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.     

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.     

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.     

Acritarch distribution along an inshore–offshore transect in the Gorstian (lower Ludlow) of Gotland, Sweden

To better understand palaeoecological signatures in Palaeozoic acritarch assemblages, the distribution of palynomorphs has been quantitatively studied in eight localities from the Gorstian, lower Ludlow (Late Silurian) of Gotland, Sweden. The localities are situated along an inshore–offshore transect comprising shallow marine lagoonal environments to distal shelf facies. Process-bearing acritarchs and sphaeromorphs constitute the main components within the palynomorph assemblages. The lateral distribution of palynomorphs exhibits characteristic features at three different levels as follows. (1) With regard to the overall composition of the palynomorph assemblages, the abundance of process-bearing acritarchs increases towards the distal shelf, while the abundance of sphaeromorphs decreases. (2) At the generic level, the acritarchs Micrhystridium and Dilatisphaera are more abundant in the proximal facies, while Evittia, Percultisphaera and Oppilatala are more common in distal environments. (3) At an infrageneric level, Micrhystridium morphotypes with shorter processes are mainly present in proximal environments, while those with longer and ramified processes occur in more distal shelf environments. The palynomorph distribution along the inshore–offshore transect highlights the potential of acritarchs and prasinophytes as palaeoenvironmental indicators.     

Cryptospore Assemblages from the Ordovician/silurian Boundary in the Puna Region, North-west Argentina

Palynomorph assemblages have been recovered from deposits believed to straddle the Ordovician/Silurian boundary, from the upper member of the Salar del Rincón Formation, in the Puna region of north-west Argentina. The palynomorph assemblages are dominated by terrestrial cryptospores, but also contain marine elements (acritarchs, prasinophycean algae and chitinozoans). The cryptospore assemblages are similar in composition to those described from coeval deposits worldwide, suggesting that the producers were cosmopolitan and tolerated a wide range of climatic conditions. They are correlated with the Imperfectotriletes spp. Interval Biozone (sub-biozone α) of the Imperfectotriletes spp.- Laevolancis divellomedia Assemblage Biozone, indicating a Hirnantian (latest Ordovician) age. Acritarchs include late Ordovician species such as Eupoikilosusa striata and Villosacapsula cf. setosapellicula , that coexist with the Llandovery species Dactylofusa estillis. Based on lithological and palynological evidence, an early Llandovery, or a late Hirnantian (post-glacial) age is proposed for the basal part of the upper member of the Salar del Rincón Formation.     

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 latest Tremadocian messaoudensis-trifidum acritarch assemblage from the upper part of the Lierneux Member (Salm Group, Stavelot Inlier, Belgium)

The acritarch assemblage occurring in the upper part of the Lierneux Member (Jalhay Formation, Salm Group, Stavelot Inlier, Belgium) has recently been attributed to the messaoudensis-trifidum microflora and dated as latest Tremadocian. It contains most of the characteristic species of the English Lake District sub-assemblage 3 except for the two galeate species Cymatiogalea deunfii and C. messaoudensis. The reinvestigation of all available material has established a very diverse association of over 40 species. It shows a great potential from both stratigraphic and palaeogeographic points of view. A formal Stelliferidium trifidum–Veryhachium lairdii Biozone defined by the appearance of 11 species is proposed. Athabascaella playfordii has been observed in all samples. Previously known only from areas of low to intermediate latitudes, the species is shown for the first time to occur also at higher latitudes. Most of the identified species, including Dactylofusa velifera, Kladothecidium eligmosum and Palaiosphaeridium kamax are illustrated. Some are briefly described. The synonymy between Priscogalea chevronensis and Stelliferidium trifidum is confirmed.     

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.     

High-energy sedimentary events in lagoonal successions of the Upper Silurian of Podolia, Ukraine

Three Late Silurian carbonate profiles of the Malynivtsy and Skala Formations from Podolia (western Ukraine) are discussed in terms of sedimentation dynamics. Their common feature is the appearance of thick, stromatoporoid-rich beds within fine-grained peritidal deposits. These intercalations are composed of fossils typical of offshore sedimentary environments. In the most spectacular case, a channel, several tens of metres wide and infilled with stromatoporoids, is incised in a peritidal cyclic complex. The successions investigated exhibit sedimentary features that are diagnostic of onshore redeposition. Independently of the scale of the recorded sedimentary events, the onshore redeposition was caused by factors with energy levels exceeding those of average storms, probably by hurricanes or even tsunami waves. The dynamic nature of some of the stromatoporoid beds has to be taken into account when constructing the curves illustrating bathymetrical and facies development of the Silurian succession of Podolia. The genesis of lens-shaped stromatoporoid beds, elongated depending on their origin, either parallel or perpendicular to the facies belts, should be considered an important factor in reconstructions of the depositional architecture of sedimentary hydrocarbon collectors.     

