Shell microstructure and mineralogy of the Littorinidae: ecological and evolutionary significance

An examination of the shell microstructure and mineralogy of species from 30 of the 32 genera and subgenera of the gastropod family Littorinidae shows that most species have a shell consisting of layers of aragonitic crossed-lamellar structure, with minor variations in some taxa. However, Pellilitorina, Risellopsis and most species of Littorina have partly or entirely calcitic shells. In Pellilitorina the shell is made entirely of calcitic crossed-foliated structure, while in the other two genera there is only an outer calcitic layer of irregular-prismatic structure. A cladistic analysis shows that the calcitic layers have been independently evolved in at least three clades. The calcite is found only in the outermost layers of the shell and in species inhabiting cooler waters of both northern and southern hemispheres. Calcium carbonate is more soluble in cold than warm water and, of the two polymorphs, calcite is about 35% less soluble than aragonite. We suggest that calcitic shell layers are an adaptation of high latitude littorinids to resist shell dissolution.