The Shells of Pteropods

While math­e­mat­i­cally speaking we are entitled to look upon the bivalve shell of the Lamellibranch as consisting of two distinct elements, each comparable to the entire shell of the univalve, we have no biological grounds for such a statement; for the shell arises from a single embryonic origin, and afterwards becomes split into portions which constitute the two separate valves. We can perhaps throw some indirect light upon this phenomenon, and upon several other phenomena connected with shell-growth, by a consideration of the simple conical or tubular shells of the Pteropods. The shells of the latter are in few cases suitable for simple math­e­mat­i­cal in­ves­ti­ga­tion, but nevertheless they are of very considerable interest in connection with our general problem.

Fig. 295. Pteropod shells: (1) Cuvierina columnella; (2) Cleodora chierchiae; (3) C. pygmaea. (After Boas.)

The morphology of the Pteropods is by no means well understood, and in speaking of them I will assume that there are still grounds for believing (in spite of Boas’ and Pelseneer’s arguments) that they are directly related to, or may at least be directly compared with, the Cephalopoda.

The simplest shells among the Pteropods have the form of a tube, more or less cylindrical (Cuvierina), more often conical (Creseis, Clio); and this tubular shell (as we have already had occasion to remark, on p. ), frequently tends, when it is very small and delicate, to assume the character of an unduloid. (In such a case it is more than likely that the tiny shell, or that portion of it which constitutes the unduloid, has not grown by successive {571} increments or “rings of growth,” but has developed as a whole.) A thickened “rib” is often, perhaps generally, present on the dorsal side of the little conical shell. In a few cases (Limacina, Peraclis) the tube becomes spirally coiled, in a normal logarithmic spiral or helico-spiral.

Fig. 296. Diagrammatic transverse sections, or outlines of the mouth, in certain Pteropod shells: A, B, Cleodora australis; C, C. pyramidalis; D, C. balantium; E, C. cuspidata. (After Boas.)

Fig. 297. Shells of thecosome Pteropods (after Boas). (1) Cleodora cuspidata; (2) Hyalaea trispinosa; (3) H. globulosa; (4) H. uncinata; (5) H. inflexa.

In certain cases (e.g. Cleodora, Hyalaea) the tube or cone is curiously modified. In the first place, its cross-section, originally {572} circular or nearly so, becomes flattened or compressed dorso-ventrally; and the angle, or rather edge, where dorsal and ventral walls meet, becomes more and more drawn out into a ridge or keel. Along the free margin, both of the dorsal and the ventral portion of the shell, growth proceeds with a regularly varying velocity, so that these margins, or lips, of the shell become regularly curved or markedly sinuous. At the same time, growth in a transverse direction proceeds with an acceleration which manifests itself in a curvature of the sides, replacing the straight borders of the original cone. In other words, the cross-section of the cone, or what we have been calling the generating curve, increases its dimensions more rapidly than its distance from the pole.

Fig. 298. Cleodora cuspidata.

In the above figures, for instance in that of Cleodora cuspidata, the markings of the shell which represent the successive edges of the lip at former stages of growth, furnish us at once with a “graph” of the varying velocities of growth as measured, radially, from the apex. We can reveal more clearly the nature of these variations in the following way which is simply tantamount to converting our radial into rectangular coordinates. Neglecting curvature (if any) of the sides and treating the shell (for simplicity’s sake) as a right cone, we lay off equal angles from the apex O, along the radii Oa, Ob, etc. If we then plot, as vertical equidistant ordinates, the magnitudes Oa, Ob ... OY, and again on to Oa ′, we obtain a diagram such as the following (Fig. ); by {573} help of which we not only see more clearly the way in which the growth-rate varies from point to point, but we also recognise much better than before, the similar nature of the law which governs this variation in the different species.

Fig. 299. Curves obtained by transforming radial ordinates, as in Fig. , into vertical equidistant ordinates. 1, Hyalaea trispinosa; 2, Cleodora cuspidata.

Furthermore, the young shell having become differentiated into a dorsal and a ventral part, marked off from one another by a lateral edge or keel, and the inequality of growth being such as to cause each portion

Fig. 300. Development of the shell of Hyalaea (Cavolinia) tridentata, Forskal: the earlier stages being the “Pleuropus longifilis” of Troschel. (After Tesch.)

to increase most rapidly in the median line, it follows that the entire shell will appear to have been split into a dorsal and a ventral plate, both connected with, and projecting from, {574} what remains of the original undivided cone. Putting the same thing in other words, we may say that the generating figure, which lay at first in a plane perpendicular to the axis of the cone, has now, by unequal growth, been sharply bent or folded, so as to lie ap­prox­i­mate­ly in two planes, parallel to the anterior and posterior faces of the cone. We have only to imagine the apical connecting portion to be further reduced, and finally to disappear or rupture, and we should have a bivalve shell developed out of the original simple cone.

In its outer and growing portion, the shell of our Pteropod now consists of two parts which, though still connected together at the apex, may be treated as growing practically independently. The shell is no longer a simple tube, or simple cone, in which regular inequalities of growth will lead to the development of a spiral; and this for the simple reason that we have now two opposite maxima of growth, instead of a maximum on the one side and a minimum on the other side of our tubular shell. As a matter of fact, the dorsal and the ventral plate tend to curve in opposite directions, towards the middle line, the dorsal curving ventrally and the ventral curving towards the dorsal side.

