<Emphasis Type="Italic">Oredezhosteus</Emphasis>, a new psammosteid genus (Heterostraci,Psammosteiformes) from the Lower Frasnian (Upper Devonian) of the Main Devonian Field

A new psammosteid, Oredezhosteus kuleshovi gen. et sp. nov., from the Amata Regional Stage (Lower Frasnian) of the Leningrad Region is described. In addition, Psammosteus livonicus Obruchev from beds of the same age from Latvia is included in the new genus. The major diagnostic characters distinguishing Oredezhosteus gen. nov. from Psammosteus Agassiz are the large rounded striated tubercles, which cover more than half width of the branchials, and the long base of the branchial plates.     

A new pachyosteomorph arthrodire (Pisces: Placodermi) from the Upper Frasnian of the Central Devonian Field,Russia

A new genus and species of pachyosteomorph arthrodire, Omalosteus krutoensis gen. et sp. nov. from shallow-water marine deposits of the Evlanovo Regional Stage (Upper Frasnian, Upper Devonian) of the Central Devonian Field is described. It is tentatively referred to the family Trematosteidae Gross, 1932, which was earlier believed to be endemic to central Europe. The new genus is characterized by a rounded, dorsoventrally compressed cross section of the pectoral region and smooth head shield and postcranial membrane bones. The centrale is elongated, forming a narrow lateral lobe. The praeorbitale, postorbitale, and centrale come in contact in the center of the orbital region. The mediodorsale is wide, its carinal process projects considerably posteriorly. The anterior margin of the anterior ventrolaterale lacks articular facets for the interlaterale.     

The placoderm <Emphasis Type="Italic">Plourdosteus livonicus</Emphasis> (Eastman) in the early Frasnian of the Central Devonian Field and the trophic structure of the Mikhailovskii Fish Assemblage

The placoderm Plourdosteus livonicus (Eastman) (Euarthrodira, Coccosteina) from the Upper Timanian Mikhailovskii Fish Assemblage of the Central Devonian Field is described. The composition and structure of the assemblage are discussed and compared with later Frasnian fish assemblages of the Central Devonian Field.     

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.     

Psammosteiforms (Agnatha,Heterostraci) from the Lower Frasnian of the Mikhailovskii Mine,Kursk Region,Russia

Psammosteiforms, including Tartuosteus (?) zheleznogorskensis, sp. nov. and Psammosteus cf. P. praecursor Obruchev, from the Lower Frasnian (Upper Devonian) of the Mikhailovskii Mine (Kursk Region) are described. Agnathans, placoderms, crossopterygians, and acanthodians from this locality are determined. The similarity of the Mikhailovskii Fish Assemblage to the assemblage of southern Timan suggests it be assigned to the Upper Timan Regional Stage.     

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.     

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.     

Indigenous demosponge spicules in a Late Devonian stromatoporoid basal skeleton from the Frasnian of Belgium

This paper records the first example of a demosponge spicule framework in a single specimen of a Devonian stromatoporoid from the Frasnian of southern Belgium. The small sample (2.5 × 2 cm) is a component in a brecciated carbonate from a carbonate mound in La Boverie Quarry 30 km east of Dinant. Because of the small size of the sample, generic identification is not confirmed, but the stromatoporoid basal skeleton is similar to the genus Stromatopora. The spicules are arranged in the calcified skeleton, but not in the gallery space, and are recrystallized as multi‐crystalline calcite. The spicules fall into two size ranges: 10–20 μm diameter and 500–2000 μm long for the large ones and between 5–15 μm diameter and 50–100 μm length for the small ones. In tangential section, the spicules are circular, they have a simple structure, and no axial canal has been preserved. The large spicules are always monaxons, straight or slightly curved styles or strongyles. The spicules most closely resemble halichondrid/axinellid demosponge spicules and are important rare evidence of the existence of spicules in Palaeozoic stromatoporoids, reinforcing the interpretation that stromatoporoids were sponges. The basal skeleton may have had an aragonitic spherulitic mineralogy. Furthermore, the spicules indicate that this stromatoporoid sample is a demosponge.     

