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As well as most hypogean taxa [ 31 ], troglobiomorphic spiders represent filtered subsamples of regional epigean species pools [ 4 ]. However, there are genera and families without epigean relatives, like Trogloraptoridae [ 32 ]. In terms of foraging guilds sensu [ 31 ] , a functional disharmony is often observed, with entire guilds being scarcely represented, if not absent, in caves—e.
Aside from the notable exception of the Iberian Peninsula [ 4 ], there are no extensive compilations about functional groups in subterranean spiders, making it difficult to draw any general pattern. As for other taxonomic groups [ 33 ], the core of the diversity is found in temperate regions—especially in the Northern Hemisphere. In this regard, cave fauna in the tropics is probably understudied [ 47 ], and it is likely that the number of tropical troglobiont spiders will grow in the near future [ 40 ].
Subterranean adaptations Species under the same selective pressure evolve convergent adaptive traits. Being subjected to strong environmental filters, hypogean habitats are a remarkable example of this phenomenon [ 1 ]. The primary limiting factors for subterranean species are the lack of solar radiation and the scarcity of food resources—but see [ 48 ] for an in-depth discussion. The process of adaptation implies the evolution of well-defined and often convergent biological traits [ 49 ].
Indeed, troglobiomorphic spiders share adaptive features found in other groups of cave animals, supposedly due to the strong and similar selective pressures acting underground [ 1 ]. We here discuss the details on these adaptations, grouping them according to morphology, physiology and behaviour. They include reduction or loss of cuticular pigments, eye regression or loss, thinning of integuments, leg elongation and heavy spination [ 38 , 39 , 50 — 59 ]. Data referring to the European species of Troglohyphantes Linyphiidae suggest that pigmentation is the first character undergoing regression in the process of subterranean adaptation [ 51 ], whereas the loss of pigment around the eyes and the progressive eye reduction represent a second step in the adaptation process [ 51 , 59 ].
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