In scarcity of light and primary producers, subterranean ecosystems are generally extremely oligotrophic habitats, receiving poor supplies of degradable organic matter from the surface. Human direct impacts on cave ecosystems mainly derive from intensive tourism and recreational caving, causing important alterations to the whole subterranean environment. In particular, artificial lighting systems in show caves support the growth of autotrophic organisms (the so-called lampenflora), mainly composed of cyanobacteria, diatoms, chlorophytes, mosses and ferns producing exocellular polymeric substances (EPSs) made of polysaccharides, proteins, lipids and nucleic acids. This anionic EPSs matrix mediates to the intercellular communications and participates to the chemical exchanges with the substratum, inducing the adsorption of cations and dissolved organic molecules from the cave formations (speleothems). Coupled with the metabolic activities of heterotrophic microorganisms colonising such layer (biofilm), this phenomenon may lead to the corrosion of the mineral surfaces. In this review, we investigate the formation of biofilms, especially of diatom-dominated ones, as a consequence of artificial lighting and its impacts on speleothems. Whenever light reaches the subterranean habitat (both artificially and naturally) a relative high number of species of diatoms may indeed colonise it. Cave entrances, artificially illuminated walls and speleothems inside the cave are generally the preferred substrates. This review focuses on the diatom flora colonising subterranean habitats, summarizing the information contained in all the scientific papers published from 1900 up to date. In this review we provide a complete checklist of the diatom taxa recorded in subterranean habitats, including a total of 363 taxa, belonging to 82 genera. The most frequent and abundant species recorded in caves and other low light subterranean habitats are generally aerophilic and cosmopolitan. These are, in order of frequency: Hantzschia amphioxys, Diadesmis contenta, Orthoseira roeseana, Luticola nivalis, Pinnularia borealis, Diadesmis biceps and Luticola mutica. Due to the peculiarity of the subterranean habitats, the record of rare or new species is relatively common. The most important environmental factors driving species composition and morphological modifications observed in subterranean populations are analysed throughout the text and tables. In addition, suggestions to prevent and remove the corrosive biofilms in view of an environmentally sustainable cave management are discussed.

Tjuv-Antes grotta (Tjuv-Ante's Cave) located in northern Sweden is a round-abraded sea cave ('tunnel cave'), about 30 m in length, formed by rock-water abrasion in a dolerite dyke in granite gneiss. Abundant speleothems are restricted to the inner, mafic parts of the cave and absent on granite parts. The speleothems are of two types: cylindrical (coralloid, popcorn-like), and flowstone (thin crusts). Coralloids correspond to terrestrial stromatolite speleothems in which layers of light calcite alternate with dark, silica-rich laminae. The dark laminae are also enriched in carbon and contain incorporated remains of microorganisms. Two types of microbial communities can be distinguished associated with the speleothems: an Actinobacteria-like biofilm and a fungal community. Actinobacteria seem to play an important role in the formation of speleothem while the fungal community acts as both a constructive and a destructive agent. A modern biofilm dominated by Actinobacteria is present in the speleothem-free parts of the dolerite and located in cave ceiling cracks. These biofilms may represent sites of early speleothem formation. Because of its unusual position in between two types of host rock, Tjuv-Ante's Cave represents a unique environment in which to study differences in microbe-rock interactions and speleothem genesis between the granite and dolerite host rock. Our study shows that the mafic rock is superior to the granite in hosting a microbial community and to support formation of speleothems.

The karst cave ‘Vlychada’of Diros, one of the oldest show caves in Peloponnese, sustains extended phototrophic biofilms on various substrata – on rocks inside the cave including speleothems, and especially near the artificial lighting installation (‘Lampenflora’). After a survey of the main abiotic parameters (Photosynthetically Active Radiation -PAR, Temperature -T, Relative Humidity -RH, Carbon Dioxide -CO2) three clusters of sampling sites were revealed according to Principal Component Analysis (PCA): i) the water gallery section predominately influenced by CO2, ii) the dry passages influenced by RH and PAR, and iii) the area by the cave exit at the dry section influenced by temperature. The collected samples from the water gallery section and the dry passages of the cave revealed a total of 43 taxa of Cyanobacteria, with the unicellular/colonial forms being the most abundant. The applied non-metric Multi-dimensional Scaling Ordination (nMDS) of the cumulative species composition showed a clear distinction between the water gallery section and the dry passages of the cave. Further comparison with previous data from other wild caves of Peloponnese (‘Kastria’, ‘Francthi’, and ‘Selinitsa’) was conducted revealing a distinction between the show cave and the wild ones. Apart from the human impact on cave ecosystems – through aesthetic alteration (‘greening’) of cave decorations by the ‘Lampenflora’, and by the cleaning treatments and restoration projects on the speleothems – identification of the organisms constituting the ‘Lampenflora’ might provide taxonomically and ecologically significant taxa.

Helictites—an enigmatic type of mineral structure occurring in some caves—differ from classical speleothems as they develop with orientations that defy gravity. While theories for helictite formation have been forwarded, their genesis remains equivocal. Here, we show that a remarkable suite of helictites occurring in Asperge Cave (France) are formed by biologically-mediated processes, rather than abiotic processes as had hitherto been proposed. Morphological and petro-physical properties are inconsistent with mineral precipitation under purely physico-chemical control.

Instead, microanalysis and molecular-biological investigation reveals the presence of a prokaryotic biofilm intimately associated with the mineral structures. We propose that microbially-influenced mineralization proceeds within a gliding biofilm which serves as a nucleation site for CaCO3, and where chemotaxis influences the trajectory of mineral growth, determining the macroscopic morphology of the speleothems. The influence of biofilms may explain the occurrence of similar speleothems in other caves worldwide, and sheds light on novel biomineralization processes.