Revision Of Late Devonian (Famennian) And Some Early Carboniferous (Tournaisian) Crinoids And Blastoids From The Type Devonian Area Of North Devon

The crinoids and blastoids from the Pilton (Beds) Formation of the type Devonian of north Devonshire are revised. These fossils were monographed by the Rev. G. F. Whidborne in 1898, but have not been studied since that time. Recent studies on various groups of fossils from the Pilton and related rocks in North Devon confirm that the great majority of these fossils are Famennian, although three specimens from Fremington are probably Early Carboniferous (Tournaisian). We identify four blastoid taxa from a fauna that is sparse and poorly preserved; two spiraculates, one fissiculate, and one taxon unidentifiable at the ordinal level. Mesoblastus cf. M. crenulatus from the Gattendorfia Zone (Lower Carboniferous) near Fremington is the oldest known representative of this genus. The crinoid fauna is somewhat more diverse, but the preservation is equally poor. No changes are made in the flexible crinoids. Among camerate crinoids, one species is reassigned to Eumorphocrinus and one is retained in Actinocrinites. Specimens of some crinoids, such as Rhodocrinites and Megistocrinus, are so poorly preserved that certain identification was not possible. The hexacrinoid Adelocrinus, relegated to uncertainty for 150 years, is here shown to be a valid genus that is very similar to Arthroacantha, but not synonymous with it. Among the cladid crinoids, the dominant groups are those within the Superfamily Scytalocrinacea, which includes Bridgerocrinus, Sostronocrinus, and Scytalocrinus, all of which are placed in the new family Sostronocrinidae. One new species, Glossocrinus whidbornei, is named. Non‐pinnulate cladids, common in older Devonian rocks, do not occur. The fauna shows considerable similarity with faunas from eastern North America and Germany. It shows less resemblance to the extensive Famennian crinoid and blastoid fauna of north‐western China, despite some remarkable congruencies, especially the occurrence of very similar species of Actinocrinites in these widely separated areas.     

Discovery of Lower Famennian Stromatoporoidea in the Kuznetsk Basin

Stromatoporoidea recorded for the first time from the Lower Famennian of the Kuznetsk Basin are described. The assemblage studied includes three new species, Actinostroma quasifenestratum sp. nov., Kyklopora tomiensis sp. nov., and Coenostroma rarum sp. nov., and three previously known species of different genera.     

Late Famennian gastropoda from south-west England

The gastropod fauna of the Upper Devonian Baggy and Pilton formations in south‐west England is revised and includes some 30 taxa. The topmost part of the Upper Famennian succession in Devon is represented by clastic near‐shore and shallow shelf sediments, indicating a short‐term transgressive phase (‘Strunian Transgression’). The sequence yields a highly diverse fauna dominated by brachiopods and ostracodes, locally supplemented by crinoids, bryozoans, trilobites and molluscs. The taxa ‘Patellostium’britannicum sp. nov., Angyomphalus (Angyomphalus) junius sp. nov. and Dictyotomaria eurocapillaria sp. nov. are erected; a junior homonym is replaced by Macrochilina? piltonensis nom. nov. The gastropod fauna displays an independent character, where latest Devonian faunal elements overlap with Late Palaeozoic taxa expressing a transition similar to that of the bivalves, brachiopods, echinoderms and corals, without a sharp faunal break at the Devonian/Carboniferous boundary. Apart from the Caenogastropoda, all subclasses of gastropods are represented. Members of the bellerophontoids, pleurotomarioids and loxonematoids are most abundant, followed by murchisonioids, naticimorphs, euomphalomorphs and platyceratoids. The various gastropod groups represent different ecological demands and trophic categories, and together with the accompanying fauna indicate that nearly all habitats and niches were occupied in the shallow South Laurussian Shelf.     

New bryozoans (Bryozoa) from the Upper Devonian (Lower Famennian) of the Kuznetsk Basin

New bryozoans from the Upper Devonian (Famennian) of the borderlands of the Kuznetsk Coal Basin are described: Orthopora tomensis sp. nov. and Minussina incrustata sp. nov.     

