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Reported are descriptions of twelve samples representing a variety of sideritic structures, including nodules mostly from roof shale of the Middle Pennsylvanian Sydney Coalfield, Canada. The co-occurrence of fossiliferous nodules and compression fossils in the shaley roof rocks at Point Aconi enhance greatly palaeontological information. Newly discovered in a coal seam, and part of the sample, is a 40 mm thick continuous sheet-like layer of siderite with abundant permineralized-like-compressed small to micron-sized structures in a rather evenly-sized sideritic matrix, probably indicating a genetic origin different from that of the nodules. Methods include some thin-section, and two X-Ray analysis. However, large systematic sampling is a prerequisite to explore that situation, which additionally could provide faunal information for Euromerican correlation.

Reported are results of an initial approximate imitation of a Carboniferous fern frond, i.e., marattialean Acitheca polymorpha (Schimper), Middle Pennsylvanian Sydney Coalfield, Canada. The simulation experiment is based on the analysis of 14 infrared spectra obtained by means of Fourier Transform Infrared spectroscopy from four detached fragments of sterile polymorphic penultimate-pinna compressions. The calculated relative, semi-quantitative, chemical data from the infrared spectra are the input for principal component analysis deriving a 3D (three-dimensional) chemometric model. To interpret it, the four specimens are placed in hypothetical-frond positions simulating a tripinnate frond, based on diminishing penultimate-rachial widths from 1-mm (distal) to 10-mm (proximal). Hypothetical conclusions include position-dependent chemistries, specifically that of opposing trends of aromaticity vs. aliphaticity in pinnules-rachises. This, in turn, would suggest potential for (i) fern-frond reconstruction, and (ii) for determination of a most likely frond position of fragmentary specimens by “chemical classification”; the predictive aspect. However, further experimental refinement is necessary particularly based on larger frond segments to confirm or disconfirm the overall hypothetical results.

A shaley slab (65 x 45 x 7 cm) from the Sydney Coalfield, Canada, Cantabrian age, on splitting apart revealed 2 – 3 layers each entombing thousands of abscised pinnules of Linopteris obliqua and eight dispersed compound-synangial structures. The campanulary-ventral-sporal micromorphology of the best preserved structure of these compares sufficiently well with previously reported structures from the Sydney Coalfield named Potoniea krisiae. Earlier studies involving larger sampling suites furthermore contributed to the observation that Hexagonocarpus sp. (female organ) and P. krisiae (male organ) usually co-occur with abscised L. obliqua pinnules; however, these two organs do not co-occur on isochronous bedding planes. In the absence of confirmatory organic attachments, the presented data provide as yet the strongest support for the hypothesis of the organs’ connectivity, but whether female-male trees existed or not, and the mode of attachment of the organs remain unknown. Hypothesized for the latter is pinnate attachment.

The Wuda Tuff Flora from the Taiyuan Formation of Inner Mongolia is a palaeobotanical lagerstatte that is key to our current understanding of the composition and ecology of the earliest Permian Cat...

Premise of research. Climbing plants are important components of modern-day tropical forests and primarily comprise angiosperms, but in Paleozoic tropical forests, pteridosperms were the dominant c...

The most ancient macroscopic plants fossils are Early Silurian cooksonioid sporophytes from the volcanic islands of the peri-Gondwanan palaeoregion (the Barrandian area, Prague Basin, Czech Republic). However, available palynological, phylogenetic and geological evidence indicates that the history of plant terrestrialization is much longer and it is recently accepted that land floras, producing different types of spores, already were established in the Ordovician Period. Here we attempt to correlate Silurian floral development with environmental dynamics based on our data from the Prague Basin, but also to compile known data on a global scale. Spore-assemblage analysis clearly indicates a significant and almost exponential expansion of trilete-spore producing plants starting during the Wenlock Epoch, while cryptospore-producers, which dominated until the Telychian Age, were evolutionarily stagnate. Interestingly cryptospore vs. trilete-spore producers seem to react differentially to Silurian glaciations—trilete-spore producing plants react more sensitively to glacial cooling, showing a reduction in species numbers. Both our own and compiled data indicate highly terrestrialized, advanced Silurian land-plant assemblage/flora types with obviously great ability to resist different dry-land stress conditions. As previously suggested some authors, they seem to evolve on different palaeo continents into quite disjunct specific plant assemblages, certainly reflecting the different geological, geographical and climatic conditions to which they were subject.

