A Silurian myodocope with preserved soft-parts: cautioning the interpretation of the shell-based ostracod record

Ostracod crustaceans are the most abundant fossil arthropods. The Silurian Pauline avibella gen. et sp. nov., from the Herefordshire Lagerstätte, UK, is an extremely rare Palaeozoic example with soft-part preservation. Based on its soft-part morphology, especially the exceptionally preserved limbs and presence of lateral eyes, it is assigned to the myodocopid myodocopes. The ostracod is very large, with an epipod on the fifth limb pair, as well as gills implying the presence of a heart and an integrated respiratory–circulatory system as in living cylindroleberidid myodocopids. Features of its shell morphology, however, recall halocyprid myodocopes and palaeocopes, encouraging caution in classifying ostracods based on the carapace alone and querying the interpretation of their shell-based fossil record, especially for the Palaeozoic, where some 500 genera are presently assigned to the Palaeocopida.     

Upper Silurian microplankton of the Leintwardine Group, Ludlow Series, in the type Ludlow area and adjacent regions

The acritarchs and prasinophyte algae from the Upper Bringewood, Lower Leintwardine, Upper Leintwardine and Lower Whitcliffe formations of the Ludlow type area and surrounding regions are described. The following new taxa are proposed: Cymatiosphaera pumila sp. nov., Melikeriopalla pustula sp. nov., Cheleutochroa beechenbankensis sp. nov., Cymbosphaeridium molyneuxii sp. nov., Flammulasphaera bella gen. et sp. nov., Percultisphaera incompta sp. nov., Salopidium aldridgei sp. nov. and Umbellasphaeridium? wicanderi sp. nov. A holotype is defined for Lophosphaeridium galeatum Hill and a further 34 new taxa are described under open nomenclature. Two biozones, identified by the first appearance of the zone taxon, are defined in the Sunnyhill section, Ludlow [Global Stratotype Section and Point (GSSP) for the base of the Ludfordian Stage, Ludlow Series]. The base of the Leoniella vilis Biozone is identified at 4.52m below the top of the Upper Bringewood Formation at Beechenbank, Aymestrey. The base of the Triangulina sanpetrensis Biozone is identified at 15.49m above the base of the Lower Leintwardine Formation and Ludfordian Stage at the Sunnyhill section. These biozones allow correlation with sections in north–west Spain, Podolia and Gotland. The possibility of using Visbysphaera whitcliffense and U.? wicanderi as biozonal indicators is suggested.     

Chitinozoan biostratigraphy of the basal Wenlock Series (Silurian) Global Stratotype Section and Point

Diverse and abundant assemblages of chitinozoans allow the recognition of the upper part of the Margachitina margaritana Biozone in the uppermost Llandovery and lowermost Wenlock series of the Hughley Brook section. The Cingulochitina bouniensis and Salopochitina bella biozones are identified in the lower part of the Buildwas Formation, Wenlock Series. The chitinozoan data indicate that the base of the Wenlock Series most probably correlates with a level in the upper centrifugus or lower murchisoni graptolite biozones. Chitinozoans also indicate that the base of the riccartonensis graptolite Biozone may occur within the Buildwas Formation and not the overlying Coalbrookdale Formation.     

Silurian bolbozoids and cypridinids (Myodocopa) from Europe: pioneer pelagic ostracods

Abstract: Bolbozoid and cypridinid myodocope ostracods from France, the Czech Republic and Sardinia consist of four genera (one new: Silurocypridina) and at least ten species (eight new: Bolbozoe acuta, B. rugosa, B. parvafraga, Parabolbozoe armoricana, Silurocypridina retroreticulata, S. variostriata, S. calva and Calocaria robusta). They have a functional design, palaeogeographical distribution, depositional setting and faunal associates that suggest that they were swimmers in the water column, living in dim light, relatively deep environments, above near bottom dysoxic/anoxic conditions. These myodocope faunas include some possible pelagic species sensu stricto, with widespread palaeogeographical (transoceanic) distribution, and some species that occupied hyperbenthic niches and were capable of making short incursions to the sediment to scavenge on carrion. These data support the model that Silurian myodocopes were pioneer pelagic ostracods.     

