Pattern of some Silurian bioevents

The best‐known Silurian bioevent occurred at the end of the Wenlock: the lundgreni event, together with the nassa‐ludensis crisis, was established among planktic graptolites. The East Baltic data show several peaks of high diversity (especially triangulatus, turriculatus, scanicus zones) and three levels of low diversity of graptolites (antennularius, radians, ludensis zones). These are more or less coincident with sea‐level changes. Energetic innovation of the shallow shelf corals started in the early Silurian. The late Wenlock regression seems not to affect them seriously but the late Silurian decline coincides with the aridization of the climate and a regression of the shelf seas. The Agnatha had strong radiations in the Wenlock (ludensis Zone) and Ludlow (leintwardinensis Zone), the fishes in the Pridoli. Many Silurian vertebrates were long‐ranging and extinction rate was relatively low; only at the late leintwardinensis level and in the latest Ludlow did considerable extinctions occur.     

Faunal and facies changes through the mid Homerian (late Wenlock,Silurian) positive carbon isotope excursion in Podolia,western Ukraine

The late Wenlock is characterized by two global regressive‐transgressive eustatic cycles in association with a double‐peaked positive carbon isotope excursion. The onset of the excursion coincides with an extinction event affecting graptolites (the lundgreni event) and proposed to affect conodonts (the Mulde Event) and proliferation of non‐skeletal carbonates. In order to test the hypothesized relationships between eustatic and ecological changes, the tropical carbonate Homerian succession in Podolia has been examined with respect to conodont, sequence and δ¹³C stratigraphy. Four depositional sequences (DS) have been identified. The onset of the δ¹³C excursion occurs at the boundary between DS1 and DS2, corresponding to a forced regression of proposed glacioeustatic origin. The following rapid eustatic transgression associated with the highest δ¹³C values of 5.2‰ includes a higher‐order shallowing episode recorded in Podolia as normal regression and a boundary between DS2 and DS3. This interval is distinguished by the presence of oncoids and thrombolitic buildups. The latest Wenlock eustatic fall and the corresponding second peak of the δ¹³C excursion corresponds in Podolia to a stratigraphic gap. The first δ¹³C peak (top of DS1 and DS2) corresponds to the O. bohemica longa conodont Zone, the interval between the two peaks (DS3) – to K. ortus absidata and C. murchisoni zones, and DS4 is tentatively placed in the lowermost Ludlow Series. The record of relative sea‐level changes in Podolia is consistent with reconstructions based on successions in England and Sweden. The moderate drop in conodont taxonomic richness may reflect the primary depositional control over their proposed extinction.     

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.     

The diachronism of the Wenlock Limestone

A quick transgressive event occurs near the top of the Wenlock Limestone. The initiation of this is thought to be synchronous, proving the top of the Wenlock Limestone to be diachronous and younging southwards. As a result the top of this rock unit cannot be used satisfactorily as a time line. It is suggested that the initiation of the transgression be taken as the practical means for the correlation of the Wenlock-Ludlow Series boundary. The Birmingham Siltstone Formation is erected for the strata between Dudley and Ludlow which are younger than Wenlock Limestone and older than the transgression. The Wenlock Limestone of Usk was deposited after the transgression and is unrelated to the main Wenlock Limestone. Thus it is treated as a separate entity and named here the Usk Limestone Formation.     

Paleoecology,biostratigraphy and biogeography of late Silurian to early Devonian acritarchs and prasinophycean phycomata in well A161, Western Libya,North Africa