Scolecopteris libera n. sp. is established on partial three-dimensionally preserved materials containing both sterile and fertile fronds collected from the volcanic tuff at the top of the Taiyuan Formation at the Wuda Coalfield, Inner Mongolia, China. It is characterized by tripinnate fronds with rachises over 200 mm in diameter; and long lanceolate or falciform pinnules with thick veins. Pinnules are of the pecopterid-type, with those at the base of the ultimate pinnae usually divided into small lobes. Eight to twelve circular synangia are arranged in two rows along the midvein. Synangia are borne on a short pedicel attached to the middle of lateral veins. A synangium is composed of 7–10 fusiform exannulate sporangia with pointed apices. The sporangia are free among each other beyond the base of synangia. In situ microspores of the Cyclogranisporites leopoldii type differ from all other in situ Paleozoic marattialean spores.

The morphology and anatomy of a new zygopterid fern are described from the Middle Pennsylvanian (Bolsovian) of Floha, SE Germany. The fossils occur as allochthonous remains within the basal Schweddey Ignimbrite, which preserves a diverse wetland plant community of cordaitaleans, ferns, pteridosperms, calamitaleans, and lycophytes. Pinnae of Alloiopteris loecsei sp. nov. exhibit a combined carbonized and anatomical preservation type due to rapid entombment in fine-grained pyroclastics. The newly collected Alloiopteris fossils show typical gross morphology, but also reveal the anatomy of penultimate and ultimate rachides. The affinity of the new fern is discussed amongst zygopterids in general and the famous zygopterid stems Zygopteris primaria (von Cotta) Corda and Asterochlaena laxa Stenzel from the Floha site in particular. Our investigation highlights the striking potential of pyroclastic deposits for preserving and yielding both morphological and three-dimensional anatomical aspects of plant fossils.

Significance There were two heterosporous lignophyte lineages of which only one, the seed plants, survived the Permian–Triassic mass extinction. Based on exceptionally complete fossil trees from a 300-My-old volcanic ash, the enigmatic Noeggerathiales are now recognized as belonging to the other lineage. They diversified alongside the primary seed plant radiation and constitute seed plants’ closest relatives. Noeggerathiales are reconstructed as members of a plexus of free-sporing woody plants called progymnosperms, extending their age range by 60 My. Following the origin of seed plants, progymnosperms were previously thought to have become gradually less abundant before dying out in Carboniferous. We show they diversified and evolved complex morphologies including cone-like structures from modified leaves before going extinct at the Permian–Triassic extinction., Noeggerathiales are enigmatic plants that existed during Carboniferous and Permian times, ∼323 to 252 Mya. Although their morphology, diversity, and distribution are well known, their systematic affinity remained enigmatic because their anatomy was unknown. Here, we report from a 298-My-old volcanic ash deposit, an in situ, complete, anatomically preserved noeggerathialean. The plant resolves the group’s affinity and places it in a key evolutionary position within the seed plant sister group. Paratingia wuhaia sp. nov. is a small tree producing gymnospermous wood with a crown of pinnate, compound megaphyllous leaves and fertile shoots each with Ω-shaped vascular bundles. The heterosporous (containing both microspores and megaspores), bisporangiate fertile shoots appear cylindrical and cone-like, but their bilateral vasculature demonstrates that they are complex, three-dimensional sporophylls, representing leaf homologs that are unique to Noeggerathiales. The combination of heterospory and gymnospermous wood confirms that Paratingia, and thus the Noeggerathiales, are progymnosperms. Progymnosperms constitute the seed plant stem group, and Paratingia extends their range 60 My, to the end of the Permian. Cladistic analysis resolves the position of the Noeggerathiales as the most derived members of a heterosporous progymnosperm clade that are the seed plant sister group, altering our understanding of the relationships within the seed plant stem lineage and the transition from pteridophytic spore-based reproduction to the seed. Permian Noeggerathiales show that the heterosporous progymnosperm sister group to seed plants diversified alongside the primary radiation of seed plants for ∼110 My, independently evolving sophisticated cone-like fertile organs from modified leaves.

A relatively rich association of six species of embryophyte plants is known from the upper Silurian of the Prague Basin (Bohemian Massif, Czech Republic). All stratigraphically controlled specimens come from the Neocolonograptus parultimus-Neocolonograptus ultimus Zone of the Přidoli. A new genus and species, Tichavekia grandis Psenicka, Sakala et Kraft, is established. The new combination Aberlemnia bohemica (Schweitzer) Sakala, Psenicka et Kraft, comb. nov. illustrates its Lycophytina affinity. The distribution and taphonomy of these plants in the Prague Basin indicates the proximity of exposed land interpreted to be islands of volcanic origin. Two local associations show primary differences in vegetated areas of the islands and apparent ecological responses of the primitive vascular plants to their habitats. Suitable environments in the coastal zones of volcanic islands in the Prague Basin, which was situated at the outer periphery of a broad Gondwanan shelf, likely represent significant transfer points for an initial dispersion and a first expansion of land plants. Repeated vegetation of the islands is inferred from the discrete distribution of land plants at different stratigraphic levels. These changes are explained partly in context of local changes of the island relief, and partly as responses to global climatic and sea level fluctuations, and fit environmental dynamics of islands of volcanic origin in general. This Initial Plant Diversification and Dispersal Event (IPDDE) is of considerable significance in the evolution of plants documented from the Bohemian Massif.

