Fossil fungi with suggested affinities to the Endogonaceae from the Middle Triassic of Antarctica.

Documented fossil evidence of zygomycetous fungi is rare. A conspicuous fungal fossil, Jimwhitea circumtecta gen. et sp. nov., occurs in permineralized peat from the Middle Triassic of Antarctica. The fossil is interpreted as a mantled zygosporangium that buds from a macrogametangium subtended by a sac-like macrosuspensor. The macrogametangium is united at its tip with a microgametangium which is subtended by a micro-suspensor. This configuration is strikingly similar to the zygosporangium-gametangia complexes seen in certain modern Endogonaceae. Co-occurring with J. circumtecta are isolated propagules closely resembling the zygosporangium of J. circumtecta and a portion of a sporocarp containing zygosporangia embedded in a gleba. Several of the sporangia are borne on ovoid or elongate structures, which we interpret as gametangia. These fossils offer an exceptionally detailed view of the morphology and reproductive biology of early Mesozoic zygomycetes.

The fossil record of the Peronosporomycetes (Oomycota).

Evidence of fossil Peronosporomycetes has been slow to accumulate. In this review various fossils historically assigmed to the Peronosporomycets are dicussed briefly and an explanation is provided as to why the fossil record of this grouop has remained inconsistent. In recent year there has been several new reports of fossil peronosporomycetes based on structurally preserved oogonium-antheridium complexes from Derovonian and Carboniferous rocks that demonstrate the existence of these organisms as fossils and refute the long-standing assumption that they are too delicate to be preserved. Among these are serral tyoes characterized by oogonial surface members of the group. To date at last three groups of fossil vascular plants (i.e. lycophytes, ferns and seed ferns) are known to host peronosporomycetes aas endophytes; however only one form has been identified as a parasite.

A fossil peronosporomycete oogonium with an unusual surface ornament from the Carboniferous of France.

A new fossil peronosporomycete from the upper Visean (Mississippian) of France occurs as a globose oogonium at the tip of a thin-walled hypha. The oogonium surface is prominently ornamented by densely spaced, long and subtle, straight or once to several times furcated thread-like extensions; many possess an opaque, bulb-like swelling at base. Antheridia adpressed to the oogonium are clavate and paragynous. This fossil represents only the third record of an unequivocal peronosporomycete from the Carboniferous, and thus provides important details about the evolutionary history of this group of organisms.

Combresomyces cornifer gen. sp. nov., an endophytic peronosporomycete in Lepidodendron from the Carboniferous of central France.

Structurally preserved periderm of the lycophyte Lepidodendron rhodumnense from the Visean (Mississippian) of central France contains a peronosporomycete (Combresomyces cornifer gen. sp. nov.) that occurs in the form of pyriform to subglobose terminal oogonia. On the surface is a conspicuous ornamentation, which may have formed through condensation of a mucilaginous extra-oogonial wall secretion. Some oogonia contain thin-walled spherules, which may represent (walled) oospheres or spores of an endoparasitic fungus (?chytrid), whereas single, large spheres in the interior are interpreted as oospores. Antheridia adpressed to several of the specimens are clavate and paragynous. This discovery sheds light on the morphology and biology of peronosporomycetes in a terrestrial ecosystem some 330My ago. Although the organism occurs exclusively in the periderm of L. rhodumnense, it is not known whether it represents a symptomless endophyte, pathogen, or saprotroph.

Fungal endophytes in a 400-million-yr-old land plant: infection pathways, spatial distribution, and host responses.

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.

An alternative mode of early land plant colonization by putative endomycorrhizal fungi.

Rhizomatous axes of Nothia aphylla, a land plant from the 400-myr-old Rhynie chert, host a fungus that closely resembles Glomites rhyniensis (Glomeromycota), the endomycorrhizal fungus of the Rhynie chert plant Aglaophyton major. However, G. rhyniensis is an intercellular endophyte that becomes intracellular exclusively within a well-defined region of the cortex, while the fungus in N. aphylla initially is intracellular but later becomes intercellular in the cortex. We hypothesize that N. aphylla displays an alternative mode of colonization by endomycorrhizal fungi, perhaps related to the peculiar internal anatomy of the lower portion of the rhizomatous axis, in which the radial arrangement of cells, along with the virtual absence of intercellular spaces, provides no intercellular infection pathway into the cortex.