A new species of the conodont genus <Emphasis Type="Italic">Polygnathus</Emphasis> from the Tournaisian of the northern Urals,Chernyshev Ridge and Pai-Khoi

A new conodont species, Polygnathus postvogesi sp. nov., from the Tournaisian deepwater deposits of the northern Urals, southeastern part of the Chernyshev Ridge, and central Pai-Khoi is described. The new species continue the phylogenetic lineage of the Late Devonian polygnathid P. marginvolutus Gedik, P. vogesi Ziegler.     

New Middle Devonian conodont data from the Dong Van area,NE Vietnam (South China Terrane)

Middle Devonian conodonts from the Si Phai section in NE Vietnam are described. The section ranges from the Middle Devonian ensensis to timorensis conodont zones to the Late Devonian rhomboidea conodont Zone. A rich overall assemblage is described, including 27 taxa of species or subspecies rank and 11 taxa described in an open nomenclature. Among the dominant Polygnathus forms, four new taxa are described: Polygnathus linguiformis saharicus subsp. nov., Polygnathus linguiformis vietnamicus subsp. nov., Polygnathus rhenanus siphai subsp. nov., and Polygnathus xylus bacbo subsp. nov. Conodont assemblages are attributed to polygnathid, polygnathid-klapperinid, and klapperinid conodont biofacies representing hemipelagic to pelagic environments. The klapperinid biofacies, unreported in the previous literature, are here attributed to offshore areas of the external shelf. The taxonomic compositions of the studied conodont assemblages, as well as their CAI characteristics (CAI 4–5), suggest a palaeogeographic affinity of the studied strata to the Chinese Devonian Guangxi Basin, and the South China Terrane in general. Furthermore, the conodont biofacies and the palaeogeographic distribution of the fauna are discussed.     

Frasnian conodonts from the eastern Russian Platform

The Frasnian conodonts of the eastern Russian Platform from the Timan in the north to the Orenburg Region in the south are characterized in detail both paleontologically and stratigraphically. This paper is a continuation of a previously published monograph (Ovnatanova and Kononova, 2001) on the Frasnian conodonts of the shallow-water strata of the central Russian Platform, where polygnathids prevail. The conodont assemblages from the shallow-water and basinal sections of the Volga-Ural province and Southern Timan are analyzed. Problems and difficulties associated with the correlation of the shallow-water and deep-water Frasnian sections of the eastern Russian Platform and their correlation with the existing zonal conodont scales are discussed. The correlation between the deep-water Mendym deposits and the shallow-water Rechitza and Voronezh strata of the Volga-Ural province with the Vetlasyan and Sirachoi deposits of the Southern Timan is substantiated. The stratotype of the Semiluki Regional Stage of the Central Devonian Field contains equivalents of the lowermost beds of the Domanik Formation (unit 1), based entirely on polygnathids. Phylogenetic reconstructions for Palmatolepis and Polygnathus are suggested based on the ontogenetic series for some species of these two genera and the presence of transitional forms between some of the species. Based on these phylogenetic reconstructions, conodont zonal scales for the shallow-water and deep-water sections of the Frasnian Stage of the Russian Platform are proposed and their correlation with the existing conodont zonal scales is also adduced. Some aspects of biofacies control are considered based on the distribution of conodonts in the sections studied. In Systematic Paleontology, 91 conodont species of the genera Ancyrodella, Mesotaxis, Palmatolepis, and Polygnathus are described, including the new species Palmatolepis menneri, P. kaledai, P. acutangularis, and Polygnathus reitlingerae. DEDICATION To the blessed memory of the outstanding geologist Vladimir Vladimirovich Menner     

Integrated radiolarian and conodont biostratigraphy of the Middle Permian Gufeng Formation (South China)

Radiolarians are usually abundant in chert sequences and they have thus been widely used for the biostratigraphy of deep-water sediments. However, there are many difficulties in the correlation of radiolarian biostratigraphic schemes with the standard conodont zones. In this study, 21 radiolarian species were extracted from the Gufeng Formation that crops out in the Luojiaba (LJB) section (western Hubei, China), together with 5 co-occurring conodont species. In this way, it is the first time that the Pseudoalbaillella globosa, Follicucullus monacanthus and F. scholasticus radiolarian zones can be directly correlated with the Jinogondolella nankingensis gracilis, J. aserrata and J. postserrata conodont zones. Accordingly, the 3 radiolarians zones are now firmly correlated with the Roadian to middle Capitanian interval (Middle Permian).     

