New palynological results from the Gafo Formation (southern sector of the Pulo do Lobo Domain, South Portuguese Zone) are integrated with recently studied sections and drillholes from the Portuguese and Spanish sectors. A total of 44 samples were studied, 27 of which were positive for palynomorph taxonomy. This research revealed well preserved palynological assemblages, including 73 spore species allocated to 28 genera, four acritarch genera, three prasinophyte algae genera plus common chitinozoan remains. Some additional forms were retained under open nomenclature. From this, the first complete age determination for the Gafo Formation in Portugal and Spain was achieved, indicating a middle Famennian (Grandispora gracilis–Grandispora famenensis, GF Biozone) to a late Famennian (Grandispora echinata, VH Biozone) age. A greywacke sample from the same Gafo Formation was dated by U–Pb zircon geochronology and a maximum depositional age of 369 ± 2.5 Ma was obtained. A correlation between these palynological and U–Pb zircon data and the palynoflora assemblages of previous authors was made, as well as with the ages of felsic volcanic rocks found intercalated in the Gafo Formation, confirming the complex stratigraphy of Pulo do Lobo Domain. The results are consistent with stratigraphic mapping and structural interpretations, allowing a middle–late Famennian age (GF and VH Biozone) to be assigned to the Gafo Formation sedimentary rocks. This work has also contributed to a reinterpretation of Gafo Formation depositional facies correlatives (e.g. the Santa Barbara Group in Spain) as being the same lithological unit.

The Devonian/Carboniferous Boundary (DCB) interval is associated with mass extinction, isotope excursions and a short glacial episode. This study investigates how boundary extinction and environmental change is expressed in the glacial high-palaeolatitudinal record of the Bolivian Altiplano (western Gondwana). A latest Devonian and early Carboniferous section has been investigated using sedimentology, palynology, total organic carbon and bulk δ13Corganic. The Colpacucho Formation is a Late Devonian shelfal–marine siliciclastic sequence. It is overlain in the study area by a unit of coarse sandstones and sandy diamictites, interpreted as glaciomarine. This distinctive glaciomarine unit is at least 7 km wide and 60–120 m thick with a variably incisive basal contact (<100 m). It is of very latest Famennian age and is a stratigraphic equivalent of proven glacigenic deposits across central South America. The offshore marine Kasa Formation overlies the glacigenic unit above a basal flooding surface. The DCB is 12 m above this flooding surface on the last occurrence of Retispora lepidophyta and significant palynological assemblage changes. This includes the loss of the Umbellasphaeridium saharicum phytoplankton bioprovince, endemic to Gondwana. Marine and terrestrial palynological extinctions are synchronous with a 2 ‰ positive carbon isotope excursion interpreted to be reflective of changes in organic matter delivery and preservation during an interval of environmental stress. These results inform wider debates on global environmental change and mass extinction at the DCB.

The Upper Famennian (Upper Devonian) Strud locality has yielded very abundant and diversified flora as well as vertebrate and arthropod faunas. The arthropod fauna, mostly recovered from fine shales deposited in a calm, confined floodplain habitat including temporary pools, has delivered a putative insect and various crustaceans including eumalacostracans and notostracan, spinicaudatan and anostracan branchiopods. Here we present the Strud eurypterids, consisting of semi-articulated juvenile specimens assigned to Hardieopteridae recovered from the pool and floodplain deposits, as well as larger isolated fragments of potential adults recovered from stratigraphically lower, coarser dark sandy layers indicative of a higher-energy fluvial environment. The Strud fossils strongly suggest that, as proposed for some Carboniferous eurypterids, juvenile freshwater eurypterids inhabited sheltered nursery pools and migrated to higher-energy river systems as they matured.

