Cave exploitation by an usual epigean species: a review on the current knowledge on fire salamander breeding in cave

The fire salamander (Salamandra salamandra) is a relatively common epigean amphibian, widely distributed throughout Europe, which usually gives birth to aquatic larvae. Even if epigean streams represent the most common places in which the species breeds, in some countries caves with underground waters are also used. To improve our understanding of the habitat features allowing successful breeding of salamanders in underground sites, we combined an exhaustive review of the available literature, especially the grey one, with direct observations performed from 2008 to 2017 in several natural and artificial caves of Lombardy, Liguria and Tuscany (Italy), Ariège and Provence (France). We provide a synthesis of published and unpublished caves in which the fire salamander breeding has been observed, along with a synthesis of the investigated ecological, behavioural and morphological traits. The use of underground sites is reported in several published papers and appears to be a common phenomenon not limited to single karst areas. The absence of predators, the relative stability of the aquatic habitats and the possibility to exploit new ecological resources are environmental factors that favour the breeding of the fire salamander. Our synthesis suggests that breeding of fire salamanders in caves is not a random event, but a widespread phenomenon that may be linked to specific biogeographical factors. Further insights may be obtained by performing genetic analyses on both cave and epigean populations, and considering larger landscape scales for ecological studies as well. Gene flow between salamanders that breed in caves and in streams probably occurs, but on the other hand, assortative mating might limit it, thus allowing the conservation of local adaptations driving successful cave colonisation