Terms employed to denote various Parts of the Univalve Shell.—The apex is the extreme top of the spire, and generally consists of the embryonic shell, which may often be recognised by its entire want of sculpture. When the embryonic shell happens to be large, the apex is often mammillated, e.g. in Fusus, Neptunea, and some Turbinella; in the Pyramidellidae it is sinistral.

The suture is the line of junction between any two successive whorls. It may be deep, and even channelled, or very shallow, as in Fig. 150 B (p. 246).

The spire is the whole series of whorls except the last or body whorl. A whorl is a single revolution of the spiral cone round the axis. The spire may be subulate (as in Terebra, Fig. 150 C), turreted (Scalaria), depressed (Polygyratia, Fig. 150 A), conical (Trochus), globose (Ampullaria, Natica, Fig. 150 B), with almost all conceivable gradations between these types. The number of whorls is best counted by placing the shell mouth downwards, and reckoning one for every suture that occurs between the extreme anterior point of the shell and the apex.

The mouth or aperture may be (a) entire, as in Helix, Natica, Ampullaria, when its peristome or margin is not interrupted by any notch or canal, or (b) prolonged at its anterior and sometimes also at its posterior end into a canal. The anterior canal serves as a protection to the siphon,[347] the posterior canal is mainly anal in function, and corresponds, in part, to the hole of Fissurella, the slit in Pleurotoma and Emarginula, and the row of holes in Haliotis. The mouth presents every variety of shape, from the perfect circle in Cyclostoma and Trochus, to the narrow and prolonged slit in Conus and Oliva.

The right margin of the mouth (the left, in sinistral shells) is termed the outer lip or labrum, the left margin the inner lip, labium, or columella lip.[348] In young shells the outer lip is usually thin and unfinished, while in the adult it is generally thickened into a rib, or furnished with more or less prominent teeth, or given an inward or outward curve. In some genera, especially the Strombidae, the outer lip of the adult develops long finger-like processes, which sometimes attain an extraordinary size (chap. xiv.). As growth proceeds, these marginal teeth and ribs are either dissolved and disappear, or are permanently incorporated, in the shape of varices, with the framework of the shell. Some shells, e.g. Natica, Turritella, Actaeon, have a permanently unfinished outer lip, even in the adult stage. The columella lip varies in shape with the mouth as a whole; thus it may be straight, as in Conus, or excavated, as in Sigaretus, Struthiolaria, and Bulla. Frequently it is continued by part of the body whorl, as in Ficula, Dolium, and Fasciolaria.

The folds or plaits on the columella, which are often characteristic of the genus or even family (e.g. Fasciolariidae, Mitridae, Turbinellidae) are not merely external, but continue down the whole spire (see Fig. 177, which also shows how successive fresh growths have thickened the columella).

The whorls may be wound in a spiral, which is either hollow, as in Solarium, or quite compact, as in Oliva, Terebra, Cypraea, with every possible intermediate grade. This concavity, which varies in depth and width, is known as the umbilicus, and shells are accordingly spoken of as deeply (e.g. most Trochidae and Naticidae), narrowly (e.g. Lacuna, Littorina), or widely (e.g. Solarium) umbilicated. When the spiral is quite flat, as in Planorbis and some Helix, the umbilicus vanishes entirely. Shells in which the whorls are so compactly coiled on an ascending spiral that there is no umbilicus, are termed imperforate.

The Slit.—Many shells are furnished with a slit in the last whorl, which opens, in most cases, on the outer lip, and is sometimes of considerable depth, at others a mere notch. In the patelliform shells it is always in front of the apex. The function of the slit appears to be mainly anal, the excretory products being thus allowed to escape by a passage of their own, without soiling the clean water taken in by the branchiae. The posterior canal of some Gasteropoda probably performs a similar function. In the adult Fissurella the slit becomes an apical hole (see Fig. 178 F), in the allied genera it is either immediately in front of the spire (Puncturella), or half-way between the spire and the anterior margin (Rimula), or on the margin and well marked (Emarginula), or a mere indentation of the margin (Hemitoma). In Pleurotomaria it is exceptionally long, and is well marked in Bellerophon, Schismope, Scissurella, Murchisonia, and Pleurotoma (where it is sutural). In Haliotis and Polytremaria it is replaced by a series of holes, which are closed up as the animal grows past them. Some of these holes (at least in Haliotis) certainly serve the purpose of admitting water to the branchiae, while others are anal. In Trochotoma there are only two holes, united by a narrow fissure.

