A morphogram for silica‐witherite biomorphs and its application to microfossil identification in the early earth rock record

Archean hydrothermal environments formed a likely site for the origin and early evolution of life. These are also the settings, however, were complex abiologic structures can form. Low‐temperature serpentinization of ultramafic crust can generate alkaline, silica‐saturated fluids in which carbonate–silica crystalline aggregates with life‐like morphologies can self‐assemble. These “biomorphs” could have adsorbed hydrocarbons from Fischer–Tropsch type synthesis processes, leading to metamorphosed structures that resemble carbonaceous microfossils. Although this abiogenic process has been extensively cited in the literature and has generated important controversy, so far only one specific biomorph type with a filamentous shape has been discussed for the interpretation of Archean microfossils. It is therefore critical to precisely determine the full distribution in morphology and size of these biomorphs, and to study the range of plausible geochemical conditions under which these microstructures can form. Here, a set of witherite‐silica biomorph synthesis experiments in silica‐saturated solutions is presented, for a range of pH values (from 9 to 11.5) and barium ion concentrations (from 0.6 to 40 mmol/L BaCl₂). Under these varying conditions, a wide range of life‐like structures is found, from fractal dendrites to complex shapes with continuous curvature. The size, spatial concentration, and morphology of the biomorphs are strongly controlled by environmental parameters, among which pH is the most important. This potentially limits the diversity of environments in which the growth of biomorphs could have occurred on Early Earth. Given the variety of the observed biomorph morphologies, our results show that the morphology of an individual microstructure is a poor criterion for biogenicity. However, biomorphs may be distinguished from actual populations of cellular microfossils by their wide, unimodal size distribution. Biomorphs grown by diffusion in silica gel can be differentiated by their continuous gradient in size, spatial density, and morphology along the direction of diffusion.     

Garnet-filled trails associated with carbonaceous matter mimicking microbial filaments in Archean basalt

The study of the earliest traces of life on Earth can be complicated by abiotically formed biomorphs. We report here the finding of clustered micrometer-sized filaments of iron- and calcium-rich garnets associated with carbonaceous matter in an agate amygdale from a 2.7-billion-year-old basalt of the Maddina Formation, Western Australia. The distribution of carbonaceous matter and the mineral phases composing the filaments were analyzed using a combination of confocal laser scanning microscopy, laser-Raman micro-spectroscopy, focused ion beam sectioning and transmission electron microscopy. The results allow consideration of possible biogenic and abiotic processes that produced the filamentous structures. The filaments have a range of sizes, morphologies and distributions similar to those of certain modern iron-mineralized filamentous bacteria and some ancient filamentous structures interpreted as microfossils. They also share a high morphological similarity with tubular structures produced by microbial boring activity. However, the microstructures and the distribution of carbonaceous matter are more suggestive of an abiotic origin for the filaments. They are characteristic features of trails produced by the displacement of inclusions associated with local dissolution of their silica matrix. Organic compounds found in kerogen or bitumen inclusions may have contributed significantly to the dissolution of the quartz (or silica gel) matrix driving filamentous growth. Discriminating the products of such abiotic organic-mediated processes from filamentous microfossils or microbial borings is important to the interpretation of the scarce Precambrian fossil record and requires investigation down to the nanoscale.     

Microbial life associated with low‐temperature alteration of ultramafic rocks in the Leka ophiolite complex

Water–rock interactions in ultramafic lithosphere generate reduced chemical species such as hydrogen that can fuel subsurface microbial communities. Sampling of this environment is expensive and technically demanding. However, highly accessible, uplifted oceanic lithospheres emplaced onto continental margins (ophiolites) are potential model systems for studies of the subsurface biosphere in ultramafic rocks. Here, we describe a microbiological investigation of partially serpentinized dunite from the Leka ophiolite (Norway). We analysed samples of mineral coatings on subsurface fracture surfaces from different depths (10–160 cm) and groundwater from a 50‐m‐deep borehole that penetrates several major fracture zones in the rock. The samples are suggested to represent subsurface habitats ranging from highly anaerobic to aerobic conditions. Water from a surface pond was analysed for comparison. To explore the microbial diversity and to make assessments about potential metabolisms, the samples were analysed by microscopy, construction of small subunit ribosomal RNA gene clone libraries, culturing and quantitative‐PCR. Different microbial communities were observed in the groundwater, the fracture‐coating material and the surface water, indicating that distinct microbial ecosystems exist in the rock. Close relatives of hydrogen‐oxidizing Hydrogenophaga dominated (30% of the bacterial clones) in the oxic groundwater, indicating that microbial communities in ultramafic rocks at Leka could partially be driven by H₂ produced by low‐temperature water–rock reactions. Heterotrophic organisms, including close relatives of hydrocarbon degraders possibly feeding on products from Fischer–Tropsch‐type reactions, dominated in the fracture‐coating material. Putative hydrogen‐, ammonia‐, manganese‐ and iron‐oxidizers were also detected in fracture coatings and the groundwater. The microbial communities reflect the existence of different subsurface redox conditions generated by differences in fracture size and distribution, and mixing of fluids. The particularly dense microbial communities in the shallow fracture coatings seem to be fuelled by both photosynthesis and oxidation of reduced chemical species produced by water–rock reactions.     