Microplankton associations, biofacies and palaeoenvironment of the type lower Ludlow Series, Silurian

The acritarchs and prasinophyte algae from the type lower Ludlow Series of the Goggin Road section, Ludlow, England, are resolved into seven recurrent associations comprising taxa with similar environmental preferences. Endemic and environmentally sensitive associations of acritarchs and prasinophytes are identified and high-resolution fluctuations in the early Ludlow palaeoenvironment are established. An early Ludlow crisis in the acritarchs is recognized in the lower part of the Middle Elton Formation, when an abrupt palaeoenvironmental change in the Ludlow area resulted in a large decline in the abundance of the acritarchs, but allowed Tasmanites and retiolitid graptolites to flourish briefly. Cymbosphaeridium sp. A, Pulvinosphaeridium ludlowense and Multiplicisphaeridium arbusculum forma A are taxa possibly specialized, or produced as a response, to a stressed palaeoenvironment, as they are most abundant when other acritarchs and prasinophytes are uncommon. The low abundance of acritarchs and prasinophytes in the Upper Elton Formation may be related to high sedimentation rates and to the slumping of sediments caused by instability on the shelf of the Welsh Basin, or to lower plankton productivity.     

Silurian Chitinozoa from the type Wenlock and Ludlow of Shropshire,England

Chitinozoa recorded from the Wenlock and Ludlow from the type areas in Shropshire and other localities in the Welsh Basin are listed, and the ranges of 35 taxa of stratigraphic value plotted.     

Silurian acritarchs from the type Wenlock and Ludlow of Shropshire,England

Acritarch taxa are recorded and listed from the Wenlock and Ludlow from the type areas in Shropshire, England, and other localities in the Welsh Basin of South Wales and the Welsh Borderland of England. The ranges of 141 taxa of stratigraphic value are plotted. Five new genera are described: Hogklintia, Lepto brachion, Rhaco brachion, Salopidium and Wrensnestia. 36 new species are described, and 32 new combinations proposed. Seven acritarch zones are recognised in the Wenlock and Ludlow.     

Sea level and faunal changes during the latest Llandovery and earliest Ludlow (Silurian)

It is now recognized that the late Telychian and early Gorstian sea level changes are, like many others in the Silurian, of world‐wide extent. The 30–50 m deepening events at these times were between 1 and 2 Ma in duration, so melting continental ice caps appear to be the most probable cause. The faunal changes associated with the two events are respectively very close to the base and the top of the Wenlock Series and thus segregate many of the faunas diagnostic of the Llandovery, Wenlock, and Ludlow series. Permanent extinctions (often followed by radiations) are more prevalent in the graptolites, conodonts, and acritarchs, while benthic faunas are more affected by regional shifts in community distributions. This means that, in the benthos, slowly evolving lineages are the only reliable guides to time. Such phyletic evolution, however, appears to have been unaffected by sea‐level events.     

Miocene ostracods from the Tremiti Islands and Hyblean Plateau: biostratigraphy and description of new and poorly known species