Diverse and well preserved acritarch and prasinophycean phycomata assemblages were recovered from the late Silurian to Lower Devonian strata of well A161 in western Libya, and four distinct acritarch biozones are recognized, based on the stratigraphic distribution of 156 species. The palynoflora is independently dated by means of chitinozoans, and allows discussion of the evolution of acritarchs and prasinophyte phycomata across the Ludlow–Přı́dolı́ boundary in relation to probable major climatic change, as well as in the early and middle Přı́dolı́, and the lower Lochkovian. Correlations are proposed with the British Isles, Baltica, and Algerian Sahara. Sedimentation occurred in shallow high-energy conditions throughout, but with periodic rise of sea level. The changes in marine to terrestrial palynomorph ratios through the section document the relationship between marine palynomorph assemblages and sea surface conditions in these marginal marine environments. The major drift of Gondwana towards low latitudes during the Ludlow–Přı́dolı́ transition seems to have been the driving force behind homogenization of assemblages on the two sides of the Rheic ocean, and explains the similarities between phytoplanktonic assemblages of the north Gondwanan margin and the South of Baltica. The data suggest that the Rheic ocean was almost closed by the late Silurian, and had become restricted to a moderately deep sea. In the Lochkovian the microflora are strongly facies-dependent and delineate more restricted provinces such as the Ibarmaghian domain in the sense of Plusquellec (1987) including the Maghreb and Ibero–Armorican areas.Nine new species are described: Arkonia nova, Arkonia paulumstriata, ?Cymatiosphaera florida, Cymatiosphaera nimia, Dactylofusa hispidusa, Disparifusa quasibernesgae, Evittia areolata, Multiplicisphaeridium verticisum, and ?Villosacapsula steemansii. In addition, three new combinations are suggested: Visbysphaera bonita (Cramer) comb. nov., Visbysphaera jardinei (Cramer) comb. nov. and Visbysphaera albanega (Cramer et al.) comb. nov.     

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.     

Late Wenlock (Middle Silurian) global Bioevent: Possible chemical cause for mass graptolite mortalities

Graptolites nearly became extinct in the latest Wenlock in all preserved stratigraphic sequences of this age. Graptolite mortalities occurred along the western coast of Laurentia and at sites that surrounded the Proto‐Tethys. Graptolite mass mortalities took place among deep‐water, open ocean dwelling organisms. After the mass mortalities, only the Pristiograptus dubius group and retiolids surface or near‐surface dwellers, survived. For a period of time, little speciation or diversification occurred. The base of the Ludlow is marked by diversification, with appearances of S. colonus, M. nilssoni and other groups which occur in near surface waters. None of the extensive plate movements postulated for the Silurian readily explain the mass extinctions that occurred. During the Silurian, global temperatures were warmer than present and atmospheric oxygen concentrations were lower, creating extensive oceanic anoxia. Below the oxygenated surface layers of the ocean, was an anoxic, non‐sulfidic zone (i.e. nitrate‐reducing) above a sulfidic zone. Graptolites lived over a range of depth from the oxygenated zone to either near or in the nitrate‐reducing zones. As the oxygen concentration declined through the Silurian, the depth of the oxic zone would have become shoaler with expanding anoxia. Late Wenlock graptolites that were unable to migrate to shallower depths, living in borderline oxygen conditions, could have been killed, resulting in the mortalities of the late Wenlock. Only those graptolites that were surface dwellers survived, adapted and reradiated.     

内蒙古达茂旗巴特敖包地区包尔汉图剖面牙形刺生物地层

内蒙古达茂旗(达尔罕茂明安联合旗)巴特敖包地区志留纪、泥盆纪碳酸岩相地层发育,本区珊瑚、腕足类、层孔虫等底栖大化石的研究工作已有一定基础。但志留纪、泥盆纪地层在时代确定和对比上还存在很多问题,必须用主导化石门类牙形刺加以解决。研究表明：包尔汉图剖面的顶部属泥盆系无疑,应归入阿鲁共组,而不是西别河组;巴特敖包地区的海侵,始于罗德洛统卢德福德阶(Ludfordian)早期。本剖面没有发现文洛克世和罗德洛世早期的海相沉积。本文描写了一个志留纪牙形刺新种：Ozarkodina uncrispa sp．nov．。     

A conodont, thelodont and acanthodian fauna from the Lower Přídolí (Silurian) of the Much Wenlock area, Shropshire