Geospatial insights into the controls of microbialite formation at Laguna Negra,Argentina

Microbialites provide a record of the interaction of microorganisms with their environment constituting a record of microbial life and environments through geologic time. Our capacity to interpret this record is limited by an incomplete understanding of the microbial, geochemical, and physical processes that influence microbialite formation and morphogenesis. The modern system Laguna Negra in Catamarca Province, Argentina contains microbialites in a zone of carbonate precipitation associated with physico-chemical gradients and variable microbial community structure, making it an ideal location to study how these processes interact to drive microbialite formation. In this study, we investigated the geospatial relationships between carbonate morphology, geochemistry, and microbial community at the macro- (decimeter) to mega- (meter) scale by combining high-resolution imagery with field observations. We mapped the distribution of carbonate morphologies and allochtonously-derived volcaniclasts and correlated these with sedimentary matrices and geochemical parameters. Our work shows that the macroscale distribution of different carbonate morphologies spatially correlates with microbial mat distributions—a result consistent with previous microscale observations. Specifically, microbialitic carbonate morphologies more commonly occur associated with microbial mats while abiotically derived carbonate morphologies were less commonly associated with microbial mats. Spatial variability in the size and abundance of mineralized structures was also observed, however, the processes controlling this variability remains unclear and likely represent a combination of microbial, geochemical, and physical processes. Likewise, the processes controlling the spatial distribution of microbial mats at Laguna Negra are also unresolved. Our results suggest that in addition to the physical drivers observed in other modern environments, variability in the spatial distribution of microbialites and other carbonate morphologies at the macro- to megascale can be controlled by microbial processes. Overall, this study provides insight into the interpretation of microbialite occurrence and distributions in the geologic record and highlights the utility of geospatial statistics to probe the controls of microbialite formation in other environments.     

Rock geochemistry induces stress and starvation responses in the bacterial proteome

Interactions between microorganisms and rocks play an important role in Earth system processes. However, little is known about the molecular capabilities microorganisms require to live in rocky environments. Using a quantitative label‐free proteomics approach, we show that a model bacterium (Cupriavidus metallidurans CH34) can use volcanic rock to satisfy some elemental requirements, resulting in increased rates of cell division in both magnesium‐ and iron‐limited media. However, the rocks also introduced multiple new stresses via chemical changes associated with pH, elemental leaching and surface adsorption of nutrients that were reflected in the proteome. For example, the loss of bioavailable phosphorus was observed and resulted in the upregulation of diverse phosphate limitation proteins, which facilitate increase phosphate uptake and scavenging within the cell. Our results revealed that despite the provision of essential elements, rock chemistry drives complex metabolic reorganization within rock‐dwelling organisms, requiring tight regulation of cellular processes at the protein level. This study advances our ability to identify key microbial responses that enable life to persist in rock environments.     