The main taxonomic and stratigraphical results of a study of ostracods from the middle-late Miocene of the Tremiti Islands and the Hyblean Plateau are presented. The faunas are mostly typical of deep thermospheric or psychrospheric bathyal environments. Excluding a few, mostly juvenile shallow water contaminants, a total of 127 species were recorded. Twenty-four taxa are discussed herein; of these, 13 species (Cytherella dissimilis nov. sp., Cytherella parvula nov. sp., Cytherella subtilis nov. sp., Argilloecia triangularis nov. sp., Retibythere (Retibythere)? claudii nov. sp., Retibythere (Retibythere)? gibba nov. sp., Cytheropteron (Cytheropteron) joachinoi nov. sp., Cytheropteron (Aversovalva) consuetum nov. sp., Cytheropteron (Aversovalva) trimerumense nov. sp., Cytheropteron (Aversovalva) turgidum nov. sp., Loxoconcha julianii nov. sp., Buntonia whatleyi nov. sp., Pterygocythere nuda nov. sp.) and a subspecies (Costa tricostata dorsoarcuata nov. subsp.) are described as new, while six are left in open nomenclature. The stratigraphical distribution of ostracods is expressed in terms of planktonic foraminiferal zones. Information obtained from this study enriches biostratigraphical data on Mediterranean Miocene deep-sea ostracods, especially with respect to the Langhian.     

Integrated Silurian Chitinozoan and Graptolite Biostratigraphy of the Banwy River Section, Wales

The succession of 38 upper Llandovery–lower Wenlock chitinozoan taxa from graptolitic horizons in the Banwy River section (Powys, Wales) is described. Five new species are named: Bursachitina nestorae Conochitina leviscapulae Conochitina mathrafalensis Belonechitina cavei Belonechitina meifodensis . A further ten taxa are described under open nomenclature. Seven chitinozoan biozones are recognized in the Banwy River section, three of which ( Cingulochitina bouniensis Conochitina acuminata Salopochitina bella ) are new. The base of each biozone is correlated with the graptolite biostratigraphical scheme as follows: Angochitina longicollis Biozone — upper spiralis Biozone; Conochitina acuminata Biozone — lowermost lapworthi Biozone; Margachitina banwyensis Biozone — upper lapworthi Biozone; Margachitina margaritana Biozone — lowermost insectus Biozone; Cingulochitina bouniensis Biozone — upper murchisoni Biozone; Salopochitina bella Biozone — upper firmus Biozone. The succession of chitinozoan biozones in the Banwy River section is compared with that in other sections which have graptolite biostratigraphical control. This has highlighted the following: (1) the correlation of the base of the dolioliformis Biozone with the graptolite biozonation is imprecise; (2) E. dolioliformis is recorded only from levels after the first A. longicollis in Sweden (although this may reflect previously unrecognized synonymies); (3) the longicollis Biozone may be diachronous, its base correlating with levels low in the Telychian in Sweden, Norway and Estonia and with the upper Telychian spiralis Biozone in Wales and the Prague Basin; (4) data herein and from the Prague Basin indicate that the base of the margaritana Biozone correlates with a level low in the insectus Biozone.     

Sea level and faunal change during the late Wenlock and earliest Ludlow (Silurian): A point of view from the Algerian Sahara

A late Wenlock transgression (C. rigidus zone) affecting a large area of the Algerian Sahara has been known for the last thirty years; in this case there is no break at the Wenlock‐Ludlow boundary and the sedimentation is continuous to the earliest Ludlow. But in many other areas of Sahara a late Wenlock regression is observed. This regression may or may not be followed by a latest Wenlock or an early Ludlow transgression. A sedimentary model is proposed in despite of several problems. In order to compare what happens in each region a schematic diagram is used showing the tendencies to shallowing and deepening. Differences observed between the North Sahara basin and the other parts of the Algerian Sahara and comparison between local sea‐level curves and the global sea‐level curve proposed by Johnson et al. (1991) lead to reconsider the evidence of an eustatic early Ludlow event. To end the problem of the disappearance of nearly all graptolites at the end of Wenlock is discussed. The influence of land emergence in the evolution of marine fossil groups in the disturbance that it can bring to the development of the microplankton and the chemistry of the sea seems to have been too neglected.     

