The advent of wood (secondary xylem) is a major event of the Paleozoic Era, facilitating the evolution of large perennial plants. The first steps of wood evolution are unknown. We describe two small Early Devonian (407 to 397 million years ago) plants with secondary xylem including simple rays. Their wood currently represents the earliest evidence of secondary growth in plants. The small size of the plants and the presence of thick-walled cortical cells confirm that wood early evolution was driven by hydraulic constraints rather than by the necessity of mechanical support for increasing height. The plants described here are most probably precursors of lignophytes.

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.

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