The fate of the homoctenids (Tentaculitoidea) during the Frasnian–Famennian mass extinction (Late Devonian)

The homoctenids (Tentaculitoidea) are small, conical‐shelled marine animals that are among the most abundant and widespread of all Late Devonian fossils. They were a principal casualty of the Frasnian–Famennian (F‐F, Late Devonian) mass extinction, and thus provide an insight into the extinction dynamics. Despite their abundance during the Late Devonian, they have been largely neglected by extinction studies. A number of Frasnian–Famennian boundary sections have been studied, in Poland, Germany, France, and the USA. These sections have yielded homoctenids, which allow precise recognition of the timing of the mass extinction. It is clear that the homoctenids almost disappear from the fossil record during the latest Frasnian ‘Upper Kellwasser Event’. The coincident extinction of this pelagic group, and the widespread development of intense marine anoxia within the water column, provides a causal link between anoxia and the F‐F extinction. Most notable is the sudden demise of a group, which had been present in rock‐forming densities, during this anoxic event. One new species, belonging to Homoctenus is described, but is not formally named here.     

Microconchid tubeworms across the upper Frasnian – lower Famennian interval in the Central Devonian Field,Russia

Abstract: Microconchid tubeworms (Tentaculita) encrusting brachiopod shells have been investigated from the upper Frasnian – lower Famennian (Upper Devonian) deposits of the Central Devonian Field, Russia. The condition of microconchids and associated encrusting taxa is reported for the first time from the early Famennian recovery interval (crepida Chron) following the Frasnian–Famennian mass extinction. Two species, one new (Palaeoconchus variabilis sp. nov.) and the second one in open nomenclature (Palaeoconchus sp.), are described. Compared to lower Famennian specimens, they seem to be preferentially grouped on the anterior parts of the brachiopod host shells, which are interpreted as the most suitable sites away from the sea‐bottom and sediment. During the late Frasnian (Late rhenana Chron), microconchids, outnumbered by cornulitids and as abundant as foraminifers, were also associated with trepostome bryozoans, tabulates, rugose corals and various problematic encrusters. During the early Famennian recovery interval encompassing the crepida Chron, microconchids greatly outnumbered all associated encrusters, including the previously dominant cornulitids, while foraminifers, tabulates and rugose corals vanished. Early Famennian microconchids, represented by the single, albeit very abundant, species Palaeoconchus variabilis sp. nov., were opportunists that rapidly colonised the environment during the ongoing transgression following the regression‐driven biotic crisis in the area of the Central Devonian Field. In comparison to their late Frasnian predecessors and even other Middle Devonian specimens, no size reduction (the so‐called Lilliput effect) of early Famennian microconchid tubes was observed. It is probable that microconchids either rapidly attained their ‘normal’ sizes or they did not suffer any dwarfism following the Frasnian–Famennian event.     

Ostracodes du Dévonien moyen et supérieur du Tafilalt (Maroc)

Forty-four ostracod species are recognised from the upper part of the Eifelian to the base of the Famennian of the Djebel Mech Irdane, Bou Tchrafine and El Atrous sections, in the Tafilalt. The majority are in open nomenclature, and one is new: Tubulibairdia tafilaltensis nov. sp. Ostracods belong principally to the Eifelian Mega-Assemblage and they are generally indicative of poorly oxygenated relatively deep marine environments below storm wave base. However, in the base of the Famennian of the Bou Tchrafine section, ostracods belong to the Myodocopid Mega-Assemblage indicative of strong hypoxic water conditions. They confirm that the hypoxic water conditions linked to the “Upper Kellwasser Horizon”, subsisted in North Africa, in the extreme base of the Famennian. Twenty-four ostracod species are recognized in the Djebel Mech Irdane Eifelian/Givetian stratotype, where the most important change in the ostracod fauna is related to the 40 cm-thick level of grey silty-marl located 10 cm below the boundary. However, this change is not indicative of an extinction event at this level. Three zones established on ostracods are recognized in the Frasnian and in the base of the Famennian of the Tafilalt.     

