Chemosynthetic sediment and planktonic community composition and sizes, aqueous geochemistry and sediment mineralogy were determined in 15 non-photosynthetic hot springs in Yellowstone National Park (YNP). These data were used to evaluate the hypothesis that differences in the availability of dissolved or mineral substrates in the bulk fluids or sediments within springs coincides with ecologically differentiated microbial communities and their populations. Planktonic and sediment-associated communities exhibited differing ecological characteristics including community sizes, evenness and richness. pH and temperature influenced microbial community composition among springs, but within-spring partitioning of taxa into sediment or planktonic communities was widespread, statistically supported (P < 0.05) and could be best explained by the inferred metabolic strategies of the partitioned taxa. Microaerophilic genera of the Aquificales predominated in many of the planktonic communities. In contrast, taxa capable of mineral-based metabolism such as S(o) oxidation/reduction or Fe-oxide reduction predominated in sediment communities. These results indicate that ecological differentiation within thermal spring habitats is common across a range of spring geochemistry and is influenced by the availability of dissolved nutrients and minerals that can be used in metabolism.

The concept that life emerged where alkaline hydrogen-bearing submarine hot springs exhaled into the most ancient acidulous ocean was used as a working hypothesis to investigate the nature of precipitate membranes. Alkaline solutions at 25-70°C and pH between 8 and 12, bearing HS(-)±silicate, were injected slowly into visi-jars containing ferrous chloride to partially simulate the early ocean on this or any other wet and icy, geologically active rocky world. Dependent on pH and sulfide content, fine tubular chimneys and geodal bubbles were generated with semipermeable walls 4-100 μm thick that comprised radial platelets of nanometric mackinawite [FeS]±ferrous hydroxide [∼Fe(OH)(2)], accompanied by silica and, at the higher temperature, greigite [Fe(3)S(4)]. Within the chimney walls, these platelets define a myriad of micropores. The interior walls of the chimneys host iron sulfide framboids, while, in cases where the alkaline solution has a pH>11 or relatively low sulfide content, their exteriors exhibit radial flanges with a spacing of ∼4 μm that comprise microdendrites of ferrous hydroxide. We speculate that this pattern results from outward and inward radial flow through the chimney walls. The outer Fe(OH)(2) flanges perhaps precipitate where the highly alkaline flow meets the ambient ferrous iron-bearing fluid, while the intervening troughs signal where the acidulous iron-bearing solutions could gain access to the sulfidic and alkaline interior of the chimneys, thereby leading to the precipitation of the framboids. Addition of soluble pentameric peptides enhances membrane durability and accentuates the crenulations on the chimney exteriors. These dynamic patterns may have implications for acid-base catalysis and the natural proton motive force acting through the matrix of the porous inorganic membrane. Thus, within such membranes, steep redox and pH gradients would bear across the nanometric platelets and separate the two counter-flowing solutions, a condition that may have led to the onset of an autotrophic metabolism through the reduction of carbon dioxide.

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Thermodynamic calculations provide valuable insights into the reactions that drive the profound fluid transformations during serpentinization, where surface fluids are transformed into some of the most reduced and alkaline fluids on Earth. However, environmental observations usually deviate from thermodynamic predictions, especially those occurring at low temperatures where equilibrium is slowly reached. In this work, we analyzed 138 low‐temperature (<40°C) fluids from the Samail ophiolite in Oman to test thermodynamic predictions with environmental observations. Four fluid types were identified through this work. (i) Type 1 circumneutral (pH 7–9) fluids result from fluid interactions with serpentinized rocks common in the shallow subsurface. (ii) Fluids with pH ranging from 9 to 11 and low Si concentrations are products of intermediate stages of serpentinization. (iii) Type 2 hyperalkaline (pH > 11) fluids approach equilibrium with diopside, and with serpentine and brucite actively forming during advanced stages of serpentinization. Lastly, (iv) most fluids sampled in this work deviate from predicted equilibrium compositions and depict various degrees of mixing between Type 1 and 2 fluids. Mixed fluids fall within the same pH range but have considerably higher dissolved Si than intermediate‐type fluids. Hyperalkaline fluids exhibit variable degrees of mixing despite maintaining pH > 11, implying strong buffering capacity of serpentinization‐generated fluids. Overall, this work demonstrates that predicted and measured compositions of serpentinization‐derived fluids can be reconciled using a combination of equilibrium and fluid‐transport simulations. This work substantiates these calculations as useful tools in exploring serpentinization reactions in continents and perhaps in other low‐temperature environments on Earth and beyond.

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