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     

Evidence of parallel evolution in the dental elements of Sweetognathus conodonts

The repeated emergence of similar morphologies in the dental elements of Permian Sweetognathus conodonts has been a hypothesized example of parallel evolution. To test if morphological parallelisms occur between isolated Sweetognathus lineages, this study uses two-dimensional-based geometric morphometrics combined with a revised and expanded phylogeny of Permian Sweetognathus conodonts to quantify dental element trait distributions and compare the phenotypic trajectories between lineages. A hierarchical clustering method was used to identify recurrent species pairs based on principal component scores describing their morphological variation, with the further incorporation of widely used ecological metrics such as limiting similarity and morphological overlap. Our research implies that a major contributor to conodont diversity in Palaeozoic marine trophic networks is the emergence of recurrent parallel morphologies via disruptive and directional selection. This study illustrates the mechanisms through which conodonts achieved their status as hyper-diverse predators and scavengers, contributing substantially to the complexity of Palaeozoic marine communities.     

Katian (Upper Ordovician) conodonts from Wales

Middle and upper Katian conodonts were previously known in the British Isles from relatively small collections obtained from a few localities. The present study is mainly based on 17 samples containing more than 17 000 conodont elements from an approximately 14‐m‐thick succession of the Sholeshook Limestone Formation in a road cut near Whitland, South Wales, that yielded a diverse fauna of more than 40 taxa. It is dominated by representatives of Amorphognathus, Aphelognathus/Plectodina and Eocarniodus along with several coniform taxa. Representatives of Decoriconus, Istorinus and Sagittodontina are reported from the Ordovician of UK for the first time. The fauna is a typical representative of the British Province of the Atlantic Realm and includes a mixture of taxa of North American, Baltoscandic and Mediterranean affinities along with pandemic species. Based on the presence of many elements of Amorphognathus ordovicicus and some morphologically advanced specimens of Amorphognathus superbus, the Sholeshook Limestone Formation is referred to the lower A. ordovicicus Zone. Most of the unit is also coeval with Zone 2 of the Cautleyan Stage in the British regional stage classification, and stage slice Ka3 of the middle Katian Stage in the global stratigraphical classification, an age assignment consistent with data from trilobites, graptolites and chitinozoans. The unusually large collection of M elements of Amorphognathus provides insight into the complex morphological variation in this element of some Katian species of this genus. The Sholeshook conodont fauna is similar to those of the Crûg and Birdshill limestones, but differs in several respects from the slightly older ones from the Caradocian type area in the Welsh Borderland. Although having some species in common, the Sholeshook conodont fauna clearly differs from coeval Baltoscandic faunas and is even more different in composition compared with equivalent North American Midcontinent faunas.     

Testing hypotheses of element loss and instability in the apparatus composition of complex conodonts: articulated skeletons of Hindeodus

Knowledge of the conodont skeleton, in terms of the morphology of the elements and the positions they occupy, provides the foundation for understanding of homology, taxonomy and evolutionary relationships in conodonts. This knowledge also underpins analyses of conodont functional morphology and feeding. Direct evidence of skeletal anatomy and apparatus architecture comes from natural assemblages: fossils that preserve together the articulated remains of the conodont apparatus, either collapsed onto a bedding plane or as clusters of elements in which juxtaposed and overlapping elements have been fused together by diagenetic minerals. Here we describe six clusters of the biostratigraphically important conodont Hindeodus parvus from the Lower Triassic Shangsi section, Sichuan Province, South China. Five of these clusters represent the partial remains of articulated skeletons, providing direct evidence of the number and arrangement of elements in the apparatus. Combined with data from previously published natural assemblages this provides a test of the hypothesis that Triassic conodonts had a reduced dentition. Hindeodus parvus possessed a complete raptorial array of two M and nine S elements (unpaired S₀; symmetrically paired S₁, S₂, S₃, S₄); the paired P₁ locations were occupied by carminiscaphate elements, but the apparatus lacked P₂ elements. This is consistent with broader evidence for a particularly high degree of integration and constraint operating on the S–M array of morphologically complex conodonts, leading to conserved architecture of the array over a period of more than 250 million years. The loss of elements from the P domain implies a change in food processing ability and, given the predominance of data from P elements in conodont taxonomy and biostratigraphy, the hypothesis of element loss from the P domain has significant implications for the broader understanding of conodont diversity and evolutionary patterns.     