The root of the problem: palaeoecology of distinctive crinoid attachment structures from the Silurian (Wenlock) of Gotland

The holdfast (attachment structure) is the most understudied aspect of the palaeoecology of the endoskeleton of fossil crinoids. A new collection of well-preserved holdfasts from a recently reopened quarry at Hunninge, Gotland, in Homerian (upper Wenlock) strata includes several morphologies. The most common are terminal dendritic radicular holdfasts (TDRHs) that may be derived from the cladid Ennallocrinus d'Orbigny. These have a consistent morphology of five, equally spaced, long radices that spread across the sea floor. These crinoids were gregarious, and TDRHs in a group commonly show the same radice orientations. The radices have a large axial canal compared with those of modern crinoids; each included, at least, nervous tissues. Taken together, these features suggest that, apart from attachment, these distinctive TDRHs may have served a sensory function. Other holdfasts in this assemblage also show monospecific aggregations, perhaps suggesting biochemical attraction such as that shown by certain other sessile invertebrates such as barnacles.     

The fourth series of the Silurian System in Podolia

The Skala horizon of Podolia has been under discussion as a possible candidate for the stratotype of the fourth Silurian series (primary subdivision) in the international standard chronostratigraphical scale. After a brief survey of the history of investigation, the section through the basal boundary of the Skala horizon is fully documented. The lithostratigraphy, biostratigraphy, and correlation of the Skala horizon are then discussed.     

Integrated conodont and radiolarian biostratigraphy of the upper Norian in Baoshan Block,Southwestern China

Two significant stratigraphical microfossils, conodonts and radiolarians, are usually used for the Upper Triassic chronostratigraphy. The Baoshan Block was located in eastern Tethys during the Late Triassic where the biostratigraphical data of Upper Triassic are still poorly known. We collected new samples from the Hongyan section (HY) for biostratigraphical study. This 24-m-thick section in Dabaozi Village, Baoshan City, is mainly composed of thin-layered limestones, sandstone and siltstone. The conodont fauna is referred to Sevatian 1 (late Norian), in which the species Mockina englandi, Mockina carinata and Mockina mosheri morphotype B are first recognized in the Baoshan Block, and thus eastern Tethys. The Norian radiolarian associations are first reported in the Baoshan Block, which correlate with the biozonation of North America and also that proposed for Central Japan. The radiolarian assemblages found in the analysed samples in HY section can be referred to the Sevatian Betraccium deweveri Zone. The Baoshan Block is a key area for conodont and radiolarian-based correlations between the Tethys, Japan and North American domains.     

Homology of Holocene ostracode biramous appendages with those of other crustaceans: the protopod, epipod, exopod and endopod

Unambiguously biramous appendages with a proximal precoxa, well-defined coxa and basis, setose plate-like epipod originating on the precoxa, and both an endopod and exopod attached to the terminal end of the basis are described from several living Ostracoda of the order Halo-cyprida (Myodocopa). These limbs are proposed as the best choice for comparison of ostracode limbs with those of other crustaceans and fossil arthropods with preserved limbs, such as the Cambrian superficially ostracode-like Kunmingella and Hesslandona. The 2nd maxilla of Metapolycope (Cladocopina) and 1st trunk limb of Spelaeoecia, Deeveya and Thaumatoconcha (all Halocypridina) are illustrated, and clear homologies are shown between the parts of these limbs and those of some general crustacean models as well as some of the remarkable crustacean s.s. Orsten fossils. No living ostracodes exhibit only primitive morphology; all have at least some (usually many) derived characters. Few have the probably primitive attribute of trunk segmentation (two genera of halocyprid Myodocopa, one order plus one genus of Podocopa, and the problematic Manawa); unambiguously biramous limbs are limited to a few halo-cyprids. Homologies between podocopid limbs and those of the illustrated primitive myodocopid limbs are tentatively suggested. A setose plate-like extension, often attached basally to a podocopid protopod, is probably homologous to the myodocopid epipod, which was present at least as early as the Triassic. Somewhat more distal, less setose, and plate-like extensions, present on some podocopid limbs (e.g., mandible), may be homologous instead to the exopod (clearly present on myodocopid mandibles). The coxa (or precoxa) is by definition the most basal part of the limb. A molar-like tooth is present proximally on the mandibular protopod of many ostracodes; it is the coxal endite and projects medially from the coxa (or proximal protopod). The Ostracoda is probably a monophyletic crustacean group composed of Myodocopa and Podocopa. All have a unique juvenile (not a larva) initially with three or more limbs. Except that juveniles lack some setae and limbs, they are morphologially similar to the adult. Thus the following suite of characters in all instars may be considered a synapomorphy uniting all Ostracoda: (1) Each pair of limbs is uniquely different from the others. (2) The whole body is completely enclosed within a bivalved carapace that lacks growth lines. (3) No more than nine pairs of limbs are present in any instar. (4) The body shows little or no segmentation, with no more than ten dorsally defined trunk segments. No other crustaceans have this suite of characters. A probable synapomorphy uniting the Podocopa is a 2nd antenna with exopod reduced relative to the endopod.     