Famennian conodonts from the Hongguleleng Formation at the Bulongguoer stratotype section,western Junggar,Northwest China

The Hongguleleng Formation, the highest and most important Devonian marine carbonate horizon in western Junggar, contains an endemic shallow-water Icriodus–Polygnathus conodont fauna with rare palmatolepids and other genera. The conodont faunas from the Bulongguoer and the Wulankeshun sections are similar, with high abundance (about 40%) of endemic taxa (13 taxa), indicating isolation of the Junggar Basin during early Famennian. Non-endemic species in the faunas suggest that the Lower Member of the Hongguleleng Formation is assignable to the Pa. rhomboidea Zone to the Pa. marginifera marginifera Zone of early Famennian, not including the Frasnian–Famennian boundary. The Upper Member may be of late Famennian–early Tournaisian in age on the basis of our preliminary faunal analysis.     

New Conodont Species from the Famennian (Upper Devonian) of the Urals

New conodont species of the genus Polygnathus from the Upper Famennian of the Urals and Cis-Urals are described. Po. tsygankoi sp. nov. is distributed from the expansa‑praesulcata conodont zones. This species occupies an intermediate position in the phylogenetic lineage Polygnathus atasuensis–Po. tsygankoi sp. nov.–Po. lobatus sensu Kononova. Po. serriformis sp. nov. is known from the praesulcata Zone. Evolutionary relationships of Po. serriformis sp. nov. are unknown.     

A new Devonian species of the enigmatic Carboniferous conodont Dollymae

Arrow-shaped P₁ elements of Dollymae are characteristic for the Tournaisian conodont faunas worldwide but the phylogeny of this conodont remains obscure and its origin is cryptic. Dollymae peregrina n. sp., from the upper Famennian (upper expansa Zone) strata in the Holy Cross Mountains (central Poland), is the oldest representative of the Dollymae lineage. The morphology of the new species’ P₁ element shows primitive features in comparison to other members of this genus and is transitional in the number of element processes. Thus D. peregrina n. sp. is most probably ancestral to the remaining species. The genus appears to be monophyletic, probably rooted in Pelekysgnathus within the Icriodontidae.     

新疆石炭系杜内-韦宪阶界线层的牙形类POLYGNATHUS COMMUNIS GANCAOHUENSIS (新亚种)POLYGNA-THUS COMMUNIS BRANSON ET MEHL 1934的种系发生

本文提出和描述了产自我国西北新疆南部喀拉塔克剖面的一个牙形类新亚种Polygnathus communis gan-caohuensis,并根据以前出版的信息,命名了另一个新亚种P0lygnathus communis carmanae.种系发生研究揭示了Polygnathus communis Branson et Mehl的亚种对于法门阶最上部至韦宪阶地层的划分是至关重要的.此种的cari-na→gancaohuensis谱系,连同发现于喀拉塔克剖面的菊石、有孔虫和腕足动物,被认为是划分杜内-韦宪阶界线的潜在的标志化石.因此,喀拉塔克剖面有可能成为全球杜内-韦宪阶界线的一个候选层型剖面.     

Morphogenetic patterns of the shelf-like protuberance in the late Emsian-early Eifelian polygnathid conodonts

