The varix: Evolution, distribution, and phylogenetic clumping of a repeated gastropod innovation

A recurrent theme in evolution is the repeated, independent origin of broadly adaptive, architecturally and functionally similar traits and structures. One such is the varix, a shell-sculpture innovation in gastropods. This periodic shell thickening functions mainly to defend the animal against shell crushing and peeling predators. Varices can be highly elaborate, forming broad wings or spines, and are often aligned in synchronous patterns. Here we define the different types of varices, explore their function and morphological variation, document the recent and fossil distribution of varicate taxa, and discuss emergent patterns of evolution. We conservatively found 41 separate origins of varices, which were concentrated in the more derived gastropod clades and generally arose since the mid-Mesozoic. Varices are more prevalent among marine, warm, and shallow waters, where predation is intense, on high-spired shells and in clades with collabral ribs. Diversification rates were correlated in a few cases with the presence of varices, especially in the Muricidae and Tonnoidea, but more than half of the origins are represented by three or fewer genera. Varices arose many times in many forms, but generally in a phylogenetically clumped manner (more frequently in particular higher taxa), a pattern common to many adaptations