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Analyses by secondary ion mass spectroscopy (SIMS) of 11 specimens of five taxa of prokaryotic filamentous kerogenous cellular microfossils permineralized in a petrographic thin section of the ∼3,465 Ma Apex chert of northwestern Western Australia, prepared from the same rock sample from which this earliest known assemblage of cellular fossils was described more than two decades ago, show their δ13C compositions to vary systematically taxon to taxon from −31‰ to −39‰. These morphospecies-correlated carbon isotope compositions confirm the biogenicity of the Apex fossils and validate their morphology-based taxonomic assignments. Perhaps most significantly, the δ13C values of each of the five taxa are lower than those of bulk samples of Apex kerogen (−27‰), those of SIMS-measured fossil-associated dispersed particulate kerogen (−27.6‰), and those typical of modern prokaryotic phototrophs (−25 ± 10‰). The SIMS data for the two highest δ13C Apex taxa are consistent with those of extant phototrophic bacteria; those for a somewhat lower δ13C taxon, with nonbacterial methane-producing Archaea; and those for the two lowest δ13C taxa, with methane-metabolizing γ-proteobacteria. Although the existence of both methanogens and methanotrophs has been inferred from bulk analyses of the carbon isotopic compositions of pre-2,500 Ma kerogens, these in situ SIMS analyses of individual microfossils present data interpretable as evidencing the cellular preservation of such microorganisms and are consistent with the near-basal position of the Archaea in rRNA phylogenies.

Within the plant microbiota, mutualistic fungal and bacterial symbionts are striking examples of microorganisms playing crucial roles in nutrient acquisition. They have coevolved with their hosts since initial plant adaptation to land. Despite the evolutionary distances that separate mycorrhizal and nitrogen-fixing symbioses, these associations share a number of highly conserved features, including specific plant symbiotic signaling pathways, root colonization strategies that circumvent plant immune responses, functional host-microbe interface formation, and the central role of phytohormones in symbiosis-associated root developmental pathways. We highlight recent and emerging areas of investigation relating to these evolutionarily conserved mechanisms, with an emphasis on the more ancestral mycorrhizal associations, and consider to what extent this knowledge can contribute to an understanding of plant-microbiota associations as a whole.

The Early Devonian Rhynie and Windyfield cherts remain a key locality for understanding early life and ecology on land. They host the oldest unequivocal nematode worm (Nematoda), which may also offer the earliest evidence for herbivory via plant parasitism. The trigonotarbids (Arachnida: Trigonotarbida) preserve the oldest book lungs and were probably predators that practiced liquid feeding. The oldest mites (Arachnida: Acariformes) are represented by taxa which include mycophages and predators on nematodes today. The earliest harvestman (Arachnida: Opiliones) includes the first preserved tracheae, and male and female genitalia. Myriapods are represented by a scutigeromorph centipede (Chilopoda: Scutigeromorpha), probably a cursorial predator on the substrate, and a putative millipede (Diplopoda). The oldest springtails (Hexapoda: Collembola) were probably mycophages, and another hexapod of uncertain affinities preserves a gut infill of phytodebris. The first true insects (Hexapoda: Insecta) are represented by a species known from chewing (non-carnivorous?) mandibles. Coprolites also provide insights into diet, and we challenge previous assumptions that several taxa were spore-feeders. Rhynie appears to preserve a largely intact community of terrestrial animals, although some expected groups are absent. The known fossils are (ecologically) consistent with at least part of the fauna found around modern Icelandic hot springs. This article is part of a discussion meeting issue ‘The Rhynie cherts: our earliest terrestrial ecosystem revisited’.

Siliceous rocks served as raw materials in the production of stone tools from the Middle Paleolithic onwards. Due to migration, the provenance of archaeological artefacts can differ from their natural outcrop location. The aim of this work was the application of 1D and 2D low-field nuclear magnetic resonance (LF-NMR) relaxometry to distinguish cherts by their original source. Herein, bedded cherts and accompanying nodular cherts coming from three different outcrops of Kraków-Częstochowa Upland were investigated. 1D and 2D ( T 1 -T 2 ) experiments of water-saturated and dry rock sample states delivered T 1 , T 2 times and T 1 /T 2 ratios of distinct hydrogen populations – parameters sensitive to pore size, surface properties, and hydrogen bonding length. In-depth analysis of NMR data showed substantial differences in the porosity, pore surface and pore structure properties of investigated chert samples tested in the three different saturation levels (100% water-saturated, dried and differential). Finally, principal component analysis (PCA) was performed to reduce the number of correlations obtained and highlight the most important NMR properties specific to the particular outcrop localization.\"Please check captured corresponding author email if correct.\"\"The email is correct\"