The shelf-like protuberance on the outer side of the pit in polygnathid conodonts is a unique and easily recognizable feature usually viewed as taxonomically significant for the biostratigraphically important species Polygnathus serotinus. However, the protuberance is also developed in other late Emsian-early Eifelian polygnathids, which mainly include P. praeinversus, P. vigirei, P. pseudoserotinus and P. linguiformis bultyncki. Although several evolutionary lineages have been proposed to elucidate the phylogenesis of the protuberance, its morphogenesis has been rarely studied. In this study, intraspecific variation of P. linguiformis bultyncki is investigated on the basis of abundant specimens from South China and Germany, and two morphotypes are recognized. Moreover, three morphogenetic patterns of the shelf-like protuberance are proposed. Polygnathus praeinversus and P. pseudoserotinus are assigned to the first morphogenetic pattern, with the protuberance formed as a vestigial outer flank of the asymmetrical basal cavity. The second pattern, which can be applied to P. serotinus and P. vigirei, is characterized by a protuberance that is formed as a shelf-like extension of the pit partially buttressed by platform material in juvenile/immature specimens, and by a well-developed shelf-like extension of the pit on its inner side and a bulge in the platform proper on its outer side in mature specimens. Polygnathus linguiformis bultyncki is characterized by the third morphogenetic pattern. Its protuberance is ontogenetically acquired and though absent in juvenile specimens, in immature and mature specimens it is formed as a bulge in the platform proper fully supported by platform material. This study provides an exemplar for conodont research of other geological periods as growth histology can play an important role in investigating morphological variation of conodonts.     

A new study of Olenekian–Anisian boundary conodont biostratigraphy of the Tulong section in Himalaya Terrane,southern Tibet

The Himalaya Terrane of southern Tibet exposes successive shallow-marine carbonate deposits from the Lower to Upper Triassic, and is a key region for studying the Triassic conodont biostratigraphy at the northern margin of the Indian Plate. On the basis of newly collected samples from the Kangshare and Laibuxi formations at the Tulong section, 11 conodont species of 7 genera were identified, and four conodont zones were established, namely, the Novispathodus abruptus Zone (lower Spathian, first reported in Tibet), the Columbitella jubata Zone (middle Spathian), the Triassospathodus symmetricus Zone (upper Spathian), and the Chiosella timorensis Zone (lowermost Anisian) in ascending order. The first occurrence (FO) of Chiosella timorensis indicates the Olenekian–Anisian boundary (OAB) at Bed 25, upper part of the Kangshare Formation. The regional and global correlation of these conodont zones is synthesized.     

Emsian (Early Devonian) Yujiang Event in South China

The Yujiang Event is reported here as a newly recognized event that occurred during the deposition of the Early Devonian Yukiang Formation in Liujing, Guangxi, and its equivalents in South China. The first episode of the Yujiang Event is characterized by the extinction of biostrome that is constructed mainly by colonies of rugose coral, tabulatimorph coral and bryozoan as well as the extinction of evolved biota including some brachiopods, bivalves, etc. The age of the first episode of the Yujiang Event dated by conodonts is at the beginning or the lower part of the Polygnathus nothoperbonus Zone. The main episode of Yujiang Event is evidenced by the extinction of the Rostrospirifer tonkinensis Fauna (sensu lato) at the top of Yukiang Formation and by the sudden occurrences of the dolostones and dolomitic limestone of the overlying Moding Formation. At the basal part of the Moding Formation, the new finding of conodont Polygnathus nothoperbonus and dacryconarids Nowakia (N.) barrandei indicates that this episode of Yujiang Event may occur in the upper part of the Polygnathus nothoperbonus Zone. Overlying the Yukiang Formation and its equivalent beds, various lithic deposits correspondingly yielding different biota can be seen widely in many localities and sections in South China and northern Vietnam. Therefore, the Yujiang Event can be recognized not only as a biotic extinction event, but also as a geological event.     

Integrated biostratigraphy and carbon isotopes from the Cambrian Tangwangzhai section,North China

The Tangwangzhai section, western Shandong Province, North China, the type section for the Cambrian Kushan and Chaomitian formations, yielded a diverse and relatively well-preserved conodont fauna, in which we recognize the Westergaardodina orygma, Westergaardodina matsushitai, Muellerodus? erectus, and Westergaardodina aff. fossa–Prooneotodus rotundatus zones of the North China conodont zonation. The Tangwangzhai conodont succession can be correlated not only with the polymerid trilobites occurring in the section but also with the conodont zones established for South China. The first occurrence of Furnishina longibasis and Furnishina quadrata in the upper part of the Westergaardodina matsushitai Zone allows the recognition of the base of the Paibian Stage and Furongian Series in the upper part of the Kushan Formation. The base of the Jiangshanian Stage, in the uppermost Muellerodus? erectus Zone, can be recognized by the presence of Westergaardodina cf. calix close to the base of the Chaomitian Formation. Chemostratigraphic analyses of the Tangwangzhai section show the onset of a positive carbon isotope excursion, referred to the SPICE event, in the upper part of the Kushan Formation at a level corresponding to the first occurrence of F. longibasis and F. quadrata. The base of the Jiangshanian Stage in the section is close to the demise of the SPICE positive excursion.     

