It had been long known to the continental naturalists, that in certain Russian deposits, very extensively developed, there occur in considerable abundance certain animal organisms; but for many years neither their position nor character could be satisfactorily determined. By some they were placed too high in the scale of organized being; by others too low. Kutorga, a writer not very familiarly known in this country, described the remains as those of mammals;—the Russian rocks contained, he said, bones of quadrupeds, and, in especial, the teeth of swine: whereas Lamarck, a better known authority, though not invariably a safe one,—for he had a trick of dreaming when wide awake, and of calling his dreams philosophy,—assigned to them a place among the corals. They belonged, he asserted, as shown by certain star-like markings with which they are fretted, to the Polyparia. He even erected for their reception a new genus of Astrea, which he designated, from the little rounded hillock which rises in the middle of each star, the genus Monticularia. It was left to a living naturalist, M. Eichwald, to fix their true position zoologically among the class of fishes, and to Sir Roderick Murchison to determine their position geologically as ichthyolites of the Old Red Sandstone.

Sir Roderick, on his return from his great Russian campaigns, in which he fared far otherwise than Napoleon, and accomplished more, submitted to Agassiz a series of fragments of these gigantic Ganoids; and the celebrated ichthyologist, who had been introduced little more than a twelvemonth before to the Pterichthys of Cromarty, was at first inclined to regard them as the remains of a large cuirassed fish of the Cephalaspian type, but generically new. Under this impression he bestowed upon the yet unknown ichthyolite of which they had formed part, the name Chelonichthys, from the resemblance borne by the broken plates to those of the carapace and plastron of some of the Chelonians. At this stage, however, the Russian Old Red yielded a set of greatly finer remains than it had previously furnished; and of these casts were transmitted by Professor Asmus, of the University of Dorpat, to the British and London Geological Museums, and to Agassiz. “I knew not at first what to do,” says the ichthyologist, “with bones of so singular a conformation that I could refer them to no known type.” Detecting, however, on their exterior surfaces the star-like markings which had misled Lamarck, and which he had also detected on the lesser fragments submitted to him by Sir Roderick, he succeeded in identifying both the fragments and bones as remains of the same genus and on ascertaining that M. Eichwald had bestowed upon it, from these characteristic sculpturings, the generic name Asterolepis, or star-scale, he suffered the name which he himself had originated to drop. Even this second name, however, which the ichthyolite still continues to bear, is in some degree founded in error. Its true scales, as I shall by and by show, were not stelliferous, but fretted by a peculiar style of ornament, consisting of waved anastomosing ridges, breaking atop into angular-shaped dots, scooped out internally like the letter V; and were evidently intermediate in their character between the scales which cover the Glyptolepis and those of the Holoptychius. And the stellate markings which M. Eichwald graphically describes as minute paps rising out of the middle of star-like wreaths of little leaflets, were restricted to the dermal plates of the head.

Agassiz ultimately succeeded in classing the bones which had at first so puzzled him, into two divisions—interior and dermal; and the latter he divided yet further, though not without first lodging a precautionary protest, founded on the extreme obscurity of the subject, into cranial and opercular. Of the interior bones he specified two,—a super-scapular bone, (supra-scapulaire,)—that bone which in osseous fishes completes the scapular arch or belt, by uniting the scapula to the cranium; and a maxillary or upper jaw-bone. But his world-wide acquaintance with existing fishes could lend him no assistance in determining the places of the dermal bones: they formed the mere fragments of a broken puzzle, of which the key was lost. Even in their detached and irreducible state, however, he succeeded in basing upon them several shrewd deductions. He inferred, in the first place, that the Asterolepis was not, as had been at first supposed, a cuirassed fish, which took its place among the Cephalaspians, but a strongly helmed fish of that Cœlacanth family to which the Holoptychius and Glyptolepis belong; in the second, that, like several of its bulkier cogeners, it was in all probability a broad, flat-headed animal; and, in the third, that as its remains are found associated in the Russian beds with numerous detached teeth of large size,—the boar tusks of Kutorga—which present internally that peculiar microscopic character on which Professor Owen has erected his Dendrodic or tree-toothed family of fishes,—it would in all likelihood be found that both bones and teeth belonged to the same group. “It appears more than probable,” he said, “that one day, by the discovery of a head or an entire jaw, it will be shown that the genera Dendrodus and Asterolepis form but one.” As we proceed, the reader will see how justly the ichthyologist assigned to the Asterolepis its place among the Cœlacanths, and how entirely his two other conjectures regarding it have been confirmed. “I have had in general,” he concluded, “but small and mutilated fragments of the creature’s bones submitted to me, and of these, even the surface ornaments not well preserved; but I hope the immense materials with which the Old Red Sandstone of Russia has furnished the savans of that country will not be lost to science; and that my labors on this interesting genus, incomplete as they are, will excite more and more the attention of geologists, by showing them how ignorant we are of all the essential facts concerning the history of the first inhabitants of our globe.”