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.     

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.     

华南泥盆系法门阶浅海相四射珊瑚的组合序列

晚泥盆世弗拉期末F-F的生物灭绝事件,导致珊瑚以及许多浅海底栖生物灭绝,几乎整个法门期珊瑚群都处于残存阶段,华南地区目前只在湖南的个别地方发现少量的Smithiphyllum。直到法门期末珊瑚和其它一些造礁底栖生物开始复苏,出现了不少与典型泥盆纪珊瑚存在很大差别的新分子。华南泥盆纪最晚期(Strunian)的珊瑚可划分成上、下两个组合:上部Cystophrentis组合;下部Eocaninophyllum组合。泥盆—石炭纪之交的另一次生物灭绝事件,使新生的泥盆纪最晚期的珊瑚又遭灭绝。     

Were Phanerozoic mass extinctions among brachiopod superfamilies selective by taxa longevity?

All mass extinctions are characterized by certain kind of selectivity. An analysis of stratigraphic ranges of 112 brachiopod superfamilies implies that some Phanerozoic mass extinctions (Late Ordovician, Frasnian/Famennian and Devonian/Carboniferous, Early Jurassic, and Cretaceous/Paleogene) were selective by taxa longevity. They preferentially affected relatively old superfamilies and favoured a survival of relatively young superfamilies. No explanation of this selectivity as an apparent phenomenon is fully satisfactory. The Permian/Triassic mass extinction did not favour a survival of “young” superfamilies because of abnormally low rate of origination established since the Pennsylvanian and the absence of these “young” taxa. This study confirms tentatively a difference between Paleozoic and post-Paleozoic times by the importance of post-extinction recovery intervals for taxa longevity.     

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.     

Anatomical reinvestigation of Archaeopteris macilenta from the Upper Devonian (Frasnian) of South China

Abstract Archaeopteris macilenta is one of the most widespread plants in the Late Devonian. Based on fossils from the Frasnian Huangjiadeng Formation, Yichang District of Hubei Province, for the first time we study in detail the anatomy of this progymnosperm plant in South China. Ultimate axes are protostelic with three xylem sympodia and lack secondary tissue. Penultimate axes are eustelic, bearing eight sympodia and a thin band of secondary xylem. Radially symmetrical sympodia of mesarch primary xylem produce traces of appendages in a spiral arrangement. Archaeopteris macilenta and A. halliana (A. roemeriana) are dominant in the Frasnian and Famennian, respectively. Comparisons with these two species from other tectonic plates indicate consistent stelar architectures. Global spread, continuous occurrence, and identical anatomy during the Late Devonian indicate that Archaeopteris survived the Frasnian–Famennian extinction event. In this time, endemic genera and cosmopolitan taxa, including Archaeopteris, suggest the palaeogeographic isolation of South China and certain associations with other plates.     

Fluctuations in a late devonian-early Mississippian phytoplankton flora of Ohio,U.S.A

Fossil diversity and abundance fluctuations of Late Devonian (Famennian)—Early Mississippian (Tournaisian) phytoplankton assemblages were determined in a stratigraphic sequence of samples in Barberton test core 3, from northeast Ohio. Both diversity and abundance were relatively high during deposition of the lower Chagrin Shale (Upper Devonian) and both decreased during deposition of the upper Chagrin Shale. The overlying Cleveland Shale (Upper Devonian) is characterized by sharply increased abundance with a corresponding (although less marked) increase in diversity; in contrast, both abundance and diversity were at relatively low levels during deposition of the Lower Mississippian Bedford Shale, the uppermost unit of the sequence. These trends agree with the phytoplankton periodicity model presented by Tappan, which indicates large-scale extinction at the end of the Devonian.Many of the acritarch and prasinophycean algal taxa have restricted stratigraphic occurrences within the section studied, making them useful for local and perhaps worldwide correlation. The stratigraphically restricted taxa provide a basis for the proposal of five biostratigraphic zones within the Upper Famennian (Upper Devonian) and a portion of the Lower Mississippian. In ascending order occur: the Gorgonisphaeridium ohioense Assemblage Zone, the Acriora petala Assemblage Zone, the Muraticavea enteichia Range Zone, the Cymatiosphaera labyrinthica Range Zone, and the Navifusa drosera Assemblage Zone.     