Brachiopods and other fossils from the Permo-Triassic boundary beds of the Antalya Nappes (SW Taurus, Turkey)

An uppermost Permian-Lower Triassic biota of brachiopods, conodonts, algae and foraminifers from the Pamucak and Kokarkuyu formations at Çürük Da? (Antalya, Turkey) is here described. The brachiopods belong to two different assemblages: a lower assemblage, early Wuchiapingian in age, with Spinomarginifera cf. S. helica, Spinomarginifera cf. S. iranica, Alatorthotetina sp. ind., Orthothetina sp. ind., Ombonia antalyensis nov. sp. and few specimens of Pennospiriferinoidea; an upper assemblage, Changhsingian in age, comprising S. cf. S. iranica, Spinomarginifera cf. S. spinosocostata, Spinomarginifera sp. ind. and Orthothetina sp. ind., characteristic taxa of the low diversity survival brachiopod faunas of latest Permian age (Survival Fauna 1). The occurrence of the conodont Hindeodus cf. praeparvus above the brachiopod fauna confirms its Changhsingian age. The oolitic grainstones at the top of the Pamucak Formation contain Permocalculus sp., Macroporella cf. apachena, species of Hemigordius and Palaeozoic Lagenida. Coarse calcite fibrous cements pervade the oo-bioclastic grainstones, suggesting early marine cementation. The base of the Kokarkuyu Formation is characterized by the disaster forms Earlandia amplimuralis and “Cornuspira” mahajeri, gastropods and ostracods. The conodont Isarcicella lobata has been recovered 31 m above the base of the Kokarkuyu Formation, indicating the occurrence of the second Triassic conodont zone above the parvus biozone and below the staeschei biozone. The faunal content at the transition of the Pamucak and Kokarkuyu formations records the biotic survival in the aftermath of the end-Permian extinction. Facies evolution from lower energy inner platform wackestones and packstones to higher energy open platform oolitic grainstones indicates a transgression at the top of the Pamucak Formation, which continues into the Lower Triassic Kokarkuyu Formation.     

Upper Devonian conodonts of northeastern European Russia

The conodont succession in the Upper Devonian facially different sections of northeastern European Russia (Chernyshev Ridge, Subpolar and Polar Urals, Pai-Khoi) are studied. The sections are subdivided and correlated taking into account the global standards for stages. The Frasnian Montagne Noire zonation (Klapper, 1989) is used for the first time for northeastern European Russia; the authors correlated this zonation with the scale of Ziegler and Sandberg (1990). Famennian deposits are subdivided using scale of Ziegler and Sandberg (1984) and miospore zonation. The age of local geological units (formations) is specified; the formations of different regions of northeastern European Russia are correlated to each other and to the regional subdivisions of EEP and Southern Urals. A total of 93 species of Ancyrodella, Mesotaxis, Palmatolepis, Polygnathus, and Zieglerina are described. Two new species Ctenopolygnathus parallelus sp. nov. and Polygnathus masonae sp. nov. are distinguished and five species are described in the open nomenclature. Special chapter is devoted to the phylomorphogenesis of Ancyrodella.     

Revision of Famennian-Tournaisian (Late Devonian – Early Carboniferous) conodont biostratigraphy of Sardinia,Italy

Conodont biostratigraphy of the Famennian and Tournaisian of Sardinia is provided on the basis of all data available in the literature. More than 130 conodont taxa documented from this time-interval allow recognition of nineteen conodont biozones. The zonation scheme adopted here is broadly similar to the standard zonation schemes used in the Upper Devonian and Lower Carboniferous, though a few variations are needed to better accommodate the species occurrences in Sardinia: the velifer Zone and the styriacus Zone have been reintroduced, even though with slightly different meanings than previously used; the Lower praesulcata Zone is expanded to include the Middle praesulcata interval. The Upper praesulcata Zone and the sulcata Zone are missing. All zones are described and discussed and a complete list of bioevents is provided.     

