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.     

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.     

Bivalve ecology in a Silurian shelf environment

The Ludlow Series of the Welsh Borderland represents a gradient from quiet. deep shelf muds to shallow, storm-deposited silts. A fauna of 44 bivalve species shows a pattern of overlapping environmental ranges which span the full extent of the shelf gradient. Bivalve abundance is highest at the extremes of this gradient, where endobyssate forms are common in storm-deposited silts, and nuculoids locally dominate the fauna of the deep shelf muds. Species diversity of bivalves, however, is highest in low-stress. mid-shelf environments, where bivalves of small population sizes comprise a minimal proportion of the benthic fauna. Niche sizes, species diversity. and total environmental range of the bivalves is comparable to that of Ludlow brachiopods, even though the latter group is far more abundant as individuals.     

The Silurian of the Mehaigne Valley (Brabant Massif,Belgium): Biostratigraphy (Chitinozoa)

The Chitinozoa were studied in a 2100 m thick Silurian sequence of flysch-type sediments in the Mehaigne area (Brabant Massif, Belgium). Lower and middle Wenlock graptolites occur only in the middle part of the sequence. Moderately to poorly preserved Chitinozoa from 56 samples belong to 10 genera and 35 species and can be used to establish 5 assemblage zones of which some are divided in subzones. Comparison with Sweden and Great Britain permits these zones to be dated as spanning the Late Ordovician and the middle Llandovery to early or middle Ludlow. Probably the Chitinozoa were deposited originally in the outer deeper shelf and carried off shore by turbidity currents into the Mehaigne area.     

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 appearance of Baragwanathia (Lycophytina) in the Silurian

The evidence mainly involving graptolites is restated which supports an Upper Silurian (Ludlow) dating for a vascular flora containing Baragwanathia (Lycophytina) in Victoria, Australia.     

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 environmental significance of Lingula in the Ludlow Series of the Welsh Borderland and Wales

Cherns. Lesley 1979 01 15: The environmental significance of Lingula in the Ludlow Series of the Welsh Borderland and Wales. Lethaia . Vol. 12. pp. 35–46, Oslo. ISSN 0024–1164.
The inarticulate brachiopod Lingula is commonly regarded as an indicator of very nearshore environments, particularly when found in monospecific assemblages. However, neither of the two common species in the Lower Leintwardine Beds, L. lewisii and L. lata . which occur commonly in such associations, have distributions indicative of such an environment. Their Occurrence correlates closely with the Leintwardinian palaeogeography. The former species is concentrated in shelf areas, the latter is confined to the basin; the two distributions overlap little. Individuals of both species are found articulated and perpendicular to the bedding, apparently in situ . Hence, the species may be inferred to have equivalent, infaunal modes of life comparable to living representatives of the family. The basinal distribution of L. lara is an original feature. The depth of the actively subsiding basin was relatively great, judging from sedimentological and palaeogeographical evidence. Therefore, L. lata was a species adapted to 'deep-water' environments. The lingulid distributions, reviewed against the concept of brachiopod depth-related communities previously defined for the Ludlow show the latter to be inapplicable in the Leintwardinian. The depth significance postulated for the recurrent species associations typical of the shelf sequences cannot be accepted.     

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.     

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.     

Stable carbon isotope development and sea-level changes during the Late Ludlow (Silurian) of the Łysogóry region (Rzepin section,Holy Cross Mountains,Poland)

The Ludlow deposits of the Winnica Formation in the Rzepin section (Holy Cross Mountains, Poland) have been studied with respect to their facies evolution and stable carbon isotope ratios from whole-rock samples. The C-isotope curve of the Rzepin section records a distinct positive excursion with maximal values of +8.9‰. A Late Ludlow positive isotope excursion is known from different paleocontinents and thus is regarded as a global isotope event. The presence of the event allows for a chemostratigraphic correlation of the Rzepin profile with the classical, biostratigraphically well-dated Gotland section. The Ludlow deposits of the Holy Cross Mountains are interpreted in terms of sea-level changes during the isotope excursion. Because the Rzepin and Gotland sections are positioned on the opposite sides of the same foreland basin, a comparison of their sequence stratigraphy allows to test the regularities of the sea-level changes on the shelf of Baltica. In the present paper, a modified view of the recently published sequence stratigraphy of the Gotland succession is presented. Our results indicate that the positive Ludlow δ¹³C excursion is connected with prolonged low-stand conditions with small, internal transgressive pulses.     

