov., Normalograptus legrandi sp. nov., N. melchini sp. nov., N. sobolevskayae sp. nov., and Metaclimacograptus bjerreskovae sp. nov. Five evolutionary lineages are defined within the ' Glyptograptus ' tamariscus (Nicholson) plexus, modifying and extending the pioneer work of Packham (1962). The diverse ' Glyptograptus ' faunas are considered to be a replacement for the Normalograptus fauna which peaked in the late Ordovician and early Silurian. Metaclimacograptus , although reasonably diverse in its speciation, exhibits a conservative style of evolution with little fundamental change throughout the Llandovery. It is suggested that cortical bandages have species-specific dimensions and micro-features. Some problematical micro- and ultrastructural features are described, notably fibrillar textures, parallel to the fusellar margins, in several species.     

Dissepiments in the graptolite Monograptus turriculatus

An example of the graptolite Monograptus turriculatus (Barrande, 1850) from the crispus Biozone (Silurian, Llandovery) of the Girvan district, Scotland, shows the first recorded instance of dissepiment formation in graptoloid graptolites. The dissepiments are formed by the fusion of thecal spines with the dorsal wall of the adjacent part of the succeeding whorl of this spiral graptolite. The formation of these structures appears easiest to envisage using models of the graptolite colony that do not involve extrathecal tissue. □ Graptolithina , Monograptus, morphology, functional morphology, Silurian, Llandovery.     

Silurian (Wenlock–ludlow) Graptolites from Bolivia

Well preserved middle to upper Silurian (Wenlock–Ludlow) graptolites from Bolivia are described for the first time. Generally monospecific graptolite faunas, of species largely endemic to South America, are found in a few levels in the lower part of the Kirusillas, Rio Carrasco and Uncía formations. The oldest identified level yields specimens of Pristiograptus praedeubeli (Jaeger) and is referred to the upper Wenlock. Younger faunas belong to the Ludlow and include Saetograptus , Monograptus and Neodiversograptus specimens. These may be referred to the Gorstian (lower Ludlow). The fauna includes Saetograptus argentinus robustus ssp. nov. and Monograptus bolivianus sp. nov.     

Functional morphology and mode of life of Palaeozoic leiocope ostracodes

Leiocopes differ from all other major ostracode groups (e.g. Palaeocopa. Binodicopa, Podocopa, Myodocopa) mainly in having a simple rounded/convex shape and a strong (dorsal and/or ventral) asymmetry. These morphological features are highly conservative in leiocopes through the Palaeozoic (lower Ordovician-middle Devonian). Functional morphology and facies distribution suggest that typical 'dome-like' leiocopes (e.g. aparchitids) may have lived off the bottom in relatively deep water environments, contrasting with most (inferred) benthic Palaeozoic ostracodes (e.g. heavily calcified dimorphic palaeocopes). Reversely, smaller sized and both dorsally and ventrally asymmetrical leiocopes (e.g. Ordovician jaanussoniids) are considered as possible benthic dwellers with reduced mobility on or within the substratum. □ Ostracodes, Leiocopa, Aparchitidae, Jaanussoniidae. Palaeozoic. Ordovician, Silurian, Devonian, asymmetry, functional morphology.     

Aspects of recent advances in the Ordovician stratigraphy and palaeontology of China

The continuous and richly fossiliferous Ordovician succession of China (particularly South China) comprises a heterogeneous suite of litho- and bio-facies, which has been a main focus of stratigraphical and palaeontological research in recent years. Among the seven GSSPs established in China, three are within the Ordovician System, and the GSSP of the Darriwilian Stage at Huangnitang, Changshan County, western Zhejiang Province, was the first “golden spike” in China and the first for the Ordovician System. A series of case studies have revealed that: (1) the Ordovician radiation of some fossil groups on the Upper Yangtze Platform (e.g., brachiopods and graptolites) reached their first α-diversity acme in the Didymograptellus eobifidus Biozone, four zones earlier than the global trend; (2) the β-diversity peak was attained 3–4 zones later than the α-diversity peak; (3) many brachiopod communities or faunas first occurred in the central part of the Upper Yangtze Platform and subsequently expanded to both more offshore and near-shore facies; (4) diachroneity existed in many aspects of the radiation. The end-Ordovician mass extinction was a severe event in South China. Two pulses of the extinction are recognized for a number of major fossil groups, some being most strongly affected during the first pulse whereas the others suffered during the second pulse. Macroevolution during the Ordovician–Silurian transition has been investigated in detail, and the role of the Lazarus effect has been found to be less important than previously believed.     