Conodonts, thelodonts and acanthodians have previously been collected from the P?ídolí of Shropshire (Welsh Borderland) from bone beds such as the Ludlow Bone Bed. In general, specimens collected from these types of deposit are very abraded and make taxonomic studies difficult. This contribution describes well preserved thelodont microelements from two samples from the Downton Castle Sandstone Formation (P?ídolí, Silurian) of the Much Wenlock area of Shropshire. Head scales, a wide variety of transitional scales, and pore scales of the lateral line sensory canals of the thelodont Paralogania ludlowiensis (Gross) are described for the first time. A new acanthodian Nostolepis linleyensis sp. nov. and elements from the apparatus of the conodont Ozarkodina?hemensis sp. nov. are documented from the same samples. The thelodonts in both samples are typical of upper Ludlow–P?ídolí faunas from across the Welsh Borderland, and are also present in smaller numbers in the Ludlow Series of Gotland, Sweden, Saaremaa Island, Estonia, and in Latvia and Lithuania. The conodont O? hemensis sp. nov. is restricted to the Much Wenlock area of Shropshire. The co-occurrence of well preserved elements of conodonts and thelodonts suggests that they were deposited fairly rapidly with little or no re-working and that a restricted marine environment prevailed in the early P?ídolí in the Much Wenlock area of Shropshire.     

Symbiotic interactions in the Silurian of Baltica

Thirteen symbiotic associations occur in the Silurian of Baltica. Symbiosis was especially prominent among colonial animals, most commonly with stromatoporoids. These sponges hosted the most diverse fauna of endobiotic symbionts (including rugosans, Syringopora, ‘polychaetes’, cornulitids and lingulids). This pattern can be explained by the abundance of stromatoporoids in the Silurian of Baltica and their large skeletal volume, making them attractive hosts for smaller invertebrates. There is an evolutionary trend of an increasing number of different pairs of symbiotic taxa from the Llandovery to the Ludlow, with a remarkable increase in the Ludlow. This is likely related to an increase in the number of mutualistic taxa that could have had evolutionary advantages over organisms less amenable to symbiosis. The number of different pairs of symbiotic taxa also increased in the Wenlock, which may be linked to delayed recovery from the end‐Ordovician mass extinction.     

Conodontophorid biodiversification during the Ordovician in South China

Wu, R., Stouge, S. & Wang, Z. 2012: Conodontophorid biodiversification during the Ordovician in South China. Lethaia, Vol. 45, pp. 432–442. Analysis of the Ordovician conodontophorid diversity pattern for South China using normalized and total diversity measures reveals that diversity peaks occurred in the mid‐Tremadocian, mid‐late Floian, early Dapingian and mid‐Darriwilian periods. The conodontophorids radiated during the Floian, maintaining relatively high diversity into the early part of the Middle Ordovician until a significant diversity decrease occurred in the late Dapingian. A relatively low diversity level prevailed in the Late Ordovician. Three diversification intervals based on origination, extinction and turnover rates have been identified i.e. (1) Tremadocian to mid‐late Floian, (2) early Dapingian and (3) late Dapingian to early Darriwilian. Diversity curves for conodontophorids, brachiopods, graptolites, acritarchs and trilobites from South China are comparable during the Early Ordovician, although differences are apparent in the Middle and Late Ordovician. In South China, conodontophorid diversity reacted primarily to sea‐level changes during the Early and Middle Ordovician, when the peak of this biodiversification generally coincided with a transgression. Climate changes – especially the global cooling that occurred during the Late Ordovician glaciation – and sea‐water chemistry were also important controlling factors. □Biodiversification, conodonts, Ordovician, South China.     

Extinction and opportunistic evolution among late Wenlock graptolites

The late Homerian was a time of profound importance to graptolites. Following the complete extinction of Cyrtograptus and the near extinction of Monograptus (s.s.)at the end of the lundgrenitestis Zone(lower Homerian, upper Wenlock), the morphologically simple and long-ranging Pristiograptus dubius lineage underwent rapid (opportunistic) cladogenesis during the succeeding upper Homerian. Two clades, each with novel and sometimes complex sicular and thecal structures, appeared. One clade, the 'Pristiograptus' praedeubeli group, gave rise to similar but more complex species, such as ' P.' ludensis and 'P.' deubeli , which in turn gave rise to Ludlow taxa, including Saetograptus (s.l.) and Pseudomonclimacis. The other clade, the Lobograptus group, gave rise to Ludlow taxa Lobograptus, Neolobograptus, Bohemograptus , and Neocucullograptus some with very complex thecal structures. Ludlow and younger monograptid faunas therefore comprise at least three separate lineages: Monograptus (s.s.), the Lobograptus group (clade), and the 'Pristiograptus' praedeubeli group (clade). Cladogenesis, evolution, extinction, Homerian, PRISTIOGRAPTUS, LOBOGRAPTUS     

