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Sphenophyllum Brongniart constitutes a common and important element in the Carboniferous and Permian floras around the world, but its records in the Devonian Period are limited and remain to be investigated with respect to their morphology, anatomy, and distribution. Diversity of Sphenophyllum during Late Devonian in South China was significant with the presence of the well-known species including S. lungtanense Gothan and Sze, S. pseudotenerrimum Sze, and S. changxingense Huang et al. In this article, we describe a new species of the genus, S. fanwanense n. sp., from the Fanwan section of Changxing County, Zhejiang Province, characterized by having at least two orders of axes, with six spoon-shaped leaves attached at the node. A comprehensive review of previous records of Sphenophyllum from the Upper Devonian of South China shows that at least four species (S. fanwanense, S. lungtanense, S. pseudotenerrimum and S. changxingense), each with distinct leaf morphology, occur in South China in the Late Devonian, mostly from the Wutong Formation (Famennian). Given the lack of concrete evidence for anisophylly or heterophylly among the Devonian Sphenophyllum species (and more broadly, the Devonian sphenopsids), we suggest that the development of these features later in the Carboniferous and Permian periods was likely related to an increase in complexity of forest communities through time.

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.

The Devonian/Carboniferous (D/C) transition is characterized by a major transgressive/regressive cycle which led to a widespread ocean anoxia known as the Hangenberg Black Shale Event (HBSE), as well to a major sea-level fall (Hangenberg Sandstone Event, HSSE), recognized around the world. Both events are known as the Hangenberg Crisis. In order to examine the D/C transition in shallow water environment, the Mighan section in eastern Alborz was studied in terms of conodont biostratigraphy and stable isotope geochemistry. Twenty-five conodont species belonging to seven genera were identified and 5 conodont zones discriminated; namely, the Bispathodus aculeatus aculeatus Zone, Bispathodus costatus Zone, Bispathodus ultimus Zone, Siphonodella praesulcata Zone, costatus-kockeli Interregnum, and the sulcata Zone. Below the Devonian–Carboniferous boundary (DCB), the Hangenberg Black Shale and Hangenberg Sandstone equivalents were recognized, representing the Hangenberg Crisis that highly affected trilobite, ammonoid, brachiopod and conodont faunas at Mighan and worldwide. The kockeli Zone of the latest Famennian is missing at Mighan due to the lack of conodonts, probably related with the major environmental changes linked with the Hangenberg Crisis recognizable worldwide. Carbon isotopes measured of micrites from Mighan indicate a proximal depositional environment of a shallow shelf with terrestrial input and the oxygen isotope values from conodont apatite suggest warm seawater temperatures of tropical and subtropical setting in the study area.

This paper proposes a synthesis of the taxonomy, phylogeny, palaeogeographic distribution, regional biostratigraphy, and palaeobiogeographic faunal development of Carboniferous fusuline foraminifers. They appeared in the latest Tournaisian and comprised a small-sized, morphologically conservative taxonomic group during the Mississippian. Fusulines became larger and prevailed in Pennsylvanian foraminiferal assemblages. Carboniferous fusulines consist of Ozawainellidae, Staffellidae, Schubertellidae, Fusulinidae, and Schwagerinidae, in which 95 genera are considered as valid taxonomically. Upsizing their shells throughout the Pennsylvanian is likely related to symbiosis with photosynthetic microorganisms, which was accelerated by the acquisition of a keriothecal wall in Late Pennsylvanian schwagerinids. Regional fusuline succession data from 40 provinces provide a refined biostratigraphy, enabling zonation and correlation with substage- or higher-resolution precision in the Pennsylvanian. Their spatio-temporal faunal characteristics show that fusulines had a cosmopolitan palaeobiogeographic signature in Mississippian time, suggesting unrestricted faunal exchange through the palaeoequatorial Rheic Ocean. After the formation of Pangaea, Pennsylvanian fusulines started to show provincialism, and their distributions defined the Ural–Arctic Region in the Boreal Realm, Palaeotethys, Panthalassa, and North American Craton regions in the Palaeoequatorial Realm, and Western Gondwana and Eastern Peri-Gondwana regions in the Gondwana Realm. The Western Palaeotethys and East European Platform Subregions maintained higher generic diversity throughout the Pennsylvanian.

The Lough Foyle Basin is a half-graben that straddles the border between Northern Ireland and the Republic of Ireland and contains sediments that range in age from Lower Carboniferous to Holocene. The basin’s post-rift succession is represented by sediments of the Penarth Group and Lias Group. The lithostratigraphy and biostratigraphy of that interval are revised using new borehole material and existing outcrop. Palaeontological data provide a chronostratigraphic framework and aid palaeoenvironmental interpretations. Foraminifera, ostracods, palynomorphs indicate a nearshore, marginal marine depositional setting throughout much of the Rhaetian with a more marine, shelf and nearshore depositional setting for the Lias Group. The Penarth Group succession is similar to that elsewhere in Northern Ireland but the Lias Group (Waterloo Mudstone Formation) differs in that five distinct members can be recognised (Clooney Mudstone, Drummans Siltstone, Gortmore Mudstone, Tircreven Sandstone and Ballyleighery Mudstone), four of which are newly described. The deltaic and shelf sandstones of the Tircreven Sandstone Member are the only such examples preserved in the Jurassic strata of Northern Ireland and are some of the oldest in the Jurassic of the UK and Ireland, providing evidence of the proximity of the nearby Irish Landmass and representing a useful comparison for Early Jurassic sandstone reservoirs in offshore basins.

The Bashkirian (Late Carboniferous) ammonoid genus Bilinguites has a nearly global distribution in the tropical and subtropical zones and 25 species have been described up till now. Most of them are considered endemic, but some may have a wide geographic distribution. Our study on new and previously collected materials from North China (Gansu and Ningxia) shows that the stratigraphically youngest species of Bilinguites in North China and Western Europe are morphologically very similar, possibly indicating correspondence of the ammonoid faunas. Bilinguites scalpratus is revised and established as a stratigraphically important species of the genus in North China; it co-occurs with the first gastrioceratid species Cancelloceras pachygyrum. This occurrence can be correlated with the Western European co-occurrence of Cancelloceras cancellatum and Bilinguites superbilinguis at the base of the G1 Zone.

