Stromatolites are the oldest evidence for life on Earth, but modern living examples are rare and predominantly occur in shallow marine or (hyper-) saline lacustrine environments, subject to exotic physico-chemical conditions. Here we report the discovery of living freshwater stromatolites in cool-temperate karstic wetlands in the Giblin River catchment of the UNESCO-listed Tasmanian Wilderness World Heritage Area, Australia. These stromatolites colonize the slopes of karstic spring mounds which create mildly alkaline (pH of 7.0-7.9) enclaves within an otherwise uniformly acidic organosol terrain. The freshwater emerging from the springs is Ca-HCO3 dominated and water temperatures show no evidence of geothermal heating. Using 16 S rRNA gene clone library analysis we revealed that the bacterial community is dominated by Cyanobacteria, Alphaproteobacteria and an unusually high proportion of Chloroflexi, followed by Armatimonadetes and Planctomycetes, and is therefore unique compared to other living examples. Macroinvertebrates are sparse and snails in particular are disadvantaged by the development of debilitating accumulations of carbonate on their shells, corroborating evidence that stromatolites flourish under conditions where predation by metazoans is suppressed. Our findings constitute a novel habitat for stromatolites because cool-temperate freshwater wetlands are not a conventional stromatolite niche, suggesting that stromatolites may be more common than previously thought.
Estimates of the time at which life arose on Earth make use of two types of evidence. First, astrophysical and geophysical studies provide a timescale for the formation of Earth and the Moon, for large impact events on early Earth, and for the cooling of the early magma ocean. From this evidence, we can deduce a habitability boundary, which is the earliest point at which Earth became habitable. Second, biosignatures in geological samples, including microfossils, stromatolites, and chemical isotope ratios, provide evidence for when life was actually present. From these observations we can deduce a biosignature boundary, which is the earliest point at which there is clear evidence that life existed. Studies with molecular phylogenetics and records of the changing level of oxygen in the atmosphere give additional information that helps to determine the biosignature boundary. Here, we review the data from a wide range of disciplines to summarize current information on the timings of these two boundaries. The habitability boundary could be as early as 4.5 Ga, the earliest possible estimate of the time at which Earth had a stable crust and hydrosphere, or as late as 3.9 Ga, the end of the period of heavy meteorite bombardment. The lack of consensus on whether there was a late heavy meteorite bombardment that was significant enough to prevent life is the largest uncertainty in estimating the time of the habitability boundary. The biosignature boundary is more closely constrained. Evidence from carbon isotope ratios and stromatolite fossils both point to a time close to 3.7 Ga. Life must have emerged in the interval between these two boundaries. The time taken for life to appear could, therefore, be within 200 Myr or as long as 800 Myr. Key Words: Origin of life-Astrobiology-Habitability-Biosignatures-Geochemistry-Early Earth. Astrobiology 18, 343-364.
Grazing and burrowing organisms usually homogenise microalgal mats that form on benthic sediments of many aquatic ecosystems. In the absence of this disruption, microalgal mats can accrete laminated deposits (stromatolites). Stromatolites are rare in modern coastal ecosystems, but persist at locations where metazoans are largely excluded. This study aimed to assess the trophic structure at stromatolite locations where metazoans co-occur, to determine the grazing influence exerted by the metazoans on the stromatolite-forming microalgae (cyanobacteria and diatoms). Stable isotope signatures (δ(13)C and δ(15)N) were used as food-web tracers and dietary composition of consumers was calculated using source mixing models. Results clearly demonstrate that the dominant macrofaunal grazers do not utilise stromatolite material as a food resource, but rather subsist on autochthonous macroalgae. For instance, the mean (±SD) dietary composition of two of the most abundant grazers, Melita zeylanica (Amphipoda) and Composetia cf. keiskama (Polychaeta), consisted of 80 ± 11% and 91 ± 7% macroalgae, respectively. This suggests that the stromatolite-forming benthic microalgae are not disrupted significantly by grazing pressures, allowing for the layered mineralisation process to perpetuate. Additionally, grazers likely have a restrictive influence on pool macroalgae, maintaining the competitive balance between micro- and macroalgal groups.