The ‘Island Rule’ Acting on Anuran Populations (Bufonidae: Rhinella ornata) of the Southern Hemisphere

Darwin and Wallace, in the mid‐nineteenth century, were the first to document examples of natural selection acting on island dwellers. A century later a pattern of morphological differences among organisms on islands was coined the ‘island rule’, which states that on islands species with small individuals tend toward gigantism and large individuals tend toward dwarfism. Selective pressures such as limited resources and increased intraspecific competition modulate the size of organisms in these environments. Of the several works that have tested vertebrates for adherence to the island rule only two have addressed amphibians. This work is the third record of body size variation of island amphibian populations, and the first for the Southern Hemisphere. The islands investigated were once continuous with mainland, and now are isolated as a result of sea level fluctuations that took place in the Pleistocene and Holocene. This study compared morphometric variation in populations of Rhinella ornata (Bufonidae) occurring on three islands of the Costa Verde to populations on five continental areas in Rio de Janeiro, Brazil. We measured 18 morphometric variables of 177 individuals. There was a shift toward smaller body size (dwarfism) in two of the three island populations studied. We attribute this general pattern to geographic factors, verifying the expression of the island rule in tropical frogs populations (insular dwarfism) operating inversely in relation to those of temperate environments (island gigantism).     

The geographic structure of morphological variation in eight species of fiddler crabs (Ocypodidae: genus Uca) from the eastern United States and Mexico

Species with larger geographic distributions are more likely to encounter a greater variety of environmental conditions and barriers to gene flow than geographically‐restricted species. Thus, even closely‐related species with similar life‐history strategies might vary in degree and geographic structure of variation if they differ in geographic range size. In the present study, we investigated this using samples collected across the geographic ranges of eight species of fiddler crabs (Crustacea: Uca) from the Atlantic and Gulf coasts of North America. Morphological variation in the carapace was assessed using geometric morphometric analysis of 945 specimens. Although the eight Uca species exhibit different degrees of intraspecific variation, widespread species do not necessarily exhibit more intraspecific or geographic variation in carapace morphology. Instead, species with more intraspecific variation show stronger morphological divergence among populations. This morphological divergence is partly a result of allometric growth coupled with differences in maximum body size among populations. On average, 10% of total within‐species variation is attributable to allometry. Possible drivers of the remaining morphological differences among populations include gene flow mediated by ocean currents and plastic responses to various environmental stimuli, with isolation‐by‐distance playing a less important role. The results obtained indicate that morphological divergence among populations can occur over shorter distances than expected based on dispersal potential. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 248–270.     

Behavioural response of Kootenai white sturgeon (Acipenser transmontanus,Richardson, 1836) early life stages to gravel,pebble, and rubble substrates: guidelines for rearing substrate size

Guidelines for creating rearing substrate for sturgeon early life stages are needed for restoration programmes creating habitats for spawning and rearing of early life stages. To determine the effects of rock size on motile early life stages, experiments were conducted in artificial streams to observe the behaviour of free embryos and larvae of Kootenai River white sturgeon (Acipenser transmontanus) relative to rock size. Most (≥90%) of the free embryos in replicate test streams with 100% gravel, 100% pebble, or 100% rubble hid under rocks, with few moving downstream. There was no difference in downstream movement of free embryos among rock treatments, therefore all rock types provided cover habitat. Similarly, in rock mixture tests, with a variable percentage of pebble, small rubble, or large rubble in different tanks, even fewer free embryos moved downstream. With increasing age, larvae increasingly used the open bottom and velocity refuges downstream of or alongside rocks of any size while drift feeding. Downstream movement of larvae in both rock regime tests was affected by rock size, with significantly reduced movement relative to increasing abundance of large rock (rubble). However, in all rock mixtures, free embryos (and later, larvae when they stopped dispersing) preferred the smallest rock size available (pebble; P = 0.0001). This suggests a strong innate preference of both life stages for small substrate that is likely related to increased survival. A rock mixture of 10% gravel (16–32 mm diameter) and 30–40% pebble (diameter, 30–60 mm) should provide adequate rearing substrate for free embryos and early‐larvae. The remaining 50–60% should be mixed rubble and boulders for spawning and egg rearing.     

The Search for Biomarkers and Microbial Fossils in Antarctic Rock Microhabitats

The survival of lithobiontic microbial communities in Antarctica's extreme cold, dry environment is conditioned by changes in external climatic conditions that can lead to the death of these microorganisms. In the present study, granite samples from maritime Antarctica and sandstone from the Antarctic continental desert were collected with the aim of searching for biomarkers and microbial fossils at the microscopic level of observation. The results reveal the presence of inorganic biomarkers in the form of physicochemical bioweathering mineral patterns, and inorganic deposits such as calcium oxalates and silica. The presence of fossilised algae and other microorganisms within the sandstone rock was also confirmed. Identifying the internal cell structure within the fossilised cells is proposed as a new criterion for the biogenicity of biomorphs.     