We report on a sedimentological and biostratigraphic evaluation that was carried out on an approximately 1 km thick coal-bearing Pennsylvanian succession encountered in the boreholes Hema 82 and 85 in the Amasra coalfield of the Zonguldak – Amasra Coal Basin, Turkey.Results of the study include a refined interpretation of sedimentary environments of the coal-bearing strata and their cyclic patterns, improved constraints on macrofloral biozones as well as of boundaries and thickness of lithostratigraphic units. A paleosol-based climatic trend is also proposed, and the tectono-sedimentary history of the Amasra coalfield is compared with other coal basins in Europe. Stratigraphically constrained macroflora confirms a recently suggested discontinuous sedimentary record that contrasts with the previously widely accepted opinion that deposition was continuous throughout the Pennsylvanian. Hiatuses were identified between each adjacent formation. An early Asturian hiatus between the Karadon and Kizilli formations has its equivalent in other Variscan basins of Europe, namely in the Dobrudzha, Upper Silesian, Intra-Sudetic and central and western Bohemian basins. It is in agreement with the former paleogeographic position of the Zonguldak – Amasra Basin and its basement in the southern margin of Laurussia, in the eastern continuation of these central European basins. This conclusion is further supported by paleosol morphology-based climatic trend to increasing aridity around the Westphalian/Stephanian boundary observed in the studied boreholes, and identical with a similar trend in the above-mentioned central European basins.

A new species of adpressed marattialean fern with vegetative and fertile pinnules, Pectinangium xuanweiense sp. nov., is described from the Lopingian (upper Permian) in eastern Yunnan and western Guizhou provinces, Southwest China. The fronds of this new species are at least bipinnate, with linear to lanceolate ultimate pinnae. The pinnules are alternately arranged and pecopterid-type with pinnate venation. Fertile pinnules show recurved margins and abaxially bear a row of synangia on each side of the midvein. Synangia are 0.7 mm long and 0.4 mm wide on average, arranged in a density of approximately 16 per centimeter. Each synangium consists of four sporangia that are completely fused in well-regulated rectangular/ovoid shapes in the distal view. Sporangia are exannulate, 370 μm long and 200 μm wide on average. The sporangial walls are composed of at least three layers of elongated rectangular to polygonal cells. In situ spores are circular to oval, 25–(30)–35 μm in diameter, with trilete marks and microverrucate sculptures. Ultrastructural investigations indicated that the exine encompasses a perine, sexine, and nexine. In situ spores correspond to Verrucosisporites if found as sporae dispersae, and thus indicate that P. xuanweiense sp. nov. belongs to the Acitheca miltonii Group (palynological grouping) of Paleozoic marattialeans. Our study provides insight into the diversity of late Permian marattialeans in the tropics of the eastern Paleo-Tethys ocean.

An early Permian tuff bed between Coal 7 and Coal 6 of the Wuda Coalfield in Inner Mongolia bears T0 peat-forming vegetation preserved in situ. Documentation of the fossil record uncovered in an area of about 80 m2 shows 14 morphotaxa representing 11 whole-plant species with a wide variety of growth forms. The uncovered phytocoenosis is interpreted as a forest dominated by cordaites as the tallest trees, the remains of which covered over 50% of the excavated area. Marattialean tree ferns represent the lower canopy that was not taller than about 3 m. This storey fills gaps between cordaites and covers over 20% of the area. A rare element of lower canopy taxa was the cycadophyte genus Pterophyllum represented by a single plant. Poorly developed herbaceous groundcover includes Sphenophyllum oblongifolium, and four small fern species mostly with sphenopteroid type of foliage. Although members of the herbaceous layer covered only 7% of the excavated area, it was the most diverse storey of the peat-forming forest. Considering the small area excavated, along with the higher diversity known from the same tuff bed in the adjacent, former opencast mine, it appears that species richness in the forest was comparable to some of the less diverse Westphalian peat-forming floras in Euramerica.

Plants and insects are the two most diverse groups among terrestrial macro-organisms. They interact with each other to various extents, and constitute the most significant part of complex terrestrial ecosystems. The early Permian tuff flora in Wuda, China, buried by volcanic ash in its growth position, represents an intricate peat-forming forest in the subtropics of the eastern Tethys. However, the paleoecology of the community, especially the relationships between plants and their herbivorous insects, remains poorly understood. We report evidence for plant–insect interactions in the flora based on a thorough investigation of insect-mediated damage on the plant specimens. In total, 8 types of insect-mediated damage, belonging to 5 functional feeding groups, were recognized in 11 plant species. The insect damage recognized in the flora comprises oviposition on calamitalean axes; external leaf feeding, galling, oviposition, and wood boring on marattialean ferns; external leaf feeding, galling, and oviposition on noeggerathialeans; external leaf feeding, piercing and sucking, galling, and oviposition on cycadophytes. Damage occurrence indicates that the cycadophyte taxon Pterophyllum sp. cf. P. daihoense is likely the most frequently targeted plant in the flora. Our results may be a subset of the broader pattern of plant–insect interaction in the early Permian subtropics, due to the unusual environmental conditions, but they do provide a glimpse into the peat-forming swamp ecosystems of the late Paleozoic Cathaysia.