Early Carboniferous granitoids, distributed in the Yining and Central Tianshan blocks, Chinese western Tianshan, are important for deciphering their tectonic contexts and evolutionary history of the South Tianshan Ocean. Here we present LA-ICP-MS zircon U-Pb ages, in situ Hf isotopic data, and whole-rock geochemical data of the monzogranites and syenogranites from the Kalejun Mountains, southern part of the Yining Block, aiming to constrain their petrogenesis and geodynamic background. The results reveal that these monzogranites and syenogranites were emplaced at 356 Ma and 344 Ma, respectively. Geochemically, they are characterized by high SiO2 (68.74-75.76 wt.%) and K2O (4.58-6.18 wt%), low MgO (0.14-0.76 wt.%) and P2O5 (0.04-0.24 wt.%) contents. In addition, they are depleted in Ba, Sr, Eu, Ti, Nb, and Zr, and enriched in Rb, Cs, Th, and U. These features indicate that the Kalejun granites are high-silica I-type granites. The low positive epsilon(Hf)(t) values (+1.1 to +8.2) and old T-DM2(Hf) model ages (1294 to 827 Ma) of the studied zircons imply that the parental magma of these granites were derived from the reworking of Proterozoic basement with minor involvement of juvenile materials. Combined with the Late Devonian to Early Carboniferous magmatic activities in the Yining and Central Tianshan blocks (e.g. the discovery of A-type bubble rhyolites and high-Mg andesites), it is suggested that the Kalejun granites were likely related to the roll-back of the South Tianshan oceanic slab. Finally, we argue that the subduction of South Tianshan Ocean probably terminated at the end of the Early Carboniferous.

The quarries located to the south of Vise, in southeastern Belgium, correspond to historical type area of the Visean Stage (Carboniferous, Mississippian) and yielded an extremely rich macrofauna (known as the Vise fauna), including mainly brachiopods and mollusks (bivalves, gastropods, cephalopods), and also corals, trilobites and bryozoans, that are found in institutional collections all over the world. The Vise fauna is of late early Warnantian (late Asbian) age and was essentially recovered from a microbial buildup of Cracoean affinities that was exploited only in the quarries F and G sensu Horion and Gosselet’s (1892) terminology. Only the rugose corals have been revised, and the rest requires a complete taxonomic reassessment. Brachiopods are particularly diverse, and many genera (Productida, Orthida, Rhynchonellida, Terebratulida) were established in the past on the basis of type species from Vise. Productides are by far the most common elements of the brachiopod association, with the development of aberrant morphologies (Proboscidella, Striatifera). Some terebratulides and mollusks display ‘colour’ bandings that have been preserved. Two so-called Carboniferous species described by de Koninck (1887), namely Rhynchonella solitaris and Spirifer capillaris, are middle or late Frasnian in age.

The upper member of the Tagnana Formation of the Djebel Bechar Group (informally the Tagnana-3 or Tagnana Ill member) yielded fusulinoideans and other related foraminifera of the genera Pseudoendothyra (rare), Pseudostajfella (Semistaffella) (rare), Millerella, Pseudonovel/a, Plectostaffella, Eostaffella and Mediocris. The presence of Eostaffellina (rare) and Endostaffella is doubtful. The difference between Millerella and Eostaffella cannot be adequately defined by evolute versus involute coiling. Pseudonovella Kireeva, 1949 is redefined to include Seminovella Rauzer-Chernousova, 1951. The foraminifera are from two samples, one from close to the top and the other from near the base of the Tagnana III Member. Conodonts from the same samples, in comparison with the fusulinoidean foraminifera indicate similar or slightly younger ages; combined evidence from conodonts and foraminifera shows that the Tagnana-III Member at its upper boundary is Askynbashky or Akavassky (probably late Akavassky), while its lower boundary is Akavassky or Siuransky (probably late Siuransky) in age.

Xiphosura, the so-called horseshoe crabs, are a group of extant marine chelicerates that have a fossil record extending back to the Lower Ordovician. Their temporal range, coupled with a supposed record of bradytelic evolution has made them a focus of palaeontological consideration for over two centuries. As such, there are an array of taxa erected during the 20th century that have remained unexamined in light of new material and revised phylogenetic theories. Stem xiphosurids in particular require further research to uncover the diversity and disparity of non-crown group horseshoe crabs. To align with the recent increase in mid to late Paleozoic horseshoe crab research, we reconsider the stem xiphosurids “Kasibelinurus” randalli and “Bellinurus” alleganyensis from the Late Devonian of Pennsylvania and New York State. We conclude that they are synonyms and belong in neither Kasibelinurus nor Bellinurus, and the new genus, Patesia, is erected to accommodate them. Reconsidering stem xiphosurids highlights that particularly in the Late Devonian there was a high disparity of form. One such stem xiphosurid may represent the morphological stock that gave rise to the Carboniferous taxa and the first xiphosurid radiation.