Many of the most diverse clades of Late Palaeozoic echinoids (sea urchins) originated in the Devonian period. Our understanding of diversity dynamics of these Late Palaeozoic clades are thus informed by new systematic descriptions of some of their earliest members. The Proterocidaridae are a diverse and morphologically distinct clade of stem group echinoids with flattened tests and enlarged adoral pore pairs, which are first known from the Upper Devonian. We herein report on a new species of Hyattechinus, Hyattechinus anglicus n. sp., from the Upper Devonian of the North Devon Basin, Devon, UK. This is the first Devonian Hyattechinus known from outside of the Appalachian Basin, USA, and provides novel information regarding the palaeogeographic and stratigraphic distribution of proterocidarids in Late Devonian times. We additionally update the stratigraphic distribution of Devonian Hyattechinus from the Appalachian Basin, following recent biostratigraphic resolution of their occurrences. Hyattechinus appears to have been present in the Rheic echinoderm fauna during Late Devonian times, and comparison of the palaeoenvironmental setting of Hyattechinus anglicus with that of other Hyattechinus from the Famennian of the Appalachian Basin suggests that the genus may have preferred siliciclastic settings. Furthermore, this new taxon increases the diversity of echinoids from the Upper Devonian of Devon to three species.

Archaeopterid trees were the main components of most Late Devonian forests. Their aerial axes characterised by a eustele with mesarch primary xylem strands, leaf traces departing radially from cauline bundles and secondary xylem tracheids with radial pits arranged in groups, are referred to the genus Callixylon Zalessky. The nineteen species of Callixylon described to date from North America, North Africa, Europe, Xinjiang and several parts of Russia range from the late Givetian to the Mississippian. In this paper, we describe a new species of Callixylon from two specimens collected in the Famennian locality of Mader el Mrakib in eastern Anti-Atlas. Callixylon wendtii sp. nov. is characterised by the presence of sclerotic nests in the pith, a new character for the genus. Its wood shows narrow rays of variable height, with unevenly distributed ray tracheids. Small vascular traces crossing the wood within the two innermost growth rings are interpreted as evidence for short-lived leaves. This discovery adds to the diversity of the genus Callixylon in an area of northern Gondwana that may have been favourable to the establishment of a diverse community of archaeopterid trees.

Arthropods were the first known animals to colonise land habitats, with myriapods and arachnids having done so at least by the Silurian. Much later, several lineages of Pancrustacea (hexapods and the paraphyletic crustaceans) also ventured onto land; the hexapods by the Early Devonian, and later at least four other groups of crustaceans, namely isopods, amphipods, ostracods and decapods, most of which generally colonised the continental water bodies. All faced a series of challenges (in particular: gas exchange; desiccation; reproduction; osmoregulation; and exposure to ultraviolet radiation), resulting in many morphological, physiological and ecological adaptations. Nonetheless, whether they reached land via saltwater or freshwater remains poorly documented, mainly because relevant localities are few. The Famennian (Late Devonian) Strud locality in Belgium provided an exceptional source of information on early aquatic continental ecosystems and their plant, vertebrate and arthropod colonisers at a crucial step in the terrestrialisation process. Here, we review and update its crustacean fauna, which inhabited floodplain and temporary pool waters. New anatomical details of the notostracan Strudops goldenbergi Lagebro et al., 2015, as well as a new genus and species of spinicaudatan, are described. We also discuss the ecology of this unique, early continental ecosystem and the insights it gives into the terrestrialisation process.