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.     

New species of stromatoporoidea from the Upper Ordovician and Silurian of the Subpolar Urals

Stromatoporoidea are widespread in the Upper Ordovician and Silurian beds of the Kozhim River (western slope of the Subpolar Urals). Five new species Cystostroma prodigiosum sp. nov., Stylostroma flabellatum sp. nov., Labechiina arguta sp. nov., Ecclimadictyon faveolatum sp. nov., and Araneosustroma astroplexum sp. nov. are described.     

The early Ludfordian leintwardinensis graptolite Event and the Gorstian–Ludfordian boundary in Bohemia (Silurian,Czech Republic)

The late Gorstian to early Ludfordian hemipelagic succession of the south‐eastern part of the Prague Synform preserves a rich fossil record dominated by 28 species of planktic graptoloids associated with pelagic myodocopid ostracods, pelagic and nektobenthic orthocerid cephalopods, epibyssate bivalves, nektonic phyllocarids, rare dendroid graptolites, brachiopods, crinoids, trilobites, sponges and macroalgae. Faunal dynamics have been studied with particular reference to graptolites. The early Ludfordian leintwardinensis graptolite extinction Event manifests itself as a stepwise turnover of a moderate diversity graptolite fauna rather than an abrupt destruction of a flourishing biota. The simultaneous extinction of the spinose saetograptids Saetograptus clavulus, Saetograptus leintwardinensis and the rare S. sp. B. at the top of the S. leintwardinensis Zone was preceded by a short‐term acme of S. clavulus. Cucullograptus cf. aversus and C. rostratus vanished from the fossil record in the lower part of the Bohemograptus tenuis Biozone. No mass proliferation of Bohemograptus has been observed in the postextinction interval. Limited indigenous speciation gave rise to Pseudomonoclimacis kosoviensis and Pseudomonoclimacis cf. dalejensis. Egregiograptus (the only novel element of cryptic origin) made its earliest occurrence in association with the latest cucullograptids. A postextinction graptolite assemblage of moderate diversity, composed for the most part of long‐ranging taxa, persisted through the middle and upper tenuis Biozone until new rediversification in the Neocucullograptus inexpectatus Biozone. Unlike the graptoloids, the planktic, epiplanktic, nektonic and nektobenthic shelly fauna did not suffer significant extinction in the early Ludfordian. The Gorstian–Ludfordian boundary is placed at the lowest occurrence of S. leintwardinensis in spite of the very modest graptolite record available from the Ludfordian GSSP where no graptolites occur below the basal bed of the Lower Leintwardine Formation. Elsewhere, the Gorstian–Ludfordian boundary has been placed at the base of the Saetograptus linearis Biozone which has been considered roughly correlative with the leintwardinensis Biozone. Indeed, our morphometric study places the worldwide biozonal index species S. linearis in synonymy with S. leintwardinensis and thus considerably enhances the biostratigraphical utility of the latter index species. Pseudomonoclimacis antiqua sp. nov. is described.     

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.     

Biogeographic associations of Silurian bryozoan genera in North America, Baltica and Siberia

Six distinct bryozoan paleobiogeographical provinces existed during the Silurian of North America, Baltica and Siberia. These provinces are based on parsimony analysis of endemicity (PAE), each supported by a number of endemic genera only present in that assemblage. The area cladogram displays a nested pattern, permitting the clustering of assemblages into larger units that might be indicative of their areas of origin. Paired-group cluster analysis supports the existence of provincial assemblages. Relationships indicate the action of both vicariance and geodispersal. Geographic boundaries between the provinces include land, deep sea, and climate zone boundaries, including barriers created by the Iapetus Ocean and the Tornquist Sea.     

Early Silurian (Telychian) rugose coral fauna of Daguan area, northeast Yunnan province, China

Study on rugose coral fauna of the Sifengya Formation (early Telychian) and Daluzhai Formation (mid-late Telychian) in Daguan area, northeast Yunnan Province, China was carried out. Rugose coral fauna of the Sifengya Formation included 18 genera and 34 species, while Daluzhai Formation with nine genera, ten species. We described rugose coral fauna (12 genera, 19 species) including one new genus and five new species, i.e. Protoketophyllum daguanense gen. et sp. nov., Crassilasma huanggexiense sp. nov., Pseudophaulactis heae sp. nov., P. convolutus sp. nov., and Shensiphyllum minor sp. nov. The characteristics and geological significance of rugose coral fauna of Sifengya Formation and Daluzhai Formation were analyzed. Particularly, rugose coral fauna of the Sifengya Formation represent early Telychian rugosan fauna in the Upper Yangtze region and improve the sequences of early Silurian (Llandovery) rugose coral assemblages in Yangtze region. It is therefore very meaningful to further analyze radiation period of rugose coral fauna in such epoch. __________ Translated from Acta Palaeontologica Sinica, 2005, 44(2): 229–246 [译自: 古生物学报, 2005, 44(2): 229–246]     

