Mollusc Shell - Evolution

Evolution

The fossil record shows that all molluscan classes evolved some 500 million years ago from a shelled ancestor looking something like a modern monoplacophoran, and that modifications of the shell form ultimately led to the formation of new classes and lifestyles. However, a growing body of molecular and biological data indicate that at least certain shell features have evolved many times, independently. The nacreous layer of shells is a complex structure, but rather than being difficult to evolve, it has in fact arisen many times convergently. The genes used to control its formation vary greatly between taxa: under 10% of the (non-housekeeping) genes expressed in the shells that produce gastropod nacre are also found in the equivalent shells of bivalves: and most of these shared genes are also found in mineralizing organs in the deuterostome lineage. The independent origins of this trait are further supported by crystallographic differences between clades: the orientation of the axes of the deposited aragonite 'bricks' that make up the nacreous layer is different in each of the monoplacophora, gastropods and bivalves.

Mollusc shells (especially those formed by marine species) are very durable and outlast the otherwise soft-bodied animals that produce them by a very long time (sometimes thousands of years even without being fossilized). Most shells of marine molluscs fossilize rather easily, and fossil mollusc shells date all the way back to the Cambrian period. Large amounts of shells sometimes form sediment, and over a geological time span can become compressed into limestone deposits.

Most of the fossil record of molluscs consists of their shells, since the shell is often the only mineralised part of a mollusc (however also see Aptychus and operculum). The shells are usually preserved as calcium carbonate – usually any aragonite is pseudomorphed with calcite. Aragonite can be protected from recrystalization if water is kept away by carbonaceous material, but this did not accumulate in sufficient quantity until the Carboniferous; consequently aragonite older than the Carboniferous is practically unknown: but the original crystal structure can sometimes be deduced in fortunate circumstances, such as if an alga closely encrusts the surface of a shell, or if a phosphatic mould quickly forms during diagenesis.

The shell-less aplacophora have a chitinous cuticle that has been likened to the shell framework; it has been suggested that tanning of this cuticle, in conjunction with the expression of additional proteins, could have set the evolutionary stage for the secretion of a calcareous shell in an aplacophoran-like ancestral mollusc.

The molluscan shell has been internalized in a number of lineages, including the coleoid cephalopods and many gastropod lineages. Detorsion of gastropods results in an internal shell, and can be triggered by relatively minor developmental modifications such as those induced by exposure to high platinum concentrations.