Operculum (gastropod)

The operculum (plural: opercula or operculums), meaning little lid, is a corneous or calcareous anatomical structure like a trapdoor which exists in many (but not all) groups of sea snails and freshwater snails, and also in a few groups of land snails; the structure is found in some marine and freshwater gastropods, and in a minority of terrestrial gastropods, including the families Helicinidae, Cyclophoridae, Aciculidae, Maizaniidae, Pomatiidae, etc.Example of a concentric operculum (Filopaludina martensi)Example of a polygyrous spiral with eccentric nucleus in the operculum of Marstonia comalensis, scale bar: 200 μmExample of an oligogyrous spiral in Cremnoconchus syhadrensisFossil spiral opercula (both sides) from the Pliocene of Cyprus The operculum (plural: opercula or operculums), meaning little lid, is a corneous or calcareous anatomical structure like a trapdoor which exists in many (but not all) groups of sea snails and freshwater snails, and also in a few groups of land snails; the structure is found in some marine and freshwater gastropods, and in a minority of terrestrial gastropods, including the families Helicinidae, Cyclophoridae, Aciculidae, Maizaniidae, Pomatiidae, etc. The operculum is attached to the upper surface of the foot and in its most complete state, it serves as a sort of 'trapdoor' to close the aperture of the shell when the soft parts of the animal are retracted. The shape of the operculum varies greatly from one family of gastropods to another. It is fairly often circular, or more or less oval in shape. In species where the operculum fits snugly, its outline corresponds exactly to the shape of the aperture of the shell and it serves to seal the entrance of the shell. Many families have opercula that are reduced in size, and which are not capable of closing the shell aperture. Opercula have sometimes been modified: in the Strombidae the operculum is claw-shaped and is used to push into the substrate in a leaping form of locomotion. Virtually all pulmonate snails are inoperculate, i.e. they do not have an operculum, with the exception of the Amphiboloidea. However, some terrestrial pulmonate species are capable of secreting an epiphragm, a temporary structure that can in some cases serve some of the same functions as an operculum. The epiphragm may be distinguished from the true operculum by its homogeneity and want of growth marks. In ammonites (extinct shelled cephalopods), a calcareous structure known as the aptychus (plural aptychi) existed. When these were first described they were thought to be valves of a bivalve species, then for many years after that they were considered to be a form of paired or single operculum-like structures belonging to ammonites. More recently the aptychus or paired aptychi have been hypothesized to be a jaw apparatus of ammonites. Perhaps the most essential function of the operculum in gastropods is to allow snails to resist drying out, or desiccation. This is very important in intertidal marine snails during low tide, and this also enables operculate freshwater and land snails to survive periods of drought and dry weather. In those marine species where the operculum completely seals the shell, it can also serve as a protection against predators when the snail body is retracted. In life, the operculum is attached at the ending of the columellar muscle with an opercular disc dorsally to the upper surface of the posterior part of the foot. However, in Buccinum a layer of long cylindrical epithelial cells, with mostly distinct nuclei, and long divided processes enter between the muscular fibres. The operculum, a cuticular development of these cells, is composed, as may be seen in the corneous opercula of Murex,Purpura, Triton, of very thin superimposed layers. The cylindrical cells are attached with their head to the lowermost layer. The operculum grows in size as the shell grows, such that the operculum remains in proportion to the apertural size. In many species, when the animal is active and crawling, part of the underside of the shell rests on the outer surface of the operculum.