A new plant of Late Devonian (Famennian) age, Melvillipteris quadriseriata gen. et sp. nov., is reported from Melville Island, Nunavut, Canadian Arctic Archipelago. Main axes and two orders of lateral branches are recognized. The main axes are monopodial in habit, with a zigzag and/or upright appearance and dense adventitious roots attached on some internodes. First-order branches are inserted on the main axes in distichous pairs, showing a quadriseriate pattern (i.e. alternate pairs); they bear one or two sterile ultimate appendages on the proximal portions, and then distally bear alternate second-order branches. Second-order branches bear alternate ultimate appendages which may be fertile or sterile; the sterile ultimate appendages are composed of one or two successive dichotomies, terminating in oppositely recurved tips, and fertile appendages show an elaborate branching system. Each fertile appendage has an initial dichotomy forming two sister branches, each of which further divides three or four times to produce ultimate divisions terminated by sporangia. Two to four small fusiform sporangia are grouped in a truss which is supplied by penultimate division of the fertile appendage. The sporangia have a pointed tip and a longitudinal dehiscence. This new plant resembles other plants with a quadriseriate branching pattern (e.g. Rhacophyton Crépin, Cephalopteris Nathorst, Protocephalopteris Ananiev, Ellesmeris Hill, Scheckler & Basinger and Protopteridophyton Li & Hsü), and is assigned to the Rhacophytales. Its similarity to and divergence from the rhacophytaleans, cladoxylopsids and other related plants are discussed.

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.

The biostratigraphic significance of selected uppermost Famennian (Upper Devonian) and lower Tournaisian (Mississippian) brachiopod genera, belonging to the orders Rhynchonellida (e.g. Araratella), Spiriferida (e.g. Sphenospira, Prospira), Spiriferinida (Syringothyris) and Productida (except Chonetidina), is discussed. Owing to the difficulties of identifying productidine and strophalosiidine genera, in contrast to rhynchonellides and spiriferides, the biostratigraphic potential of the former has generally been overlooked. Brachiopods flourished in neritic environments that were unfavourable for conodonts and ammonoids. In the absence of the latter traditional marker fossils, they are potentially important for locating the Devonian–Carboniferous boundary in shallow water depositional settings in conjunction with rugose corals and foraminifers. On a worldwide scale, further work is required to reach a better assessment of the aftermath of the Hangenberg biological Crisis on brachiopods, notably in revising the faunas from the classical areas of the Famennian and Tournaisian stages in Western Europe.

New cranial material of the ‘rhynchodipterid’ lungfish Soederberghia groenlandica from Upper Devonian (Famennian) strata in East Greenland is described. Previously unknown structures identified here include components of the hyoid arch (ceratohyal, hypohyal) and the lower jaw (prearticular, dentary). Earlier interpretations of the cheek and mandible of Soederberghia are reconsidered in the light of new fossil specimens. Some of the difficulties in assessing the homologies between cheek bones in Soederberghia and those of other lungfishes stem from confusion over the arrangement in Rhynchodipterus, and a revised interpretation of this genus is proposed. The single infraorbital bar found in Soederberghia probably originates, in part, from an expanded bone 10 (quadratojugal) of the kind found in Griphognathus. Hypotheses that posit ‘rhynchodipterid’ polyphyly seem unlikely in light of a set of derived cranial characters that define a coherent radiation of long-snouted, denticle-bearing lungfishes known from the Late Devonian. The hypothesis presented here places ‘rhynchodipterids’ as a paraphyletic grade with respect to fleurantiids. Rhynchodipterus, Soederberghia, and fleurantiids form a clade to the exclusion of the species of Griphognathus. G. minutidens is the sister taxon to this apical group, while G. sculpta and G. whitei are more remote from it.

Restoration of the morphology of Angustidontus seriatus Cooper, 1936 based on complete specimens from the Famennian of Nevada and Poland, supports its affinity to the coeval alleged decapod Palaeopalaemon and suggests eocarid (possibly also peracarid) affinities. Predatory adaptation of the thoracopods and the relatively short pereion make this crustacean only superficially resemble the archaeostomatopod hoplocarids, because the large grasping appendages of Angustidontus represent the first, rather than second, maxillipeds and acted in the opposite direction: downward. Another similar adaptation of the antennae in the Viséan Palaemysis suggests a widespread adaptation to predation among early eumalacostracans. The large sample collected from the Woodruff Formation of Nevada permits biometric characterisation of the grasping maxillipeds of Angustidontus, showing that their highly variable morphology should not be used to define species. All previously described species are therefore here synonymised with A. seriatus. Differences in gnathobases of mandibles found in articulated specimens in Nevada, and associated with isolated maxillipeds and articulated specimens possibly representing another unnamed species in Poland, suggest that such mandibles may eventually prove to be taxonomically more significant.