Dolomitic limestones of the upper Piaui Formation of the Parnaiba Basin, Northern Brazil, preserve a rich and diversified invertebrate fauna of Middle Pennsylvanian age. The fauna shows affinities with the Pennsylvanian Itaituba Formation of the Amazon Basin, Brazil, and with Upper Paleozoic Andean faunas of the Cerro Prieto Formation and Tarma Group of Peru, and the Cano Indio Formation of Venezuela. In North America similar faunas are present in the Mississippian-Pennsylvanian Amsden Formation and in the Desmoinesian of Missouri. The Piaui fauna thus shows a clear "Tethyan" (boreal) affinity in contrast with other Late Paleozoic assemblages from southern South America. Bivalves comprise the majority of the fauna (60%, 31 species), followed by gastropods (17%, 9 species) and brachiopods (17%, 9 species). Other invertebrates including a cephalopod, a trilobite and a bryozoan make up the remainder of the fauna (6%, 3 species). Three species of protobranch bivalves, Polidevcia bellistriata, Nuculavus levatiformis and Solemya (Janeia) radiata are described here in. Obesity and elongation indexes as well as the external ornamentation and muscle scars of the nuculoid species here recovered are consistent with their interpretation as rapid (P. bellistriata) and slow, shallowly burrowing (N. levatiformis) deposit feeders. S. (Janeia) radiata, a slow shallow burrower, deposit feeder, is preserved in life position. This is the first of three publications that aim to ameliorate the present lack of paleontologic and paleobiogeographic knowledge on the Carboniferous of northern South America.

Upper Carboniferous deposits of the Rod El-Hamal Formation are well exposed at Wadi Araba, north Eastern Desert, Egypt. The upper part of the Formation includes late Moscovian (Middle Pennsylvanian) mounds formed by filamentous microbes and phylloid algae. Phylloid algae are loosely accumulated and supported by micritic encrustations that were formed by filamentous microbes. The filamentous microbes growing perpendicularly to the phylloid algal surfaces produced concentrated areas of peloids-A (peloid-A aggregations). The spaces between the phylloid algal-microbial frameworks were filled with concentrated areas of peloids-B (peloid-B aggregations) oriented perpendicular to the phylloid algae surfaces. Microbial communities as well as phylloid algae constitute significant components in the late Moscovian buildups in the study area. Below and above the mound facies, the non-reefs carbonates of Unit (IV) of the Rod El-Hamal Formation are composed of molluscan wackestone, ferruginous-molluscan packstone, and bioclastic rudstone facies, reflecting shallow subtidal depositional environment. Pennsylvanian phylloid algal-bearing buildups were divided into Tethyan and Pangean buildups. The Tethyan buildups are widely distributed in the circum-equatorial tropical paleolatitudes. The Egyptian buildups exhibit similarity in the building organisms to those of the Tethyan, but with a higher proportion of microbialites. Therefore, the subtropical Egyptian buildups are regarded as a sub-type of the tropical Tethyan buildups.

The late Paleozoic temnospondyl Sclerocephalus formed an aquatic top predator in various central European lakes of the late Carboniferous and early Permian. Despite hundreds of specimens spanning a wide range of sizes, knowledge of the endocranium (braincase and palatoquadrate) remained very insufficient in Sclerocephalus and other stereospondylomorphs because even large skulls had unossified endocrania. A new specimen from a stratigraphically ancient deposit at St. Wendel in southwestern Germany is recognized as representing a new taxon, S. concordiae new species, and reveals a completely ossified endocranium. The sphenethmoid was completely ossified from the basisphenoid to the anterior ethmoid region, co-ossified with the parasphenoid, and the basipterygoid joint was fully established. The pterygoid bears a slender, S-shaped epipterygoid, which formed a robust pillar lateral to the braincase. The massive stapes was firmly sutured to the parasphenoid. In the temnospondyl endocranium, character evolution involved various changes in the epipterygoid region, which evolved distinct morphologies in each of the major clades.UUID: http://zoobank.org/5e6d2078-eacf-4467-84cf-a12efcae7c0b

The vertebrate transition to land is one of the most consequential, yet poorly understood periods in tetrapod evolution. Despite the importance of the water-land transition in establishing modern ecosystems, we still know very little about the life histories of the earliest tetrapods. Bone histology provides an exceptional opportunity to study the biology of early tetrapods and has the potential to reveal new insights into their life histories. Here, we examine the femoral bone histology from an ontogenetic series of Greererpeton, an early tetrapod from the Middle-Late Mississippian (early Carboniferous) of North America. Thin-sections and micro-CT data show a moderately paced rate of bone deposition with significant cortical thickening through development. An interruption to regular bone deposition, as indicated by a zone of avascular tissue and growth marks, is notable at the same late juvenile stage of development throughout our sample. This suggests that an inherent aspect to the life history of juvenile Greererpeton resulted in a temporary reduction in bone deposition. We review several possible life history correlates for this bony signature including metamorphosis, an extended juvenile phase, environmental stress, and movement (migration/dispersal) between habitats. We argue that given the anatomy of Greererpeton, it is unlikely that events related to polymorphism (metamorphosis, extended juvenile phase) can explain the bony signature observed in our sample. Furthermore, the ubiquity of this signal in our sample indicates a taxon-level rather than a population-level trait, which is expected for an environmental stress. We conclude that movement via dispersal represents a likely correlate, as such events are a common life history strategy of aquatically bound vertebrates.

The Olistostrome member of the Sana-Una Paleozoic complex of the Ljubija ore mine in Bosnia and Herzegovina contains limestone fragments of pebble to block size that have been examined paleontologically. The recovered conodont fauna of the first sample is characterized by the species Declinognathodus lateralis, Idiognathoides sulcatus sulcatus and Idiognathodus sp. confirming its mid-Bashkirian age. This report is the first on the occurrence of these taxa in the area. The second sample with chaetetid demosponges yields an abundant diversified microbiota consisting of cyanobacteria, algae and foraminifera. Chlorophyts are marked by the common siphonoclad occurrence of Donezella lutugini and D. lunaensis, whereas rhodophyts include rare representatives of Stacheia, Stacheoides, Pseudoungdarella and Masloviporidium. The presence of Asphaltinella horowitzi and Aphralysia carbonaria of unclear taxonomic position is also documented. Pseudostaffellids, eostaffellids and other foraminifera, mostly endothyrids are present. The examined associations of fossils point to the Bashkirian age of the primary rock that originated in a very shallow habitat most probably linked to a high-energy reef environment.