Microorganisms are ubiquitous in modern environments, where they have a variety of essential functions but little is known about their diversity and roles in early terrestrial environments. The earliest direct evidence of filamentous microorganisms associated with plants occurs around 407 million years ago in landscapes dominated by an herbaceous flora. 100 million years later, forests were well established and associations had increased in diversity. After more than a century since the first descriptions, the need for better understood fossils has renewed interest in their study. New discoveries and advances in imaging methods are beginning to reveal the importance of Cyanobacteria, Fungi and Oomycota and their interactions with plants in early floras.
The use of metals as biosignatures in the fossil stromatolite record requires understanding of the processes controlling the initial metal(loid) incorporation and diagenetic preservation in living microbialites. Here, we report the distribution of metals and the organic fraction within the lithifying microbialite of the hypersaline Big Pond Lake (Bahamas). Using synchrotron-based X-ray microfluorescence, confocal, and biphoton microscopies at different scales (cm-μm) in combination with traditional geochemical analyses, we show that the initial cation sorption at the surface of an active microbialite is governed by passive binding to the organic matrix, resulting in a homogeneous metal distribution. During early diagenesis, the metabolic activity in deeper microbialite layers slows down and the distribution of the metals becomes progressively heterogeneous, resulting from remobilization and concentration as metal(loid)-enriched sulfides, which are aligned with the lamination of the microbialite. In addition, we were able to identify globules containing significant Mn, Cu, Zn, and As enrichments potentially produced through microbial activity. The similarity of the metal(loid) distributions observed in the Big Pond microbialite to those observed in the Archean stromatolites of Tumbiana provides the foundation for a conceptual model of the evolution of the metal distribution through initial growth, early diagenesis, and fossilization of a microbialite, with a potential application to the fossil record.
Chlorophyll f, the most far-red (720-740 nm) absorbing chlorophyll species, was discovered in cyanobacterial isolates from stromatolites and subsequently in other habitats as well. However, the spatial distribution and temporal dynamics of Chl f in a natural habitat have so far not been documented. Here we report the presence of Chl f in cyanobacterial beachrock biofilms. Hyperspectral imaging on cross-sections of beachrock from Heron Island (Great Barrier Reef, Australia), showed a strong and widely distributed signature of Chl f absorption in an endolithic layer below the dense cyanobacterial surface biofilm that could be localized to aggregates of Chroococcidiopsis-like unicellular cyanobacteria packed within a thick common sheath. HPLC-based pigment analyses showed in-situ ratios of Chl f to Chl a of 5% in brown-pigmented zones of the beachrock, with lower ratios of ~0.5% in the black and pink-pigmented biofilm zones. Enrichment experiments with black beachrock biofilm showed stimulated synthesis of Chl f and Chl d when grown under NIR (740 nm), with a Chl f to Chl a ratio increasing four-fold to 2%, whereas the Chl d to Chl a ratio went from 0% to 0.8%. Enrichments grown under white light (400-700 nm) produced no detectable amounts of either Chl d or Chl f. Beachrock cyanobacteria thus exhibited characteristics of far-red light photoacclimation (FaRLiP), enabling Chl f -containing cyanobacteria to thrive in optical niches deprived of visible light when sufficient NIR is prevalent. This article is protected by copyright. All rights reserved.
Modern decimeter-scale columnar stromatolites from Lake Joyce, Antarctica, show a change in branching pattern during a period of lake level rise. Branching patterns correspond to a change in cyanobacterial community composition as preserved in authigenic calcite crystals. The transition in stromatolite morphology is preserved by mineralized layers that contain microfossils and cylindrical molds of cyanobacterial filaments. The molds are composed of two populations with different diameters. Large diameter molds (>2.8 μm) are abundant in calcite forming the oldest stromatolite layers, but are absent from younger layers. In contrast, <2.3 μm diameter molds are common in all stromatolites layers. Loss of large diameter molds corresponds to the transition from smooth-sided stromatolitic columns to branched and irregular columns. Mold diameters are similar to trichome diameters of the four most abundant living cyanobacteria morphotypes in Lake Joyce: Phormidium autumnale morphotypes have trichome diameters >3.5 μm, whereas Leptolyngbya antarctica, L. fragilis, and Pseudanabaena frigida morphotypes have diameters <2.3 μm. P. autumnale morphotypes were only common in mats at <12 m depth. Mats containing abundant P. autumnale morphotypes were smooth, whereas mats with few P. autumnale morphotypes contained small peaks and protruding bundles of filaments, suggesting that the absence of P. autumnale morphotypes allowed small-scale topography to develop on mats. Comparisons of living filaments and mold diameters suggest that P. autumnale morphotypes were present early in stromatolite growth, but disappeared from the community through time. We hypothesize that the mat-smoothing behavior of P. autumnale morphotypes inhibited nucleation of stromatolite branches. When P. autumnale morphotypes were excluded from the community, potentially reflecting a rise in lake level, short-wavelength roughness provided nuclei for stromatolite branches. This growth history provides a conceptual model for initiation of branched stromatolite growth resulting from a change in microbial community composition.