New crinoids from the Baltic region (Estonia): fossil tip‐dating phylogenetics constrains the origin and Ordovician–Silurian diversification of the Flexibilia (Echinodermata)

This study documents previously unknown taxonomic and morphological diversity among early Palaeozoic crinoids. Based on highly complete, well preserved crown material, we describe two new genera from the Ordovician and Silurian of the Baltic region (Estonia) that provide insight into two major features of the geological history of crinoids: the early evolution of the flexible clade during the Great Ordovician Biodiversification Event (GOBE), and their diversification history surrounding the end‐Ordovician mass extinction. The unexpected occurrence of a highly derived sagenocrinid, Tintinnabulicrinus estoniensis gen. et. sp. nov., from Upper Ordovician (lower Katian) rocks of the Baltic palaeocontinent provides high‐resolution temporal, taxonomic and palaeobiogeographical constraints on the origin and early evolution of the Flexibilia. The Silurian (lower Rhuddanian, Llandovery) Paerticrinus arvosus gen. et sp. nov. is the oldest known Silurian crinoid from Baltica and thus provides the earliest Baltic record of crinoids following the aftermath of the end‐Ordovician mass extinction. A Bayesian ‘fossil tip‐dating’ analysis implementing the fossilized birth–death process and a relaxed morphological clock model suggests that flexibles evolved c. 3 million years prior to their oldest fossil record, potentially involving an ancestor–descendant relationship (via ‘budding’ cladogenesis or anagenesis) with the paraphyletic cladid Cupulocrinus. The sagenocrinid subclade rapidly diverged from ‘taxocrinid’ grade crinoids during the final stages of the GOBE, culminating in maximal diversity among Ordovician crinoid faunas on a global scale. Remarkably, diversification patterns indicate little taxonomic turnover among flexibles across the Late Ordovician mass extinction. However, the elimination of closely related clades may have helped pave the way for their subsequent Silurian diversification and increased ecological role in post‐Ordovician Palaeozoic marine communities. This study highlights the significance of studies reporting faunas from undersampled palaeogeographical regions for clade‐based phylogenetic studies and improving estimates of global biodiversity through geological time.     

Restudy of some Ordovician–Silurian boundary graptolites from Anticosti Island, Canada, and their biostratigraphic significance

The Ellis Bay Formation on Anticosti Island has long been recognized for the biostratigraphic importance of its latest Ordovician conodont, palynomorph, and shelly fossil assemblages. However, a sparse record of graptolites has made it difficult to correlate these assemblages with the graptolite biozonation. Restudy of some previously described and examination of newly collected normalograptid specimens from the Ellis Bay and lower Becscie formations, however, shows that biostratigraphically important taxa are present and these species provide important constraints on the age of the strata. Normalograptus parvulus and N. minor have been identified in the upper half of the Ellis Bay Formation, suggesting that these strata are late Hirnantian in age ( N. persculptus Biozone). Normalograptus imperfectus and Normalograptus sp. aff. N. acceptus occur in the basal beds of the Becscie Formation, indicating that these strata are earliest Silurian. These graptolite data support the hypothesis that the positive carbon isotope excursion seen in the uppermost strata of the Ellis Bay Formation is isochronous with that seen in the Hirnantian strata at Dob's Linn, Scotland, and does not span much of the lower to mid-Hirnantian, as is the case in Arctic Canada and Nevada, USA.     Anticosti Island , Hirnantian, graptolites, Ordovician, Silurian.     