Early Silurian(Telychian)rugose coral fauna of Daguan area, northeast Yunnan Province,China

Study on rugose coral fauna of the Sifengya Formation(early Telyehian)and Daluzhai Formation (mid-late Telychian)in Daguan area,northeast Yunnan Province,China was carded out.Rugose coral fauna of the Sifengya Formation included 18 genera and 34 species,while Daluzhai Formation with nine genera,ten species.We described rugose coral fauna(12 genera,19 species)including one new genus and five new species,i.e.Protoketophyllum daguanense gen.et sp.nov..Crassilasma huanggexiense sp.nov.,Pseudophaulactis heae sp.nov.,P.convolutus sp.nov.,and Shensiphyllum minor sp.nov..The characteristics and geological significance of rugose coral fauna of Sifengya Formation and Daluzhai Formation were analyzed.Particularly,mgose coral fauna of the Sifengya Formation represent early Telychian rugosan fauna in the Upper Yangtze region and improve the sequences of early Silurian(Llandovery)mgose coral assemblages in Yangtze region.It is therefore very meaningful to further analyze radiation period of rugose coral fauna in such epoch.     

Enantiomorphous dimorphism in Silurian and Devonian bivalves; Maminka Barrande, 1881 (Lunulacardiidae, Silurian) - the oldest known example

The genus Maminka Barrande, 1881 (Lunulacardiidae) from the Wenlock and Ludlow (Silurian, Lower Paleozoic) of Europe is the oldest known bivalve with enantiomorphous dimorphism which is common among the Silurian and Devonian Lunulacardiidae and Antipleuridae. It represents the result of a special adaptation to the reclining mode of life combined with byssal attachment when early in ontogeny either the left or right valve became the lower valve; later, the bivalve develops as two enantiomorphous sets of shells. Distribution, ontogeny and evolution of this interesting feature in the genera Maminka Barrande, 1881, Stolidotus Hede, 1915, Mila Barrande, 1881, Dualina Barrande, 1881, Antipleura Barrande, 1881, Silurina Barrande, 1881 and Hercynella Kayser, 1878 are discussed.     

Regular mode of increase, and constrained but variable growth, in the Silurian rugose coral Stauria favosa

Stauria favosa exhibits a typical pattern of axial division (tripartite or quadripartite). The four protosepta first appear with a definite polarity in offset corallites, and metasepta are inserted serially, following Kunth's rule. There are no variations in essential modes of asexual reproduction. However, at the corallum periphery, up to four offset corallites appear to have been derived by discrete offsetting within the parent calices. Detailed observations of vertical morphological changes, however, indicate that this apparently unique mode of increase is derived from axial division at the sites of the four protosepta, as usual in this species. Just after the initiation of division, the contraction of polyps occurred in response to somewhat accidental, deteriorating habitat conditions. The combination of asexually divided corallites and rejuvenescence resulted in phaceloid daughter corallites, which subsequently underwent partial mortality at the corallum periphery. Stauria favosa always exhibits not only regular modes of asexual reproduction and septal insertion under the strict control of phylogenetic and developmental constraints, but also variable growth forms of corallites and coralla. This variability, though within limited options, is palaeoecologically significant for successfully coping with unpredictable habitats.     

Late Silurian trilobites from the Nuratau and Turkestan ranges, Uzbekistan and Tajikistan

Silurian trilobites of Ludlow and P?ídolí age are described from sections in southern Uzbekistan and from adjacent parts of Tajikistan. They belong to 22 species (10 named and 12 under open nomenclature) distributed among 13 genera belonging to the families Proetidae, Tropidocoryphidae, Aulacopleuridae, Scharyiidae, Cheiruridae, Encrinuridae and Odontopleuridae. The following new species are described: Interproetus pentaxus, Paleodechenella turkestanica, P. zaaminicus, Cromus tamchii and Leonaspis nuratensis. The generic associations from the mid Ludlow bear a striking resemblance to coeval ones from Bohemia, and include genera otherwise known only from that region or from adjacent parts of central Europe. The trilobite faunas from the late Ludlow and P?ídolí series are much less diverse and are dominated by genera of the Warburgellinae (Tropidocoryphidae), which include one species common to Vaigatch Island in the Russian Arctic, and Podolia, Ukraine. The closest links of the diverse central Asian mid Ludlow faunas are therefore to central Europe, whilst the sparser younger Ludlow and P?ídolí faunas, such as they are, suggest closer links to Baltica.