The youngest African clymeniids (Ammonoidea, Late Devonian) – failed survivors of the Hangenberg Event

A co-occurrence of the clymeniid Postclymenia evoluta Schmidt, 1924 and the goniatite Acutimitoceras hilarum Korn, 2002 is reported from the Anti-Atlas of Morocco. Both species occur in the same limestone horizon within the Acutimitoceras prorsum Zone, that has yielded an exclusive conodont fauna of the Upper praesulcata Zone (latest Devonian, above the Hangenberg Black Shale). This record is firm evidence that some clymeniids survived the global Hangenberg Event, but soon later became extinct without descendants.     

Fossil plants with spores in the sporangia from the Upper Devonian (Frasnian) deposits of northern Timan

Fossil plants from the Upper Devonian (Frasnian) of northern Timan are studied. The sporangia contain well-preserved spores, which were studied in transmitted light using a scanning electron microscope. The genus Gutzeitia S. Snigirevsky, gen. nov. is established. Macroremains and in situ spores of G. timanica (Petros.) S. Snig., comb. nov. and ?Cephalopteris mirabilis (Nath.) Nath. are described. Microspores in Dimeripteris gracilis Schmalh. are identified, and microspores and megaspores in ?Cephalopteris mirabilis are studied for the first time.     

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.     

A late Devonian (Frasnian) coral-bafflestone reef at Houshan in Guilin, South China

Summary Givetian to early Carboniferous sediments of South China are characterized by carbonates. Middle and Late Devonian strata are best developed in the Guilin area. Reefs and organic shoals are recorded by various lithofacies types indicating the existence of an extended carbonate platform and a change of the composition of reef communities in time. Starting in the late Devonian, stromatoporoids and corals were replaced by algae that subsequently played an important role together with stromatoporoids, receptaculitids and fasciculate rugose corals in reef communities. In Houshan, 5 km west of Guilin, a coral-bafflestone reef occurs in the Frasnian strata, situated near an offshore algal-stromatoporoid reef. The coral reef was formed in a back-reef area adjacent to the inner platform margin. The coral-bafflestone reef is unique among the late Devonian reefs of South China with regard to the biotic composition. The reef is composed of fasciculate colonies ofSmithiphyllum guilinense n. sp. embedded within in packstones and wackestones. The height of colonies reaches 1 m. The community is low-diverse. The species ofSmithiphyllum occurring in the Frasnian reef complexes of Guilin exhibit a distinct facies control:Smithiphyllum guilinense occurs in or near to margin facies and formed bafflestone, constituting a coral reef whereasSmithiphyllum occidentale Sorauf, 1972 andSmithiphyllum sp.—characterized by small colonies with thin corallites—are restricted to the back-reef and marginal slope facies. The bush-like coral colonies baffled sediments. Algae and stromatoporoids (mainlyStachyodes) are other reef biota. Reef-dwelling organisms are dominated by brachiopods. The reefs are composed from base to top of five lithofacies types: 1) cryptalgal micrite, 2) peloidal packstone, 3) stromatactis limestone, 4) coral-bafflestone, and 5) pseudopeloidal packstone. The reef complex can be subdivided into back-reef subfacies, reef flat and marginal subfacies, and marginal fore-slope subfacies. The Houshan coral-bafflestone reef is not a barrier reef but a coral patch reef located near the inner margin of a carbonate platform.     

Stromatoporoid palaeoecology in the Frasnian (Upper Devonian) Belgian platform,and its applications in interpretation of carbonate platform environments

Abstract: Stromatoporoid faunas in the Frasnian of southern Belgium are abundant in the carbonate platform environments present in this area. Stromatoporoids dominate the large skeletal organisms and occur principally in biostromes. The stromatoporoid assemblage is represented by a small number of taxa. Stromatoporoid genera include Actinostroma, Amphipora, Atelodictyon, Clathrocoilona, Salairella, Stachyodes, Stictostroma, Stromatopora and Trupetostroma which are present in environments ranging from the outer, outer intermediate, inner intermediate and inner zones and associated biostromes. Most large skeletal stromatoporoids are low profile, which reinforces the conclusions of previous studies that low‐profile growth forms were the most successful stromatoporoid forms. These low‐profile forms are likely to have been important sediment stabilisers that may have led to expansion of the carbonate factory. Growth forms vary between facies, indicating some degree of environmental control on form; for example, laminar in the intermediate zone, bulbous and domical in the inner and outer zones. Stromatoporoid taxa vary in occurrence across the environmental gradient from shallow to deep. There is some taxonomic control on growth forms, with some taxa showing more variability than others in different environments.     