Foliomena Fauna (Brachiopoda) from the Upper Ordovician of Sardinia

The late Ordovician brachiopod assemblage from Sardinia is one of the youngest members of the deep-water Foliomena fauna and is characterized by the following core taxa: Christiania , Cyclospira , Dedzetina and Foliomena . The fauna also contains Epitomyonia , Leangella , Glyptorthis and Skenidioides , which are more typical of shallower-water environments during the late Ordovician but occupied deeper-water niches during the Silurian following the termination of the Foliomena fauna. The suprafamilial placement of the family Chrustenoporidae is discussed and the new species Dedzetina serpaglii and Leangella ( Leangella ) fecunda are established. In common with many mid-Ashgill Foliomena faunas the Sardinian assemblage shows significant differences from other faunal developments of this type, reflecting its geographical position and shallower water conditions than those of the classic early Ashgill Foliomena faunas. The brachiopods occur with abundant trilobites belonging to a variant of the cyclopygid fauna. The faunas developed on part of a complex of microcontinents derived from peri-Gondwana during the Ordovician.     

川东南中奥陶统大沙坝组的腕足动物群

四川东南部长宁双河及其附近地区的中奥陶统上部大沙坝组以特殊的岩相和生物相有别于黔北的可比地层十字铺组和鄂西的牯牛潭组,地层中漂浮相化石(笔石)与底栖壳相化石(如腕足动物和三叶虫)多次交互出现,笔石动物属于上层下垂对笔石动物群(Didymograptus artus Fauna),时代为中奥陶世Darriwilian(相当于Arenig最晚期至Llanvirn)。在所采集的3300余枚化石标本中有近90％是腕足动物化石,经研究共鉴定出19属21种,分属于4目7超科15科,正形贝目和扭月贝目分别有8属,占全部属群的84％。根据属的生物地理分布,该动物群中广布型分子7属、区域性分子6属,仅限于华南的土著性分子占总量的31．6％,显示出此期华南腕足动物群具有较强的地方性色彩,进一步证实中奥陶世海洋生物强分区性的推断。该腕足动物群宜称作Saucrorthis-Glyptorthis动物群。在统计分析的基础上,识别出Saucrorthis-Glyptorthis群落,包括三个以腕足动物占优势的化石群集(association),即Glyptorthis?simplex群集、Saucrorthis obscura群集和Calyptolepta chengkouensis群集,并结合沉积岩石学以及其它共生化石门类进行群落生态分析,认为该群落所生活的环境以正常浅海砂泥质底域为主(相当于BA2-3)。与上扬子台地贵州遵义十字铺组和重庆城口厚坪组两个相近腕足动物群相比,长宁地区中奥陶世的Saucrorthis-Glyptorthis动物群更接近于厚坪组所产者。与世界上其它一些同期且具相似生态位的动物群相比,除与哈萨克斯坦南部的同期动物群比较接近外,与澳大利亚、北美、英国等地的同期动物群几乎毫无联系,指示华南中奥陶世腕足动物群可能代表一个独特的生物地理小区。     

The relation of Late Ordovician glaciation to the Ordovician-Silurian changeover in North American brachiopod faunas

Sheehan, P. M.: The relation of Late Ordovician glaciation to the Ordovician-Silurian changeover in North American brachiopod faunas.
The Ordovician-Silurian changeover of brachiopod faunas in North American epicontinental seas involved the abrupt extinction of endemic Late Ordovician stocks and subsequent repopulation of North American seas by Old World taxa. The Late Ordovician Gondwanaland glaciation may have lowered sea levels sufficiently to place severe stress on the widespread shallow marine faunas in North America, resulting in their eventual extinction. The Late Ordovician depositional history in North America is not well enough known to establish the presence of a latest Ordovician regression, but the earliest Silurian was an interval of off-lap in North America. Therefore, the glacial lowering of sea level is considered to be the most likely cause of the faunal changeover.     