Archean and Proterozoic stromatolites are sparry or fine-grained and finely laminated; coarse-grained stromatolites, such as many found in modern marine systems, do not appear until quite late in the fossil record. The cause of this textural change and its relevance to understanding the evolutionary history of stromatolites is unclear. Cyanobacteria are typically considered the dominant stromatolite builders through time, but studies demonstrating the trapping and binding abilities of cyanobacterial mats are limited. With this in mind, we conducted experiments to test the grain trapping and binding capabilities of filamentous cyanobacterial mats and trapping in larger filamentous algal mats in order to better understand grain size trends in stromatolites. Mats were cut into squares, inclined in saltwater tanks at angles from 0 to 75° (approximating the angle of lamina in typical stromatolites), and grains of various sizes (fine sand, coarse sand, and fine pebbles) were delivered to their surface. Trapping of grains by the cyanobacterial mats depended strongly on (i) how far filaments protruded from the sediment surface, (ii) grain size, and (iii) the mat’s incline angle. The cyanobacterial mats were much more effective at trapping fine grains beyond the abiotic slide angle than larger grains. In addition, the cyanobacterial mats actively bound grains of all sizes over time. In contrast, the much larger algal mats trapped medium and coarse grains at all angles. Our experiments suggest that (i) the presence of detrital grains beyond the abiotic slide angle can be considered a biosignature in ancient stromatolites where biogenicity is in question, and, (ii) where coarse grains are present within stromatolite laminae at angles beyond the abiotic angle of slide (e.g., most modern marine stromatolites), typical cyanobacterial-type mats are probably not solely responsible for the construction, giving insight into the evolution of stromatolite microfabrics through time.
Extant marine stromatolites act as partial analogues of their Archean counterparts, but are rare due to depleted ocean calcium carbonate levels and suppression by eukaryotic organisms. Unique, peritidal tufa stromatolites at the interface between marine and freshwater inputs were discovered in South Africa in the past decade. Our aim was to investigate the benthic microalgal community (green algae, diatoms and cyanobacteria) of these stromatolites to assess succession and dominance patterns using real-time, in situ measurements of algal concentrations and composition. These biological measurements were modelled using generalised linear modelling (GLM) multivariate statistics against water physical and chemical parameters measured at regular monthly intervals, from January to December 2014. Salinity peaked and temperature dipped in winter, with both correlated to microalgal community change (GLM: p < 0.01). Diatoms and cyanobacteria, which construct the stromatolites, were consistently the dominant groups within the algal community, with minimal green algae present throughout the year. Importantly, this demonstrates a unique, relatively stable microalgal stromatolite community as opposed to those of other marine stromatolites, which likely require seasonal and stochastic disturbance to persist. This has implications in terms of interpreting community succession and differential layering in modern and fossilised stromatolites, respectively.
Abstract Raman spectrometers are being miniaturized for future life-detection missions on Mars. Field-portable Raman spectrometers, which have similar spectral parameters to the instruments being developed for Mars rovers, have been used to examine extant biosignatures, but they have not yet been used to examine ancient biosignatures. Here, a portable Raman spectrometer was used to analyze an Ordovician stromatolite at the outcrop, revealing both its mineralogy and the presence of sp(2) carbonaceous material. As stromatolites are often used as proof of the presence of life in Archean rocks and are searched for on Mars, the ability to analyze them in the field with no sample preparation has important ramifications for future Mars missions. However, these results also reveal that a 785 nm excitation source, rather than the 532 nm excitation source planned for future missions, might be a better choice in the search for fossil biosignatures. Key Words: Raman spectroscopy-Portable/miniaturized Raman instruments-Stromatolites-Carbonaceous material-Life detection. Astrobiology 13, xxx-xxx.