High‐resolution conodont biostratigraphy from the Darriwilian Stage (Middle Ordovician) of the Argentine Precordillera and biodiversity analyses: a CONOP9 approach

Ordovician conodonts have been extensively documented in the Argentine Precordillera, providing a robust database for a diverse set of palaeontological studies. Despite the numerous studies, the published taxonomy and stratigraphical ranges, data remain contradictory for particular time intervals. Data from a new conodont collection from the Gualcamayo Formation exposed at the Cerro La Chilca section, and new information on the occurrence and ranges of species from the Las Chacritas and Las Aguaditas sections are presented herein. We used a computer‐assisted numerical sequencing program (CONOP9) to construct a composite stratigraphical range chart from data of 57 conodont species in four sections from the Argentine Precordillera. The identified zones of Lenodus variabilis, Yangtzeplacognathus crassus, Eoplacognathus pseudoplanus and Eoplacognathus suecicus allowed us to verify and adjust the biostratigraphical scheme for the Darriwilian of the Central Precordillera. Additionally, species of the genera Histiodella, namely Histiodella sinuosa, Histiodella holodentata, Histiodella kristinae and Histiodella bellburnenisis, enable a reasonable correlation between the Histiodella‐based zonation and the Baltoscandian zonation. Conodont diversity is evaluated using conventional measures (total diversity and normalized diversity) and an interval‐free approach with CONOP9 software. Our data show a positive pattern in conodont diversification, increasing rapidly through the L. variabilis to the Y. crassus zones and reaching a peak in the E. pseudoplanus Zone. When analysing diversity fluctuations with respect to the environmental changes within the depositional basin, migrations in and out of the basin related to local sea level fluctuations appear to be an important process driving the conodont diversity pattern in the Precordillera.     

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

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

Conodont chronostratigraphical resolution and Declinognathodus evolution close to the Mid‐Carboniferous Boundary in the Barcaliente Formation type section,NW Spain

The Barcaliente Formation in its type section provides significant conodont occurrences in the Mid‐Carboniferous Boundary interval (between the Mississippian and Pennsylvanian sub‐periods). A sequence of Declinognathodus morphotype appearances (considered here as species) is recognized from the uppermost Serpukhovian to the lower Bashkirian. These morphotypes belong to Declinognathodus noduliferus (Ellison & Graves) sensu lato, the primary marker for the worldwide correlation of the Mid‐Carboniferous Boundary. Among them, D. inaequalis (Higgins) appears to have been the first species to occur at the Mid‐Carboniferous Boundary bed in the Arrow Canyon GSSP. A morphocline between D. bernesgae (Sanz‐López et al.) and D. inaequalis is studied here to identify the bed correlated with the GSSP. Other diagnostic first occurrences of conodonts (Rachistognathus minutus (Higgins & Bouckaert), D. noduliferus, D. lateralis (Higgins & Bouckaert), Idiognathoides asiaticus Nigmadganov & Nemirovskaya and I. corrugatus (Harris & Hollingsworth)) are evaluated as chrono‐markers throughout their worldwide distribution, particularly for the Eurasian faunas. The available secondary markers provide a better resolution and support chrono‐correlation in sections other than that at the GSSP. The worldwide first and late occurrences of studied taxa are considered within the background of the Mid‐Carboniferous Extinction Event and the conodont overturn, where crises progenitor Declinognathodus gave rise to members of the Family Idiognathodontidae. Firstly, progenitor taxa are good biostratigraphic tools, whereas local extinction of genera considered Mississippian in age and new taxa arising later in the post‐crises radiation suggest an early Bashkirian palaeogeography and climatic conditions confining temporarily the biogeographic distribution of conodonts.     

New data from Oman indicate benthic high biomass productivity coupled with low taxonomic diversity in the aftermath of the Permian–Triassic Boundary mass extinction

