Perithecial ascomycetes from the 400 million year old Rhynie chert: an example of ancestral polymorphism.

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 interpreted 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.

Cuticles of Mariopteris occidentalis White nov. emend. from the Middle Pennsylvanian of Oklahoma (USA), and a new type of climber hook for mariopteroid pteridosperms.

Cuticles of Mariopteris occidentalis are described from the Desmoinesian (Middle Pennsylvanian) of Oklahoma (USA). This species, like other mariopteroids, had a vine- to liana-like growth habit and climbed with specialized climber hooks. However, M. occidentalis is different from other mariopteroids in having small recurved hooks on the abaxial surfaces of the pinna axes. The diagnosis for M. occidentalis White 1899 is emended based on additional macroscopical observations and data on the epidermal anatomy; a lectotype is designated. M. occidentalis is compared with Pseudomariopteris cordato-ovata from the Stephanian and Autunian of Europe and North America, a taxon which is considered very similar and may be related. Although the two taxa indeed display similarities, significant evidence for a closer relationship could not be found. Finally, some features, e.g. marginal water pits and the stomatal structure, are considered with regard to their palaeoautecological significance where they are interpreted as adaptations to special physiological requirements of a vine- to liana-like life form.

Reconstruction of Pseudomariopteris busquetii, a vine-like Late Carboniferous-Early Permian pteridosperm.

The growth habit of the Late Carboniferous-Early Permian pteridosperm Pseudomariopteris busquetii is reconstructed based on compression material from the upper Stephanian of the Blanzy-Montceau and Commentry Basins (Massif Central, France), and the upper Rotliegend of the Saar-Nahe Basin (Nahe Group, N 4, Rheinland Pfalz, Germany). Pseudomariopteris busquetii was a medium-sized, vine- to liana-like plant with slender stems to which small bipartite fronds were attached. What is most interesting is that the species used at least two different strategies to both anchor and support the plant body. Most specimens possess specialized climber hooks developed from apical extensions of the pinna axes, indicating that the fronds were used to attach the plant. A few specimens suggest that the stem may also have had some capacity for attachment. In the absence of suitable supports, however, P. busquetii was apparently able to grow in dense stands or thickets in which the individual plants supported each other.

A contribution to the knowledge of the pteridosperm genera Pseudomariopteris Danzé-Corsin nov. emend. and Helenopteris nov. gen.

The macromorphology and epidermal anatomy of three Late Carboniferous-Early Permian pteridosperm species, conventionally assigned to the form-genus Pseudomariopteris Danzé-Corsin, are described from the Stephanian of the Blanzy-Montceau Basin (Central France). The generic diagnosis of Pseudomariopteris is emended, and P. busquetii is designated as the type species of the genus. The combination P. cordato-ovata (Weiss) Gillespie et al. is validated to replace the illegitimate name P. ribeyronii (Zeiller) Danzé-Corsin. The species diagnoses of P. busquetii and P. cordato-ovata are emended. On the basis of its epidermal anatomy, P. paleaui, another species orininally described from Central France, is transferred to the newly established genus Helenopteris Krings et Kerp, and the new combination Helenopteris paleaui is introduced. Lectotypes of P. busquetii and H. paleaui are selected. In addition, specimens from other basins are (re-)examined and illustrated. Seeds attached to a P. busquetii specimen suggest a callistophytalean relationship for Pseudomariopteris. Compression fossils and cuticles of Pseudomariopteris and Helenopteris are interpreted in terms of palaeoecology: Pseudomariopteris busquetii and P. cordato-ovata had climbing/scrambling growth habits; specialized climber hooks can be demonstrated for both species, either in compression or in cuticular preservation. Although the growth habit of H. paleaui is not yet entirely clear, the size of its fronds suggests that this taxon most probably was also a scrambler/climber. Finally, based on epidermal features, a tentative reconstruction of stages of the pinnule ontogeny of H. paleaui is provided.

Transpiration and assimilation of early Devonian land plants with axially symmetric telomes-simulations on the tissue level.