Uppermost Devonian–Lower Carboniferous conodonts from the Indert Formation in the Shine Jinst area,South Mongolia

In southwestern Mongolia, conodonts from the Indert Formation at its type locality in Shine Jinst were investigated to improve regional and global correlation of the uppermost Devonian–lowermost Carboniferous. The abundance and diversity of the conodont fauna is quite low in general, whereas representatives of Siphonodella, Protognathodus, Pseudopolygnathus, Polygnathus and Bispathodus have been collected from the Indert-Uul section. Five conodont biointervals, from the Bi. costatus-Pr. kockeli Interregnum (ckI) to the Siphonodella sandbergi Zone, could be recognized from the top Heermorit Member and the lower part of the Shombon Member, while the middle and upper parts of the Shombon Member cannot be zoned based on the conodonts. Absence of index fossils Si. sulcata, Pr. kuehni and Pr. kockeli hampered precise recognition of the Devonian–Carboniferous boundary (DCB) at Indert-Uul, but this transition is well delineated by typical uppermost Famennian and Tournaisian conodont assemblages. The DCB is tentatively placed at the base of the Shombon Member, consistent with the lithological boundary and the resuming of carbonate production during the DCB transitional interval.     

Sedimentary and biofacies records in calciturbidites at the Devonian-Carboniferous boundary in Moravia (Moravian-Silesian Zone, Middle Europe)

Summary At the Devonian/Carboniferous boundary, major climatic and oceanographic changes influenced sedimentation on carbonate platforms and in peri-platfrom asreas. Three deep-water carbonate successions in Moravia, which were selected to represent different paleotectonic settings, have been studied with the aim of testing the influence of eustatic, climatic and tectonic controls on sedimentation and conodont paleoecology and taphonomy. On the slopes of the wide carbonate platforms of the Moravian Karst Development (Lesní lom and Grygov sections), an exemplary highstand shedding systems developed in the upper Famennian (expansa Zone), marked by a pronounced thickness of their respective calciturbidite successions and an abundance of shallow-water skeletal grains.Palamatolepis— andBispathodus-dominated conodont assemblages contain an admixture ofPolygnathus representing a transported, near-shore component. The eustatic sea-level fall in the praesulcata Zone and the lowstand conditions at the D/C boundary resulted in a decline of carbonate platform production and condensed deposition or nondeposition. In the Lesní lom section, a condensed sequence of turrbiditic calcarenites and shales (Middle praesulcata—lowermost sulcata Zone) was followed by lime mud calciturbidites (sulcata and duplicata Zones). In the conodont assemblages, the first event in the Lower praesulcata Zone was associated with the reduction of ‘mesopelagic’Palmatopic and a bloom of epipelagicPolygnathus communis. The second event in the Middle praesulcata Zone corresponds to the onset of polygnathidprotogranthodid biofacies, indicating a carbonate slope environment. In the Grygov section, a pronounced thickening and upward-coarsening succession of tubiditic calcilutites through calcarenites and intraclast breccias, with poor palmatolepid-bispathodid connodont assemblages (expansa Zone), indicates a progradation of the calciturbidite system associated with sea-level highstand. After a break in sedimentation, covering the interval from the Lower praseulcata to the base of Lower crenulata Zone, thick-bedded, fine-grained calciturbidites were deposited in the Lower crenulata Zone, and are associated with poor, mixed assemblages where siphonodellids and polygnathids predominate. At the isosticha-Upper crenulata/Lower typicus boundary, coasre grained, turbiditic calcarenites and breccias rich in clastic quartz grains and mixed conodont assemblages with reworked Frasnian and Famennian conodonts indicate a deep erosion of the source area, presumably due totectonic uplift (relative lowstand). In the Jesenec section, on the flanks of the volcanic seamount (the Drahany Development), a deep-water Upper Famennian condensed succession of calciturbidites and presumably winnowed pelagic limestones is marked by conodont assemblages of palmatolepid-bispathodid biofacies. More proximal calciturbidites with mixed deep-water and shallowwater conodonts prograde at the top of the Upper Famennian succession (Middle to Upper expansa Zone). A striking hiatus, covering the interval from the Early preaesulcata to the base of Lower crenulata Zone, resulted from extreme condensation and submarine bottom current erosion due to sea-level lowstand in the late Famennian and early Tournaisian. The renewed middle Tournaisian calciturbidite sedimentation with strong evidence of erosion at the source area indicates global eustatic rise and tectonic uplift of the Drahany Development seamounts (relative lowstand). The earlier occurrence of the uplift in the Jesenec area, relative to the Grygov section, shows the advance of tectonic processes over time in the Moravian-Silesian basin (orogenic polarity) as a consequence of Variscan orogenic movements.     