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.     

Spiriferide brachiopods from the Famennian (Late Devonian) Hongguleleng Formation of western Junggar,Xinjiang, northwestern China

Junggar is one of the most important areas in the study of the Upper Devonian and Lower Carboniferous in the Junggar-Hinggan Stratigraphic Region. Nevertheless, abundant benthic fossils await modern taxonomic studies. In this paper, six genera and eight species of Famennian spiriferide brachiopods are described from the Duguer Member and its equivalents of the Hongguleleng Formation in western Junggar, adding new data to the Rugaltarostrum Subassemblage of the “Palaeospirifer”–Megalopterorhynchus Brachiopod Assemblage of this area. Five previously recognized species within this subassemblage are revised, including: Cyrtospirifer junggarensis F.M. Zhang [now Ulbospirifer? junggarensis (F.M. Zhang)]; Tenticospirifer tenticulum transversus F.M. Zhang [now Cyrtospirifer transversus (F.M. Zhang)]; Tenticospirifer koketekensis F.M. Zhang [now Cyrtospirifer koketekensis (F.M. Zhang)]; Mucrospirifer quadratus F.M. Zhang (now “Mucrospirifer” quadratus F.M. Zhang); and Mucrospirifer bouchadi (Muir-Wood) (now Tylothyris cf. novamexicana Stainbrook). Three additional species, Cyrtiorina houi n. sp., Goungjunspirifer sinicus F.M. Zhang, and Cyrtospirifer procumbens Simorin in Litvinovich et al., are described. Ulbospirifer? junggarensis and Goungjunspirifer sinicus are characterized by a prismatic layer. Cyrtiorina houi shows a moderately high and triangular ventral interarea, with delthyrium covered by pseudodeltidium with a rounded foramen. Cyrtospirifer procumbens can be distinguished from other species of this genus by its shallow, clearly limited sulcus with prominent primary plications, and high ventral interarea. Cyrtospirifer transversus has an alate to subpentagonal outline. Cyrtospirifer koketekensis has rounded and distinct flank plications. The ventral delthyrium of “Mucrospirifer” quadratus is covered by a pseudodeltidium. Tylothyris cf. novamexicana develops an angular sulcus in cross-section. Comparisons with the coeval brachiopod faunas from various regions indicate that, biogeographically, western Junggar has a close relationship with the Tarbagatai Mountain Range, northeastern Kazakhstan, central Kazakhstan, and Karaganda Basin, and a certain affinity with North America.     

Biostratigraphical value of some Strunian (Devonian, uppermost Famennian) Productidina, Rhynchonellida, Spiriferida, Spiriferinida brachiopods

The space of Devonian time considered in this paper corresponds to the uppermost part of the Famennian or the latest Famennian as a fourfold subdivision of the stage, usually called by many authors Strunian in neritic facies and Wocklumeria Stufe or Wocklumian in pelagic facies. Here, we examine the biostratigraphical value of certain brachiopod genera and species as bio-markers of the uppermost Famennian throughout the world (Europe, the former USSR, Middle East, Asia, North America, North Africa, and Australia). We have focused our study for species which stratigraphical range has been firmly established in correlation with the conodont biozones (Upper expansa, Early, Middle and Late praesulcata), the ammonoid biozones (do VI = “Wocklumeria Stufe”), or the foraminifera biozones (kobeitusana Zone). Other data for which the stratigraphical range is not based on standard biozonation are provided in the annex. The brachiopod bio-markers discussed in this paper belong to 14 Productidina genera (Acanthatia, Araksalosia, Ericiata, Hamlingella, Mesoplica, Nigerinoplica, Orbinaria, Ovatia, Rugauris, Semiproductus, Sentosia, Spinocarinifera, Steinhagella, Whidbornella), 7 Rhynchonellida genera (Araratella, Centrorhynchus, Hadyrhyncha, Megalopterorhynchus, Novaplatirostrum, Rozmanaria, Tchanakhtchirostrum), 13 Spiriferida genera (Brachythyris, Cyrtospirifer, Dichospirifer, Eochoristites, Imbrexia, Parallelora, Prospira, Rigauxia, Sphenospira, Tenisia, Toryniferella, Tylothyris, Voiseyella) and one Spiriferinida genus (Syringothyris). Other orders have not been studied in this paper. The main features of the uppermost Famennian brachiopod taxa represented in this paper are commented. All these taxa are listed as completely as possible throughout the world (with complements in the annex). A quick summary of the geographic distribution of the more represented taxa is given in conclusion.     