Upper Devonian Sponges from the Holy Cross Mountains, Central Poland

The rich fauna of Late Devonian (Late Frasnian) siliceous sponges from the Holy Cross Mountains, Poland is composed of 15 species and 11 genera. Both astylospongid demosponges (lithistids) and hexactinosan hexactinellids are present. The following new genera and/or species are proposed: D regulara Rigby and Pisera sp. nov., Jazwicella media Rigby and Pisera gen. et sp. nov., Astyloscyphia irregularia Rigby and Pisera gen. et sp. nov., A. turbinata Rigby and Pisera gen. et sp. nov., Astylotuba modica Rigby and Pisera gen. et sp. nov., Paleoregulara cupula Rigby and Pisera gen. et sp. nov., Paleoramospongia bifurcata Rigby and Pisera gen. et sp. nov., Cordiospongia conica Rigby and Pisera gen. et sp. nov., Paleocraticularia elongata Rigby and Pisera gen. et sp. nov., P gigantia Rigby and Pisera gen. et sp. nov., Polonospongiadevonica Rigby and Pisera gen. et sp. nov., P fistulata Rigby and Pisera gen. et sp. nov., Urnospongia modica Rigby and Pisera gen. et sp. nov., and Conicospongia annulata Rigby and Pisera gen. et sp. nov. The investigated fauna contains the youngest astylospongiids known and the oldest well-preserved, and most diversified Palaeozoic hexactinosans. The sponge fauna constituted a significant element of a brachiopod-coral-sponge assemblage that inhabited a deep slope of the local Dyminy Reef structure, during its final phase of growth, in a clearly hemipelagic setting. This fauna is limited to the intrashelf depression within an incipiently drowned carbonate platform.     

Patterns of evolution and extinction in proetid trilobites during the late Devonian mass extinction event,Canning Basin,Western Australia

Abstract: In the early Late Devonian, terminal Frasnian proetid trilobites have previously only been known from Europe and North Africa. For the first time, a rich fauna of late Frasnian proetids is described from the Virgin Hills Formation, Canning Basin, Western Australia. Seventeen species in six genera are described, of which three are new: Rudybole gen. nov., Palpebralina gen. nov. and Canningbole gen. nov. A new subgenus, Chlupaciparia (Australoparia) subgen. nov. is also described. Fourteen of the species are new: Palpebralia initialis sp. nov., P. pustulata sp. nov., ?P. sp. nov. A, Rudybole depressa sp. nov., Palpebralina pseudopalpebralis sp. nov. (comprising the subspecies P. pseudopalpebralis pseudopalpebralis subsp. nov. and P. pseudopalpebralis ultima subsp. nov.), P. minor sp. nov., P. ocellifer sp. nov., Canningbole latimargo sp. nov., C. henwoodorum sp. nov., C. macromma sp. nov., Pteroparia extrema sp. nov., Chlupaciparia (Chlupaciparia) planiops sp. nov., Chlupaciparia (Australoparia) australis sp. nov. and C. (Australoparia) lata sp. nov. The subspecies Rudybole adorfensis angusta subsp. nov. is also described. The proetids range through conodont Zones 11–13b and terminate at the Upper Kellwasser Event, which marks the terminal Frasnian mass extinction event. Three of the six proetid lineages, Palpebralia, Canningbole and Pteroparia, show evolutionary trends of eye reduction. Two of the remaining lineages, Rudybole and Palpebralina, consist exclusively of blind taxa. The last, Chlupaciparia, also comprises forms with reduced eyes. The proetids show a stepped pattern of extinction during the late Frasnian, which correlate with two Kellwasser biocrises documented in European/North African Frasnian sections. The highest diversity preceded the Lower Kellwasser event that occurred at the end of conodont Zone 12 and saw the extinction of all species present in that zone. However, only one genus, Pteroparia, locally became extinct. A major higher‐level taxonomic mass extinction at the top of Zone 13b initiated the Upper Kellwasser extinction event. This included extinction at the generic level, with all five remaining genera becoming extinct, and at the family level, with the loss of the Tropidocoryphidae.     