A high-precision global biostratigraphy of myodocope ostracods for the Silurian upper Wenlock Series and Ludlow Series

The wide, trans-oceanic geographical distribution of myodocope ostracods during the Silurian (especially during the Ludlow and Pridoli epochs), and their widespread preservation in rocks of that age, permits the establishment of a transcontinental biostratigraphy of comparable resolution to coeval graptolite/chitinozoan/conodont biozones. Seven myodocope biozones, extending from the Homerian Stage, upper Wenlock Series Cyrtograptus lundgreni graptolite biozone to the middle part of the Ludfordian Stage of the Ludlow Series, enable a time-resolution for each biozone of circa 1 million years. These biozones can provide high-resolution correlation across Europe into Arctic Russia and Central Asia. There is also the potential for a myodocope biostratigraphy applicable from the uppermost Silurian (Pridoli) to the Carboniferous.     

Strontium stratigraphy of the Burdigalian transgression in the Western Mediterranean

Brandano, M. & Policicchio, G. 2011: Strontium stratigraphy of the Burdigalian transgression in the Western Mediterranean. Lethaia, Vol. 45, pp. 315–328. A ⁸⁷Sr/⁸⁶Sr analysis of selected pectinid shells has been performed to refine the stratigraphical framework of the Burdigalian transgressive event. Samples were collected from shallow water successions of the Western Mediterranean area (North‐Eastern Sardinia, Southern Corsica, Central Apennines). The Western Mediterranean is the result of the interaction between the European and the African plates. Sardinia and Corsica were extended and rifted apart from the Iberian Peninsula during the opening of the Liguro‐Provencal basin, between 30 and 15 Ma. In the outcrops of Northern Sardinia and Southern Corsica, Burdigalian deposits nonconformably overlie Varisican granites. In the central Apennines the Burdigalian deposits paraconformably overlie the Cretaceous platform. These are transgressive deposits, characterized by the presence of Miogypsina globulina, which points out early to middle Burdigalian age. The Sr isotope ages indicate that marine transgression in Northern Sardina and Southern Corsica took place between 18.6 and 18.3 Ma, which could correspond to the eustatic sea level rise of the Bur3 third‐order sequence. In the Central Apennines the Burdigalian transgression was anticipated (18.8 Ma) by subsidence input linked to the eastward migration of the orogenic system. This input allowed the Miocene carbonate ramp to develop on the Cretaceous platform substrate, while successive eustatic rise controlled the evolution of the stratigraphical architecture. The following regressive phase is recorded in both the Apennine and Corsica successions. The end of this phase is dated between 17.6 and 17.5 Ma and corresponds to sea level drop corresponding to the occurrence of the Mi 1b event. □Burdigalian, pectinids, strontium stratigraphy, transgression, Western Mediterranean     

The effects of the final stages of the Late Ordovician glaciation on marine palynomorphs (chitinozoans, acritarchs, leiospheres) in well Nl-2 (NE Algerian Sahara)