Extinction of nanicellid foraminifera during the Frasnian–Famennian biotic crisis: some far‐reaching evolutionary consequences

This study evaluates the severity of the poorly known and mostly underestimated foraminiferal extinction during the Frasnian–Famennian biotic crisis and its evolutionary aftermath. During this global event, worldwide, truly plurilocular planispiral (Nanicellidae) and uniseriate, palmate (Semitextulariidae) foraminifera associated with metazoan reefs died out entirely. Highly advanced test morphology such as that of nanicellids did not reappear in the earth's history until the Late Triassic. Moreover, morphotype comparable to that of the Devonian bilaterally flattened and palmate semitextularids appeared again until the Middle Jurassic (Frondicularia, Lagenida). In terms of the degree of test septation and chamber arrangement as well as general test shape, these foraminifera were ‘very far ahead of their time’. In consequence, foraminifera suffered a significant collapse during the F‐F biodiversity crisis, leading to an amazingly long evolutionary time lag in the case of plurilocular foraminifera lasting at least 150 million years.     

Non-calcified macroalgae from the lower Famennian (Upper Devonian) of the Holy Cross Mountains,Poland

Non-calcified algal remains were collected from the lower Famennian deposits at the Kowala quarry in the Holy Cross Mountains, central Poland. Each specimen is represented solely by vegetative organs. On the basis of the material collected, a new algal genus and species has been formally erected. Vittella dixii nov. gen., nov. sp. (Order ?Bryopsidales) includes thalli consisting of long bundles of hair-like filament structures arranged in parallel fashion to the thallus length. Some other algal remains are regarded as Inocaulis-like forms. They possess long, flat blades with entire margins, and are ornamented with very dense, hair-like projections. The rest of the specimens found so far has been illustrated and tentatively classified as probably belonging to either Bryopsidales or Dasycladales algae. The exceptional preservation of the non-calcified algal thalli investigated suggests that special taphonomic conditions have prevailed during sedimentation of the lower part of the Famennian Kowala section. These may have involved rather rapid burial in oxygen-restricted (dysoxic and anoxic) sea-bottom environment, inhibiting the activities of scavengers and bioturbators, as well as oxidative degradation. The preservation of the algae, together with sedimentological and geochemical characteristics of the host rocks indicate that they rather represent parautochthonous assemblage, buried close to their natural habitat.     

Phyllocarid remains (Malacostraca, Crustacea) from the basal Famennian (Late Devonian) of the Montagne Noire (Southern France)

From the early Famennian of the Montagne Noire 19 different morphotypes of probably phyllocarid (Crustacea) affinities are described. Apart from formerly unknown enamel-like structures that veil the uppermost tips of the denticles on most of the gnathal lobe morphs, the stratigraphic implications concerning extinction behaviour across the Kellwasser mass extinction event and during early Famennian diversification are discussed. In addition, phyllocarids in the Famennian seem to form a useful stratigraphic tool for biostratigraphical correlation. Therefore, three basal Famennian phyllocarid assemblage zones are distinguished.      