Famennian (Late Devonian) ammonoids from the Ouarourout section (Saoura Valley,Algeria)

In North Africa, the ammonoids constitute an important part of the Devonian marine macro-invertebrates. New material has been recently collected in the ‘Argiles de Marhouma’ Formation from several sections located near Béni-Abbès in the Saoura Valley (Ougarta Mountains, Algerian Sahara). Red nodular limestones (i.e., ‘griotte’ facies) characterize this formation rich in ammonoids. The assemblages from the Ouarourout section are composed of Goniatitida and Clymeniida constituting 30 species and 19 genera belonging to 11 families, with Cyrtoclymenia, Erfoudites, Kosmoclymenia, Planitornoceras, Platyclymenia, and Prionoceras, as the most abundant genera. The taxonomic comparison and the biostratigraphic correlations are made by considering the assemblages recorded in the Anti-Atlas of Morocco, which are very close to those observed in Algeria. In the Ouarourout area, the ‘Argiles de Marhouma’ Formation contains middle and late Famennian ammonoid assemblages and three biostratigraphic intervals can be identified: Planitornoceras, Platyclymenia and Medioclymenia.     

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.     

Palynostratigraphy of the Frasnian-Famennian boundary deposits from the Central Devonian Field, western Russia and comparisons with adjacent areas

Detailed palynological research carried out on samples from the three quarries at Khlevnoye and Kamenka (Voronezh region) permitted recognition of the Frasnian-Famennian boundary in the lower Kamenka quarry. Two miospore zones, Cristatisporites deliquescens-Verrucosisporites evlanensis (DE) from the Frasnian and Cyrtospora cristifera-Diaphanospora zadonica (CZ) from the Famennian, have been recognized. Some faunistic data were also obtained from the F-F boundary deposits. Research confirmed the existence of a sedimentological gap between the Livny and Zadonsk horizons, including all of the Volgograd horizon. These new palynological results are compared with data from adjacent areas: Pripyat Depression (Belarus), Timan-Pechora Province and the Volgograd Volga region (Russia), western Pomerania and the Holy Cross Mountains (Poland).     

Severe extinction and rapid recovery of mammals across the Cretaceous–Palaeogene boundary,and the effects of rarity on patterns of extinction and recovery

The end‐Cretaceous mass extinction ranks among the most severe extinctions of all time; however, patterns of extinction and recovery remain incompletely understood. In particular, it is unclear how severe the extinction was, how rapid the recovery was and how sampling biases might affect our understanding of these processes. To better understand terrestrial extinction and recovery and how sampling influences these patterns, we collected data on the occurrence and abundance of fossil mammals to examine mammalian diversity across the K‐Pg boundary in North America. Our data show that the extinction was more severe and the recovery more rapid than previously thought. Extinction rates are markedly higher than previously estimated: of 59 species, four survived (93% species extinction, 86% of genera). Survival is correlated with geographic range size and abundance, with widespread, common species tending to survive. This creates a sampling artefact in which rare species are both more vulnerable to extinction and less likely to be recovered, such that the fossil record is inherently biased towards the survivors. The recovery was remarkably rapid. Within 300 000 years, local diversity recovered and regional diversity rose to twice Cretaceous levels, driven by increased endemicity; morphological disparity increased above levels observed in the Cretaceous. The speed of the recovery tends to be obscured by sampling effects; faunas show increased endemicity, such that a rapid, regional increase in diversity and disparity is not seen in geographically restricted studies. Sampling biases that operate against rare taxa appear to obscure the severity of extinction and the pace of recovery across the K‐Pg boundary, and similar biases may operate during other extinction events.