Palynomorph assemblages, especially chitinozoans and acritarchs, from the Upper Ordovician of well Nl-2 (north-east of the Algerian Sahara) are studied in order to precisely date the ultimate effects of the Late Ordovician glaciation and to document the impact of this major climatic stress on the diversity of the palynoplankton. References are made to stable isotope excursions and to global eustatic sea level variations in order to improve the local age determination. The Hassi el Hadjar Formation, i.e. glacio-marine diamictites, is interpreted as a transgressive event resulting from the melting of the northern Gondwana ice cap. It yields poorly preserved and moderately diverse chitinozoans of late Hirnantian age. Acritarchs are more abundant in the lower part of these "microconglomeratic clays", but display a low diversity and are badly preserved throughout the whole formation. Reworked individuals are recorded in both groups. The marine sediments of the M'Kratta Formation of latest Hirnantian age contain better preserved, more abundant and more diverse palynomorph assemblages, especially in the Upper Member. The composition of this palynoplankton indicates a fairly good faunal and phytoplankton recovery after the early Hirnantian climatic stress.The extinction of the Ordovician forms, and the appearance of Silurian type taxa occur only in the uppermost Hirnantian, i.e. following with a slight delay the glacial event. The overlying black shales of Wenlock age (lower part of the Oued Mehaiguène Formation) are indicative of marine anoxic environments. They yield a virtually exclusive, but enormous number of Tasmanacea. The very peculiar composition of this palynoplankton seems to be independent of the Late Ordovician glaciation and is most likely related to the factors that, later, generated and maintained anoxic conditions in this area.A new species of chitinozoan, Spinachitina oulebsiri sp. nov. from the latest Hirnantian M'Kratta Formation, is described and illustrated.     

Central Saharan populations of Locusta migratoria cinerascens (Orthoptera: Acrididae) in irrigated perimeters: is it a recent colonisation event?

Locusta migratoria cinerascens became a potential pest in irrigated cereals under pivots, in the Touat region (Algerian Central Sahara), since the indication of the first crowding in June 1994. Morphometrics and bioclimatic approaches revealed the homogeneity between North and South Algerian populations, suggesting a recent colonisation of Central Sahara, aided by the development of large acreage perimeters. We studied the populations of this acridian from 1995 to 2004 and show that they belonged to the solitarious or transiens phase and never reached the gregarious one. The absence of this last phase is discussed in the light of heterogeneous densities observed in the successive generations.     

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.     

Microconchid tubeworms across the upper Frasnian – lower Famennian interval in the Central Devonian Field,Russia

Abstract: Microconchid tubeworms (Tentaculita) encrusting brachiopod shells have been investigated from the upper Frasnian – lower Famennian (Upper Devonian) deposits of the Central Devonian Field, Russia. The condition of microconchids and associated encrusting taxa is reported for the first time from the early Famennian recovery interval (crepida Chron) following the Frasnian–Famennian mass extinction. Two species, one new (Palaeoconchus variabilis sp. nov.) and the second one in open nomenclature (Palaeoconchus sp.), are described. Compared to lower Famennian specimens, they seem to be preferentially grouped on the anterior parts of the brachiopod host shells, which are interpreted as the most suitable sites away from the sea‐bottom and sediment. During the late Frasnian (Late rhenana Chron), microconchids, outnumbered by cornulitids and as abundant as foraminifers, were also associated with trepostome bryozoans, tabulates, rugose corals and various problematic encrusters. During the early Famennian recovery interval encompassing the crepida Chron, microconchids greatly outnumbered all associated encrusters, including the previously dominant cornulitids, while foraminifers, tabulates and rugose corals vanished. Early Famennian microconchids, represented by the single, albeit very abundant, species Palaeoconchus variabilis sp. nov., were opportunists that rapidly colonised the environment during the ongoing transgression following the regression‐driven biotic crisis in the area of the Central Devonian Field. In comparison to their late Frasnian predecessors and even other Middle Devonian specimens, no size reduction (the so‐called Lilliput effect) of early Famennian microconchid tubes was observed. It is probable that microconchids either rapidly attained their ‘normal’ sizes or they did not suffer any dwarfism following the Frasnian–Famennian event.     

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.     

A Wenlock ancestor of the micropoma phylogenetic lineage

A new graptolite species, Monoclimacis praemicropoma , from the Upper Wenlock of Poland is described. It is an ancestor of the Lower Ludlow M. micropoma (Jaeckel). The phytogeny of the praemicropoma-micropoma lineage is discussed. M. micropoma nannopoma (Jaeger) has been assigned to a separate evolutionary lineage. Graptolites, Wenlock, phytogeny .