A late-devonian (Famennian)Renalcis-Epiphyton reef at Zhaijiang, Guilin, South China

Summary Microbial reefs, together with stromatolitic mounds and ooid shoals, constitute massive limestones in Famennian platform marginal strata in Guilin, in sharp contrast to the well-known coral-stromatoporoid reefs in the Givetian and Frasnian. Microbes played a significant and important role as stabilizers in the Famennian carbonate deposits of Guilin. A reef at Zhaijiang was constructed byEpiphyton andRenalcis, and is representative of such carbonate buildups. The reef is situated 10 km west of Guilin and corresponds to a microbe-dominated platform margin carbonate complex. Organisms in the Zhaijiang microbial reef are low diversity and dominated by ostracods and two genera of microbes,Epiphyton andRenalcis. Other microbial genera such asSphaerocodium andWetheredella occur in most of reef facies in Guilin, but their role as reef builder is doubtful because they occur only in minor amounts. The same four genera occur in volumetrically significant amounts in the upper Devonian carbonate complexes of Alberta. Canada and Western Australia. However.Epiphyton is more abundant in the Guilin reefs. The Zhaijiang microbial reef developed above Famennian proximal slope faices, as suggested by reef architecture and paleogeographic setting. The facies sequence of the microbial reef can be divided into three parts. The lower part is composed of medium-bedded bioclastic grainstones with a few microbial framestone lithoclasts, representing a proximal slope facies. The middle part consists of thin-bedded mudstone and shale with limestone lenses that are thought to be low stand deposits. In some cross sections, mudstone and shale infilled tidal channels that developed in the bioclastic grainstones.Renalcis-Epiphyton framestone constitutes the upper part with massive stacking patterns. The reef is 35 m thick and over 50 m in width. Nine litho- and biofacies are recognized. Zhaijiang reef provides an example of a binder guild-dominated buildup in the almost vacant reef ecosystem of the Famennian and represents a characteristic kind of reef after the Frasnian/Famennian extinction.     

Epizoans immured in the heterocoral Oligophylloides maroccanus Weyer, 2017: a unique record from the Famennian (Upper Devonian) of Morocco

Exceptionally well-preserved proximal parts of colonies of the late Famennian heterocoral Oligophylloides maroccanus Weyer, 2017 with immured epizoans are described here from the Jebel Bou Ifarherioun ridge of the eastern Anti-Atlas, Morocco. This is the first evidence of unique syn vivo interactions between a heterocoral and numerous epizoans, that is the colonial tabulate corals Zemmourella? sp. and Aulopora? sp.; solitary rugose corals Czarnockia? sp. and Gorizdronia? sp.; and other juvenile, undeterminable solitary rugosans, crinoids, as well as microbial structures. Detailed analysis of sectioned specimens allowed mutual interactions to be traced during growth of the host and infesting encrusting taxa. The studied associations show that the epizoans must have settled on the proximal parts of Oligophylloides in places devoid of soft tissue. The encrusted proximal portions of the colonies show that soft tissue of the heterocoral could have retracted and expanded again, fouling and immuring dead epizoans. The process of bioimmuration is perfectly marked by thin layers of microbial encrustations developing on the epibiont skeletons prior to their overgrowth by the skeleton of the heterocoral. The studied encrustations seem to be widespread, but the association provided the epizoans with additional substrate for encrustation, as well as an elevated position above the seafloor, beneficial for access to nutrients. We suggest that the host bioimmured the encrusters after their death, because of noticeable microbial encrustations and veneers of sediment, a lack of malformations, relatively rapid growth and aggressive activity of epizoans.     

Evidence for shallow‐water ‘Upper Kellwasser’ anoxia in the Frasnian–Famennian reefs of Alberta,Canada