Angustidontidae (Crustacea: Malacostraca) from the Upper Devonian of Ukraine and Belarus

Previously unknown specimens of angustidontid crustaceans from the Frasnian and Famennian of Ukraine and from the Famennian of Belarus are described.     

Anatomical reinvestigation ofArchaeopteris macilenta from the Upper Devonian (Frasnian) of South China

Archaeopteris macilenta is one of the most widespread plants in the Late Devonian.Based on fossils from the Frasnian Huangjiadeng Formation,Yichang District of Hubei Province,for the first time we study in detail the anatomy of this progymnosperm plant in South China.Ultimate axes are protostelic with three xylem sympodia and lack secondary tissue.Penultimate axes are eustelic,bearing eight sympodia and a thin band of secondary xylem.Radially symmetrical sympodia of mesarch primary xylem produce traces of appendages in a spiral arrangement.Archaeopteris macilenta and A.halliana (A.roemeriana) are dominant in the Frasnian and Famennian,respectively.Comparisons with these two species from other tectonic plates indicate consistent stelar architectures.Global spread,continuous occurrence,and identical anatomy during the Late Devonian indicate that Archaeopteris survived the Frasnian-Famennian extinction event.In this time,endemic genera and cosmopolitan taxa,including Archaeopteris,suggest the palaeogeographic isolation of South China and certain associations with other plates.     

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.     

The history of Devonian-Carboniferous reef communities: Extinctions,effects, recovery

Summary Analysis of the taxonomic composition, diversity and guild structure of five “typical” reef and mud mound communities ranging in age from Late Devonian-Early Carboniferous indicates that each of these aspects of community organization changed dramatically in relation to three extinction events. These events include a major or mass extinction at the end of the Frasnian; reef communities were also effected by less drastic end-Givetian and mid-late Famennian extinctions of reef-building higher taxa. Peak Paleozoic generic diversities for reef-building stromatoporoids and rugose corals occurred in the Eifelian-Givetian; reef-building calcareous algal taxa were longranging with peak diversity in the Devonian. These three higher taxa dominated all reef-building guilds (Constructor, Binder, Baffler) in the Frasnian and formed fossil reef communities with balanced guild structures. The extinction of nearly all reef-building stromatoporoids and rugose corals at the end of the Frasnian and the survival of nearly all calcareous algac produced mid-late Famennian reef communities dominated by the Binder Guild. Despite the survival of most calcareous algae and tabulate corals, the mid-late Famennian extinction of all remaining Paleozoic stromatoporoids and nearly all shelf-dwelling Rugosa brought the already diminished Devonian reef-building to a halt. These Devonian extinctions differ from mass extinctions by the absence of a statistically significant drop in taxonomic diversity and by their successional and cumulative effects on reef communities. Tournaisian mud mounds contain communities markedly different from the frame-building communities in Late Devonian and Visean reefs. Mound-building biotas consist of an unusual association dominated by erect, weakly skeletonized members of the Baffler Guild (chiefly fenestrate Bryozoa; Pelmatozoa) and laterally expanded, mud-binding algae/stromatolites and reptant Bryozoa. The initial recovery to reefs with skeletal frameworks in the Visean was largely due to the re-appearance of new species of abundant colonial rugose corals (Constructor Guild) and fenestrate Bryozoa. This Frasnian-Visean evolution in the taxonomic composition and structure of the reef-building guilds is also expressed by abrupt changes in biofacies and petrology of the reef limestones they produced. Thus, “typical” Frasnian reef limestones with balanced guild structures are framestones-boundstones-bafflestones, Famennian reefs are predominantly boundstones, Tournaisian mud mounds are bafflestones and Visean reefs are bafflestones-framestones.     

