After terrestrialization, the diversification of arthropods and vertebrates is thought to have occurred in two distinct phases, the first between the Silurian and the Frasnian stages (Late Devonian period) (425-385 million years (Myr) ago), and the second characterized by the emergence of numerous new major taxa, during the Late Carboniferous period (after 345 Myr ago). These two diversification periods bracket the depauperate vertebrate Romer's gap (360-345 Myr ago) and arthropod gap (385-325 Myr ago), which could be due to preservational artefact. Although a recent molecular dating has given an age of 390 Myr for the Holometabola, the record of hexapods during the Early-Middle Devonian (411.5-391 Myr ago, Pragian to Givetian stages) is exceptionally sparse and based on fragmentary remains, which hinders the timing of this diversification. Indeed, although Devonian Archaeognatha are problematic, the Pragian of Scotland has given some Collembola and the incomplete insect Rhyniognatha, with its diagnostic dicondylic, metapterygotan mandibles. The oldest, definitively winged insects are from the Serpukhovian stage (latest Early Carboniferous period). Here we report the first complete Late Devonian insect, which was probably a terrestrial species. Its 'orthopteroid' mandibles are of an omnivorous type, clearly not modified for a solely carnivorous diet. This discovery narrows the 45-Myr gap in the fossil record of Hexapoda, and demonstrates further a first Devonian phase of diversification for the Hexapoda, as in vertebrates, and suggests that the Pterygota diversified before and during Romer's gap.
The Famennian (Upper Devonian, c. 372 to 359 Ma) strata of Belgium have recently received much attention after the discoveries of early tetrapod remains and outstandingly preserved continental arthropods. The Strud locality has yielded a diverse flora and fauna including seed-plants, tetrapods, various placoderm, actinopterygian, acanthodian and sarcopterygian fishes, crustaceans (anostracans, notostracans, conchostracans and decapods) and a putative complete insect. This fossil assemblage is one of the oldest continental – probably fresh-water – ecosystems with a considerable vertebrate and invertebrate diversity. The study of the palaeoenvironment of the Strud locality is crucial because it records one of the earliest and most important phases of tetrapod evolution that took place after their emergence but before their terrestrialization. It raises the question of environmental and ecological conditions for the Devonian aquatic ecosystem and the selection pressures occurring at the onset of tetrapod terrestrialization. The present study characterized the fluvial facies of the Upper Famennian sedimentary rocks of Strud and the surrounding areas. The exceptional preservation of arthropods and plants in the main fossiliferous layers is explained by rapid burial in the fine-grained sediment of the quiet and confined flood plain environment. Newly investigated fossiliferous sections in the Meuse–Samson area led to the description and correlation of key sections (Strud, Wierde and Jausse sections, complemented by the less continuous Haltinne, Huy and Coutisse sections). Moreover, the investigated sections allowed a review of the age of the fossiliferous horizon, which is now definitely considered to be Late Famennian in age.
ABSTRACT. Recent refinements of the palynological stratigraphic scheme around the Devonian-Carboniferous Boundary in southern Belgium result in new questions on the effect of the Hangenberg Biocrisis on spores and on the validity of some 'biozones'. Verrucosisporites nitidus, the guide taxa of the last Devonian palynozone (LN zone) is lacking in many sections and its presence is only acknowledged in proximal settings. Hence the LN zone is considered as an ecozone rather than a biozone. Moreover, investigation of the uppermost part of the uppermost Famennian shows that the palynological assemblages is dominated by abnormal forms of Retispora lepidophyta, notably by R. lepidophyta var. tener which seems to be characteristic of this interval in Western Europe. It is thus proposed to introduce the global tener event as a potential marker of the Hangenberg Biocrisis on land and in proximal marine settings. This profusion of abnormal spores is thought to be related to climatic variation possibly in link with global cooling below the Devonian-Carboniferous Boundary.
The Eifelian lithostratigraphic succession of southern Belgium is here revised based on new field investigations and reconsideration of biostratigraphic data. The Couvin Formation is divided into four members: the Villers-la-Tour Member recorded the onset of stromatoporoid biostromes, the Petigny Member is a dark fine-grained limestone that recorded the Choteč event, the Cul d'Efer Member is dominated by stromatoporoid-coral biostromes. These first three members were previously united in the Foulerie Member and correspond to the transgressive and highstand system tracts of the Middle Devonian 3 rd -order sequence MD1. The argillaceous and bioclastic successive units of the Abîme Member recorded the sequence MD2. Laterally, this sequence is only composed of siliciclastic deposits of the Vieux Moulin, Station and Cimetière members of the Jemelle Formation. The highstand system tract is the newly introduced carbonate Vierves Member. Large bioherms, newly described as the Wancennes Formation, recorded the first two sequences MD1 and MD2. A major sequence boundary caps the Couvin Formation and is overlaid by the transgressive and diachronous Chavées Member of the Jemelle Formation with a depositional gap increasing eastwards. The highstand system tract of sequence MD3 is recorded in the bioherms of the Tienne Sainte-Anne Member (upper part of the Jemelle Formation), also capped by a sequence boundary. These bioherms probably acted as highs around which the sandy deposits of the Lomme Formation accumulated as lowstand system tract of sequence MD4. The Hanonet Formation and its crinoidal Wellin Member form the transgressive system tracts of sequence MD4 that terminates in the bioclastic and biohermal Trois-Fontaines Formation (Givetian). The Kačák event is identified in the transgressive system tract of this sequence. The geographic distribution of facies and of reefs in particular led to the recognition of at least six sedimentation areas corresponding most probably to synsedimentary faulted blocks here defined as the Eau Blanche, Viroin, Lesse, Ourthe, Condroz and Sambre blocks. These sedimentation areas structured the Namur-Dinant Basin and recorded distinct depositional history and were probably active during the Devonian and Carboniferous interval.
A rich and diverse coral fauna collected from the Livian (Viséan, Mississippian) of Zonguldak and Bartın (North-western Turkey) contains numerous specimens of Dorlodotia and related forms. The most common species, D. delepinei Charles, 1933, is redetermined as Dorlodotia briarti, an European species. A new species of Dorlodotia is described for very large, phaceloid forms: D. euxinensis. The genus Ceriodotia is created for species close to Dorlodotia with a cerioid habit. It comprises two species: C. bartinensis and C. petalaxoides, both common in the Viséan of Zonguldak and Bartın. Based on comparison of morphological characters and features of the astogeny, a phylogenetic lineage is proposed for Dorlodotia and the new taxa. The evolution of D. euxinensis from D. briarti is peramorphic. Subsequent paedomorphosis resulted in the lineage D. euxinensis -C. bartinensis -C. petalaxoides. Ceriodotia probably includes Viséan taxa from Asia previously described as 'Acrocyathus'.
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