The Tubed Land Operculates.—A group of the Cyclophoridae, which is restricted to Further India and the great Malay Islands, has developed a remarkable sutural tube on the exterior of the last whorl, near the aperture, A similar tube, but more obscure, exists in Alycaeus. Several stages in the development of this tube may be noticed, beginning with the elevation of part of the peristome into a simple irregular shelly plate, which is continued, first into a short, and then into a long tube, which becomes soldered to the shell; finally, the tube becomes free, and the anterior part of the last whorl is disconnected from the spire (Fig. 180 A-D).

It is singular that the tube does not appear to be of any use to the animal, since its internal extremity, in the complete form, is closed, and does not communicate with the interior of the whorl. It may be presumed, however, that in origin the tube served as a means of conveying air to the animal when the operculum was closed. The holes in the peristome of Pupina, Cataulus, and Anostoma (Fig. 154) may be compared.

The Operculum.—The operculum is a cuticular development of a group of cells situated on the dorsal side of the foot, exactly over the terminal point of the fibres of the columellar muscle. It is so situated that in crawling it is generally carried free of the shell, sometimes at the extreme upper end of the foot, more usually somewhat nearer to the shell (Fig. 181). In Pterocyclus it is pushed back into the umbilicus when the animal is in motion.

The operculum is present in nearly all land, fresh-water, and marine Prosobranchiata, absent in all Opisthobranchiata in the adult state, except Actaeon, and in all Pulmonata, except Amphibola. It has been lost in the following marine Prosobranchiata: many Cancellariidae and Conidae, Oliva (though present in Olivella and Ancilla), Harpidae, Marginellidae, Voluta proper (though present in V. musica), nearly all Mitridae, Cypraeidae, Doliidae, Ianthinidae; and, of land genera, in Proserpinidae. It is evident, therefore, that its presence or absence is of limited value in classification. In some species of Ampullaria and Natica it is horny, in others shelly. Dall found that in a number of specimens of Volutharpa ampullacea, 15 p.c. had opercula, 10 p.c. traces of the operculigenous area, but no operculum, the rest no trace of either. Monstrosities of Buccinum undatum sometimes occur, which have two, or in rare case three opercula.

As a rule, the operculum exactly fits the mouth of the shell. But in cases where the mouth is very large (e.g. Conus, Strombus, Concholepas, some Bullia), it only covers a very small portion and is quite inadequate as a protection (Fig. 62, p. 155). Again, when the shell has assumed a more or less limpet-shaped form, and habitually adheres to flat surfaces without much occasion for locomotion, the operculum becomes degraded and is probably on the way to being lost altogether. This is the case with Navicella (a modified Nerita, see Fig. 13, p. 17), Concholepas (a modified Purpura), Sigaretus (a modified Natica). Probably the more completely patelliform shells of Crepidula, Haliotis, Fissurella, and Patella have reached the stage at which the operculum has been lost entirely. In Navicella, besides becoming degraded, the operculum has actually become partly internal, and apparently serves the purpose of separating the viscera from the upper part of the foot, something like the shelly plate in Crepidula. This explains why the operculum in this genus is polished on both sides.[349]

Some authors have imagined that the operculum is homologous (a) to the second valve in Pelecypoda, (b) to the byssus. It differs, however, morphologically from the former in the essential point of not being produced by the mantle, and from the latter in not being produced by a special gland.

