Nematophytes, created by Strother (1993), were a group of strange organisms in Ordovician, Silurian, and Devonian sediments. They tend to have algal-fungal characteristics that make it difficult to place them in any one category. The "body" of these organisms is a meshwork of various-sized filamentous tubes (nemato means "thread"; phyte means "plant"). Some of these organisms appear to be lichens, fungal-algae symbionts, or fungi-adjacent groups with no modern analogs.
Ordovician (?) - Devonian
Eukaryotes
└Nematophytes
Dawson, 1859
Found in Ordovician, Silurian, and Devonian sediments,
These fossils can be immensely large (8m long, 1.3m wide).
They exhibit a pseudoparenchymatous construction, with horizontally arranged tubes or hyphae.
This taxon has been placed in many wide-ranging taxonomic groups
Phaeophyceae : Penhallow, 1889; Pia, 1927; Arnold, 1947; Gothan & Weyland, 1954; Taylor, 1981; Meyen, 1987; Schweitzer, 1990
Rhodophyceae: Jonker, 1979; Schweitzer, 1983
Chlorophyta: Carruthers, 1872
Streptophyta: Dawson, 1859, 1873, 1875; Lang, 1937; Graham et al., 2010
Fungi: Carruthers, 1872; Church, 1919; Selosse, 2002a; Selosse & Strullu-Derrien, 2015; Berbee et al., 2020
Ascomycota: Edwards et al., 2013; Honegger et al., 2018; Retallack, 2019
Basidiomycota: Hueber, 2001; Retallack, 2019
Mucoromycota: Retallack & Landing, 2014; Retallack, 2019
An analysis of the evidence argues against an agaricomycete sporocarp or ascomycete lichen association and that the taxonomic identity remains unclear (Nelson & Boyce, 2022)
If Prototaxites was a fungus, it may represent part of an extinct lineage lacking extant descendants (Nelson & Boyce, 2022)
Vajda et al. (2022) propose that Prototaxites represents a complex hyphal aggregation (rhizomorph) that may have grown horizontally, similar to modern complex aggregated mycelial growth forms, such as cords and rhizomorphs.
Anatomical similarities with modern fungal rhizomorphs, Armillaria mellea
Their main function was possibly to redistribute water and nutrition from nutrient-rich to nutrient-poor areas, facilitating the expansion of early land plant communities
Loron et al. (2025) propose that Prototaxites cannot be assigned to any known group of eukaryotes, including fungi, lichens, algae, plants, animals, etc.
While the tube-like nature of its cells makes the fungal connection seem apparent, there is no evidence of chitin, and "...its anatomy was fundamentally distinct from all known extant or extinct fungi."
Diversity
P. caledonianus
P. clevelandensis
P. crassus
P. forfarensis
P. hicksii
P. lafontii
P. laxus
P. loganii
P. ortonii
P. psygmophylloides
P. saharianus
P. storriei
P. tenuis
Above: The microscopic structure of Prototaxites
Below: Reconstruction of Prototaxites thriving in a Devonian landscape (Vajda et al. 2022)
Above: Reconstruction of Prototaxites during the Silurian
Above: Fran Hueber, in front of fossil of Prototaxites
Dawson, 1886
Found in Late Devonian sediments, these fossils exhibit a thalloid appearance with short dichotomous branching. The branches in the largest species were as much as one centimeter across.
Interpretations seem to narrow this organism to a photosynthetic plant, either a fossil liverwort or brown alga.
No definitive brown algae have been identified from before the Cenozoic era
Lignin and cutin have been found in the thallus, similar to plants and charophytes
Embedded in the tissues of the thallus are chambers in which spores (200 micrometre diameter) were produced by meiosis, similar to conceptacles
Spores show no features in common with living groups of brown algae, and have sporopollenin in the spore walls, similar to streptophytes
Diversity
P. arnoldii
Bharadwaj & Venkatachala, 1960
Late Devonian of Kentucky
P. bilobata
Dawson, 1886
P. braziliensis
Dawson, 1886
P. furcata
Arnold, 1954
P. huronensis
Dawson, 1886
P. punctata [type]
(Newton) Dawson (1886)
P. ravenna
White & Stadnichenko, 1923
Above: Protosalvinia sp.
Below: Compression fossils of Protosalvinia furcata from Devonian shale of Kentucky
Miller, 1857
Late Silurian-Early Devonian of Scotland
Compression fossils that are circular or elliptic in shape
The form is 0.5-2.0 cm in diameter, showing a reticulated, mesh-like structure on the surface
Small discs present in the meshes of the reticulum, which appear to be sporangia with 30K+ spores
Some specimens show a radial pattern of striations and a "holdfast" in the center of the disk. These latter structures were probably involved in the attachment to the soil
The systematic place of Parka is uncertain
Fossils resemble both extant liverworts and green algae, although chemical composition indicates a relationship with green algae
This was probably a group of early plant-algae organisms, which colonized near-shore, barren land
Above & Below: Compressions of Parka decipiens
This form taxon is found in the Middle Silurian and the Early Devonian;
Nematothallus fossils resemble sheets of cells, such as the cuticle of a plant
Fossils can appear as irregularly shaped black patches measuring 0.5 to 6 cm in diameter
It is not developed in the way of cell division in higher plants; therefore, it is called pseudoparenchymatous
Under the cuticle is a mat of thin threads or tubes, which vary in diameter (3-40 µm). Spores have also been found in the mats.
Some researchers have suggested an affinity with lichens or red algae
Diversity
N. pseudovasculosa
Lochkovian of the UK
N. radiata
Lochkovian of the UK
Above& Below: Nematothallus
Kräusel, 1954
Dichotomous branching, and a flattened/elliptical cross-section with a thick (20–80 μm) upper cuticular surface.
It is also perforated with pores resembling those of some liverworts.
Spongiophyton has been mistakenly interpreted as tree resin and lycopod cuticle, and was later identified as the cuticle of a thalloid plant.
It has most recently been interpreted on morphological and isotopic grounds as a lichen (Taylor et al., 2004)
Spongiophyton is the earliest evidence of lichens, from the Early Devonian of Brazil (Becker-Kerber et al., 2025).
This fossil had internal hyphae, algal cells, and possible reproductive structures, confirming that it was one of the oldest known fossilized lichens.
Spongiophyton abundance and wide paleo-geographic distribution in Devonian successions reveal an ecologically prominent presence of lichens during the late stages of terrestrial colonization
Diversity
S. articulatum
Kräusel, 1954
Early Devonian of Brazil
S. hirsutum
Kräusel, 1954
Early Devonian of Brazil
S. lenticulare
Kräusel, 1954
S. minutissimum
Kräusel, 1954
Early Devonian of Brazil
S. nanum
Kräusel, 1954
Early Devonian of Brazil