A southward extension of the Silurian Tuvaella brachiopod fauna

The co-occurrence of Tuvaella gigantea Chernyshev, Tannuspirifer pedaschenkoi (Chernyshev), Leptos-trophia elegestica Chernyshev and Stegerhynchus sp. in the Upper Hongliuxia Formation, Barkol, NE Xinjiang, China, shows that this is a representative of the well-known Tuvaella Fauna in China. The correlation of the Tuvaella gigantea Assemblage Zone of the Upper Hongliuxia Formation at its type locality is discussed to document the correlation of this formation with the Pichishui Beds of the Upper upper Chergak Formation in Tuva, the Woduhe Formation m the Lesser Khingan Mountains, and other equivalents based on the presence in some abundance of T. gigantea , and without T. rackovskii. The Tuvaella gigantea-bearing strata are tentatively assigned to the Ludlow. With regard to the paleozoogeographic affinity of the Tuvaella Fauna it is at present suspected by some that the area inhabited by Tuvaella represents a peculiar zoogeographic province or region during the Silurian. However, the information available indicates that the Tuvaella occurrences in east-central Asia represent nothing more than a specialized environmental situation in the light of the presence of a single endemic genus, i.e. Tuvaella itself, among 25 articulate brachiopod genera examined. The evidence of both lithology and brachiopods investigated has important implications for suggesting a shallow, commonly rough-water environment between low intertidal and high subtidal areas.     

Development of facies and C/O-isotopes in transects through the Ludlow of Gotland: Evidence for global and local influences on a shallow-marine environment

