How did the carrier shell Xenophora crispa (König, 1825) build its shell? Evidence from the Recent and fossil record

The genus Xenophora comprises species of marine gastropods (Cretaceous-Recent) able to add fragments of various origins to their shell surface. Agglutination potentials vary, from species lacking attachments to species completely covered by agglutinated materials, as in the Mediterranean species Xenophora crispa. Here, we analyse Recent and fossil specimens of Xenophora crispa from the Mediterranean area using SEM and XRD, to better understand their biomineralization patterns and the mechanisms leading to the agglutination of shells, bioclasts and lithoclasts, and their evolution in time. We also provide new data on poorly studied gastropod shell microstructures. We conclude that: (1) most of the Xenophora crispa shell consists of an aragonitic crossed lamellar fabric, but fibrous to spherulitic prismatic fabrics, seemingly of calcite, have been found in the columella and peripheral edge (the thickest parts of the shell); (2) attachment of objects is mediated by a prismatic microstructure, indicating that this may be the most functional fabric in attachment areas in molluscs; and (3) the functional purpose of the agglutination in Xenophora crispa may be related to a snowshoe strategy to successfully colonize muddy substrates, coupled with tactile and olfactory camouflage. Indeed, this species secretes in the columella and peripheral edge a less dense and a more organic rich calcitic fabric, possibly to lighten the shell thickest parts in order not to sink in soft sediments and to facilitate the shell raising from the substrate to create a protected feeding area. This behaviour seems to have been maintained by X. crispa over 2 My time span.