Early terrestrial ancestors of the land flora are characterized by a simple, axially symmetric habit and evolved in an atmosphere with much higher CO(2)concentrations than today. In order to gain information about the ecophysiological interrelationships of these plants, a model dealing with their gaseous exchange, which is basic to transpiration and photosynthesis, is introduced. The model is based on gas diffusion inside a porous medium and on a well-established photosynthesis model and allows for the simulation of the local gas fluxes through the various tissue layers of a plant axis. Necessary parameters consist of kinetical properties of the assimilation process and other physiological parameters (which have to be taken from extant plants), as well as physical constants and anatomical parameters which can be obtained from well-preserved fossil specimens. The model system is applied to an Early Devonian land plant, Aglaophyton major. The results demonstrate that, under an Early Devonian CO(2)concentration, A. major shows an extremely low transpiration rate and a low, but probably sufficiently high assimilation rate. Variation of the atmospheric CO(2)concentration shows that the assimilation is fully saturated even if the CO(2)content is decreased to about one-third of the initial value. This result indicates that A. major was probably able to exist under a wide range of atmospheric CO(2)concentrations. Further applications of this model system to ecophysiological studies of early land plant evolution are discussed.

Epidermal anatomy of Barthelopteris germarii from the Upper Carboniferous and Lower Permian of France and Germany.

The epidermal anatomy of Barthelopteris germarii, a late Paleozoic seed fern, is described on the basis of material from the Upper Stephanian of Central France. A number of features are described for the first time for this species. Some have never been reported for late Paleozoic pteridosperms. Although our material is in most respects very similar to previously described material of B. germarii from Germany, there are also some striking differences, especially the peltate glandular trichomes that are very common in the French but completely absent in the German material. Their presence is most probably an ecological adaptation as in many modern plants. This and other epidermal and gross-morphological features (e.g., the presence of papillae on subsidiary cells of the stomata, anastomosing venation) indicate that B. germarii was well adapted to stressed conditions. Of more general interest is the preservation of the material. There appears to exist a clear relationship between the preservation of anticlinal walls and the thickness of the parenchymatic mesophyll; anticlinal walls are very well preserved where parenchyma was thin. Therefore, the absence of anticlinal walls in fossil cuticles, which has often been used as a taxonomic character, is not necessarily a primary feature.

A cyanolichen from the Lower Devonian Rhynie chert.

The 400 million-year-old Rhynie chert has provided a wealth of information about various types of fungal interactions that existed in this Early Devonian paleoecosystem. In this paper we report the first unequivocal evidence of a lichen symbiosis from the Rhynie chert. Specimens of a new genus, Winfrenatia, consist of a thallus of superimposed layers of aseptate hyphae and, on the upper surface, numerous uniform depressions. Extending into the base of each depression are hyphae that form a three-dimensional netlike structure. Enclosed within each of the net spaces is a coccoid cyanobacterium, each cell of which is surrounded by a thick sheath. These photobiont cells divide in three planes, resulting in cell clusters of up to perhaps 64 individuals. The photobiont is parasitized by the fungus in the base of each net as new cyanobacterial cells are formed distally. Reproduction is by endospores and soredia. Affinities of the mycobiont appear closest to members of the Zygomycetes, while the photobiont is most similar to coccoid cyanobacteria of the Gloeocapsa and Chroococcidiopsis types. We speculate that this cyanobacterial symbiosis was well adapted to exploit and colonize new ecological niches, especially in the periodically desiccated environment postulated for the Rhynie chert paleoecosystem.

Four hundred-million-year-old vesicular arbuscular mycorrhizae.

The discovery of arbuscules in Aglaophyton major, an Early Devonian land plant, provides unequivocal evidence that mycorrhizae were established >400 million years ago. Nonseptate hyphac and arbuscules occur in a specialized meristematic region of the cortex that continually provided new cells for fungal infection. Arbuscules are morphologically identical to those of living arbuscular mycorrhizae in consisting of a basal trunk and repeatedly branched bush-like tuft within the plant cell. Although interpretations of the evolution of mycorrhizal mutualisms continue to be speculative, the existence of arbuscules in the Early Devonian indicates that nutrient transfer mutualism may have been in existence when plants invaded the land.