The foregoing problem of contact, or intersection, of the successive whorls, is a very simple one in the case of the discoid shell but a more complex one in the turbinate. For in the discoid shell contact will evidently take place when the retardation of the inner as compared with the outer whorl is just 360°, and the shape of the whorls need not be considered.
As the angle of retardation diminishes from 360°, the whorls will stand further and further apart in an open coil; as it increases beyond 360°, they will more and more overlap; and when the angle of retardation is infinite, that is to say when the true inner edge of the whorl does not grow at all, then the shell is said to be completely involute. Of this latter condition we have a striking example in Argonauta, and one a little more obscure in Nautilus pompilius.
In the turbinate shell, the problem of contact is twofold, for we have to deal with the possibilities of contact on the same side of the axis (which is what we have dealt with in the discoid) and {547} also with the new possibility of contact or intersection on the opposite side; it is this latter case which will determine the presence or absence of an umbilicus, and whether, if present, it will be an open conical space or a twisted cone. It is further obvious that, in the case of the turbinate, the question of contact or no contact will depend on the shape of the generating curve; and if we take the simple case where this generating curve may be considered as an ellipse, then contact will be found to depend on the angle which the major axis of this ellipse makes with the axis of the shell. The question becomes a complicated one, and the student will find it treated in Blake’s paper already referred to.
When one whorl overlaps another, so that the generating curve cuts its predecessor (at a distance of 2π) on the same radius vector, the locus of intersection will follow a spiral line upon the shell, which is called the “suture” by conchologists. It is evidently one of that ensemble of spiral lines in space of which, as we have seen, the whole shell may be conceived to be constituted; and we might call it a “contact-spiral,” or “spiral of intersection.” In discoid shells, such as an Ammonite or a Planorbis, or in Nautilus umbilicatus, there are obviously two such contact-spirals, one on each side of the shell, that is to say one on each side of a plane perpendicular to the axis. In turbinate shells such a condition is also possible, but is somewhat rare. We have it for instance, in Solarium perspectivum, where the one contact-spiral is visible on the exterior of the cone, and the other lies internally, winding round the open cone of the umbilicus[521]; but this second contact-spiral is usually imaginary, or concealed within the whorls of the turbinated shell. Again, in Haliotis, one of the contact-spirals is non-existent, because of the extreme obliquity of the plane of the generating curve. In Scalaria pretiosa and in Spirula there is no contact-spiral, because the growth of the generating curve has been too slow, in comparison with the vector rotation of its plane. In Argonauta and in Cypraea, there is no contact-spiral, because the growth of the generating curve has been too quick. Nor, of course, is there any contact-spiral in Patella or in Dentalium, because the angle α is too small ever to give us a complete revolution of the spire. {548}
The various forms of straight or spiral shells among the Cephalopods, which we have seen to be capable of complete definition by the help of elementary mathematics, have received a very complicated descriptive nomenclature from the palaeontologists. For instance, the straight cones are spoken of as orthoceracones or bactriticones, the loosely coiled forms as gyroceracones or mimoceracones, the more closely coiled shells, in which one whorl overlaps the other, as nautilicones or ammoniticones, and so forth. In such a succession of forms the biologist sees undoubted and unquestioned evidence of ancestral descent. For instance we read in Zittel’s Palaeontology[522]: “The bactriticone obviously represents the primitive or primary radical of the Ammonoidea, and the mimoceracone the next or secondary radical of this order”; while precisely the opposite conclusion was drawn by Owen, who supposed that the straight chambered shells of such fossil cephalopods as Orthoceras had been produced by the gradual unwinding of a coiled nautiloid shell[523]. To such phylogenetic hypotheses the mathematical or dynamical study of the forms of shells lends no valid support. If we have two shells in which the constant angle of the spire be respectively 80° and 60°, that fact in itself does not at all justify an assertion that the one is more primitive, more ancient, or more “ancestral” than the other. Nor, if we find a third in which the angle happens to be 70°, does that fact entitle us to say that this shell is intermediate between the other two, in time, or in blood relationship, or in any other sense whatsoever save only the strictly formal and mathematical one. For it is evident that, though these particular arithmetical constants manifest themselves in visible and recognisable differences of form, yet they are not necessarily more deep-seated or significant than are those which manifest themselves only in difference of magnitude; and the student of phylogeny scarcely ventures to draw conclusions as to the relative antiquity of two allied organisms on the ground that one happens to be bigger or less, or longer or shorter, than the other. {549}
At the same time, while it is obviously unsafe to rest conclusions upon such features as these, unless they be strongly supported and corroborated in other ways,—for the simple reason that there is unlimited room for coincidence, or separate and independent attainment of this or that magnitude or numerical ratio,—yet on the other hand it is certain that, in particular cases, the evolution of a race has actually involved gradual increase or decrease in some one or more numerical factors, magnitude itself included,—that is to say increase or decrease in some one or more of the actual and relative velocities of growth. When we do meet with a clear and unmistakable series of such progressive magnitudes or ratios, manifesting themselves in a progressive series of “allied” forms, then we have the phenomenon of “orthogenesis.” For orthogenesis is simply that phenomenon of continuous lines or series of form (and also of functional or physiological capacity), which was the foundation of the Theory of Evolution, alike to Lamarck and to Darwin and Wallace; and which we see to exist whatever be our ideas of the “origin of species,” or of the nature and origin of “functional adaptations.” And to my mind, the mathematical (as distinguished from the purely physical) study of morphology bids fair to help us to recognise this phenomenon of orthogenesis in many cases where it is not at once patent to the eye; and also, on the other hand, to warn us, in many other cases, that even strong and apparently complex resemblances in form may be capable of arising independently, and may sometimes signify no more than the equally accidental numerical coincidences which are manifested in identity of length or weight, or any other simple magnitudes.
I have already referred to the fact that, while in general a very great and remarkable regularity of form is characteristic of the molluscan shell, that complete regularity is apt to be departed from. We have clear cases of such a departure in Pupa, Clausilia, and various Bulimi, where the enveloping cone of the spire is not a right cone but a cone whose sides are curved. It is plain that this condition may arise in two ways: either by a gradual change in the ratio of growth of the whorls, that is to say in the logarithmic spiral itself, or by a change in the velocity of {550} translation along the axis, that is to say in the helicoid which, in all turbinate shells, is superposed upon the spiral. Very careful measurements will be necessary to determine to which of these factors, or in what proportions to each, the phenomenon is due. But in many Ammonitoidea where the helicoid factor does not enter into the case, we have a clear illustration of gradual and marked changes in the spiral angle itself, that is to say of the ratio of growth corresponding to increase of vectorial angle. We have seen from some of Naumann’s and Grabau’s measurements that such a tendency to vary, such an acceleration or retardation, may be detected even in Ammonites which present nothing abnormal to the eye. But let us suppose that the spiral angle increases somewhat rapidly; we shall then get a spiral with gradually narrowing whorls, and this condition is characteristic