As regards shape and formation, the operculum has usually a more or less well-marked nucleus which may be central (e.g. Livona), sub-central (Ampullaria), lateral (Purpura), or terminal (Pyrula). As a rule, both the inner and outer surfaces are fairly flat, but in Torinia, Cyathopoma, and Pterocyclus the outer surface is elevated and conically spiral, in some Turbo (e.g. Sarmaticus) it is covered with raised tubercles resembling coral, while in others (e.g. Callopoma) it is scored with a deep trench. Aulopoma, a land genus peculiar to Ceylon, has a paucispiral operculum with hollow whorls, deceptively like a Planorbis; it fits over the aperture instead of into it. In Livona and most Trochidae the operculum is cartilaginous and multispiral. In Strombus it is narrow, curved, and often serrated like a leaf on one of the edges; in Conus it is narrowly oblong and rather featureless; in Littorina, paucispiral and always cartilaginous. In many cases (e.g. Paludina) there is no true spiral form, but the striae are concentric to a nearly central nucleus, and thus give the appearance of a spiral. The evolution of the operculum in Navicella from Nerita has already been illustrated (p. 10). Neritopsis has a very remarkable operculum, the striated appendage of which locks behind the columella of the shell, like the tooth in the opercula of the Neritidae.

Terms employed to denote various parts of the Bivalve Shell.—The umbo, or beak, is the apex of the hollow cone, of which each valve may be regarded as consisting. This apex is usually more or less twisted: it is markedly spiral in Isocardia, Diceras, some Chama, and especially Requienia, while in Pecten, Lepton, and others the spiral is altogether absent. As a rule the umbones point forward, i.e. towards the anterior end of the shell. In Donax, Nucula, and Trigonia, however, they point backward. The umbones are generally more or less approximated, but in Arca they are widely separated.

An equilateral shell is one in which the umbones are more or less central with regard to its anterior and posterior portion, while in an inequilateral shell the umbones are much nearer one end than the other. On the other hand, equivalve and inequivalve are terms used to express the relation of the two valves to one another as a whole. Thus nearly all bivalve shells are more or less inequilateral, but a comparatively small proportion are inequivalve.

The dorsal margin is adjacent to, the ventral margin opposite to, the umbones. The anterior and posterior margins are respectively the front and hinder edges of the shell.

The muscles which serve to close the valves leave impressions on the inner surface of each valve. These, when both muscles are present, are known as the anterior and posterior adductor impressions. The impression produced by the muscular edge of the mantle, which curves downwards and backwards from the anterior adductor impression, is known as the pallial line. In shells with only one muscle it is represented by an irregular row of small marks, or disappears altogether (Ostrea). The pallial sinus is produced by the muscles which retract the siphons, and is most marked in those genera in which the muscles are powerful and the siphons large (e.g. Tellina, Mya). It is entirely absent in genera possessing no retractile siphons.

Right and Left Valve.—The simplest way of distinguishing the valves as right and left is to hold the shell in such a way that the siphons point towards the observer, and the mouth away from him; in this position the valve to the right is called the right valve, and the valve to the left the left valve. If, however, the animal is not present, it may be remembered that the ligament is nearly always behind the beaks, and that the beaks, as a rule, point forward, thus the right and left valves can generally be named by observation of the beaks and ligament. When the ligament is median to the valves (e.g. Ostrea, Pecten), and the beaks are not curved, the valves may be recognised by noting the fact that the impression of the adductor muscle (in these cases always single) is nearer to the posterior than to the anterior side. In a similar way the pallial impression, which only forms a sinus on the posterior side, furnishes a guide to the valves of Donax, in which the beaks point backward, and of Tellina, in which the beaks are frequently central.

In the fixed inequivalves (e.g. Chama) it is sometimes the right, sometimes the left valve which is undermost, but the fixed valve, whether right or left, is always deep, and the free valve flat. Ostrea and Anomia are always fixed by the left valve.

The lunule is a well-marked area in front of and close to the umbones, usually more or less heart-shaped, and limited by a ridge. Generally it is shallow, but sometimes, as in Dosinia, Opis, and some Cardium, deeply impressed. A corresponding area behind the umbones, enclosing the ligament, is called the escutcheon (Fig. 186), but it seldom occurs.