The Qijiagou section on the southern margin of the Junggar Basin preserves thick and extensive Carboniferous carbonate rocks, and is the type section of the Upper Carboniferous in the Junggar Basin. Here, we studied the lithological assemblages and brachiopod fossils of the Upper Carboniferous Qijiagou Formation on the southern margin of the Junggar Basin. Outcrop observations indicate laterally variable thicknesses of this unit, comprising thick-bedded, fossiliferous, channelised, sandy, and calcareous limestone. Petrographic observations revealed four microfacies, indicating deposition of the formation in diverse depositional environments, including lagoonal to delta-dominated middle shelf settings. In addition, we identified 15 species of brachiopod fossils in 11 genera which belong to Spiriferida Waagen, Productida Sarycheva et Sokolskaya, and Strophomenida Opik. Thus, based on the characteristics of the lithological assemblage and brachiopod fossils, we infer that the Qijiagou area had a relatively warm environment in the middle and late Late Carboniferous. This included a turbulent and shallow water body with two regressions which occurred in a carbonate platform sedimentary environment.

The ancestry of chimaeriform chondrichthyan fishes can be traced back to the Late Triassic (∼220 Ma). To date, only one chimaeriform suborder, the Echinochimaeroidei, has been recognized from the P...

The Devonian-Carboniferous transition is marked by major perturbations to the Earth’s climate system and marine redox conditions, a first-order mass extinction, and the succeeding marine ecosystem recovery, recording a sequence of environmental and biotic events. In South China, there exist numerous well-preserved and continuous Devonian-Carboniferous boundary successions that were formed in low-latitude passive margin basin, and this presents a unique opportunity to decipher the strata along a proximal to basinal transect using integrated biostratigraphical, event-stratigraphical, and geochemical approaches. Our geochemical data (δ13Ccarb, δ15Nbulk, δ238U and Mo/U enrichments, I/Ca ratio) indicated local and global C-N cycling perturbations and marine anoxia in the main phase of the Hangenberg Crisis interval (Middle Siphonodella praesulcata Zone), which coincided with a major regression in South China. Stromatoporoid biostromes and most typical Devonian faunas did not survive into the interval, and the deep-water black shales yield only opportunistic survivors, such as ammonoid Postclymenia cf. evoluta and bivalve Guerichina, along with miospores belonging to the LN Zone. The upper Hangenberg Crisis interval (Upper Si. praesulcata Zone, equal to the Protognathodus kockeli Zone) is marked by initial post-glacial transgression, a global δ13Ccarb spike, and opportunistic faunal blooms, coincident with conodont biofacies shift to the polygnathid-protognathodid biofacies. The Protognathodus fauna, nearly absent from platform facies, is often found in the condensed upper crisis interval of the basin, slope, and platform margin facies within the Youjiang Basin, although the abundance is extremely low and the phylogenetic lineage of Pr. meischneri-Pr. collinsoni-Pr. kockeli cannot be recognised in most successions. In South China, the first occurrence of Pr. kockeli seems to correspond to lithofacies changes, recorded in the first level of transgression just above the Hangenberg Sandstone event. Multiple lines of evidence, including great spatial heterogeneity of δ13Ccarb and δ15Nbulk records, extremes of redox proxies, and lack of macrofossils, suggest that marine habitats remained in a critical/turbulent state during the Upper Si. praesulcata Zone, triggering the opportunistic blooms and minor extinctions. Following the crisis intervals, radiation/recovery of many fossil groups did not occur until the upper part of the Siphonodella sulcata Zone to the Si. duplicata Zone. In particular, a negative shift of δ13Ccarb to pre-crisis values occurring near the base of the Si. sulcata/Pr. kuehni Zone could be recognised worldwide. The research history, current status, and achievement of Devonian-Carboniferous boundary in China are briefly reviewed.

The Carboniferous Foraminifera are composed of representatives of three classes: Fusulinata, Miliolata and Nodosariata. Despite ample literature on Paleozoic Allogromiata and Textulariata, the real presence of these classes remains questionable during the Carboniferous and they are thus excluded herein. The main biostratigraphical markers belong to the superfamilies Archaediscoidea, Lasiodiscoidea and Bradyinoidea, even if many genera among the archaediscoids still have a controversial nomenclature, as well as do some lasiodiscids and bradyinoids. Secondary biostratigraphical markers belong to Lituotubelloidea (= ‘Tournayelloidea’ of the authors), Endothyroidea and Loeblichioidea (these latter giving rise to the primitive Fusulinida). The Miliolata appear at the Visean/Serpukhovian boundary interval. The typical Carboniferous miliolates are primitive nubeculariins and cornuspirinins. Tubiphytids might be miliolate and cyanobacterium consortia, derived from the nubeculariin Palaeonubecularia . The most primitive nodosariates (syzraniids) appeared in the Moscovian; and gave rise, in the latest Carboniferous, to Protonodosaria , Nodosinelloides and possibly Polarisella , Paravervilleina and the oldest Geinitzinoidea. Palaeobiological data are mainly provided by the genera Bradyina , Tetrataxis and Climacammina . Palaeobiogeographical distributions of Pojarkovella , Janischewskina, Eosigmoilina , Brenckleina , Spireitlina , Hemigordius and Syzrania testify to the successive foraminiferal migrations between the Palaeotethys, Ural and Panthalassan oceans. Two taxa are created: Eoparastaffellidae and Banffellinae.

Whip spiders (Amblypygi), as their name suggests, resemble spiders (Araneae) in some aspects, but differ from them by their heart-shaped (prosomal) dorsal shield, their prominent grasping pedipalps, and their subsequent elongate pair of feeler appendages. The oldest possible occurrences of whip spiders, represented by cuticle fragments, date back to the Devonian (c. 385 mya), but (almost) complete fossils are known from the Carboniferous (c. 300 mya) onwards. The fossils include specimens preserved on slabs or in nodules (Carboniferous, Cretaceous) as well as specimens preserved in amber (Cretaceous, Eocene, Miocene). We review here all fossil whip spider specimens, figure most of them as interpretative drawings or with high-quality photographs including 3D imaging (stereo images) to make the three-dimensional relief of the specimens visible. Furthermore, we amend the list by two new specimens (resulting in 37 in total). The fossil specimens as well as modern whip spiders were measured to analyse possible changes in morphology over time. In general, the shield appears to have become relatively broader and the pedipalps and walking appendages have become more elongate over geological time. The morphological details are discussed in an evolutionary framework and in comparison with results from earlier studies.

