Frutexites encrustations on rugose corals (Middle Devonian, southern Morocco): complex growth of microbial microstromatolites

Microbially induced Frutexites microstromatolites developed on corallites of the Givetian rugose coral Amplexus in the sedimentary cover of a submarine volcanic intrusion in the eastern Anti-Atlas of southern Morocco. The corals lived in proximity to submarine hydrothermal fluid emissions. Frutexites form irregular shrubs encrusting external walls of corallites. The shrubs, consisting of alternations of calcitic and hematitic laminae, grew predominantly on abandoned corallites. Some Frutexites grew within the sediment, whereas others developed entirely above the seafloor, or started their accretion in water and continued to grow during burial. Three types of Frutexites encrustations have been distinguished. They look similar macroscopically, but differ significantly in their microstructure and mineralogical characteristics, resulting primarily from spatial and temporal variations in redox conditions. Both hematitic and calcitic laminae are primary features of Frutexites. The shrubs accreted as a result of mineralization of microbial biofilms under fluctuating environmental conditions, caused by changes in pH, Fe2+-supply and/or oxygenation, presumably related to discharges of reducing hydrothermal fluids. The calcitic laminae are interpreted to have formed due to activity of heterotrophic (mainly sulphate-reducing) microbes, whereas the hematitic laminae developed as a result of enhanced activity of nitrate-reducers or due to periodical passive mineralization of biofilms with iron. Cathodoluminescence data provide evidence that the nitrate and sulphate reduction preceded the Mn(IV) and Fe(III) reduction, presumably due to increased accumulation of organic matter and a high stability of the iron oxides present.