Survival patterns of macrobenthic marine assemblages during the end-Permian mass extinction in the western Tethys (Dolomites, Italy)

The ecological competition between brachiopods and bivalves is analysed by means of a quantitative palaeoecologic method applied on four assemblages located within a short stratigraphic interval, approximately 2 m thick, in the lower Tesero Member of the Werfen Formation (in the Southern Alps). The assemblages originate from the Tesero, Bulla and Sass de Putia sections. The analysed stratigraphic interval, uppermost Changhsingian in age, is located between the early and heaviest phase of the end-Permian mass extinction, which occurred across the Bellerophon/Werfen formational boundary (Event Boundary), and the Permian/Triassic boundary (Chronological Boundary), when nearly all the Permian stenotopic holdovers disappeared.These assemblages are characterised by small sized skeletons (“Lilliput effect”), which represent an adaptive survival strategy in stressed and harsh habitats resulting from the climatic and palaeoceanographic changes connected with the mass extinction. The Tesero assemblages are dominated by rhynchonelliform brachiopod Orbicoelia (bed CNT10) or Streptorhynchus (bed CNT11A), which were mostly attached at the top of shallow microbialitic mounds. These assemblages are again dominated by Permian stenotopic taxa and show a Palaeozoic structure. The Tesero habitat, which again permitted the survival of brachiopods, represented one of the last refuges in the western Tethys. On the contrary, the Bulla (BU9-10) and Sass de Putia (wPK13A) assemblages are bivalve-dominated, and thus show an ecologic structure typical of Early Triassic post-extinction marine benthic communities or Palaeozoic stressed marine communities. The bivalve-dominated assemblages proliferated in prevailing muddy siliciclastic substrates, with brief episodes of microbial algal growth. The most important environmental limiting factors and leading causes of end-Permian mass extinction are discussed in terms of palaeoautecologic and palaeosynecologic analysis.The different taxonomic composition and ecologic structure of the assemblages is related to palaeogeography, including water depth and connections with the open sea. The brachiopod-dominated assemblage, exclusive of the Tesero section, proliferated in microbial carbonate habitats in near-shore environments. The bivalve-dominated assemblages, which were more widespread than the brachiopod assemblages in the Dolomites and also occurred in other western Tethys localities, occur in more open and deeper marine environments. In the western Tethys margins, the local distribution of mixed faunas suggests that the extinction of Permian stenotopic taxa was caused by the onset of poisonous water on the shelves originating from deep marine environments.This extinction pattern appears to be a regional phenomenon and does not seem be applicable on a global scale. The extinction events were controlled by a complex network of interactive factors and the survival of faunal elements was probably stochastic.     

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.     

Late Devonian–Early Carboniferous palynology of the CSDP-2 Borehole in the southern Yellow Sea,China

Late Devonian–Early Carboniferous spores recovered from the depths of 2140 m to 2031.3 m in the CSDP-2 Borehole in the southern Yellow Sea are systematically documented to characterize palynological associations for the stratigraphic correlation. The taxonomic study identifies 96 species belonging to 48 genera, including a re-examination of 33 previously described and indeterminate species. Three palynological associations are established for the Wutong Formation in the CSDP-2 Borehole. In ascending order, these are the late Famennian Aneurospora asthenolabrata–Geminospora lemurata (AL) Association from the proposed Guanshan Member, the latest Famennian Cymbosporites circinatus–Asperispora acuta (CA) Association from the lower part of the proposed Leigutai Member, and the Tournaisian Auroraspora macra–Lophozonotriletes involutus (MI) Association of the upper part of the latter member. The AL and CA associations are correlated with the palynological assemblages in the Guanshan Member, and those in the lower to middle parts of the Leigutai Member of the Wutong Formation that outcrops in the suburbs of Nanjing and southern Jiangsu. The MI Association is similar to those derived from the uppermost beds of the subsurface Wutong Formation and is dated as early Tournaisian.