In the case of the Lamellibranch or the Brachiopod, it is quite possible for both valves to grow into more or less pronounced spirals, for the simple reason that they are hinged upon one another; and each growing edge, instead of being brought to a standstill by the growth of its opposite neighbour, is free to move out of the way, by the rotation about the hinge of the plane in which it lies.

But where, as in the Pteropod, there is no such hinge, the dorsal and ventral halves of the shell (or dorsal and ventral valves, if we may call them so), if they curved towards one another (as they do in a cockle), would soon interfere with one another’s progress, and the development of a pair of conjugate spirals would become impossible. Nevertheless, there is obviously, in both dorsal and ventral valve, a tendency to the development of a spiral curve, that of the ventral valve being more marked than that of the larger and overlapping dorsal one, exactly as in the two unequal valves of Terebratula. In many cases (e.g. Cleodora cuspidata), the dorsal valve or plate, {575} strengthened and stiffened by its midrib, is nearly straight, while the curvature of the other is well displayed. But the case will be materially altered and simplified if growth be arrested or retarded in either half of the shell. Suppose for instance that the dorsal valve grew so slowly that after a while, in comparison with the other, we might speak of it as being absent altogether: or suppose that it merely became so reduced in relative size as to form no impediment to the continued growth of the ventral one; the latter would continue to grow in the direction of its natural curvature, and would end by forming a complete and coiled logarithmic spiral. It would be precisely analogous to the spiral shell of Nautilus, and, in regard to its

Fig. 301. Pteropod shells, from the side: (1) Cleodora cuspidata; (2) Hyalaea longirostris; (3) H. trispinosa. (After Boas.)

ventral position, concave towards the dorsal side, it would even deserve to be called directly homologous with it. Suppose, on the other hand, that the ventral valve were to be greatly reduced, and even to disappear, the dorsal valve would then pursue its unopposed growth; and, were it to be markedly curved, it would come to form a logarithmic spiral, concave towards the ventral side, as is the case in the shell of Spirula. Were the dorsal valve to be destitute of any marked curvature (or in other words, to have but a low spiral angle), it would form a simple plate, as in the shells of Sepia or Loligo. Indeed, in the shells of these latter, and especially in that of Sepia, we seem to recognise a manifest resemblance to the dorsal plate of the Pteropod shell, as we have it (e.g.) in Cleodora or Hyalaea; {576} the little “rostrum” of Sepia is but the apex of the primitive cone, and the rounded anterior extremity has grown according to a law precisely such as that which has produced the curved margin of the dorsal valve in the Pteropod. The ventral portion of the original cone is nearly, but not wholly, wanting. It is represented by the so-called posterior wall of the “siphuncular space.” In many decapod cuttle-fishes also (e.g. Todarodes, Illex, etc.) we still see at the posterior end of the “pen,” a vestige of the primitive cone, whose dorsal margin only has continued to grow; and the same phenomenon, on an exaggerated scale, is represented in the Belemnites.

It is not at all impossible that we may explain on the same lines the development of the curious “operculum” of the Ammonites. This consists of a single horny plate (Anaptychus), or of a thicker, more calcified plate divided into two symmetrical halves (Aptychi), often found inside the terminal chamber of the Ammonite, and occasionally to be seen lying in situ, as an operculum which partially closes the mouth of the shell; this structure is known to exist even in connection with the early embryonic shell. In form the Anaptychus, or the pair of conjoined Aptychi, shew an upper and a lower border, the latter strongly convex, the former sometimes slightly concave, sometimes slightly convex, and usually shewing a median projection or slightly developed rostrum. From this “rostral” border the curves of growth start, and course round parallel to, finally constituting, the convex border. It is this convex border which fits into the free margin of the mouth of the Ammonite’s shell, while the other is applied to and overlaps the preceding whorl of the spire. Now this relationship is precisely what we should expect, were we to imagine as our starting-point a shell similar to that of Hyalaea, in which however the dorsal part of the split cone had become separate from the ventral half, had remained flat, and had grown comparatively slowly, while at the same time it kept slipping forward over the growing and coiling spire into which the ventral half of the original shell develops. In short, I think there is reason to believe, or at least to suspect, that we {577} have in the shell and Aptychus of the Ammonites, two portions of a once united structure; of which other Cephalopods retain not both parts but only one or other, one as the ventrally situated shell of Nautilus, the other as the dorsally placed shell for example of Sepia or of Spirula.

In the case of the bivalve shells of the Lamellibranchs or of the Brachiopods, we have to deal with a phenomenon precisely analogous to the split and flattened cone of our Pteropods, save only that the primitive cone has been split into two portions, not incompletely as in the Pteropod (Hyalaea), but completely, so as to form two separate valves. Though somewhat greater freedom is given to growth now that the two valves are separate and hinged, yet still the two valves oppose and hamper one another, so that in the longitudinal direction each is capable of only a moderate curvature. This curvature, as we have seen, is recognisable as a logarithmic spiral, but only now and then does the growth of the spiral continue so far as to develop successive coils: as it does in a few symmetrical forms such as Isocardia cor; and as it does still more conspicuously in a few others, such as Gryphaea and Caprinella, where one of the two valves is stunted, and the growth of the other is (relatively speaking) unopposed.