Based on the analysis of data on bryozoan localities in territory of modern Eurasia, changes of the bryozoan richness were calculated in detail for six time slices in the interval from the early Famennian to the late Tournaisian. Highest extinction tempo was observed for the latest Famennian, apparently linked to the Hangenberg Event. However, in the richness of bryozoans during the late-latest Famennian, an abrupt increase in the origination tempo is observed, as well as a slight decrease in the total bryozoan richness at the Devonian-Carboniferous transition. Origination of abundant narrowly specialised bryozoan genera in the orders Trepostomata and Cryptostomata at the end of the Devonian and in the beginning of the Carboniferous was caused by the combined influence of extrinsic and intrinsic factors. An observed increase in the morphological specialisation of bryozoans during the Late Devonian and early Carboniferous coincides with patterns observed in other benthic organisms such as brachiopods and rugose corals. These patterns can be explained by vacation of ecological niches and changes in the structure of the marine biota in the aftermath of the Hangenberg Event.

Although the Centre Grégoire Fournier of the Maredsous Abbey is especially famous for the fossils and minerals from the Carboniferous (Viséan) ‘black marble’ of Denée, a marine conservation-Lagerstätte, its palaeontological collections likewise include some types and illustrated specimens of invertebrates (cystoids, goniatites, ostracods, trilobites) and fishes from the Ordovician, Devonian and Carboniferous of Belgium. These specimens are discussed and/or illustrated as is the case of the fragments of two Belgian meteorites (Lesves and Tourinnes-la-Grosse chondrites) that are part of the CGF mineralogical and petrological collections. Moreover, the type material of 30 species and subspecies of Upper Devonian–Pennsylvanian linguliformean and rhynchonelliformean brachiopods (described by de Koninck (1847), J. Fraipont (1888a), Ch. Fraipont (1908), Demanet (1923, 1934), Demanet (in Demanet & Van Straelen, 1938), and Grimm (1998)) and that of two Middle–Upper Devonian species of trilobites (Stainier, 1887; Richter & Richter, 1933), almost all from the Namur–Dinant Basin (southern Belgium), are re-investigated and/or fully figured for the first time in order to facilitate future taxonomic revision. The obscure Tournaisian genus Anomianella de Ryckholt (1851) is rejected from the bivalves and transferred to the brachiopods (Craniida). It is probably related to Petrocrania Raymond, 1911. The lectotype (hereby selected) of Orthis latissima M‘Coy, 1844 and that of Producta corrugata M‘Coy, 1844, both from the Viséan of Ireland, are photographically illustrated (for the first time for the former). The lectotype of Productus murchisonianus de Koninck, 1847 from the Upper Palaeozoic of Tasmania (Australia) is also designated.

Recently, the end-Devonian mass extinction (Hangenberg Crisis, 359 Ma) was identified as a first-order mass extinction, albeit not one of the “Big Five” events. Many marine and terrestrial organisms were affected by this crisis. The cause of this mass extinction is still conjectural and widely discussed. Here we report anomalously high mercury (Hg) concentrations from the South Tian Shan (Uzbekistan), together with correlation using conodont biostratigraphic data. Hg enrichment (to 5825 ppb) was detected in marine deposits encompassing the Hangenberg Crisis. In the Novchomok section, the Hangenberg Crisis interval does not contain typical Hangenberg Black Shales; however, by means of inorganic geochemistry (enrichment of redox-sensitive elements such as Mo, V, and U) we detected an equivalent level despite the lack of marked facies changes. This is the first record of Hg and Hg/total organic carbon anomalies in marly shales, marls and carbonates that are totally independent of facies changes, implying that volcanism was the most probable cause of the Hangenberg Crisis. This conclusion is confirmed by the presence of a negative δ13C excursion, which may reflect massive release of isotopically light carbon from volcanogenic and thermogenic devolatilization likely combined with increased arc-volcanism activity worldwide at the end of the Devonian.

A taxonomic revision of shallow-water Carboniferous (Pennsylvanian) red and green algae observed in Ponga, Picos de Europa and Pisuerga-Carrion Units in NW Spain is presented. Some microproblematics are included. Thirty-two genera are described; three genera and four species are new: Anatolipora cantabriensis n. sp., Pelayella multiporosa n. gen., n. sp., Kasimophyllum demuesensis n. gen., n. sp. and Penella pongaensis n. gen., n. sp. The microflora is typically Western Paleotethyan and remarkably similar to that observed in the Carnic Alps and Central Urals.

On the basis of a detailed study of 39 sections in the Djangdjir and Kokshaal ranges of the South Tianshan (Kyrgyzstan), the conodont zonation of deep-water facies of the Tournaisian, Visean, and Serpukhovian stages was specified. Nine conodont zones, including three new ones, and one unit in the rank of layers with conodonts were distinguished. The deep-water character of studied facies determines the predominant occurrence of cosmopolitan conodonts, which enable the reliable correlation with coeval sections in many regions worldwide. The comparison of conodont assemblages at the specific level indicates that conodont assemblages from the South Tianshan sections are the most similar to those from the Southern Urals and South China.

Equisetales represents a diverse and abundant group of articulate plants with a wide geographical distribution that had their greatest development in late Palaeozoic and early Mesozoic times. The o...