The Frasnian–Famennian extinction witnessed the global devastation of both level‐bottom and reef communities in low latitudes. Marine extinctions in offshore level‐bottom communities are associated with two widespread, transgressive, anoxic ‘Kellwasser Events’ that support an anoxia–extinction link. Typical Kellwasser facies of bituminous limestones and shales are not obviously recorded in shallow‐water settings, and thus, it is unclear whether anoxia played a role in reef losses. We evaluate geochemical, petrographic and facies evidence for oxygen restriction from an extremely shallow‐water carbonate platform in Alberta. Sequence stratigraphy places the Frasnian–Famennian boundary at a sequence boundary that tops a laminated mudstone and interrupts carbonate platform deposition. Two transgressive pulses have been identified, one of which is associated with the second, major transgression of T‐R cycle IId of the Devonian eustatic sea‐level curve. Geochemical proxies indicate that these transgressions were accompanied by influx of dysoxic or anoxic waters. Organic carbon and U enrichment in the Frasnian, particularly just below the Frasnian–Famennian boundary, points to episodic dysoxic conditions that probably persisted into the basal Famennian and were coincidental with the global Upper Kellwasser Event. This study provides the first evidence for the smoking gun of an anoxia‐driven extinction in very shallow waters, implicating this potent killer in the demise of the Devonian reefs.     

Eustatic trends in conodont diversity across the Frasnian–Famennian boundary in the stratotype area, Montagne Noire, Southern France

Trends in generic diversity of successive conodont communities are analysed in sections of different environmental settings across the Frasnian–Famennian (F–F) boundary in the stratotype area, Montagne Noire, France. The evolution of conodont biofacies and abundances matches the overall pattern already observed in many sections elsewhere in the world and supports the interpretation of an important eustatic sea-level fall during the Upper Kellwasser event. The change from late Frasnian deep-water palmatolepid–polygnathid biofacies to shallower-water polygnathid-icriodid biofacies during the Upper Kellwasser event occurred in all sections studied. The shallowing trend culminated at the end of the Kellwasser Event as indicated by the substantial increase of formerly poorly represented icriodids, whereas palmatolepids concomi-tantly diminished. This event occurred earlier on oxygenated outer platform submarine rises than in oxygen-depleted depressions. The sudden sea-level fall prior to the Frasnian–Famennian boundary was followed, at the beginning of the Famennian, by a deepening trend when palmatolepids dominated again. These changes in conodont generic associations and abundances occurred rapidly and synchronously. As a result, the stratigraphic resolution obtained with the evolution of biofacies is higher: it permits not only a more accurate location of the base of the Upper Kellwasser event in environments where it cannot be distinguished lithologically, but it also allows the recognition of intrazonal gaps.     

New discoveries of tetrapods (ichthyostegid‐like and whatcheeriid‐like) in the Famennian (Late Devonian) localities of Strud and Becco (Belgium)

The origin of tetrapods is one of the key events in vertebrate history. The oldest tetrapod body fossils are Late Devonian (Frasnian–Famennian) in age, most of them consisting of rare isolated bone elements. Here we describe tetrapod remains from two Famennian localities from Belgium: Strud, in the Province of Namur, and Becco, in the Province of Liège. The newly collected material consists of an isolated complete postorbital, fragments of two maxillae, and one putative partial cleithrum, all from Strud, and an almost complete maxilla from Becco. The two incomplete maxillae and cleithrum from Strud, together with the lower jaw previously recorded from this site, closely resemble the genus Ichthyostega, initially described from East Greenland. The postorbital from Strud and the maxilla from Becco do not resemble the genus Ichthyostega. They show several derived anatomical characters allowing their tentative assignment to a whatcheeriid‐grade group. The new tetrapod records show that there are at least two tetrapod taxa in Belgium and almost certainly two different tetrapod taxa at Strud. This locality joins the group of Devonian tetrapod‐bearing localities yielding more than one tetrapod taxon, confirming that environments favourable to early tetrapod life were often colonized by several tetrapod taxa.     