The Lower Devonian Rhynie chert formed as silica sinter entombed an early terrestrial ecosystem. Silica sinter precipitates only from water flowing from alkali-chloride hot springs and geysers, the surface expression of crustal-scale geothermal systems that form low-sulfidation mineral deposits in the shallow subsurface. Active alkali-chloride hot springs at Yellowstone National Park create a suite of geothermally influenced environments; vent pools, sinter aprons, run-off streams, supra-apron terrace pools and geothermal wetlands that are habitats for modern hot-spring ecosystems. The plant-rich chert, which makes Rhynie internationally famous, probably formed in low-temperature environments at the margins of a sinter apron where frequent flooding by geothermal water and less frequent flooding by river waters created ephemeral to permanent wetland conditions. Here, the plants and associated microbes and animals would be immersed in waters with elevated temperature, brackish salinity, high pH and a cocktail of phytotoxic elements which created stresses that the fossil ecosystem must have tolerated. The environment excluded coeval mesophytic plants, creating a low-diversity hot-spring flora. Comparison with Yellowstone suggests the Rhynie plants were preadapted to their environment by life in more common and widespread environments with elevated salinity and pH such as coastal marshes, salt lakes, estuaries and saline seeps. This article is part of a discussion meeting issue ‘The Rhynie cherts: our earliest terrestrial ecosystem revisited’.

The Pilbara craton of northwestern Australia is known for what were, when reported, the oldest known microfossils and paleosols on Earth. Both interpretations are mired in controversy, and neither remain the oldest known. Both the microfossils and the paleosols have been considered hydrothermal artefacts: carbon films of vents and a large hydrothermal cupola, respectively. This study resampled and analyzed putative paleosols within and below the Strelley Pool Formation (3.3 Ga), at four classic locations: Strelley Pool, Steer Ridge, Trendall Ridge, and Streckfuss, and also at newly discovered outcrops near Marble Bar. The same sequence of sedimentary facies and paleosols was newly recognized unconformably above the locality for microfossils in chert of the Apex Basalt (3.5 Ga) near Marble Bar. The fossiliferous Apex chert was not a hydrothermal vein but a thick (15 m) sedimentary interbed within a sequence of pillow basalts, which form an angular unconformity capped by the same pre-Strelley paleosol and Strelley Pool Formation facies found elsewhere in the Pilbara region. Baritic alluvial paleosols within the Strelley Pool Formation include common microfossil spindles (cf. Eopoikilofusa ) distinct from marine microfossil communities with septate filaments ( Primaevifilum ) of cherts in the Apex and Mt Ada Basalts. Phosphorus and iron depletion in paleosols within and below the Strelley Pool Formation are evidence of soil communities of stable landscapes living under an atmosphere of high CO 2 (2473 ± 134 ppmv or 8.8 ± 0.5 times preindustrial atmospheric level of 280 ppm) and low O 2 (2181 ± 3018 ppmv or 0.01 ± 0.014 times modern).

There are two general types of rooting systems in extant land plants: gametophyte rhizoids and sporophyte root axes. These structures carry out the rooting function in the free-living stage of almost all land plant gametophytes and sporophytes, respectively. Extant vascular plants develop a dominant, free-living sporophyte on which roots form, with the exception of a small number of taxa that have secondarily lost roots. However, fossil evidence indicates that early vascular plants did not develop sporophyte roots. We propose that the common ancestor of vascular plants developed a unique rooting system—rhizoidal sporophyte axes. Here we present a synthesis and reinterpretation of the rootless sporophytes of Horneophyton lignieri, Aglaophyton majus, Rhynia gwynne-vaughanii and Nothia aphylla preserved in the Rhynie chert. We show that the sporophyte rooting structures of all four plants comprised regions of plagiotropic (horizontal) axes that developed unicellular rhizoids on their underside. These regions of axes with rhizoids developed bilateral symmetry making them distinct from the other regions which were radially symmetrical. We hypothesize that rhizoidal sporophyte axes constituted the rooting structures in the common ancestor of vascular plants because the phylogenetic positions of these plants span the origin of the vascular lineage. This article is part of a discussion meeting issue ‘The Rhynie cherts: our earliest terrestrial ecosystem revisited’.

