Diatom flora in subterranean ecosystems: a review

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 diatomdominated 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.