Latitudinal and depth zonation of early Ordovician graptolites

Early Ordovician graptolites are inferred to have lived in two primary depth zones of the ocean waters. A shallow, epipelagic zone biotope contained species found preserved in all depth facies, whereas a deep, mesopelagic, zone biotope contained species now found only in deep-water facies (isograptid biofacies). An inshore biotope contained a few species that were apparently restricted to shelf-platform waters and which are now found in shelf sediments, along with members of the epipelagic biotope (didymograptid biofacies). The effects of surface water temperature on graptolite distribution, as measured by differences between high palaeolatitude (Europe/S. Britain - 'Atlantic province') and low palaeolatitude (Australasia/N. America-'Pacific province') regions, was less marked than that of depth. A survey of over 80 species and species groups shows that a surprisingly large number of graptolites were pandemic and eurythermic. In the early Ordovician, individual species were more widespread than those of any other invertebrate group, hence their great value for long-distance correlation. Biotopes, early Ordovician, depth facies, graptolite provinces, palaeolatitude.     

The earliest vascular land plants: continuing the search for proof

New collections from Bringewoodian strata (Silurian, Ludlow Series, Saetograptus leintwardinensis incipiens Zone) in Wales have yielded the oldest known specimens of the plant Cooksonia . From evidence at younger (Eř;éolí/Downton) horizons this genus is considered to be a vascular plant; although no tracheids have been isolated from the Bringewoodian specimens, on the basis of comparative morphology they are concluded to be the oldest recorded tracheophytes. The types of evidence relevant to the identification of vascular plants are briefly reviewed, and the relationships of the Welsh Cooksonia flora to other late Silurian floras are discussed. Confident stratigraphical dating is a prerequisite for any discussion of early plant evolution, emphasising the desirability of an integrated approach incorporating both geologial and palaeobotanical investigations. The dating of two published records of diverse Silurian floras, including lycopods, is questioned; recent dating of the Australian Baragwanathia flora as early Ludlow on the basis of graptolites is not supported by the shelly faunal evidence, which indicates a correlation within the Lower Devonian.     

Morphologically complex plant macrofossils from the Late Silurian of Arctic Canada

In addition to vegetative remains, fertile remains from ten plants, representing seven distinct taxa whose size and complexity are much greater than most contemporaneous fossils, are reported from late Ludlow (Ludfordian) sediments of Bathurst Island in Nunavut, Canada. Evidence for the age of these beds is gathered from stratigraphic relationships and index fossils including conodonts, graptolites, and brachiopods. Zosterophylls dominate the collection, some of which constitute the earliest record of fertile structures arranged in dense clusters and longitudinal rows along axes. Representatives include a plant that resembles Bathurstia, one species of Zosterophyllum, and two specimens that bear affinity to this genus. Distichophytum is also represented, as is a new zosterophyll named Macivera gracilis. The prevalence of sporangial clustering and reduced sporangial stalks in this flora leads to a discussion of the origins and significance of these morphological features. Following a review of some of the other Silurian floras, particularly the Baragwanathia-bearing Lower Plant Assemblage of Victoria, Australia, which also shows morphological advancement over the rhyniophytoid-dominated floras common to Laurussia, it is concluded that the Bathurst Island flora presents the best evidence to date of substantial morphological diversity, complexity, and stature of vascular land plants in this period.     

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     

Llandovery sporomorphs and graptolites from the Manbo Formation,the Mojiang County,Yunnan, China

Llandovery sporomorphs and graptolites have been recovered from the Manbo Formation, Mojiang area, western Yunnan, China, which belonged to the Indo-China Palaeoplate in the Palaeozoic. The graptolite fauna, including 8 genera and 12 species, is considered Mid Telychian, Llandovery in geological age, confirming the existence of Llandovery rocks in this region. The Silurian stratigraphical sequence of this area is reconsidered as in the ascending order: Manbo Formation (Llandovery–early Wenlock), Shuiqing Formation (late Wenlock–Ludlow) and unnamed formation (Pridoli?). The sporomorph assemblage from the Manbo Formation includes seven species in five genera. The dominant members of this assemblage are Tetrahedraletes medinensis and Laevolancis chibrikovae. Based on the low diversity and low abundance, the geological age of this assemblage is considered to be approximately the Telychian, Llandovery. The parent plants of sporomorphs probably inhabited the landmass near Mojiang area, these sporomorph-producing plants probably included bryophyte-like land plants and primitive vascular land plants. The sporomorph evidence shows that the South China and Indo-China palaeoplates may have been in close proximity (maybe with some continental bridges linking them) to each other at least in the Llandovery. They were also closely related with Gondwanaland in the Llandovery.     

