b. Osseous centrum of Raja clavata.
(Nat. size.)
The most ancient brain-bearing craniums that have come down to us in the fossil state, are those of the Ganoids of the Lower Old Red Sandstone; and in these fishes the true skull appears to have been as entirely a simple cartilaginous box, as that of the Placoids of either the Silurian period or of the present time, or of those existing Ganoids, the sturgeons. In the Lower Old Red genera Cheiracanthus and Diplacanthus, though the heads are frequently preserved as amorphous masses of colored matter, we detect no trace of internal bone, save perhaps in the gill-covers of the first-named genus, which were fringed by from eighteen to twenty minute osseous rays. The cranium seems to have been covered, as in the shark family, by skin, and the skin by minute shagreen-like scales; and all of the interior cerebral framework which appears underneath exists simply as faint impressions of an undivided body, covered by what seem to be osseous points,—the bony molecules, it is probable, which encrusted the cartilage. The jaws, in the better specimens, are also preserved in the same doubtful style, and this state of keeping is the common one in deposits in which every true bone, however delicate, presents an outline as sharp as when it occupied its place in the living animal. The dermal or skin-skeleton of both genera, which consisted, as has been shown ([pages 55, 56]) of shagreen-like osseous scales and slender spines, both brilliantly enamelled, is preserved entire; where as the interior framework of the head exists as mere point speckled impressions; and the inference appears unavoidable that parts which so invariably differ in their state of keeping now, must have essentially differed in their substance originally.
Fig. 9.
a. Portion of caudal fin of Cheiracanthus.[13]
b. Portion of caudal fin of Cheirolepis Cummingiæ.
(Mag. three diameters.)
Now, in the Cheiracanthus we detect the first faint indications of a peculiar arrangement of the dermal skeleton, in relation to certain parts of the skeleton within, which—greatly more developed in some of its contemporaries—led to important results in the general structure of these Ganoids, and furnishes the true key to the character of the early ganoid head. In such of the existing Placoids as I have had an opportunity of examining, the only portions of the dermal skeleton of bone which conform in their arrangement to portions of the interior skeleton of cartilage, are the teeth, which are always laid on a base of skin right over the jaws: there is also an approximation to arrangement of a corresponding kind, though a distant one, in those hook-armed tubercles of certain species of rays which run along the vertebral column; but in the shagreen by which the creatures are covered I have been able to detect no such arrangement. Whether it occurs on the fins, the body, or the head, or in the scale form, or in that of the prickle, it manifests the same careless irregularity. And on the head and body of the Cheiracanthus, and on all its fins save one, the shagreen-like scales, though laid down more symmetrically in lines than true shagreen, manifested an equal absence of arrangement in relation to the framework within. On that one fin, however;—the caudal,—the scales, passing from their ordinary rhomboidal to a more rectangular form, ranged themselves in right lines over the internal rays, (fig. 9, a,) and imparted to these such strength as a splint of wood or whalebone fastened over a fractured toe or finger imparts to the injured digit,—a provision which was probably rendered necessary in the case of this important organ of motion, from the circumstance that it was the only fin which the creature possessed that was not strengthened and protected anteriorly by a strong spine. In the Cheirolepis,—a contemporary fish, characterized, like its cogeners the Cheiracanthus and Diplacanthus, by shagreen-like scales, but in which the spines were wanting,—we find a farther development of the provision. In all the fins the richly-enamelled dermal-covering was arranged in lines over the rays, (fig. 9, b;) and the scale, which assumes in the fins, like the scales on the tail of the Cheiracanthus, though somewhat more irregularly, a rectangular shape, is so considerably elongated, that it assumes for its normal character as a scale, that of the joint of an external ray. A similar arrangement of external protection takes place in this genus over the bones of the head; the cartilaginous jaws receive their osseous dermal covering, and, with these, the hyoid bones, the opercules, and the cranium. And it is in these dermal plates, which covered an interior skull, of which, save in one genus,—the Dipterus,—not a vestige remains in any of the Old Red fishes thus protected, that we first trace what seem to be the homologues of the cranial bones of the osseous fishes,—at least their homologues so far as the cuticular can represent the internal. They appear for the first time, not as modified spinous processes, broadened, as in the carapace of the Chelonians, into osseous plates, but like those corneous external plates of this order of reptiles, (known in one species as the tortoise-shell of commerce,) the origin of which is purely cuticular, and which evince so little correspondence in their divisions with the sutures of the bones on which they rest, that they have been instanced, in their relation to the joinings beneath, as admirable illustrations of the cross-banding of the mechanician.
In the heads of the osseous fishes, the cranium proper, though consisting, like the skulls of birds, reptiles, and mammals, of several bones, exists from snout to nape, and from mastoid to mastoid, as one unbroken box; whereas all the other bones of the head, such as the maxillaries and intermaxillaries, the lower jaws, the opercular appendages, the branchial arches, and the branchiostegous rays, are connected but by muscle and ligament, and fall apart under the putrefactive influences, or in the process of boiling. This unbroken box, which consists, in the cod, of twenty-five bones, is the homologue of that cranial box of the Placoids which consists of one entire piece, and the homotype, according to Oken, of the bodies and spinal processes of four vertebræ; while the looser bones which drop away represent their ribs. The upper surface of the box,—that extending from the nasal bone to the nape,—is the only part over which a dermal buckler could be laid, as it is the only part with which the external skin comes in contact; and so it is between this upper surface and the cranial bucklers of the earlier Ganoids that we have to institute comparisons. For it is a curious fact, that, with the exception of the Old Red genera Acanthodus, Cheiracanthus, and Diplacanthus,[14] all the Ganoids of the period in which Ganoids first appear have dermal bucklers placed right over their true skulls, and that these, though as united in their parts as the bones proper to the cranium in quadrupeds and fishes, are composed of several pieces, furnished each with its independent centre of ossification. The Dipterians, the Cœlacanths, the Cephalaspians, and at least one genus placed rather doubtfully among the Acanths,—the genus Cheirolepis,—all possessed cranial bucklers extending from the nape to the snout, in which the plates, various, in the several genera, in form and position, were fast soldered together, though in every instance the lines of suture were distinctly marked.