Since the first fossil of Prototaxites was described in 1859, researchers have hypothesized that these organisms were giant algae, fungi, or lichens. And now the authors of a new paper in the American Journal of Botany believe they have resolved the long-standing mystery.
Prototaxites fossils have a consistent tubular anatomy, composed of primarily unbranched, non-septate tubes, arranged in concentric or eccentric rings, giving the fossils an appearance similar to that of a cross-section of a tree trunk. The fossil "trunks" vary in size and may be up to 8.8 m long and 1.37 m in diameter, making Prototaxites the largest organism on land during the Late Siluarian and Devonian periods.
Researchers hypothesized that Prototaxites fossils may be composed of partially degraded wind-, gravity-, or water-rolled mats of mixotrophic (capable of deriving energy from multiple sources) liverworts that are associated with fungi and cyanobacteria. This situation resembles the mats produced by the modern liverwort genus Marchantia.
The researchers tested their hypothesis by treating Marchantia polymorpha in a manner to reflect the volcanically-influenced, warm environments typical of the Devonian period and compared the resulting remains to Prototaxites fossils. Graham and her colleagues investigated the mixotrophic ability of M. polymorpha by assessing whether M. polymorpha grown in a glucose-based medium is capable of acquiring carbon from its substrate.
Structural and physiological studies showed that the fossil Prototaxites and the modern liverwort Marchantia have many similarities in their external structure, internal anatomy, and nutrition. Despite being subjected to conditions that would promote decomposition and desiccation, the rhizoids of M. polymorpha survived degradation, and with the mat rolled, created the appearance of concentric circles.
The fungal hyphae associated with living liverworts also survived treatment, suggesting that the branched tubes in fossils may be fungal hyphae. The very narrow tubes in the fossils resemble filamentous cyanobacteria that the researchers found wrapped around the rhizoids of the decaying M. polymorpha.
Investigations into the nutritional requirements of M. polymorpha revealed that the growth of M. polymorpha in a glucose-based medium was approximately 13 times that seen when the liverwort was grown in a medium without glucose. Stable carbon isotope analyses indicated that less than 20% of the carbon in the glucose-grown liverwort came from the atmosphere. The stable carbon isotope values obtained from M. polymorpha grown with varying amounts of cyanobacteria present span the range of values reported for Prototaxites fossils.
Taken together, these results demonstrate that the liverworts have a capacity for mixotrophic nutrition when glucose is present and that mixotrophy and/or the presence of cyanobacteria could be responsible for the stable carbon isotope values obtained from Prototaxites.
The researchers conclude that liverworts were important components of Devonian ecosystems. They say their results support previous hypotheses that microbial associations and mixotrophy are ancient plant traits, rather than ones that have evolved recently.
Citation: Graham et al., 'Structural, physiological, and stable carbon isotopic evidence that the enigmatic Paleozoic fossil Prototaxites formed from rolled liverwort mats', American Journal of Botany 97, 268-275; doi:10.3732/ajb.0900322