Biodiversity patterns of Early–Middle Ordovician marine microphytoplankton in South China

Based on new materials from six sections and all available literature data, new diversity curves are presented for the phytoplankton (acritarchs) from South China, covering the Early–Middle Ordovician interval, when the Great Ordovician Biodiversification Event took place. The total diversity curve and the origination data imply that a major radiation of the phytoplankton occurred during the analysed interval. A peak of the total acritarch diversity curve appears in the A. suecicus graptolite biozone. The diversity changes vary in the different parts of the investigated area, most probably depending on the position of the analysed sections on the carbonate shelf or the slope, reflecting diversity differences due to the position on an inshore–offshore transect.The Early–Middle Ordovician diversity pattern of the phytoplankton is compared with those of several marine invertebrate groups. Compared with the diversity curve peak of the acritarchs, the conodonts and brachiopods reached their highest diversities before the acritarchs, while the highest diversity of the chitinozoans appears slightly later. The graptolites show two peaks during the Early–Middle Ordovician, while the trilobites diversity curve shows a peak only in the Sandbian. The different fossil groups, such as chitinozoans, conodonts, graptolites, brachiopods and trilobites show therefore different evolutionary patterns to that of the acritarchs, that are not yet fully understood, and correlations are so far difficult.The acritarch diversity changes can partly be compared to the local sea-level changes from four sections in South China. At a larger scale, the acritarch radiation coincides with a general transgression. At a regional or local scale, correlations are not straightforward, pointing out that more detailed data, based on both acritarch studies and more precise sea-level investigations, are necessary.     

Fueling Plankton Production by a Meandering Frontal Jet: A Case Study for the Alboran Sea (Western Mediterranean)

A three dimensional biophysical model was employed to illustrate the biological impacts of a meandering frontal jet, in terms of efficiency and persistency of the autotrophic frontal production, in marginal and semi-enclosed seas. We used the Alboran Sea of the Western Mediterranean as a case study. Here, a frontal jet with a width of 15–20 km, characterized by the relatively low density Atlantic water mass, flows eastward within the upper 100 m as a marked meandering current around the western and the eastern anticyclonic gyres prior to its attachment to the North African shelf/slope topography of the Algerian basin. Its inherent nonlinearity leads to the development of a strong ageostrophic cross-frontal circulation that supplies nutrients into the nutrient-starved euphotic layer and stimulates phytoplankton growth along the jet. Biological production is larger in the western part of the basin and decreases eastwards with the gradual weakening of the jet. The higher production at the subsurface levels suggests that the Alboran Sea is likely more productive than predicted by the satellite chlorophyll data. The Mediterranean water mass away from the jet and the interiors of the western and eastern anticyclonic gyres remain unproductive.     

Late Permian marine ecosystem collapse began in deeper waters: evidence from brachiopod diversity and body size changes

Analysis of Permian–Triassic brachiopod diversity and body size changes from different water depths spanning the continental shelf to basinal facies in South China provides insights into the process of environmental deterioration. Comparison of the temporal changes of brachiopod diversity between deepwater and shallow‐water facies demonstrates that deepwater brachiopods disappeared earlier than shallow‐water brachiopods. This indicates that high environmental stress commenced first in deepwater settings and later extended to shallow waters. This environmental stress is attributed to major volcanic eruptions, which first led to formation of a stratified ocean and a chemocline in the outer shelf and deeper water environments, causing the disappearance of deep marine benthos including brachiopods. The chemocline then rapidly migrated upward and extended to shallow waters, causing widespread mass extinction of shallow marine benthos. We predict that the spatial and temporal patterns of earlier onset of disappearance/extinction and ecological crisis in deeper water ecosystems will be recorded during other episodes of rapid global warming.     

Trace fossil associations related to facies of an upper Ordovician low wave energy shoreface and shelf, Oslo-Asker district, Norway

Three trace fossil associations have been identified from facies interpreted as low wave energy upper, middle and lower shoreface deposits, transitional shoreface/shelf deposits and open epicontinental shelf deposits. These were developed in pre-regressive, regressive and transgressive phases associated with eustatic sea level changes caused by the upper Ordovician glaciation. The three associations are the Skolithos-Diplocraterion association, the Diplichnites-Phycodes association and the Thalassinoides association. The Skolithos-Diplocraterion association inhabited upper, middle and lower shoreface environments, the Diplichnites-Phycodes association a lowermost shoreface environment and the Thalassinoides association inhabited offshore epicontinental shelf environmcnts. The Skolithos-Diplocraferion and Diplichnites-Phycodes associations bear strong affinities with previously described Skolithos ichnofacies assemblages (indicating shoreline environments) and the Cruziana ichnofacies forms (indicating shallow water environments down to wave base) respectively. The Thalassinoides assemblage, however, closely compares with assemblages described from Mesozoic epicontinental shelf sequences. Trace fossil diversity achieves a maximum in the lowermost shoreface environment and reduces in deeper water and shallower water environments. This accords with diversity patterns identified in analogous Jurassic epicontinental palaeonvironments. The stratigraphic distribution of some trace fossil association boundaries accords with those of contemporary body fossil associations. However, several body fossil associations are included within the volume of distribution of a single trace fossil assemblage. □ Trace fossil association, shoreface and shelf, Oslo district, upper Ordovician.