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Summary• The Early Devonian Rhynie chert has been critical in documenting early land plant-fungal interactions. However, complex associations involving several fungi that enter into qualitatively different relationships with a single host plant and even interact with one another have not yet been detailed.• Here, we studied petrographic thin sections of the Rhynie chert plant Nothia aphylla.• Three fungal endophytes (co)occur in prostrate axes of this plant: narrow hyphae producing clusters of small spores; large spherical spores/zoosporangia; and wide aseptate hyphae that form intercellular vesicles in the cortex. Host responses on attack include bulging of infected rhizoids, formation of encasement layers around intracellular hyphae, and separation of infected from uninfected tissues by secondarily thickened cell walls.• A complex simultaneous interaction of N. aphylla with three endophytic fungi was discovered. The host responses indicate that some of the mechanisms causing host responses in extant plants were in place 400 million yr ago. Anatomical and life history features of N. aphylla suggest that this plant may have been particularly susceptible to colonization by fungi.

The exquisite preservation of organisms in the Early Devonian Rhynie chert ecosystem has permitted the documentation of the morphology and life history biology of fungi belonging to several major taxonomic groups (e.g., Chytridiomycota, Ascomycota, Glomeromycota). The Rhynie chert also provides the first unequivocal evidence in the fossil record of fungal interactions that can in turn be compared with those in modern ecosystems. These interactions in the Rhynie chert involve both green algae and macroplants, with examples of saprophytism, parasitism, and mutualism, including the earliest mycorrhizal associations and lichen symbiosis known to date in the fossil record. Especially significant are several types of specific host responses to fungal infection that indicate that these plants had already evolved methods of defence similar and perhaps analogous to those of extant plants. This suggests that mechanisms underlying the establishment and sustenance of associations of fungi with land plants were well in place prior to the Early Devonian. In addition, a more complete understanding of the microbial organisms involved in this complex ecosystem can also provide calibration points for phylogenies based on molecular data analysis. The richness of the microbial community in the Rhynie chert holds tremendous potential for documenting additional fungal groups, which permits speculation about further interactions with abiotic and biotic components of the environment.

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Abstract:We describe a perithecial, pleomorphic ascomycetous fungus from the Early Devonian (400 mya) Rhynie chert; the fungus occurs in the cortex just beneath the epidermis of aerial stems and rhizomes of the vascular plant Asteroxylon. Perithecia are nearly spherical with a short, ostiolate neck that extends into a substomatal chamber of the host plant; periphyses line the inner surface of the ostiole. The ascocarp wall is multilayered and formed of septate hyphae; extending from the inner surface are elongate asci interspersed with delicate paraphyses. Asci appear to be unitunicate and contain up to 16 smooth, uniseriate-biseriate ascospores. The method of ascospore liberation is unknown; however, the tip of the ascus is characterized by a narrow, slightly elevated circular collar. Ascospores appear 1-5 celled, and germination is from one end of the spore. Also present along the stems and interspersed among the perithecia are acervuli of conidiophores that are inAccepted for publication April 27, 2004. 1 Corresponding author. E-mail: tntaylor@ku.edu terpreted as the anamorph of the fungus. Conidiogenesis is thallic, basipetal and probably of the holoarthric-type; arthrospores are cube-shaped. Some perithecia contain mycoparasites in the form of hyphae and thick-walled spores of various sizes. The structure and morphology of the fossil fungus is compared with modern ascomycetes that produce perithecial ascocarps, and characters that define the fungus are considered in the context of ascomycete phylogeny.

The Carnian (Late Triassic) flora of Lunz in Lower Austria is famous for an abundance of well-preserved bennettitalean and cycadalean foliage and reproductive structures. However, only the fertile remains have been studied in detail to date. Recently completed systematic macromorphological and cuticular analyses of leaf fossils from Lunz revealed that several forms previously accommodated in the genus Macrotaeniopteris do not represent ferns but rather Bennettitales assignable to the genus Nilssoniopteris. We describe three species of Nilssoniopteris, which represent the first records for this genus from Lunz, and one of the earliest accounts of Nilssoniopteris in the Northern Hemisphere. The following new combinations are introduced: Nilssoniopteris haidingeri (Stur ex Krasser, 1909a) comb. nov., N. angustior (Stur ex Krasser, 1909a) comb. nov. and N. lunzensis (Stur ex Krasser, 1909a) comb. nov. The discovery of Nilssoniopteris further substantiates the significance of the Lunz flora as one of the richest and most diverse early Late Triassic floras from the Northern Hemisphere.

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A rachis of the fossil filicalean fern Botryopteris antiqua containing abundant septate hyphae with clamp connections is preserved in a late Visean (Mississippian; *330 Ma) chert from Esnost (Autun Basin) in central France. Largely unbranched tubular hyphae pass from cell to cell, but may sometimes produce a branch from a clamp connection. Other clamp-bearing hyphae occur clustered in individual cells or small groups of adjacent host cells. These hyphae may be tubular, catenulate with numerous hyphal swellings, or they may display a combination of both. The Visean hyphae with clamp connections predate Palaeancistrus martinii, the heretofore oldest direct fossil evidence of Basidiomycota, by some 25 Ma.