A new Early Triassic marine fauna is described from an exotic block (olistolith) from the Ad Daffah conglomerate in eastern Oman (Batain), which provides new insights into the ecology and diversity during the early aftermath of the Permian–Triassic Boundary mass extinction. Based on conodont quantitative biochronology, we assign a middle Griesbachian age to the upper part of this boulder. It was derived from an offshore seamount and yielded both nektonic and benthic faunas, including conodonts, ammonoids, gastropods and crinoid ossicles in mass abundance. This demonstrates that despite the stratigraphically near extinction at the Permian–Triassic Boundary, Crinoidea produced enough biomass to form crinoidal limestone as early as middle Griesbachian time. Baudicrinus, previously placed in Dadocrinidae, is now placed in Holocrinidae; therefore, Dadocrinidae are absent in the Early Triassic, and Holocrinidae remains the most basal crown‐group articulates, originating during the middle Griesbachian in the Tethyan Realm. Abundant gastropods assigned to Naticopsis reached a shell size larger than 20 mm and provide another example against any generalized Lilliput effect during the Griesbachian. Whereas the benthic biomass was as high as to allow the resumption of small carbonate factories, the taxonomic diversity of the benthos remained low compared to post‐Early Triassic times. This slow benthic taxonomic recovery is here attributed to low competition within impoverished post‐extinction faunas.     

Taxonomy of the proterosuchid archosauriforms (Diapsida: Archosauromorpha) from the earliest Triassic of South Africa,and implications for the early archosauriform radiation

Proterosuchidae is one of the first clades of Archosauriformes (archosaurs and closely related species) to appear in the fossil record, with the richest sample of the group coming from the Lystrosaurus Assemblage Zone (earliest Triassic) of South Africa. Four nominal proterosuchid species were described from South Africa during the twentieth century (Proterosuchus fergusi, Chasmatosaurus vanhoepeni, Chasmatosaurus alexanderi and Elaphrosuchus rubidgei), but interpretations of their taxonomy have been widely disparate. The most recent taxonomic revision concluded that P. fergusi is the only valid species and that the other nominal species are junior subjective synonyms of this taxon. This proposal was based on the interpretation that anatomical differences between the nominal species could be explained as a result of ontogenetic changes and/or post‐mortem deformation. The recent discoveries of multiple new South African proterosuchid specimens provide an impetus to revisit their taxonomy. Based upon a comprehensive re‐examination of all known specimens, as well as examination of other proterosuchid taxa in collections worldwide, we conclude that the holotype of Proterosuchus fergusi is undiagnostic. As a result, we propose a neotype (RC 846) for the species. ‘Chasmatosaurus vanhoepeni’ and ‘Elaphrosuchus rubidgei’ are considered subjective junior synonyms of P. fergusi. ‘Chasmatosaurus’ alexanderi is considered a valid species, for which we propose the new combination P. alexanderi comb. nov. A third species, P. goweri sp. nov., is erected on the basis of a single specimen (NMQR 880). All three species recognized here are taxonomically distinct from a previously described archosauriform maxilla from the lower Lystrosaurus AZ. As a result, we recognize a minimum of four archosauriform species following the Permo‐Triassic mass extinction in South Africa. Our results suggest a greater species richness of earliest Triassic archosauriforms than previously appreciated, but that archosauriform morphological disparity remained low and did not expand until the late Early Triassic – early Mid‐Triassic.     

Conodonts from the Early Triassic microbialite of Guangxi (South China): implications for the definition of the base of the Triassic System

We describe a new Early Triassic (Griesbachian) succession of conodont faunas from a high‐resolution sampling of the basal Early Triassic microbial limestone and the base of the overlying unit at the Wuzhuan section (Nanpanjiang Basin, Guangxi, South China). The microbial limestone records the earliest phase of the Early Triassic biotic recovery after the end‐Permian mass extinction. For the first time, rich conodont faunas are reported from within the microbialite. The faunas from Wuzhuan are largely dominated by anchignathodontids, including several Isarcicella species, which were previously documented only from strata above the microbialite. A total of 14 conodont species assigned to three genera is recorded from the Wuzhuan section. Starting from the base of the microbialite upwards, several species are sequentially added to the conodont assemblage. The alpha diversity peaks at the top of the microbialite. The conodont record in the considered microbialite interval at Wuzhuan is presumably unaffected by local ecological changes. It therefore more likely represents an evolutionary rather than an ecological pattern. We compare the Wuzhuan's conodont record with a well‐supported phylogenetic model and suggest that the sequence of first occurrences at Wuzhuan is the closest to the ‘true’ sequence of evolutionary events that took place during this Griesbachian radiation of anchignathodontids. Based on comparisons with the GSSP section at Meishan, we suggest further that the first occurrence of Hindeodus parvus in Meishan does not correspond to its first appearance datum.     