Sulfur isotope's signal of nanopyrites enclosed in 2.7 Ga stromatolitic organic remains reveal microbial sulfate reduction

Microbial sulfate reduction (MSR) is thought to have operated very early on Earth and is often invoked to explain the occurrence of sedimentary sulfides in the rock record. Sedimentary sulfides can also form from sulfides produced abiotically during late diagenesis or metamorphism. As both biotic and abiotic processes contribute to the bulk of sedimentary sulfides, tracing back the original microbial signature from the earliest Earth record is challenging. We present in situ sulfur isotope data from nanopyrites occurring in carbonaceous remains lining the domical shape of stromatolite knobs of the 2.7‐Gyr‐old Tumbiana Formation (Western Australia). The analyzed nanopyrites show a large range of δ³⁴S values of about 84‰ (from ?33.7‰ to +50.4‰). The recognition that a large δ³⁴S range of 80‰ is found in individual carbonaceous‐rich layers support the interpretation that the nanopyrites were formed in microbial mats through MSR by a Rayleigh distillation process during early diagenesis. An active microbial cycling of sulfur during formation of the stromatolite may have facilitated the mixing of different sulfur pools (atmospheric and hydrothermal) and explain the weak mass independent signature (MIF‐S) recorded in the Tumbiana Formation. These results confirm that MSR participated actively to the biogeochemical cycling of sulfur during the Neoarchean and support previous models suggesting anaerobic oxidation of methane using sulfate in the Tumbiana environment.     

A New Frontier for Palaeobiology: Earth's Vast Deep Biosphere

Diverse micro‐organisms populate a global deep biosphere hosted by rocks and sediments beneath land and sea, containing more biomass than any other biome except forests. This paper reviews an emerging palaeobiological archive of these dark habitats: microfossils preserved in ancient pores and fractures in the crust. This archive, seemingly dominated by mineralized filaments (although rods and coccoids are also reported), is presently far too sparsely sampled and poorly understood to reveal trends in the abundance, distribution, or diversity of deep life through time. New research is called for to establish the nature and extent of the fossil record of Earth's deep biosphere by combining systematic exploration, rigorous microanalysis, and experimental studies of both microbial preservation and the formation of abiotic pseudofossils within the crust. It is concluded that the fossil record of Earth's largest microbial habitat may still have much to tell us about the history of life, the evolution of biogeochemical cycles, and the search for life on Mars.     

Nanometer‐scale characterization of exceptionally preserved bacterial fossils in Paleocene phosphorites from Ouled Abdoun (Morocco)

Micrometer‐sized spherical and rod‐shaped forms have been reported in many phosphorites and often interpreted as microbes fossilized by apatite, based on their morphologic resemblance with modern bacteria inferred by scanning electron microscopy (SEM) observations. This interpretation supports models involving bacteria in the formation of phosphorites. Here, we studied a phosphatic coprolite of Paleocene age originating from the Ouled Abdoun phosphate basin (Morocco) down to the nanometer‐scale using focused ion beam milling, transmission electron microscopy (TEM), and scanning transmission x‐ray microscopy (STXM) coupled with x‐ray absorption near‐edge structure spectroscopy (XANES). The coprolite, exclusively composed of francolite (a carbonate‐fluroapatite), is formed by the accumulation of spherical objects, delimited by a thin envelope, and whose apparent diameters are between 0.5 and 3 μm. The envelope of the spheres is composed of a continuous crown dense to electrons, which measures 20–40 nm in thickness. It is surrounded by two thinner layers that are more porous and transparent to electrons and enriched in organic carbon. The observed spherical objects are very similar with bacteria encrusting in hydroxyapatite as observed in laboratory experiments. We suggest that they are Gram‐negative bacteria fossilized by francolite, the precipitation of which started within the periplasm of the cells. We discuss the role of bacteria in the fossilization mechanism and propose that they could have played an active role in the formation of francolite. This study shows that ancient phosphorites can contain fossil biological subcellular structures as fine as a bacterial periplasm. Moreover, we demonstrate that while morphological information provided by SEM analyses is valuable, the use of additional nanoscale analyses is a powerful approach to help inferring the biogenicity of biomorphs found in phosphorites. A more systematic use of this approach could considerably improve our knowledge and understanding of the microfossils present in the geological record.     