The ligament is a more or less elastic band, which unites the two valves along a line adjacent to the umbones. As a rule, the greater part of the ligament is external to the shell, but it may be entirely internal. It is placed, normally, behind the umbones, but in a few cases, when the hinge line is very long (Arca, Pectunculus), it extends in front of them as well. The edges of the valves, when the ligament is mainly external, are more or less excavated for its reception. When internal it is generally contained in a groove or spoon-shaped pit, known as the fossette (compare Fig. 187).

The ligament consists of two distinct parts, which may occur together or separately, the external, or ligament proper, and the internal, or cartilage. Only the external portion can be seen when the valves are closed. As a rule, the two portions are intimately connected with one another, the ligament folding over the cartilage, but in some cases, e.g. Mya, Mactra, where the cartilage is lodged within the hinge, they are completely disconnected (Fig. 187).

In Pecten the external ligament is very thin, and runs along the dorsal margin, while the internal ligament is large, solid, and situated in a shallow pit. In Perna, where the hinge is toothless, the ligament is folded into a number of transverse ridges, which fit into corresponding grooves in the shell.

The ligament proper is inelastic and insoluble in caustic potash. The cartilage is very elastic, composed of parallel fibres, slightly iridescent, and soluble in caustic potash.

The operation of the ligament—using the word as including the whole ligamental process—is in opposition to that of the adductor muscles. When the latter close the valves, they compress the ligament, an action which its elasticity resists: thus its operation tends in part towards keeping the valves open. But when ligament and cartilage are both fully developed, they work in opposition to one another, the ligament, by its resistance to compression, preventing any straining of the adductor muscles when the valves are open, and the cartilage, for the same reason, preventing the ventral margins of the shell from closing too rapidly upon one another when the valves are being shut.

The Hinge.—The valves of Pelecypoda are generally articulated, below the umbones, by a hinge which is furnished, in the majority of cases, with interlocking teeth, small pits or depressions in each valve corresponding to the teeth in the other. The teeth are distinguished as cardinal, or those immediately below the umbo, and lateral, or those to either side of the cardinals, the latter being also distinguished as anterior and posterior laterals, according as they are before or behind the umbo (Fig. 184). In shells which are tolerably equilateral there is no difficulty in distinguishing between cardinal and lateral teeth. But when they are very inequilateral, the whole hinge may share in the inequality of growth, and an anterior lateral may be thrown backward and simulate a cardinal, or a cardinal may be thrown backward and simulate a posterior lateral (e.g. Cardita, Unio, Fig. 188). In many Chama the cardinals are pushed up into the umbo and become a mere ridge, while the strong anterior lateral becomes nearly central and simulates a cardinal.

Some bivalves, e.g. Anodonta, Ostrea, Pedum, many Mytilus, have no hinge teeth at all, in others the laterals are wanting (Psammobia, Diplodonta). In the Arcadae the hinge consists of a number of very similar denticles, which are often serrated like the teeth of a comb (Fig. 189).

Hinge-teeth are probably, in origin, derived from the crenulations or ribbings of the surface of the shell, the upper ends of which impinge upon the dorsal margin and mark it in a way which is quite recognisable when the shell is thin. Similar crenulations, resulting in interlocking of the valves, are not uncommon upon the ventral margin in certain genera (Fig. 190). The mechanical effect of these continued riblets, when fitted together on the opposing valves, would be to prevent the valves sliding upon one another while closing, or after being closed. Thus there would be a probability of their surviving, even after the ribbing had disappeared from the surface of the shell, the increased strength given by the hinge compensating for, and making it possible to do without, the extra strength supplied by the ribs. It is therefore possible that the teeth of the Nuculidae and Arcadae, which have no distinction between cardinals and laterals, represent a very ancient type, from which have been evolved the various forms of hinge in which cardinals and laterals are distinguished. Even in some forms of Arcadae (comp. Pectunculus) we get a hint how the transverse teeth of the typical Arca may have become transformed into the longitudinal tooth of the normal lateral.[350]