Progressive drowning of carbonate platform in the Moravo-Silesian Basin (Czech Republic) before the Frasnian/Famennian event: facies,compositional variations and gamma-ray spectrometry

The Moravo-Silesian Basin (MSB; eastern Czech Republic and southern Poland) hosted an extensive shallow-water carbonate platform in the Middle Devonian to Frasnian interval. The platform drowned in a stepwise fashion from the Palmatolepis hassi to the Pa. linguiformis zone. Three types of drowning successions were revealed from conodont biostratigraphy, facies, microfacies and gamma-ray spectrometry data: (A) drowning to periplatform turbidite setting; (B) drowning to (hemi)pelagic seamount setting and (C) drowning associated with the stratigraphical gap. In the lower Pa. hassi zone, rapid subsidence caused the platform to drown locally along the N–S to NW–SE trending faults (type A drowning). In the upper Pa. rhenana to the Pa. linguiformis zone, the drowning accelerated in the western part of the MSB due to locally higher subsidence rates combined with the Late Frasnian biotic crisis (type B). In the southern part of the basin, the platform emerged shortly before the Frasnian/Famennian (F/F) boundary and drowned in the Early to Late Famennian (type C). The primary cause of drowning was differential subsidence at the Laurussian passive margin. Eustatic sea-level fluctuations, if any, contributed only to a minor extent to the Late Frasnian drowning, but were effective in type C drowning during the Famennian. The drowning boundaries are associated with increased contents of K and Th, reflecting the deceleration of carbonate production. Uranium contents display isolated peaks that roughly correlate with the drowning boundaries or the stratigraphic gaps associated with the F/F boundary. The uranium contents are considered to reflect local depositional conditions and are not suitable for stratigraphic correlation. On the other hand, from the K and Th contents, we can infer Late Frasnian sea-level fluctuations with duration on the order of 1 Myr. These cyclic variations in K and Th contents proved to be useful in platform-to-basin stratigraphic correlation.     

Characterization of the placoderm (Gnathostomata) assemblage from the tetrapod‐bearing locality of Strud (Belgium,upper Famennian)

The placoderm fauna of the late Famennian tetrapod‐bearing locality of Strud, Belgium, is studied on the basis of historical and newly collected material. It includes the previously described antiarch Grossilepis rikiki, the groenlandaspidid Turrisaspis strudensis sp. nov. and the actinolepidoideid Phyllolepis undulata. P. undulata is thoroughly described and joins the list of the valid Phyllolepis species confidently diagnosed. A morphometrical analysis performed on the centronuchal and anterior ventrolateral plates of the Phyllolepis material demonstrates that there is only one species of Phyllolepis in Belgium (thus, Phyllolepis konincki becomes a junior synonym of P. undulata), that P. rossimontina (Pennsylvania) is a synonym of P. undulata and that the unity of the genus Phyllolepis is strongly supported, although the characterization of several species within this genus is blurred. The strong resemblance between the faunal compositions in Strud and Red Hill (Pennsylvania, USA) suggests important faunal exchanges between these regions of the Euramerica landmass.     

The oldest peracarid crustacean reveals a Late Devonian freshwater colonization by isopod relatives

Peracarida (e.g. woodlice and side-swimmers) are, together with their sister-group Eucarida (e.g. krill and decapods), the most speciose group of modern crustaceans, suggested to have appeared as early as the Ordovician. While eucarids'' incursion onto land consists of mainly freshwater and littoral grounds, some peracarids have evolved fully terrestrial ground-crawling ecologies, inhabiting even our gardens in temperate regions (e.g. pillbugs and sowbugs). Their fossil record extends back to the Carboniferous and consists mainly of marine occurrences. Here, we provide a complete re-analysis of a fossil arthropod—Oxyuropoda—reported in 1908 from the Late Devonian floodplains of Ireland, and left with unresolved systematic affinities despite a century of attempts at identification. Known from a single specimen preserved in two dimensions, we analysed its anatomy using digital microscopy and multispectral macroimaging to enhance the contrast of morphological structures. The new anatomical characters and completeness of Oxyuropoda, together with a phylogenetic analysis with representatives of all major Eumalacostraca groups, indicate that Oxyuropoda is a crown peracarid, part of a clade including amphipods and isopods. As such, Oxyuropoda is the oldest known species Peracarida, and provides evidence that derived peracarids had an incursion into freshwater and terrestrial environments as early as the Famennian, more than 360 Ma.