Modern Subsurface Bacteria in Pristine 2.7 Ga-Old Fossil Stromatolite Drillcore Samples from the Fortescue Group,Western Australia

Background

Several abiotic processes leading to the formation of life-like signatures or later contamination with actual biogenic traces can blur the interpretation of the earliest fossil record. In recent years, a large body of evidence showing the occurrence of diverse and active microbial communities in the terrestrial subsurface has accumulated. Considering the time elapsed since Archaean sedimentation, the contribution of subsurface microbial communities postdating the rock formation to the fossil biomarker pool and other biogenic remains in Archaean rocks may be far from negligible.

Methodology/Principal Findings

In order to evaluate the degree of potential contamination of Archean rocks by modern microorganisms, we looked for the presence of living indigenous bacteria in fresh diamond drillcores through 2,724 Myr-old stromatolites (Tumbiana Formation, Fortescue Group, Western Australia) using molecular methods based on the amplification of small subunit ribosomal RNA genes (SSU rDNAs). We analyzed drillcore samples from 4.3 m and 66.2 m depth, showing signs of meteoritic alteration, and also from deeper “fresh” samples showing no apparent evidence for late stage alteration (68 m, 78.8 m, and 99.3 m). We also analyzed control samples from drilling and sawing fluids and a series of laboratory controls to establish a list of potential contaminants introduced during sample manipulation and PCR experiments. We identified in this way the presence of indigenous bacteria belonging to Firmicutes, Actinobacteria, and Alpha-, Beta-, and Gammaproteobacteria in aseptically-sawed inner parts of drillcores down to at least 78.8 m depth.

Conclusions/Significance

The presence of modern bacterial communities in subsurface fossil stromatolite layers opens the possibility that a continuous microbial colonization had existed in the past and contributed to the accumulation of biogenic traces over geological timescales. This finding casts shadow on bulk analyses of early life remains and makes claims for morphological, chemical, isotopic, and biomarker traces syngenetic with the rock unreliable in the absence of detailed contextual analyses at microscale.     

Diversity decoupled from sulfur isotope fractionation in a sulfate‐reducing microbial community

The extent of fractionation of sulfur isotopes by sulfate‐reducing microbes is dictated by genomic and environmental factors. A greater understanding of species‐specific fractionations may better inform interpretation of sulfur isotopes preserved in the rock record. To examine whether gene diversity influences net isotopic fractionation in situ, we assessed environmental chemistry, sulfate reduction rates, diversity of putative sulfur‐metabolizing organisms by 16S rRNA and dissimilatory sulfite reductase (dsrB) gene amplicon sequencing, and net fractionation of sulfur isotopes along a sediment transect of a hypersaline Arctic spring. In situ sulfate reduction rates yielded minimum cell‐specific sulfate reduction rates < 0.3 × 10^?15 moles cell^?1 day^?1. Neither 16S rRNA nor dsrB diversity indices correlated with relatively constant (38‰–45‰) net isotope fractionation (ε³⁴S_{sulfide‐sulfate}). Measured ε³⁴S values could be reproduced in a mechanistic fractionation model if 1%–2% of the microbial community (10%–60% of Deltaproteobacteria) were engaged in sulfate respiration, indicating heterogeneous respiratory activity within sulfate‐reducing populations. This model indicated enzymatic kinetic diversity of Apr was more likely to correlate with sulfur fractionation than DsrB. We propose that, above a threshold Shannon diversity value of 0.8 for dsrB, the influence of the specific composition of the microbial community responsible for generating an isotope signal is overprinted by the control exerted by environmental variables on microbial physiology.     

Cellular automaton simulation of tumour growth -- equivocal relationships between simulation parameters and morphologic pattern features.