日本志留纪至三叠纪牙形刺生物地层

本文系统地回顾了日本志留纪至三叠纪牙形刺研究的历史和现在的成果。志留纪牙形刺只有少数零星的报道,没有建立化石带;早泥盆世已建立了５个牙形刺组合,没有中、晚泥盆世的记录;石炭纪有８个牙形刺带,其中晚石炭世３个牙形刺带;二叠纪５个牙形刺带或动物群,其中,中、晚二叠世各１个带（动物群）;三叠纪可划分出１４个牙形刺带。     

Cutting the first ‘teeth’: a new approach to functional analysis of conodont elements

The morphological disparity of conodont elements rivals the dentition of all other vertebrates, yet relatively little is known about their functional diversity. Nevertheless, conodonts are an invaluable resource for testing the generality of functional principles derived from vertebrate teeth, and for exploring convergence in a range of food-processing structures. In a few derived conodont taxa, occlusal patterns have been used to derive functional models. However, conodont elements commonly and primitively exhibit comparatively simple coniform morphologies, functional analysis of which has not progressed much beyond speculation based on analogy. We have generated high-resolution tomographic data for each morphotype of the coniform conodont Panderodus acostatus. Using virtual cross sections, it has been possible to characterize changes in physical properties associated with individual element morphology. Subtle changes in cross-sectional profile have profound implications for the functional performance of individual elements and the apparatus as a whole. This study has implications beyond the ecology of a single conodont taxon. It provides a basis for reinterpreting coniform conodont taxonomy (which is based heavily on cross-sectional profiles), in terms of functional performance and ecology, shedding new light on the conodont fossil record. This technique can also be applied to more derived conodont morphologies, as well as analogous dentitions in other vertebrates and invertebrates.     

镇江地区上二叠统—下三叠统的牙形刺

详细描述了采自镇江附近１１－１，１１－２，７９－８号等钻孔的上二叠统—下三叠统的牙形刺化石共１１属２８种，建立了６个牙形刺化石带，其中上二叠统１个、下三叠统５个，并以牙形刺带为基础，进行了国内外对比。     

Siphonodella leiosa (Conodonta), a new unornamented species from the Tournaisian (lower Carboniferous) of Puech de la Suque (Montagne Noire,France)

A new conodont species, Siphonodella leiosa, is described from the lower Carboniferous pelagic limestones of the Montagne Noire (France), deposited in North Gondwana on a outer platform environment. Specimens were obtained from one level dated to the Siphonodella jii conodont Zone. The major difference from other siphonodellid conodonts known in this area is that the elements of this new species have a practically entirely smooth and unornamented platform, apart from the development of one or two low rostral ridge-like nodes. Similar morphologies were generally observed in shallow marine deposits of the same time frame from China, Russia and East and Central European areas. The new discovery reinforces the idea that ornamentation of siphonodellids is not only related to bathymetry, but that temperature could play an important role in the diversification and radiation of unornamented species during the Siphonodella jii conodont Zone.     

Where and when the earliest coccolithophores?

Gardin, S., Krystyn, L., Richoz, S., Bartolini, A. & Galbrun, B. 2012: Where and when the earliest coccolithophores? Lethaia, Vol. 45, pp. 507–523. New calcareous nannofossil analyses from the Northern Calcareous Alps of Austria are herein used to update and improve the state of knowledge about the oldest occurrence of coccolithophores reported in the literature. Previously reported Norian occurrences of coccoliths were based on an obsolete Triassic chronostratigraphy, in which the Rhaetian stage was subsumed into the Norian (‘Sevatian 2’). The oldest stratigraphical record of coccoliths spp. lies just below the Norian‐Rhaetian boundary and the first coccolith species, Crucirhabdus minutus, is recorded from the base of Rhaetian stage. The latter bio‐event is located just above the First Occurrence of the conodont Misikella posthersteni and the first occurrence of the ammonoid Paracochloceras suessi in the Steinbergkogel section (Austria), Global Stratotype Section and Point (GSSP) candidate for the Norian–Rhaetian boundary. The appearance of the coccolith Crucirhabdus minutus is seen as a robust biochronological datum that will provide useful constraints for Triassic biostratigraphy, palaeoclimatic modelling and phylogenetic reconstructions. The new calcareous nannofossil biochronology of Steinbergkogel that we present herein completes the existing biostratigraphic characterization of the Norian‐Rhaetian transition based on conodonts and ammonoids and strengthens the position of Steinbergkogel as the best GSSP proposed section for the base of the Rhaetian. The record of coccolithophores across the Norian–Rhaetian boundary at Steinbergkogel takes place along with a discernible increase in abundance of Prinsiosphaera triassica, as well as the appearance of Euconusphaera zlambachensis, which are the two most important Rhaetian pelagic carbonate producers. This succession of bio‐events is interpreted as the initiation of the pelagic carbonate production driven by the successful spreading of calcareous nannofossils in the Western Tethys during the Rhaetian. □Austria, Calcareous Alps, coccolithophores, Triassic.     