New insights into the base of the Emsian (Lower Devonian) in South China

Traditionally, the lower boundary of the Emsian (Lower Devonian) has been defined as the base of the dehiscens Zone marked by the first entry of the conodont Polygnathus dehiscens. However, following taxonomic revision of P. dehiscens, the current practice is to use the first appearance of P. kitabicus to define the base of the Emsian. As a result, specimens previously identified as P. dehiscens from South China need to be thoroughly reviewed. Careful examination of the oral and aboral features of the platform reveal that most specimens previously described as P. dehiscens from nine sections in Guangxi and Yunnan actually refer to P. excavatus excavatus or P. excavatus ssp. 114. In the South China Block, widespread and massive deposition of Devonian carbonates began during the time of the excavatus Zone, whereas the kitabicus Zone is probably located in the underlying strata, which mainly comprise marine siliciclastic rocks. Thus, the precise level of the lower boundary of the Emsian in South China is still unknown.     

Microproblematica,calcareous algae,and microbialites at the Frasnian-Famennian boundary interval in the Šumbera section (Moravian Karst,Czech Republic) and their significance in the context of the Kellwasser Crisis

The Kellwasser Crisis represents one of the most severe extinction events in the Phanerozoic. The ?umbera section (Moravo-Silesian Basin, Moravian Karst) spans the Upper rhenana to Palmatolepis minuta minuta (or younger) conodont zones and corresponds to the upper part of a carbonate ramp with material derived from shallower areas. A rich association of calcareous algae, cyanobacteria, and microproblematica, comprising renalcids (Izhella), Girvanella, Rectangulina cf. tortuosa, Rothpletzella, “solenoporaceans”, “Keega”-like microfossils, udoteacean alga Paralitanaia, volvocean and radiospherid calcispheres, palaeoberesellids, Umbellina bella, and Wetheredella, is similar in diversity to associations known from China, Australia, Canada, and Belgium. The ?umbera section represents a unique locality where changes in shallow-water biota at the Frasnian-Famennian boundary are well calibrated by detailed conodont biostratigraphy. We track in detail the evolution of shallow-water biota and the features that are often discussed in the context of “anachronistic” facies, such as flourishing microbial structures, enhanced early marine cementation, and flat-pebble conglomerates. Some features which could be indicative of environmental stress such as flat-pebble conglomerates developed during the Frasnian part of the Kellwasser Crisis (Upper rhenana to linguiformis Zones), whereas a renalcid boom started at the Frasnian-Famennian boundary.     

Lower Bashkirian conodonts from the Iraty Formation in the Alduides-Quinto Real Massif (Pyrenees,Spain)

The stratigraphy of the Carboniferous Iraty Formation is described in several sections from the southwestern Alduides-Quinto Real Massif (western Pyrenees), and biostratigraphy is updated based on conodont occurrences. The Ochaverri Member yields upper Serpukhovian conodonts, whereas the lower part of the overlying Asturreta Member is assigned to the lower Bashkirian, according to the occurrences of Declinognathodus noduliferus bernesgae and D. n. noduliferus, respectively. The upper beds of the formation (Baserdi Member) are correlated with the Kinderscoutian to lowermost Marsdenian English substages (lower Bashkirian), and new conodont occurrences are the oldest species of Neognathodus and Idiognathodus known in Spain. We discuss the status of Idiognathoides macer, several species related to I. sinuatus, I. sulcatus and Lochriea glaber. Conodont diversity is related to stratigraphic distribution of species and conodont biofacies. Finally, the lowermost Marsdenian to upper Bashkirian age of the siliciclastic rocks overlying the Iraty Formation is also discussed and compared with known data from localities in the western Pyrenees.