Organic-walled microfossils of uncertain origin, classified to an informal group named acritarchs, are most commonly interpreted as the resting cysts of marine eukaryotic phytoplankton. Some acritarchs have recently been interpreted as vegetative cells of chlorococcalean green algae, based on internal bodies that have been interpreted as their asexual reproductive structures (spores). To verify this interpretation, we applied confocal Raman spectroscopy and atomic force microscopy (AFM) to study the ultrastructure and nanostructure of exceptionally preserved acritarchs with internal bodies from the early Silurian cherts (c. 430 Ma-old) of Frankenwald (Germany). Three-dimensional Raman mapping showed the spatial distribution of carbonaceous material and other minerals in the walls of the analysed internal bodies and confirmed that these structures are comparable with spores of chlorococcalean microalgae. Our findings document therefore the oldest thus far known vegetative cells of sporulating green algae. The combination of confocal Raman and AFM techniques yielded detailed information about the nanostructure and fossilisation mode of the mineralised organic walls of both the central vesicles and the enclosed spore-like bodies.Keywords: acritarch microfossils, mineralization, Silurian, Raman, AFM 2
IntroductionThe informal taxonomic category of organic-walled microfossils named acritarchs [1] has an unclear systematic affiliation. They are traditionally interpreted as the cysts of unicellular organisms. [1,2] During the Proterozoic and Paleozoic, acritarchs formed the basis of the marine food chain and played an important role in the evolution of the global ecosystem. Acritarchs were therefore the main primary producers in the ocean for a long time and, as oxygenic microorganisms, they must have not only controlled the evolution of the marine ecosystem but also influenced planetary biochemical cycles. Acritarch-like fossils are among the oldest morphological traces of life [3] , and the elucidation of their biological relationships sheds new light on the evolution of the early biosphere. It has recently been suggested that at least some acritarchs are the sporulating vegetative stages of unicellular green microalgae because they show internal bodies that can be interpreted as spores. [4] The classification of acritarchs as green algae therefore has an important bearing on the overall understanding of basic processes in the evolution of Earth's biosphere. A precise morphological and chemical characterisation of acritarch internal bodies and of their threedimensional arrangement is difficult because they are typically studied either in thin sections or after extraction from fossil-bearing rock. A detailed chemical or morphological study of microfossils usually requires physical or chemical preparation to extract the microfossils from the rock matrix, which can cause their partial or even total destruction. However, even the maceration process does not allow for extraction and study of these microfossils and their f...