World experts of different disciplines, from molecular biology to macroecology, recognize the value of cave ecosystems as ideal ecological and evolutionary laboratories. Among other subterranean taxa, spiders stand out as intriguing model organisms for their ecological role of top predators, their unique adaptations to the hypogean medium and their sensitivity to anthropogenic disturbance. As the description of the first eyeless spider (Stalita taenaria), an array of papers on subterranean spider biology, ecology and evolution has been published, but a comprehensive review on these topics is still lacking. We provide a general overview of the spider families recorded in hypogean habitats worldwide, we review the different adaptations of hypogean spiders to subterranean life, and we summarize the information gathered so far about their origin, population structure, ecology and conservation status. Finally, we point out the limits of the knowledge we currently have regarding hypogean spiders, aiming to stimulate future research.
BackgroundReceiving poor energy inputs and being light-deficient, caves are generally considered extreme environments, characterized by low abundance and diversity of organisms. Because of their extraordinary adaptations, cave-dwelling animals offer unique study opportunities for pushing forward our current understanding of evolutionary and ecological processes [1,2]. Among other subterranean taxa, spiders are distinctive for their ecological role of top predators [3][4][5], and for the variety of functional adaptations [6], therefore representing undervalued models for the understanding the evolution of life in extreme habitats. Accordingly, subterranean spiders have served as models for physiological [7][8][9], ecological [4,5,[10][11][12], ethological [13,14] and biogeographic studies [15][16][17], among others.However, compared with other animal groups, the potential of spiders as model organisms is still under-expressed. It is possible that the paucity of studies is related to the difficulty of working in subterranean habitats and to the rarity of most subterranean species, which pose major impediments to data collection and analysis. In addition, the general lack of a state of the art on subterranean spider biology, ecology and evolution conceivably hinders advances in knowledge. In this review, we present a collection of information about spiders colonizing subterranean habitats-especially caves-and discuss their relevance in the understanding of cave life evolution and the ecology of subterranean animal communities.
(a) Terminology and acronymsWe use the term troglobiomorphism to distinguish the suite of adaptations, especially morphological, to subterranean life [18]. Despite the fact that ecological categorizations often oversimplify real cases and boundaries between categories can be vague, for the sake of this review, we use the traditional speleobiological nomenclature [19] to indicate ecological distributions of species:(1) a troglobiont is strongly bound to hypogean habitats...