The symbiotic partnership of plants and fungi was a critical means of nutrient uptake during colonization of the terrestrial surface. The Lower Devonian Rhynie Chert shows evidence for extensive phosphorus mobilization in plant debris that was pervasively colonized by fungi. Sandy sediment entrapped with fungi‐rich phytodebris contains grains of the phosphate mineral monazite which exhibit alteration to highly porous and leached surfaces. Mixed manganese‐iron oxide precipitates contain up to 2% P2O5. The mobilization of Mn, Fe, and P are all features of mycorrhizal nutrient concentration. However, the ecosystem was also exposed to toxic elements from hot spring hydrothermal activity. The oxide precipitates include titanium and iron‐titanium oxide which sequestered potentially toxic tungsten and antimony. Abundant pyrite framboids in the Rhynie Chert indicate that plant decomposition included microbial sulfate reduction. This caused the removal of some of the arsenic from the groundwaters into the pyrite, which reduced toxicity while leaving enough for putative arsenic metabolism. These relationships show the mineral component of the ecosystem modified the geochemistry of ambient waters. Plain Language Summary The symbiotic partnership of plants and fungi was a critical means of nutrient uptake during colonization of the terrestrial surface. The Lower Devonian Rhynie Chert shows evidence for extensive phosphorus mobilization in plant debris that was pervasively colonized by fungi. Sandy sediment entrapped with fungi‐rich organic matter contains grains of the phosphate mineral monazite which exhibit alteration to highly porous and leached surfaces. The ecosystem was also exposed to toxic elements from hot springs. Precipitates of oxide minerals sequestered potentially toxic tungsten and antimony. Iron sulfide allowed the removal of excess arsenic from the groundwaters. These relationships show the mineral component of the ecosystem modified the geochemistry of ambient waters. Key Points The plant‐bearing Rhynie Chert contains authigenic mineral phases, including pyrite, and manganese‐iron and titanium oxides Authigenic phases may be products of bacterial and fungal components and sequestered potentially toxic elements, including As, Sb, and W Detrital monazite was leached to supply phosphate, which was required by fungi in symbiosis with plants

The morphology of plant fossils from the Rhynie chert has generated longstanding questions about vascular plant shoot and leaf evolution, for instance, which morphologies were ancestral within land plants, when did vascular plants first arise and did leaves have multiple evolutionary origins? Recent advances combining insights from molecular phylogeny, palaeobotany and evo–devo research address these questions and suggest the sequence of morphological innovation during vascular plant shoot and leaf evolution. The evidence pinpoints testable developmental and genetic hypotheses relating to the origin of branching and indeterminate shoot architectures prior to the evolution of leaves, and demonstrates underestimation of polyphyly in the evolution of leaves from branching forms in ‘telome theory’ hypotheses of leaf evolution. This review discusses fossil, developmental and genetic evidence relating to the evolution of vascular plant shoots and leaves in a phylogenetic framework. This article is part of a discussion meeting issue ‘The Rhynie cherts: our earliest terrestrial ecosystem revisited’.

The terrestrialization of life has profoundly affected the biosphere, geosphere and atmosphere, and the Geological Magazine has published key works charting the development of our understanding of this process. Integral to this understanding – and featuring in one of the Geological Magazine publications – is the Devonian Rhynie chert Konservat-Lagerstätte located in Aberdeenshire, Scotland. Here we provide a review of the work on this important early terrestrial deposit to date. We begin by highlighting contributions of note in the Geological Magazine improving understanding of terrestrialization and Palaeozoic terrestrial ecosystems. We then introduce the Rhynie chert. The review highlights its geological setting: the Caledonian context of the Rhynie Basin and its nature at the time of deposition of the cherts which host its famous fossils. There follows an introduction to the development of the half-graben in which the cherts and host sediments were deposited, the palaeoenvironment this represented and the taphonomy of the fossils themselves. We subsequently provide an overview of the mineralization and geochemistry of the deposit, and then the fossils found within the Rhynie chert. These include: six plant genera, which continue to provide significant insights into the evolution of life on land; a range of different fungi, with recent work starting to probe plant–fungus interactions; lichens, amoebae and a range of unicellular eukaryotes and prokaryotes (algae and cyanobacteria); and finally a range of both aquatic and terrestrial arthropods. Through continued study coupled with methodological advances, Rhynie fossils will continue to provide unique insights into early life on land.