Summary The Silurian of Gotland is characterized by repeated changes in depositional facies development. The deposition of uniform sequences of micritic limestones and marls was interrupted four times by the growth of reef complexes and the formation of expanded carbonate platforms. Coinciding with these, often abrupt, facies changes extinction events occurred which predominantly affected nektonic and planktonic organisms. Ratios of carbon- and oxygen-isotopes covary with the facies development. Periods in which the deposition of limestonemarl alternations prevailed are characterized by relatively low C- and O-isotope values. During periods of enhanced reef growth isotope values are high. For these changes,Bickert et al. (1997) assume climatic changes between humid “H-periods”, with estuarine circulation systems and cutrophic surface waters with decreased salinity in marginal seas, and arid “A-periods”, with an antiestuarine circulation and oligotrophic, stronger saline surface waters. In order to separate local and regional influences on the isotopic development from the global trend, the interactions between facies formation and isotope record have to be clarified. For this purpose, the patterns of isotope values in the upper part of the Silurian sequence on Gotland (upper Wenlock —upper Ludlow) has been determined and stratigraphically correlated along four transects through different facies areas. Facies formation during this time interval was investigated by differentiation and mapping of twelve facies complexes in the southern part of Gotland. These include shelf areas, reef complexes with patch reefs and biostromes, backreef facies, and marginal-marine deposits. The good correspondence between the carbon-isotope records of the four transects suggests that local environmental conditions in the different facies areas did not influence the δ¹³C values. Therefore, a supra-regional or even global mechanism for the C-isotope variations is likely. In contrast to carbon istopes, the oxygen-isotope values of the four transects generally show parallel trends, but higher variabilities and in parts distinctly deviating developments with a trend to more negative values. These are interpreted as an effect of local warming in small shallow-water areas which developed during arid periods in reef complexes and backreef areas. The boundaries between A-periods and H-periods, as defined by δ¹³C values, which are interpreted as isochrones, can be mapped. From the upper Homerian to the Pridolian six parastratigraphic isotope zones are defined which only partly match the stratigraphic division ofHede (1942, 1960). The isotope stratigraphy results in an improved correlation between the shallow and marginal-marine areas in the eastern part of Gotland and the uniform shelf areas at the west coast of the island. Furthermore, a detailed relationship between the development of carbon and oxygen isotope ratios, the carbonate facies formation, and the succession of palaeontological events could be observed. At the transition from H-periods to A-periods, major extinction events occurred prior to the first increase of δ¹³C and δ¹⁸O values. Extinction events affected conodonts, graptolites, acritarchs, chitinozoans, and vertebrates and resulted in impoverished nektonic and planktonic communities. The reef-building benthos was less affected. Parallel to a first slight increase of isotope values, facies began to change, and reefs developed in suitable locations. The subsequent rapid increase of C- and O-isotope values occurred contemporarily with strong facies changes and a short-term drop of sea-level. Oligotrophic conditions in the later stages of A-periods led to strong reef growth and to an expansion of carbonate platforms. The transitions from A-periods to H-periods were more gradual. The δ¹³C values decreased slowly, but reef growth continued. Later the reefs retreated and were covered by the prograding depositional facies of shelf areas. The diversity of planktonic and nektonic communities increased again. The close relationship between facies formation, palaeontological events and isotope records in the Silurian suggests common steering mechanisms but gives no indication of the causes for the repeated extincion events related to H-period/A-period transitions. Especially the observation, that strong extinctions occurred prior to changes of isotope values and facies, points to causes that left no signals in the geological record. Hypothetical causes like collapse of trophical nets, anoxias, or cooling events are not evident in the sediment record or do not fit into the regular succession of period transitions.     

A taxonomic reevaluation of freshwater ostracods (Crustacea,Ostracoda) referred to the genus Cypridopsis Brady, 1868 from Southeast Asia

A taxonomic reevaluation of five species of Southeast Asian freshwater ostracods formerly assigned to the genus Cypridopsis Brady, 1868 has resulted in the retention of Cypridopsis adusta Sars, 1903, Cypridopsis exigua Sars, 1903 and Cypridopsis dubia Sars, 1903 in Cypridopsis and transfer of Cypridopsis albida (Vavra) 1898 and Cypridopsis arsenia Tressler, 1937 to the genus Plesiocypridopsis Rome, 1965.     

Seawater strontium isotope curve at the Silurian/Devonian boundary: a study of the global Silurian/Devonian boundary stratotype

Present application of ⁸⁷Sr/⁸⁶Sr chemostratigraphy to detailed stratigraphical tasks is limited by inaccurate calibration of the general seawater strontium curve to absolute as well as to relative time scales. For this reason, refinement of the general seawater strontium curve has been suggested, using mainly clearly defined global boundary stratotype sections. This study reports the first ⁸⁷Sr/⁸⁶Sr data from the global Silurian/Devonian boundary stratotype section and fills an existing 1-Ma gap in available data. Generally, the data from the stratotype fit the range interpolated from published ⁸⁷Sr/⁸⁶Sr data of the general curve, but the slight differences may suggest an existence of a high-order oscillation near the Silurian/Devonian boundary. Higher ⁸⁷Sr/⁸⁶Sr values in the Devonian part of boundary bed 20 (20-beta) may indicate an exotic material influx of recycled sediment.