In the Netherlands, Late Palaeozoic pelmatozoans – that is, stalked echinoderms – are known from building stones and cobbles in rivers, but there are no in-situ carbonate rocks from which they might be collected. Unsurprisingly, most recognisable specimens are columnals and pluricolumnals. Two small thecae, collected in the mid-1970s from silexite cobbles in the bedload of the River Maas in the Venlo-Tegelen area (province of Limburg, south-east Netherlands) are exceptional finds. One specimen, the diplobathrid camerate crinoid Rhodocrinites sp., has an unsculptured theca and some minor differences of form, yet otherwise satisfies the diagnosis of this genus. The other, the pentremitid blastoid Doryblastus? sp. is rather poorly preserved, yet is the first blastoid to be recorded from the Netherlands. Either or both of these specimens may be juveniles, particularly the blastoid. They are unlikely to be coeval, coming from separate cobbles and being of slightly different preservation. Their provenance from silexite cobbles suggest they originated from Lower Carboniferous (Tournaisian-Visean = Mississippian) carbonates in the southern Ardennes (south-central Belgium).

A symmetrical bifurcation of an Alethopteris urophylla frond from the Pennsylvanian (Upper Carboniferous) of the Northern France coalfield shows some still-attached remains of foliage. Such rare specimens contribute, step by step, to a better understanding of the frond architecture of the genus Alethopteris, which is of the bifurcatepinnate type. A few complementary general comments are given.

A revised age provided by conodonts from the Gilwern Oolite of the Clydach area of South Wales allows a clearer understanding of the palaeohydrology, palaeoclimatic history and diagenesis of previously correlated oolitic units. Earlier uncertainty over the apparent sub-regional differences in climate during the early Visean (Chadian—Arundian) lowstand is resolved. Previously a humid interval evidenced by prominent palaeo-epikarst capping the Gilwern Oolite along the northern outcrops of the South Wales synclinorium, was not recognized in what were regarded as correlative outcrops in the Chadian Gully Oolite in the southern part of the synclinorium. Conodont dating now shows that the Gilwern Oolite is much older (Courceyan) than the Chadian Gully Oolite, and during the prolonged exposure of the former there was an interval of weathering under a humid climate. This also explains the contrast between the diagenesis seen between the Gilwern and Gully oolites, which are no longer seen as correlatives.

Carboniferous conodont biostratigraphy comprises regional zonations that reflect the palaeogeographical distribution of taxa and distinct shallow-water and deep-water conodont biofacies. Some species have a global distribution and can effect high quality correlations. These taxa are incorporated into definitions of global Carboniferous chronostratigraphic units. A standard global Carboniferous zonation has not been developed. The lowermost Mississippian is zoned by Siphonodella species, excepet in shallow-water facies, where other polygnathids are used. Gnathodus species radiated during the Tournaisian and are used to define many Mississippian zones. A late Tournaisian maximum in diversity, characterized by short-lived genera, was followed by lower diversity faunas of Gnathodus species and carminate genera through the Visean and Serpukhovian. By the late Visean and Serpukhovian, Lochriea provides better biostratigraphic resolution. Shallow-water zonations based on Cavusgnathus and Mestognathus are difficult to correlate. An extinction event near the base of the Pennsylvanian was followed by the appearance of new gnathodid genera: Rhachistognathus , Declinognathodus , Neognathodus , Idiognathoides and Idiognathodus . By the middle of the Moscovian, few genera remained: Idiognathodus , Neognathodus and Swadelina. During the middle Kasimovian and Gzhelian, only Idiognathodus and Streptognathodus species were common. Near the end of the Gzhelian, a rediversification of Streptognathodus species extended into the Cisuralian.

Salthill Quarry, Clitheroe, Lancashire (Mississippian, Visean) is a Site of Special Scientific Interest (SSSI), in part based on its diverse fauna of invertebrates, particularly echinoderms. A small collection of crinoid pluricolumnals are described herein, collected from muddy horizons in the Hodder Mudstone Formation to typify the diversity and frequency of their encrusting and boring fauna. These specimens are infested by a range of episkeletozoans, namely a single occurrence of Sutherlandia parasitica (Phillips), four occurrences of Cladochonus sp. and eight of Oichnus paraboloides Bromley. Two variants of the pit (boring or embedment or both) O. paraboloides are recognised: those in which a living crinoid showed a growth reaction to pit formation; and those that did not and which were presumably dead at the time of infestation. The epizoozoan tabulate coral Cladochonus sp. is also common, including specimens intergrown between and within the columnals. Sutherlandia parasitica is relatively uncommon; the only specimen likely infested a dead pluricolumnal on the sea floor. A comparative collection of pluricolumnals infested by Cladochonus beecheri Grabau from the Mississippian Borden Group of the Midwest, USA, showed superior preservation to the Clitheroe Cladochonus sp. Cladochonus beecheri in the Borden Group infested platycrinitid crinoid stems, an association not noted from Salthill Quarry. Similar suites of borings-episkeletozoans, from two other sites – the Visean of Feltrim, Co. Dublin, Ireland, and the Permian of Timor – suggest that these were persistent associations for much of the Late Palaeozoic.

Traditionally, biostratigraphy of the terminal Famennian (Upper Devonian) and lower part of the Tournaisian (Mississippian) is based on phylogeny of the conodont genus Siphonodella Branson and Mehl. This study is focused on a Siphonodella species with a long taxonomical history. For the last three decades it was known as S. hassi Ji, 1985 and was used as a zonal index for the eponymic hassi Zone, used worldwide. However, this species faces several taxonomical problems. The main issue is the homonymy of S. hassi and S. cooperi hassi Thompson and Fellows, 1970. However, S. cooperi hassi is a subjective junior synonym of S. obsoleta Hass, 1959. The oldest junior synonym of S. hassi is S. wilberti Bardasheva, Bardashev, Weddige and Ziegler, 2004, and, according to a statement by the International Commission on Zoological Nomenclature, the latter represents the only available name for this species. An additional problem is the similar morphology of early ontogenetic stages of S. wilberti and S. quadruplicata Branson and Mehl, 1944, the youngest stratigraphically important siphonodellid species. In advanced ontogenetic stages, P1 elements that possess two rostral ridges belong to S. wilberti only. The smaller P1 elements having two or less rostral ridges comprise the S. wilberti- S. quadruplicata plexus. Maintenance of the wilberti Zone in the Tournaisian global zonation is suggested.