A new species Sphenophyllum parvifolium sp. nov. is proposed for a whole plant based on organically connected leafy shoots and cones from the Wuda locality, Inner Mongolia, North China. The most characteristic feature is the sporangiophore with lanceolate expansions bearing sporangia. Cones were studied morphologically, including cuticles and in situ spores. Cones yielded monolete spores of the Laevigatosporites or Latosporites type. Other sphenophylls from this locality include Sphenophyllum cf. oblongifolium and Sphenophyllum cf. angustifolium, both with associated cones yielding in situ spores of the same type as Sphenophyllum parvifolium, and sterile specimens of Sphenophyllum kawasakii. Some foliated axes of this type bear cones, but without in situ spores. Hook-like structures at the ends of leaf lobes of these Sphenophyllum species could have enabled the plants to climb. All species are from a volcanic tuff bed, between coal seams Nos. 6. and 7 in the Wuda Coalfield. The age of the plant-bearing tuff in the Taiyuan Formation is 298.34 ± 0.09 Ma (Asselian, earliest Permian).

Alethopteris ascendens Halle is an endemic element in the Permian Cathaysian floras and is assumed to be a member of the Medullosales based on its alethopterid type foliage. Previous accounts have documented only its pinnae and bipinnate fronds, but other parts of the plant are unknown. Here we describe new specimens from the earliest Permian (Asselian) Wuda Tuff Flora that are identified as A. ascendens, but shows additional features including an integrated structure comprising an apical hooked prolongation with two lateral rows of hooked spines. Investigation of previously published specimens from another location confirms that they also bore the same kinds of hooked appendages. Systematic comparisons demonstrate that A. ascendens is distinct from previously recognized members of the genus but that it does not conform with the generic diagnosis, leading to the establishment of Yangopteris ascendens gen. et comb. nov. The appendages in Y. ascendens are interpreted as climbing devices, and are most similar to those previously described pteridosperm Karinopteris from the Pennsylvanian of Euramerica. Similarities in the specialized climbing structures in Y. ascendens with alethopterid foliage and Karinopteris with mariopterid foliage suggest ecological convergence in unrelated pteridosperms within late Paleozoic peat-forming swamp ecosystems of Euramerica and Cathaysia. In addition, associated naked axes that co-occur with Y. ascendens are interpreted to be the main stems of the species. The growth habit of Y. ascendens is presumed to be a semi-self-supporting plant but also with a hook-climbing habit, with this similar to modern climbing palms.

A new anachoropterid fern Hansopteris uncinatus gen. et sp. nov. collected from the well-known Chinese “vegetational Pompeii” is recognized on the basis of both vegetative and fertile parts which are excellently preserved as adpressions, a few of which contain carbonized three-dimensional anatomy. Antepenultimate rachises average 3 mm in diameter and are covered by dense scales. Rachis anatomy conforms to the Anachoropteris clavata-type that is characterized by a horseshoe-shaped xylem trace with the lateral arms of its metaxylem being made up of larger tracheids than those in the medial part. Sterile fronds are tripinnately compound. Penultimate and ultimate pinnae are alternately arranged. Pinnules are sphenopterid with dentate margins. Hooked tendrils and aphlebiae are situated laterally and proximal to each penultimate rachis. On juvenile frond specimens, these structures are precocious, growing out before the laminate pinnae. The fertile frond is tripinnate with pinnules that are identical to their sterile equivalents. Round reproductive organs are about 1 mm in diameter and singly attached to each fertile pinnule. In situ spores are sub-triangular, trilete with labra and assignable to the dispersed miospore genus Leiotriletes (Naumova) Potonie et Kremp. Considering the overall preservation and the precocious hooked tendrils and aphlebiae, the growth habit of H. uncinatus is suggested to be a vine, although its stem is still unknown