The developed hinge-teeth, then, ensure the opening of the valves in one direction; they also secure their accurate closure upon one another in exactly the same plane. Exposed shells and gaping siphons matter little to animals which are protected by their burrowing propensities, but to those which live in material which can be easily penetrated by their foes, it must be of advantage to be able to buckle their armour absolutely tight. The edentulous hinge of Anodonta is a degeneration from a dentate type, which retains its teeth (in Unio, etc.) when subject to the jar of rapid streams, but tends to lose them in the stiller waters of canals, lakes, and ponds.

Other processes in the bivalve shell.—In Anatina each umbo is fissured and strengthened on the inside by a kind of umbonal plate which carries the ligament. Some forms of Liligna develop a strong internal umbonal rib, which serves as a buttress to strengthen the shell. In Pholas, the so-called falciform process serves as a point of attachment for the muscles of the foot and viscera. There is no ligament or hinge-teeth, the place of the latter being taken by the anterior adductor muscle, which is attached to the hinge-plate, the latter being reflected back into the shell.

In Septifer the anterior adductor muscle is carried on a sort of shelf or myophore, and in Cucullaea the posterior adductor is partly raised on a similar and very prominent formation.

Length and breadth of bivalve shells is variously measured. Most authorities measure length, or ‘antero-posterior diameter,’ by a straight line drawn from the extreme anterior to the extreme posterior margin, and breadth by a similar line, drawn from the umbones to a point, not very clearly marked, on the opposite ventral margin (see Figs. 184 and 185). Others, less correctly, reverse these terms. Thickness is measured by the extreme distance of the opposite faces of the closed valves. As a rule, the length exceeds, and often greatly exceeds, the thickness, but in a few cases—e.g. the Cardissa section of Cardium—this is reversed.

The periostracum.—Nearly all shells are covered, at some period of their growth, by a periostracum,[351] or surface skin, which serves the purpose of protecting the shell against the destructive effects of the chemical action set up by water or air. It also, in some cases (see p. 258), acts as a kind of base upon which the shell is deposited. In old shells it is commonly worn away, especially at those parts which are likely to become abraded.

The form and composition of the periostracum varies greatly. Sometimes (e.g. Oliva) it is a mere transparent film, at others (Zonites) it is transparent, but stout and solid. It is corneous in Solenomya, covered with fine hairs in many Helicidae, in Conus, Velutina, and Cantharus it is thick, fibrous, and persistent; in Trichotropis and some Triton it is furnished with long bristles on a thick ground (Fig. 191). In fresh-water shells it is usually rather thick, in order to protect the shell from the erosive powers of certain kinds of water. In some cases (Mya, Anatina) the periostracum is continued over the siphons, so as to form a protection throughout their whole length.

Erosion.—The fresh-water Mollusca generally, and marine mollusca in a few rare cases (Purpura, Littorina) are subject to erosion, or decay in the shell substance. In univalves erosion usually sets in near the apex (Fig. 192), where the life of the shell may be regarded as weakest, and in bivalves near the umbones. It is commonest in old shells, and rarely occurs in the very young. So long as the periostracum is present to protect the shell, erosion cannot set in, but when once it has been removed the shell is liable to the chemical changes set up in its substance by water. There is abundant evidence to show that erosion is caused by pollution of water. Out of many instances one must suffice. In a certain stream near Boston, U.S., numbers of Mollusca occurred, the shells of which were very perfect and free from disease. Some little way down stream an alkaline manufactory drained its refuse into the water. At and below this point for some distance every shell was more or less eroded, most of them seriously. Farther down, when the alkali refuse became diluted away, the shells retained their normal condition.[352]

A small percentage of lime in the water appears to produce erosion. The result of some experiments by G. W. Shrubsole, in the investigation of this point, may be tabulated as follows:[353]—