Small ammonoid assemblages are recorded from the Perapertú Formation in northern Palencia. This is a mudstone unit with local platform limestones characterised by carbonate debris flows on the limestone margins. This unit, of Late Bashkirian to Early Moscovian age, participates in a series of southwards verging thrust slicesNorth of a major fault zone which originated as the head (leading edge) of a large thrust sheet with an internal deformation consisting of thrust slices and at least one nappe structure. Opposed vergencies at the head of this major thrust sheet (Carrionas Thrust Front) and the Ruesga Fault which locally modifies its trace, mark the position where the northern branch of the Cantabric-Asturian arcuate fold belt has overridden the southern branch in early Westphalian (Langsettian) times. The chronostratigraphic significance of the Perapertú Formation is discussed in the context of marine-terrestrial correlations for the Late Bashkirian-Early Moscovian time interval. It is concluded that the evidence from NW Spain suggests a position of the base of the Moscovian at the level of basal Westphalian or even within the highest Namurian. A brief analysis of the literature shows this position to be different to some of the correlations admitted in recent publications. A newly discovered goniatite fauna from the lower part of the Perapertú Formation contains Brannerocera ssp. indicating Late Bashkirian to earliest Moscovian, and Deleshumardites cantabricus Kullmann gen. et sp. nov. This fauna is figured and described in conjunction with the new subfamily Dombaritinae Kullmann (family Delepinoceratidae).

A relatively complete conodont record from Famennian to the Mississippian/Pennsylvanian boundary was investigated in the Anarak section, Central Iran. The studied interval belongs to the Bahram, Shishtu, Ghaleh and Absheni formations. The Famennian part of the section (Bahram Formation) ranges from the Palmatolepis triangularis Zone into the Bispathodus ultimus Zone. Not all conodont zones could be defined due to the lack of indicative species. Furthermore, it seems likely that a hiatus occurs around the Devonian/Carboniferous (D/C) boundary (most probably from the Siphonodella praesulcata to the ?Siphonodella sulcata–early Siphonodella crenulata conodont zones) based on the lack of stratigraphically important conodonts as well as on sedimentological criteria. The lack of representative siphonodellids and protognathodids at the base of the Mississippian prevents detailed stratigraphic position of the D/C boundary. Lower Carboniferous (Mississippian) rocks are characterized by red nodular limestone which is unique in comparison with other studied sections of the same age in Central Iran. Within the studied section, we could define the Mississippian/Pennsylvanian boundary. The mid-Carboniferous boundary was defined by the occurrence of Declinognathus noduliferus s.l. Conodont biofacies changes (Mississippian genera Gnathodus and Lochriea have been replaced by Pennsylvanian genera Declinognathus and Idiognathodus) are recognized in this section as well., Ministry of Science Research and Technology (IR)

South-central Kentucky (USA) has one of the world’s most intensively studied karst landscape/aquifer systems, including the carbonates that hold the Mammoth Cave System. Slightly higher in the Mississippian (Lower Carboniferous) stratigraphic section, the Haney Limestone forms a locally important but much less well-studied karst aquifer. This research provides the most comprehensive synthesis to date of the karst hydrogeology of the Haney Limestone of south-central Kentucky, focusing on the distribution of and controls on caves and karst features. In contrast to drainage systems within the underlying major limestones, joints have controlled passage development in the Haney Limestone, and passage orientation is consistent with regional jointing. Bedding planes and insoluble rock at the base of the Haney Limestone also influence conduit development. Most caves developed in the Haney Limestone in the study area are single-conduit caves that receive direct allogenic recharge. Often, cave entrances are perennial springs where streams resurge, suggesting that the caves function within the contemporary landscape as drains for localized recharge areas. Most caves developed in the Haney Limestone appear to be geologically recent phenomena. Most cave passages in the study area are hydrologically active, water resurging from the sampled springs is typically undersaturated with respect to calcite, and some caves appear to be developed along stress-release fractures associated with small young valleys. This suggests that caves in the Haney Limestone were not directly influenced by the incision of the Green River over vast periods, as was the case for Mammoth Cave, but that cave development is a largely contemporary process.

Hydrasperman pteridosperms (seed ferns) were widespread and diverse in Carboniferous equatorial wetlands of Euramerica, and although present in Permian equatorial wetlands in Cathaysia, little is k...

This paper reviews the foraminiferal biostratigraphy of the Devonian–Carboniferous (D–C) boundary beds in Russia and Western Kazakhstan. The regional records of the zonal successions of Russia are summarised from sections and boreholes of the Volga-Urals Subregion of the East European Platform (Syzran no. 401 Borehole, Melekesskaya no.1 Borehole); Timan-Pechora Province of the East European Platform (Kamenka Section); South Urals (Sikaza, Zigan, Ryauzyak and Dzerzhinka sections); Middle Urals (Pershino Section); and West Siberia, Vagai-Ishim Basin (Borehole no. 1 Kurgan-Uspenskaya), and the zonal successions of Western Kazakhstan are summarised from the Mugodzhary Mountains. Three foraminiferal biozones can be defined in the interval comprising the latest Devonian and earliest Carboniferous: Quasiendothyra kobeitusana Zone, the Tournayellina pseudobeata – remnant Quasiendothyra Zone, and the Earlandia minima Zone. The D–C boundary interval is situated within the Tournayellina pseudobeata–remnant Quasiendothyra Zone. A correlation with the ammonoid and conodont zonations is proposed, which is useful for correlating deep-water and shallow-water successions.