A new whole-plant species of a calamitacean, Palaeostachya guanglongii sp. nov., is described in a volcanic ash bed from the Cisuralian (lower Permian) Wuda Coalfield, Inner Mongolia, North China. It was an approximately 2–3 m high shrub-like plant growing in a peat-forming swamp. It is characterized by three-orders of axes. The stem (first-order axis) bifurcates, resulting in two equal-sized major branches (second-order axis); both axial orders conform to the morphotaxon Calamites cf. schutzeiformis type with Arthropitys type anatomy. Each major branch bears numerous ultimate branches that are composed of vegetative and fertile branches. Vegetative branches bear the leaf Asterophyllites longifolius, while fertile branches bear A. longifolius leaves at proximal nodes and P. guanglongii sp. nov. strobili at distal nodes. Whorls of bracts and sporangiophores alternate in the strobili, with sporangiophores inserting slightly above the bract whorls. Four sporangiophores make up one whorl, with each sporangiophore bearing four sporangia, yielding in situ spores of the Calamospora type. The bifurcating stem is unique in Sphenopsida, potentially indicating the development of determinate branching in this group. Our findings also extend the knowledge of the morphological plasticity of a combination of anatomical features of the wood and interfascicular.

Sturia amoena(Stur) Němejc is a Pennsylvanian adpression true fern known from the Charbonnière de Belle et Bonne (Belgium) and from the Radnice and Kladno-Rakovník basins (Czech Republic). This revision includes a detailed study of pinna and pinnule morphology, aphlebiae and reproductive organs. Interesting details of sporangia andin situspores are described for the first time. The sporangia ofSturia amoenahave an equatorial bi-triseriate annulus and yieldedin situspores of thePunctatisporitesandApiculatisporitestypes.

Pennsylvanian fossil-fern cuticles are notoriously difficult to extract, little can be found in the palaeobotanical literature, yet they can supply significant taxonomic/systematic, phytostratigragphic, and palaeoenvironment information. This is demonstrated for cuticles from “Pecopteris” polypodioides of the coal basins in the Czech Republic and Canada. “Pecopteris” polypodioides is a marattialean tree fern which was part of the peat-forming flora, growing in a wet environment. Due to this fact, cuticles are generally very thin, and we assume that their major function was to reduce the ability of fungal spores to germinate and bacterial to grow, thereby reducing the possibility of these agents to cause disease. The secondary function was to be a major barrier to water loss and reducing the wettability of pinnules.

Detached and coalified prepollen organs with huge monolete grains are reported for the first time from the Canadian Sydney Coalfield of Pennsylvanian age. Besides regarding these organs as Dolerotheca -like, the putative question is which medullosalean seed fern produced these organs? We investigated for the first time the physicochemistry of the prepollen to determine characteristic behavior under oxidative conditions, reporting results from solid- and liquid states infrared spectroscopy, liquid 1 H magnetic resonance, high precision liquid and pyrolytic-gas chromatography, and mass/charge ratios. The organs were tripartite, campanulate (bell-shaped) with lobate margins ca. 20 mm in diameter and ca. 30 mm long, and contained hundreds of elongate sporangia that were up to 8 mm long and filled with huge up to 833 μm long but probably juvenile Monoletes prepollen. These grains were covered by an acellular sporangial layer, which in turn, was covered by a cuticle bearing the imprint of near-isodiametric cells. The principal evidence for linking the Dolerotheca -like structure to foliar Alethopteris pseudograndinioides is shared occurrences of papillate cyclocytic stomata and trichomes, and the physical association of the alethopterid foliage and the prepollen organs. The physicochemical results point to diagenetic influence on the surface morphology and taxonomic parameters, refractory nature of the prepollen (as with extant pollen), and their aliphatic characteristics that are different from the aromatic extant pollen.

The Intra-Sudetic Basin is essentially a post-orogenic (Variscan) continental basin with a depositional history that ranges from the late Visean to the Triassic. The fossil record in the Czech part of the basin, although discontinuous, spans the interval from the late Visean to the Asselian. Middle Baskhirian (late Namurian) to Asselian adpression floras collected from boreholes, coal mines and outcrops permit reconstruction of the stratigraphic ranges of individual species in considerable detail, and reveal changes in plant diversity and vegetation patterns throughout this ~ 25 Myr interval at high resolution, even to the scale of individual coal-bearing cycles. Species diversity recorded in the rock record throughout the studied interval mainly was controlled by climatically-driven potential for preservation of plant material, which was highest in poorly drained/waterlogged habitats (wetlands, lakes) concentrated in basinal lowlands and representing major windows of preservation. Low fossilization potential is typical for well-drained fluvial habitats, now represented by red beds deposited under drier (seasonal) climate. Diversity in major windows of preservation was highest during the late Langsettian and Duckmantian (> 50 species), lower in the Late Pennsylvanian (25–40 species) and the lowest in the Asselian (~ 20 species). The diversity in red bed intervals with poor preservation potential varies between 11 and 21 species. The two distinct habitat types, with high and low preservation potential, were dominated by different plant groups. Wetland habitats flourishing during the (per)humid to slightly seasonal intervals were colonized mainly by cryptogamic plants and some early gymnosperms, mainly pteridosperms and cordaitaleans. Habitats represented by red beds were dominated by cordaitaleans; from late Asturian time walchian conifers also occurred in these plant assemblages. However, conifers only became common in red beds in the Cantabrian. In the Saberian, conifers are rarely found in discrete beds associated with coal-bearing strata, however, their common occurrences start in lacustrine rocks of Asselian age. Temporal changes in vegetation patterns that show an increasing proportion of gymnosperms, in both wetland and dryland habitats are in agreement with the well known aridification trend in late Paleozoic tropical Pangea. Delays between the first appearance of walchian conifers in red beds (late Asturian) versus coal-bearing deposits (Saberian) supports the hypothesis that evolutionary innovations took place outside the windows of preservation, i.e. in well-drained, moisture-deficient areas.