OBJECTIVE: To develop an interpretation procedure which estimates simulation parameters (tumour cell motility, tumour cell adhesion, autocrine and paracrine growth control, stroma destruction) of simulated patterns solely based on morphometric features of the morphologic pattern. METHODS: A cellular automaton computer simulation program was developed which produces morphologic patterns by growth of a seed of tumour cells. At the beginning of each simulation run certain simulation parameters are assigned to the tumour cells. After the run has been completed, the resulting pattern is evaluated by a set of morphometric features. Simulation parameters and resulting morphometric features of 27,800 simulations were stored in a database and were used for the evaluation of potential relationships. RESULTS: Correlation analysis showed highly significant correlations between morphometric features on the one hand and the preset simulation parameters (tumour cell motility, tumour cell adhesion, autocrine and paracrine growth control, stroma destruction) on the other. Correlation coefficients, however, varied from 0.72 to 0.99. When only one simulation parameter varied while all others were kept constant, morphometric features yielded a highly reliable estimate of the particular simulation parameter. When variability was extended to 4 simulation parameters, morphometric features were less effective in estimating the setting of the parameters. Though in all patterns tested several possible simulation parameter constellations could be ruled out, morphometric features were usually compatible with more than one set of simulation parameters thus preventing a straightforward interpretation. CONCLUSIONS: Though simulation parameters significantly and reproducibly influence the resulting morphologic pattern as characterized by morphometric features, estimates of the simulation parameters based on morphometric features yield equivocal results.     

Limited influence of Si on the preservation of Fe mineral‐encrusted microbial cells during experimental diagenesis

The reconstruction of the history of microbial life since its emergence on early Earth is impaired by the difficulty to prove the biogenicity of putative microfossils in the rock record. While most of the oldest rocks on Earth have been exposed to different grades of diagenetic alterations, little is known about how the remains of micro‐organisms evolve when exposed to pressure (P) and temperature (T) conditions typical of diagenesis. Using spectroscopy and microscopy, we compared morphological, mineralogical, and chemical biosignatures exhibited by Fe mineral‐encrusted cells of the bacterium Acidovorax sp. BoFeN1 after long‐term incubation under ambient conditions and after experimental diagenesis. We also evaluated the effects of Si on the preservation of microbial cells during the whole process. At ambient conditions, Si affected the morphology but not the identity (goethite) of Fe minerals that formed around cells. Fe‐encrusted cells were morphologically well preserved after 1 week at 250 °C‐140 MPa and after 16 weeks at 170 °C‐120 MPa in the presence or in the absence of Si. Some goethite transformed to hematite and magnetite at 250 °C‐140 MPa, but in the presence of Si more goethite was preserved. Proteins—the most abundant cellular components—were preserved over several months at ambient conditions but disappeared after incubations at high temperature and pressure conditions, both in the presence and in the absence of Si. Other organic compounds, such as lipids and extracellular polysaccharides seemed well preserved after exposure to diagenetic conditions. This study provides insights about the composition and potential preservation of microfossils that could have formed in Fe‐ and Si‐rich Precambrian oceans.     

海底玄武岩玻璃中蚀变微结构:探索海洋深地生物圈的新材料

玄武岩玻璃中的生物蚀变微结构为微生物摄取玄武岩玻璃中营养成分,通过新陈代谢产生有机酸溶解玄武岩玻璃而形成的微米级孔洞或钻穴.生物蚀变微结构在现代海洋洋壳及代表古老洋壳残片的蛇绿岩和绿岩带中广泛存在.研究玄武岩玻璃中蚀变微结构的形态特征、形成机制及其时空分布,不仅对探索地球早期生命起源和演化具有重要启示意义,也为研究海洋...     

Tales from the tomb: the microbial ecology of exposed rock surfaces

Although a broad diversity of eukaryotic and bacterial taxa reside on rock surfaces where they can influence the weathering of rocks and minerals, these communities and their contributions to mineral weathering remain poorly resolved. To build a more comprehensive understanding of the diversity, ecology and potential functional attributes of microbial communities living on rock, we sampled 149 tombstones across three continents and analysed their bacterial and eukaryotic communities via marker gene and shotgun metagenomic sequencing. We found that geographic location and climate were important factors structuring the composition of these communities. Moreover, the tombstone‐associated microbial communities varied as a function of rock type, with granite and limestone tombstones from the same cemeteries harbouring taxonomically distinct microbial communities. The granite and limestone‐associated communities also had distinct functional attributes, with granite‐associated bacteria having more genes linked to acid tolerance and chemotaxis, while bacteria on limestone were more likely to be lichen associated and have genes involved in photosynthesis and radiation resistance. Together these results indicate that rock‐dwelling microbes exhibit adaptations to survive the stresses of the rock surface, differ based on location, climate and rock type, and seem pre‐disposed to different ecological strategies (symbiotic versus free‐living lifestyles) depending on the rock type.