Devonian and Carboniferous dendroid graptolites from Belgium and their significance for the taxonomy of the Dendroidea

Devonian and Carboniferous dendroid graptolites from Belgium are evaluated and partly revised. New finds in two different stratigraphic intervals of the ‘Carrière de Lompret’, an active quarry exploiting Frasnian limestones and shales east of Frasnes-lez-Couvin, allow the identification of Callograptus sp. and Dictyonema fraiponti, both belonging to the dendroid family Acanthograptidae. The relatively high diversity of the dendroid graptolite fauna from the Viséan Marbre noir de Denée, one of the few Carboniferous graptolite faunas in the world, can be shown to be based on astogenetic and preservational aspects. Nearly all known specimens can be included in the highly variable Dictyonema fraiponti, a fan-shaped large dendroid species with complex stipes formed from tubular thecae, possessing simple to complex bridges connecting adjacent stipes. Some of the graptolite material is well preserved and provides important information on the tubarium construction of Devonian to Carboniferous dendroid graptolites and, thus, is highly significant for a taxonomic and phylogenetic understanding of the youngest dendroid graptolite faunas worldwide. The genera Callograptus and Ptiograptus are revised based on their type species (Callograptus elegans from Quebec, Canada; Ptiograptus percorrugatus from the Silurian or Devonian of Kentucky, USA) and referred to the Acanthograptidae.     

The graptolite correlation of the North American Upper Ordovician Standard

The North American Upper Ordovician reference standard, the Cincinnatian Series, contains rich shelly microfossil faunas in its type area in the Cincinnati Region but graptolites are uncommon in most of its shallow-water calcareous sediments. Consequently, the graptolite correlation of this key sequence has remained uncertain, in part, even controversial. A review of both previously published recently discovered graptolite Occurrences in the type Cincinnatian, combined with data from the important graptolite successions in Oklahoma New York-Quebec, has not only clarified the graptolite correlation of the Cincinnatian but also added new data on the morphology taxonomy, the vertical horizontal distribution, of several taxa. The information now at hand indicates that the Edenian Stage correlates with the C. spiniferus Zone, the Maysvillian Stage with the C. pygmaeus lower middle P. manitoulinensis Zone, the Richmondian Stage with the upper P. manitoulinensis , the D. complanatus , possibly part of the C. inuiti Zone. Comparison between graptolite conodont biostratigraphic evidence reveals no apparent conflict. Correlations are proposed between Upper Ordovician North American stages, graptolite conodont zones, successions in Texas, Oklahoma, Sweden, European graptolite zones, British series.     

鄂尔多斯地台南缘中奥陶统双笔石类笔石的研究

记述陕西中部耀县、富平、泾阳三县中奥陶统平和挑启坡组双笔石类：Ｏｒｔｈｏｇｒａｐｔｕｓ，Ｇｌｙｐｔｏｇｒａｐｔｕｓ，Ｃｌｉｍａｃｏｇｒａｐｔｕｓ和Ｐｓｅｕｄｏｃｌｉｍａｃｏｇｒａｐｔｕｓ等４属中的１４种，其中包括１新种和２新亚种。建立了Ｃｌｉｍａｃｏｇｒａｐｔｕｓｂｉｃｏｒｎｉｓ带和Ｏｒｔｈｏｇｒａｐｔｕｓｌｏｎｇｉｔｈｅｃａｌｉｓ带，并讨论了桃启坡组的时代问题。     

Decay and composition of the hemichordate Rhabdopleura: implications for the taphonomy of graptolites

Although the graptolites lacked biomineralised tissue, their skeletons are abundantly preserved in deeper-water mudstones. Decay experiments and observations on the closely related living hemichordate Rhabdopleura demonstrate that the periderm and stolon are highly resistant to decay, remaining intact for months, whereas the zooids are unrecognizable within days. The extreme rarity of the preservation of traces of the zooids in graptoloids reflects their planktic lifestyle; the zooids had normally decayed before burial. Curie-point-gas-chromatography (Py-GC) and Curie-point-gas-chromatography-mass spectrometry (Py-GC-MS) of the periderm of Rhabdopleura confirms that proteinaceous organic matter is a major constituent. Ultrastructurally preserved graptolite periderm (Ordovician, Oklahoma; Silurian, Arctic Canada), on the other hand, is a highly altered kerogen-like substance rich in aliphatic biomacromolecules. The composition of the preserved graptolite periderm reflects diagenetic replacement by components probably mainly derived from algal cell walls.