Successful evolutionary forms are characterized by their longevity in the fossil record. There are many plant groups that exhibit these traits; here we have selected the ferns as one acknowledged evolutionary model. Ferns are the most successful cryptogamic plants in geologic history and are known from nearly all fossil floras since their first appearance in the Devonian. Ferns, autotrophic organisms, have colonized nearly all types of continental environments over time [U1102]. Fern groups experienced rapid radiation at the familial level in the Carboniferous and are well documented in the fossil record thereafter. Their megaphyllous leaves are easy to recognize and have reproductive organs (sporangia filled by spores) borne on the lower side of pinnules. Although growth forms show great variability in morphology and size, all have retained this simple reproductive organization for over 400 million years from when first recognized in Devonian floras.

The twining habit is a climbing strategy that helps slender plants grow upward by using circumnutation around other plants. In geological history, climbing may have already been present in the first Middle Devonian forests, as indicated by possible climbers among aneurophytalean progymnosperms [1] and lycopsids [2]. By the late Carboniferous, climbing was both more common and diverse - preserved in swamp forests with modes of attachment ranging from aerial roots to appendages modified into hooks and tendrils on the leaves [3]. However, all of these diagnoses of a climbing habit are based upon either indirect morphological characteristics of the purported climber or on direct physical contact with a host plant, but without direct preservation of twining [3,4]. Permineralized epiphytes have been preserved in the Carboniferous [5], but the interpretation of scars purported to have been caused by twiners that have been found on trunk compressions of potential host-plants has been questioned [5] (see Supplemental Information). Direct preservation of a climber engaged in true twining around a host has only been documented in the Miocene Shanwang Formation of Eastern China, albeit with the identity of the twiner difficult to establish and likely to be a self-twiner [6]. Here, we report a climbing fern engaged in left-handed twining around a seed plant from the early Permian Wuda Tuff fossil Lagerstatte of Inner Mongolia, China [7]. Moreover, the host plant is likely to also be a climber based on its overall form. Such a climber-climbing-a-climber phenomenon signals the potential ecological complexity of late Paleozoic forests.

The Pennsylvanian flora from the Italian Carnic Alps, stored in the Museo Friulano di Storia Naturale in Udine, Italy, was revised taxonomically. Plant fossils come from the Bombaso Formation and Pramollo (Auernig) Group (Late Pennsylvanian) that correspond to the lower part of the paralic to shallow marine Carboniferous–Permian Pramollo (or Pramollo-Nasfeld) Basin succession. Most of the ~ 2500 studied plant fossils and also the highest number of species come from the the Bombaso, Meledis and Pizzul formations, whereas the middle and upper part of the Pramollo Group in the Italian Carnic Alps yielded only few plant remains, contrarily to the successions on the Austrian side. In total 73 plant taxa have been identified, which represent about 59 biological species. The ferns, especially marattialen ferns, are the most diverse plant group (33 species in total) followed by pteridosperms (15 species). The stratigraphic range of the Bombaso Formation and the Pramollo Group have been re-evaluated based on presence of stratigraphically important species from both the Italian and Austrian parts of the Carnic Alps. The studied interval ranges from a middle Barruelian to middle Stephanian B sensu Wagner and Alvarez-Vazquez (2010a) and spans about 3.5 Ma. The diversity of the Carnic Alps flora is comparable with well-documented contemporaneous floras in NW Spain, French Massif Central and the Czech Republic. Floral richness and diversity together with intercalations of plant-rich horizons with fossiliferous marine limestone bands makes the Carnic Alps a potential candidate as a stratigraphically important reference section for non-marine to marine correlations.