Numerous silicified and calcareous sclerites of various sizes, recovered from the latest Famennian of Thuringia (Germany), allow the description of the first complete growth series of a blind proetoid trilobite: Drevermannia richteri. In addition, the partial ontogenetic development of Drevermannia antecurvata sp. nov. and undetermined species, Drevermannia sp. 1, are described. The proetoid anaprotaspides, associated with D. richteri, illustrate that a marked increase in larval size occurred prior to the terminal Devonian extinction event. Considering the homogeneity of larval size in older Devonian proetoids, it is interpreted as evidence that the developmental strategy of these trilobites was significantly modified. Though largely speculative, two alternative hypotheses are proposed to explain this modification. Finally, all three ontogenetic sequences show that ocular structures never develop externally in Drevermannia, but also illustrate that the development of optical nerves is not completely lost in this group. This suggests that blindness in the Drevermannia lineage followed a centripetal mode of eye reduction.

Single grains of detrital white mica from two different synorogenic sediments in the Southern Urals were analysed using the in situ ultraviolet laser ablation Ar–Ar dating technique to discriminate between age signatures associated with a high-pressure signal (phengites) from those related to muscovite only. Two disparately aged sandstone formations of Neoproterozoic (Upper Vendian) and Upper Devonian (Famennian) age were formed by the erosion of high-relief source areas with contemporaneously exhumed high-pressure rocks. A bimodal distribution of ages and chemical compositions can be detected in the two detrital populations. There is no age overlap between the two populations, reflecting completely different source areas containing high-pressure rocks of different ages. Within the Upper Vendian sandstones, detrital white mica from a 571–609 Ma age group is phengitic in composition (Si 3.3–3.41 per formula unit), while an older 645–732 Ma age group is comprised of muscovite composition grains only. The first group is compatible with the time of late exhumation and emplacement of a source area containing high-pressure rocks, the Neoproterozoic Beloretzk terrane. The older age range is compatible with a long history of cooling and the allochthonous nature of this terrane. Detrital white mica from the Famennian sandstones (Zilair Formation) comprises one age group (342–421 Ma) containing phengite (Si 3.21–3.39 per formula unit) and muscovite, and a second group (446–496 Ma) containing muscovite only. While the derivation of the second group cannot be correlated with any as yet known regional data, the first age group indicates the earliest arrival of high-pressure rocks at the surface along the suture zone after Late Devonian arc–continent collision.

Distribution patterns of eye-lenses are compared from forty-two preserved visual surfaces of the new phacopine trilobite Phacops (Phacops) tafilaltensis n. sp. from the Late Devonian of Tafilalt, Morocco. In this species, variation in the lens pattern is most common on the ventral and posterior parts of the visual surface. Post-larval specimens typically have seventeen or eighteen files of lenses, but the number present is not correlated with specimen size. Several individuals display seventeen files in one eye and eighteen in the other. The smaller number of files in one eye may reflect incomplete development of that visual surface. The length of the active section of the generative zone and its pattern of growth control the number of dorso-ventral files of lenses and their relative height. Cases of asymmetry in lens distribution are observed in six individuals. The capacity of these organisms to correct both environmental or genetic stresses appears to be weak for this character. The existence of individuals with eyes exhibiting a different number of files confirms that file number is the result of a genetically controlled developmental program. In Phacops (Phacops) tafilaltensis n. sp., the lenses are normally arranged in a regular pattern of hexagonal close packing, their size varies within the visual surface of individual specimens and their number increases during ontogeny. Variation in the timing of lens emplacement accounts for the observed irregularities in lens distribution, i.e. in ventral and posterior parts of the visual surface. Characteristic patterns of the lens arrangement on the visual surface relate to systematic categories and probably reflect evolutionary relationships.