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Prototaxites

Prototaxites is an extinct genus of large macroscopic eukaryote dating from the Late Silurian until the Late Devonian periods. Prototaxites formed large trunk-like structures up to 1 metre (3 ft) wide, reaching 8 metres (26 ft) in height, made up of tiny interwoven tubules around 50 micrometres (0.0020 in) in diameter, making it by far the largest land-dwelling organism of its time.

Morphology
With a diameter of up to , and a height reaching , Prototaxites fossils are remnants of by far the largest organism discovered from the period of its existence. Viewed from afar, the fossils take the form of tree trunks, spreading slightly near their base in a fashion that suggests a connection to unpreserved root-like structures. Concentric growth rings, sometimes containing embedded plant material, The similarity of these tubes to structures in the early plant Nematothallus has led to suggestions that the latter may represent leaves of Prototaxites. Unfortunately for this hypothesis, the two have never been found in connection, although this may be a consequence of their detachment after the organisms' death. ==History of research==
History of research
First collected in 1843, it was not until 14 years later that J.W. Dawson, a Canadian scientist, studied Prototaxites fossils, which he described as partially rotten giant conifers, containing the remains of the fungi which had been decomposing them. Carruthers faulted the name Prototaxites (loosely translated as "first yew") and insisted that the name Nematophycus ("stringy alga") be adopted, Dawson fought adamantly to defend his original interpretation until studies of the microstructure made it clear that his position was untenable, whence he promptly attempted to rename the genus himself, calling it Nematophyton ("stringy plant"), and denying with great vehemence that he had ever considered it to be a tree. Carruthers' interpretation that it was a giant marine alga was challenged just once, in 1919, when A.H. Church suggested that Carruthers had been too quick to rule out the possibility that it was a fungus. The lack of any characters diagnostic of any extant group made the presentation of a firm hypothesis difficult; This idea was received with disbelief, denial, and strong skepticism, but further evidence has emerged to support it. In 2007, isotopic analyses by a team including Hueber and K.C. Boyce of the University of Chicago concluded that Prototaxites was a giant fungus. They detected a highly variable range of values of carbon isotope ratios in a range of Prototaxites specimens. Autotrophs (organisms such as plants and algae that sustain themselves via photosynthesis) living at the same time draw on the same (i.e., atmospheric) source of carbon; as organisms of the same type share the same chemical machinery, they reflect this atmospheric composition with a constant carbon isotope trace. The inconsistent ratio observed in Prototaxites appears to show that the organism did not survive by photosynthesis, and Boyce's team deduced that the organism fed on a range of substrates, such as the remains of whichever other organisms were nearby. Other research has suggested that Prototaxites represents a rolled-up bundle of liverworts, A similar genus, Nematasketum, also consists of banded and branching tubes in axial bundles; this seems to be a fungus. In 2019, Gregory Retallack described the new species Prototaxites honeggeri from the Darriwilian stage of the Middle Ordovician Douglas Lake Member of the Lenoir Limestone, at Douglas Dam, Tennessee, which marks the earliest known appearance of the genus Prototaxites. While an Ordovician origin of the genus is mentioned by some studies, , referring to Retallack's study, maintained that the first appearance of the genus was in the Late Silurian. A 2026 paper found that the chemical composition of Prototaxites fossils from the Rhynie chert assemblage in Scotland are distinct from other organisms in the same environment, including fungi due to lacking chitin, a key structural element in extant fungi. The paper additionally found that the medullary spots of Prototaxites taiti are composed of highly dense fine branching, and identified that some tubes have internal banding. The authors suggest that both of these features are distinct from anything found in extant fungi, and may be involved in physiological functions like gas exchange and water transport that extant fungi do not have. The authors propose that Prototaxites represents a completely new extinct lineage, separate from plants, fungi and other eukaryotes. ==Ecological context==
Ecological context
The University of Chicago research team has reconstructed Prototaxites as a branchless, columnar structure. It was the tallest living organism in its day by far; in comparison, the contemporaneous plant Cooksonia reached only in height and itself towered over the "moss forests" that grew beneath it, and invertebrates were the only other land-dwelling multi-cellular life. Prototaxites could have used its tall columnar structure for spore dispersal. Alternatively, if Prototaxites contained photosynthetic structures, the height would have increased its ability to capture light. However, the variability in the ratios in the isotopic signature δ13C among specimens of Prototaxites suggest that it was heterotrophic. Intriguingly, boreholes appeared in Prototaxites long before plants developed a structurally equivalent woody stem, and it is possible that the borers transferred to plants when these evolved. == Footnotes ==
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