This paper presents a taxonomic revision of the fossil-genus Ilfeldia Remy and establishes the fossil-genus Ovulepteris nov. Three species of Ilfeldia ( Ilfeldia jejunata , Ilfeldia gregoriensis nov. sp. and Ilfeldia polysporangiata comb. nov.) have been identified in Europe from localities in the Douro Basin (Portugal), Ilfeld Basin (Germany) and at Manebach (Germany). The stratigraphic distribution of the fossil-genus varies from Gzhelian (Carboniferous) ( I. jejunata and I. gregoriensis sp. nov.) to lower Asselian (Permian) ( I. polysporangiata comb. nov.). All species of Ilfeldia are associated with larger taeniopterid leaves that could potentially be sterile equivalents due to dimorphism, and sterile and fertile leaves have been found in organic connection in I. gregoriensis sp. nov. Ilfeldia may be affiliated with the ferns or pteridosperms, but the systematic position is unclear. Ovulepteris gen. nov., which is established for leaves that bear ovule-type reproductive organs, has oblong leaves with a partly reduced leaf lamina, with reproductive organs situated on the leaf margin. The concept of Ovulepteris gen. nov. is based on material originally described by Obrhel (1957, 1965) under the names Ilfeldia robusta Obrhel and Ilfeldia lobecensis Obrhel from the Pennsylvanian of the Czech Republic. The new combinations Ovulepteris robusta (Obrhel) comb. nov. and Ovulepteris lobecensis (Obrhel) comb. nov. are established herein. Ovulepteris gen. nov. differs from Sobernheimia in the style of placement of the seeds. In Ovulepteris gen. nov., seeds are situated at the end of veins in protruding segments, whereas their attachment in Sobernheimia is uncertain but appear to be attached to the “midrib”. The diagnosis of Ovulepteris gen. nov. is not identical with the diagnoses of existing genera that have leaves with ovules, necessitating establishment of this new fossil-genus. The systematic position of all fossil-genera studied herein is uncertain, although they appear to be a member of the pteridosperms.

Boweria is a fossil-genus of Pennsylvanian true ferns whose exact taxonomic position was for many years confused. The morphology of reproductive structures (including sporangia), pinnae, pinnules and in situ spores ( Leiotriletes and Granulatisporites types) of the type species ( Boweria schatzlarensis ) are described in detail. A new species ( Boweria nowarudensis ) is distinguished by its sporangia being isolated with two types of cells and a lateral annulus, and containing microspores of the Granulatisporites minutus type. Another former species of the genus ( Boweria minor ) is transferred to a new genus ( Kidstoniopteris minor ) based its irregularly grouped free sporangia with three types of cells, and in situ microspores of the Granulatisporites type, and laevigate spores of the Leiotriletes type.

The fossil genus Perissothallus is used for adpression fossils of late Palaeozoic noncalcareous freshwater algae consisting of erect, cylindrical branches that radiate from a central holdfast and dichotomize repeatedly. The taxon currently includes P. versiformis from the Upper Pennsylvanian of North America and P. densus from the Lower Permian of Germany. In this paper we describe a third species, Perissothallus dekovensis nov. sp., from the Gzhelian (Stephanian B;Upper Pennsylvanian) of the Kladno-Rakovnik Basin, Czech Republic. The new species differs from the previously described forms with regard to thallus size and branching pattern of the erect parts. Sedimentological data and the fossils that co-occur with the algal thalli suggest that P. dekovensis lived in shallow-water areas of a lake or abandoned channel that was surrounded by tree ferns, cordaites (especially Poacordaites-type), and lycopsids.

Marattialeans were the most species-rich fern group in the Lopingian (late Permian) Cathaysian floras in the South China Block. However, most previous studies on marattialean ferns focused on their vegetative morphology; in contrast, their reproductive biology has rarely been examined. New compression materials of Zhutheca densata (Gu et Zhi) Liu, Li et Hilton from the Lopingian Xuanwei Formation in Qujing City, Yunnan Province, Southwest China, were studied and provide additional micromorphological and ultrastructural details about the reproductive organs of this marattialean plant. Reproductive organs of the Z. densata plant are characterized by radially symmetrical synangia in two or three rows on each side of the midvein, four to six exannulate sporangia laterally fused in a synangium, sporangial walls comprising three to four layers of cells, and monolete Laevigatosporites/Latosporites-type in situ spores. The spatiotemporal distribution of marattialean reproductive organs was analyzed based on a thorough comparison of 29 genera recorded in upper Paleozoic strata. Our analysis indicated that, during the Pennsylvanian and Permian periods, marattialeans with radially symmetrical synangia notably decreased in both diversification and distribution; in contrast, marattialeans with bilaterally symmetrical synangia notably increased in both diversification and distribution. After the Lopingian, marattialeans with bilaterally symmetrical synangia gradually became predominant in terms of their diversification and geographic distribution. Our study sheds new light on the evolution of marattialeans.