Research subject. Changes in the trophic structure of shallow-water pelagic ecosystems at the Devonian/Carboniferous border were investigated by studying the carbon isotope composition of conodont organic matter.Materials and methods. Two Devonian-Carboniferous shallow-water clayey-carbonate sections located in the southern part of the Pechora-Kozhva Uplift (Pechora Plate) were analysed. The Devonian-Carboniferous boundary was detected by the first occurrences of Siphonodella sulcata, S. semichatovae and Patrognathus crassus, as well as by the last occurrence of Pseudopolygnathus graulichi. The carbon isotope composition was investigated both in whole-rock carbonate samples and the conodont organic matter of two dominant species (Polygnathus parapetus and P. communis communis).Results. The distribution of stable carbon isotopes in the organic matter of conodont elements accompanied by the data on carbonate isotope composition allowed us to suggest changes in the food composition of the dominant taxa during the Late Famennian-Early Tournaisian transition. It was assumed that the latest Famennian representatives of Polygnathus parapetus and P. communis communis consumed largely phyto- and zooplankton, which is characterized by a light isotopic composition of organic carbon. The nutrition based on phyto- and zooplankton with a heavier isotopic composition of organic carbon was suggested for the early Tournaisian representatives of these species.Conclusions. The discovered variations in the carbon isotope composition of conodont organic matter in shallow-water facies may correspond to the change from the eutrophic pelagic ecosystem to the oligotrophic ecosystem, and/or global perturbation of the carbon cycle due to climatic changes. Since the available data is limited to two geological sections, it is impossible to unambiguously interpret the scale (local, regional, global) of these variations and their correlation potential.

Clam shrimp (Spinicaudata) are worldwide distributed branchiopod crustaceans specialised in ephemeral freshwater habitats. The Carboniferous is an important period for the early evolution and diversification of clam shrimp. Compared with the rare and geographically confined fossil record of the Devonian, clam shrimp in the Carboniferous have a much wider geographical distribution and higher biodiversity. Over 20 genera of clam shrimp have been recorded in the Carboniferous all over the world, but they are sparse in China. To date, five records of Carboniferous clam shrimp have been reported from China (Pruvost, 1927; Zhang et al., 1976; Wang, 1987; Zheng et al., 1988; Liu & Fan, 1995; Liao et al., 2019). Among them, four species Lioestheria? mathieui Pruvost, 1927, Protomonocarina huixianensis Wang, 1987, Retrofractus lingyuanensis Liu & Fan, 1995, and Pemphilimnadiopsis cheni Liao, Shen & Huang, 2019, are found in the Pennsylvanian Benxi Formation in North China (Pruvost, 1927; Zhang et al., 1976; Wang, 1987; Liu & Fan, 1995; Liao et al., 2019).

The Carboniferous was a period of high diversification of the polyneopteran insects of the superorder Archaeorthoptera (so-called ‘orthopteroid’ insects) (see Fossilworks site at http://fossilworks.org/bridge.pl?a=home). Some of the Late Carboniferous outcrops are very rich (Mazon Creek in North America, Commentry in France). The Konservat-Lagerstätte of Avion (Northern France) is especially rich for a Moscovian outcrop, with no less than six described genera and species: Aviogramma gracilis Prokop et al., 2014, Aviocladus pectinatus Prokop et al., 2014, Aviohapaloptera bethouxi Prokop et al., 2014, Aviologus duquesnei Coty et al., 2014, Cacurgus avionensis Schubnel et al., 2020, and cf. Tococladus sp. (Coty et al., 2014; Prokop et al., 2014; Schubnel et al., 2020). Here we describe a new genus and species, the first representative of the Carboniferous family Eoblattidae Handlirsch, 1906 from this outcrop, on the basis of a forewing.

Permian millipedes are rare, especially so considering the relative abundance of millipedes in Carboniferous rocks. We report an early Permian millipede fauna containing three new genera and species of millipedes (Oklahomasoma richardsspurense new genus new species, Karstiulus fortsillensis new genus new species, and Dolesea subtila new genus new species) found in fossil-producing pockets of the Fort Sill fissures exposed in the Dolese Quarry near Richards Spur, southwest Oklahoma, USA. These are the first new genera of invertebrates to be described from this site, one of the most prolific fossil-vertebrate sites in the world. We also comment on taxa with morphological similarities and note previously described occurrences of Permian millipedes as well as occurrences of fossil myriapods (millipedes and centipedes) in karst deposits (caves and fissure fills) in Europe, Africa, Asia, North America, and the Caribbean. In contrast with the forms found at Richards Spur, most of these previous accounts of millipedes found in caves and fissure fills are of Pleistocene forms that are closely allied to modern taxa. The taxa from Richards Spur bear some similarities to Pennsylvanian forms. Karst (cave and fissure) faunas should be ranked with concretion faunas, cannel coals, and amber faunas as a major source of fossil myriapods.UUID: http://zoobank.org/5d58e1fb-4e5b-4597-9cd9-5cc9e2096b4d

This paper contains the first descriptions of an assemblage of non-ammonoid cephalopods from the Upper Asselian-Lower Sakmarian limestones of the Shakh-Tau reef. In this paper 16 species of 15 genera belonging to orders Oncocerida (?) and Nautilida (suborders Nautilina and Liroceratina), of which seven genera and 13 species are new, are described. Two species are described in open nomenclature. Representatives of the order Oncocerida (?), as well as the nautilids of the genera Temnocheilus and Megaglossoceras, were found for the first time in the Early Permian (until now it was believed that they became extinct in the Carboniferous).

Axophyllum cavum Trautschold, 1879 (Rugosa: Stauriida: Axophyllidae), a common solitary coral in the Moscovian stage of the Moscow Basin, is briefly redescribed. It is characterized by the small size of adult individuals (a few centimeters) and a high occurrence frequency of rootlet structures on the epitheca that supported the attachment of the coral on the substrate. The extensive collection of small-sized A. cavum (up to 2 cm) with various rhizoid structures and attachment scars from the Staryi Yam locality (Moscow area, Pakhra River, abandoned Yamskoi Quarry) was studied. The morphogenesis of rhizoids was is described in detail.