EARLY ORDOVICIAN CONODONTS FROM TARUTAO ISLAND,SOUTHERN PENINSULAR THAILAND

Abstract: Early Ordovician conodont faunas of the Thung Song Formation on Tarutao Island, southern peninsular Thailand, consist of 14 known species belonging to 17 genera, and eight undescribed species. Utahconus tarutaoensis and Filodontus tenuis are new species. Three conodont zones: the Rossodus manitouensis Zone, the Utahconus tarutaoensis Zone and the Filodontus tenuis Zone, in ascending order, are defined in the study sections. These are coeval with the interval from the Rossodus manitouensis Zone to the Acodus deltatus‐Oneotodus costatus Zone of the standard zonation in the North American Midcontinent. Based on the conodonts studied here and fossils previously reported from Tarutao Island, the Thung Song Formation is early Tremadocian to middle Arenig (Ibexian) in age. This formation is lithostratigraphically subdivided into the S1 to S5 members, and our study sections consist of the S1 to S3 members. These strata accumulated on a shelf in the Early Ordovician. The depositional environments of the limestones making up the S1 and S3 members were in deeper‐shelf conditions. Limestone and shale of the S2 member formed in a shallow‐water, high‐energy environment.     

Early evolution of Declinognathodus close to the Mid‐Carboniferous Boundary interval in the Barcaliente type section (Spain)

Upper Serpukhovian to lower Bashkirian conodonts studied from the lower and middle part of the Barcaliente Formation type section (NW Spain) are not abundant, but the sedimentary record seems unusually well represented. The first occurrence of Declinognathodus bernesgae occurs more than 140 m below the first occurrence of D. inaequalis, the taxon of the D. noduliferus species group appearing in the bed of the Global Standard Stratotype‐section and Point (GSSP) for the Mid‐Carboniferous Boundary (Arrow Canyon, Nevada). P1 elements transitional between D. bernesgae and D. inaequalis are described and considered relevant to the global correlation of the Mid‐Carboniferous Boundary. In addition, D. praenoduliferus, D. cf. noduliferus, D. lateralis, Idiognathoides asiaticus, I. corrugatus and Neognathodus sp. are described and illustrated from the Barcaliente Formation. A phylogeny is proposed for the early species of these genera on the basis of the morphological changes of the P1 element, where the rostral parapet and area, the surface roughness and the length and depth of the adcarinal troughs are diagnostic characters at the specific level, while caudal ornamentation is a secondary taxonomic character.     

Morphology and Ontogenesis of Psilotrichides hawaiiensis nov. gen., nov. spec. and Molecular Phylogeny of the Psilotrichidae (Ciliophora,Hypotrichia)

The Psilotrichidae are a family of middle‐sized hypotrichs with unique morphological and ontogenetic features (e.g. the oral primordium develops in a deep pouch) that, however, did not provide a definite phylogenetic signal. Thus, we studied the 18S rRNA gene of Urospinula succisa (Müller 1786) Esteban et al., 2001 as well as the morphology and ontogenesis of Psilotrichides hawaiiensis, a new genus and species from an ephemeral swamp on Oahu Island, Hawaii. The molecular data classify the psilotrichids into the oxytrichids but without clear branching position. A brief revision, using the structure of the oral apparatus, the location of the contractile vacuole, and three ontogenetic features, showed four distinct genera: Psilotricha Stein, 1859; Urospinula Corliss, 1960; Hemiholosticha Gelei, 1954; and Psilotrichides nov. gen., which differs from the confamilials mainly by the obliquely oriented buccal cavity and the shape of the undulating membranes as well as by a distinct ridge along the right buccal margin. The pyriform species, P. hawaiiensis, is about 65 × 45 μm in size and is easily recognized by the table tennis racket‐shaped appearance due to the elongated last cirrus of the left marginal row. Refined diagnoses are provided for the family Psilotrichidae Bütschli, 1889 and the genera contained.