Two compressions of fertile Myriotheca anglica Kidston pinnae from Kidston's type collection have been studied for the first time since the publication of its protologue. The specimens came from the Lower Coal Measures Formation (middle Langsettian Substage) of the Central Pennines Basin, UK. SEM observations have shown that this species has pinnules with 17–22 Pecopteris-type sporangia, with an apical annulus consisting of four to five rows of thick-walled cells and a stomium of five to seven rows of thick-walled cells. Based on this, a new combination, Pecopteris anglica (Kidston) comb. nov. is proposed. Two different stages of maturity of the in situ microspores are recognized: sculptured forms of the Convolutispora-type are interpreted as fully matured microspores; while laevigate, Punctatisporites-type spores represent relatively immature microspores.

This paper provides documentation of the Late Carboniferous flora of the Southern Karavanke Mountains in Slovenia that is housed in the Gornjesavski Muzej (Upper Sava Museum) in Jesenice, Slovenia. The paleoflora from this area has not been systematically documented until today. 21 fossil-species, which probably represent 17 biological species, were identified. Common species include Sigillaria brardii Brongniart, Lepidodendron dissitum Sauver, Annularia carinata Gutbier, Calamites undulatus Sternberg, Sphenophyllum oblongifolium (Germar and Kaulfuss) Unger, Acitheca polymorpha (Brongniart) Schimper and Nemejcopteris feminaeformis (Schlotheim) Barthel. Stratigraphically important are Sphenophyllum oblongifolium and Nemejcopteris feminaeformis, and based on their occurrence we can assume the studied paleoflora to be Gzhelian D (Jigulites jigulensis zone on the Russian Platform), i.e. Stephanian C

Typification of Sydneia manleyi Psenicka et al. 2003 is based entirely on fertile foliage. Of late,attached sterile-fertile segments have been found which are illustrated and described, together with cuticular preparations. A new subfamily, Sydneideae subfam. nov. is erected for the monotypytic species. The synangial chemistry is compared with marattialean synangia/sporangia from the Czech Republic and from Canada. Fil: Psenicka, Josef. West Bohemian Museum in Pilsen. Palaeontological Department; República Checa Fil: Zodrow, Erwin L.. Cape Breton Universit. Palaeobiology Laboratory; Canadá Fil: D` Angelo, José Alejandro. Cape Breton Universit. Palaeobiology Laboratory; Canadá. Consejo Nacional de Investigaciones Científicas y Técnicas. Científico Tecnológico Mendoza. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales; Argentina

Described is a new species of sub-arborescent lycopsid, Omphalophloios wagneri sp. nov. from roof shale of the middle Moscovian (Bolsovian/Atokan) (Middle Pennsylvanian) Lower Block Coal in the Illinois Basin, Indiana, USA. Available material represents mostly fragments of 16 to > 60 mm wide fertile axes that bear densely spaced sporangia, which are flattened due to compaction into disc-like shape, 2.5–3.1 mm in diameter. The outer sporangial cell wall layer displays a gradual transition from isometric cells in the central parts of adaxial and abaxial sporangial valves, to a narrow belt of elongated cells in the equatorial area, interpreted as a dehiscence zone. Micro- and megasporangia alternate in irregular patches, a single sporangium being attached adaxially to an extended sporophyll base. Cyperites-like sporophyll distal laminae are ~ 25 mm long at fertile shoot apices but become progressively longer (> 110 mm) down the shoots. Based on size distribution of stems, fertile shoots are estimated to bifurcate at least five times, which suggests that O. wagneri formed a relatively dense crown. The entire habit of the plant is difficult to assess from existing material; however, the tree was probably small and interpreted as colonizing low lying to slightly raised peat swamps. Omphalophloios wagneri differs from all other adpression-based species of the genus in having the smallest sporangia and several orders of fertile-axis bifurcations. In situ megaspores of the Zonalesporites brassertii type also are different from all other Omphalophloios megaspores.

Seven compression fossils of the compound synangial organ Potoniea sensu stricto are reported for the first time from Cantabrian-aged (late Moscovian) strata in the Sydney Coalfield, Nova Scotia, Canada. They occurred at a single locality, mostly in the proximo-distal repose position clearly showing sporangia overextending the distal-proximal surfaces, and with one specimen laterally compressed. Specimens were studied under SEM and sporangia were macerated to reveal in situ microspores. The organs are characterized as a compound synangial organs consisting a very narrow stalk, vascularized proximal cap and an outer cover of sporangial parts. Long, cylindrical sporangia are grouped with synangia, terminating as free sporangial tips that extend from the distal surface of the compound synangial organ. Sporangia are densely filled with trilete, verrucate to rugulate microspores of the Verrucosisporites type. Taxonomically significant is the potential of using the cuticles from the cap of the compound synangial organ that have continuously thick bifurcating veins and stomata to link with the associated linopterid foliar taxa Linopteris and Alethopteris occurring within the Sydney Coalfield. Problems are encountered from the non-availability of linopterid cuticles and the poor preservation of Potoniea cuticles. Considering the newly discovered features, the new species Potoniea krisiae sp. nov. is proposed.