I know not what the savans of Russia have been doing for the last few years; but mainly through the labors of an intelligent tradesman of Thurso, Mr. Robert Dick,—one of those working men of Scotland of active curiosity and well-developed intellect, that give character and standing to the rest,—I am enabled to justify the classification and confirm the conjectures of Agassiz. Mr. Dick, after acquainting himself, in the leisure hours of a laborious profession, with the shells, insects, and plants of the northern locality in which he resides, had set himself to study its geology; and with this view he procured a copy of the little treatise on the Old Red Sandstone to which I have already referred, and which was at that time, as Agassiz’s Monograph of the Old Red fishes had not yet appeared, the only work specially devoted to the palæontology of the system, so largely developed in the neighborhood of Thurso. With perhaps a single exception,—for the Thurso rocks do not yet seem to have yielded a Pterichthys,—he succeeded in finding specimens, in a state of better or worse keeping, of all the various ichthyolites which I had described as peculiar to the Lower Old Red Sandstone. He found, however, what I had not described,—the remains of apparently a very gigantic ichthyolite; and, communicating with me through the medium of a common friend, he submitted to me, in the first instance, drawings of his new set of fossils; and ultimately, as I could arrive at no satisfactory conclusion from the drawings, he with great liberality made over to me the fossils themselves. Agassiz’s Monograph was not yet published; nor had I an opportunity of examining, until about a twelvemonth after, the casts, in the British Museum, of the fossils of Professor Asmus. Besides, all the little information, derived from various sources, which I had acquired respecting the Russian Chelonichthys,—for such was its name at the time,—referred it to the cuirassed type, and served but to mislead. I was assured, for instance, that Professor Asmus regarded his set of remains as portions of the plates and paddles of a gigantic Pterichthys, of from twenty to thirty feet in length. And so, as I had recognized in the Thurso fossils the peculiarities of the Holoptychian (Cœlacanth) family, I at first failed to identify them with the remains of the great Russian fish. All the larger bones sent me by Mr. Dick were, I found, cerebral; and the scales associated with these indicated, not a cuirass-protected, but a scale-covered body and exhibited, in their sculptured and broadly imbricated surfaces, the well-marked Cœlacanth style of disposition and ornament. But though I could not recognize in either bones or scales the remains of one ichthyolite more of the Old Red Sandstone, “that could be regarded as manifesting as peculiar a type among fishes as do the Ichthyosauri and Plesiosauri among reptiles,”[8] I was engaged at the time in a course of inquiry regarding the cerebral development of the earlier vertebrata, that made me deem them scarce less interesting than if I could. Ere, however, I attempt communicating to the reader the result of my researches, I must introduce him, in order that he may be able to set out with me to the examination of the Asterolepis from the same starting-point, to the Cœlacanth family,—indisputably one of the oldest, and not the least interesting, of its order.

So far as is yet known, all the fish of the earliest fossiliferous system belonged to the placoid or “broad plated” order,—a great division of fishes, represented in the existing seas by the Sharks and Rays,—animals that to an internal skeleton of cartilage unite a dermal covering of points, plates, or spines of enamelled bone, and have their gills fixed. The dermal or cuticular bones of this order vary greatly in form, according to the species or family: in some cases they even vary, according to their place, on the same individual. Those button-like tubercles, for instance, with an enamelled thorn, bent like a hook, growing out of the centre of each, which run down the back and tail, and stud the pectorals of the thorn-back, (Raja clavata,) differ very much from the smaller thorns, with star-formed bases, which roughen the other parts of the creature’s body; and the bony points which mottle the back and sides of the sharks are, in most of the known species, considerably more elongated and prickly than the points which cover their fins, belly, and snout. The extreme forms, however, of the shagreen tubercle or plate seem to be those of the upright prickle or spine on the one hand, and of the slant-laid, rhomboidal, scale-shaped plate on the other. The minuter thorns of the ray (fig. 2, a) exemplify the extreme of the prickly type; the fins, abdomen, and anterior part of the head of the spotted dog-fish (Scyllium stellare) are covered by lozenge-shaped little plates, which glisten with enamel, and are so thickly set that they cover the entire surface of the skin, (fig. 3, b,)—and these seem equally illustrative of the scale-like form. They are shagreen points passing into osseous scales, without, however, becoming really such; though they approach them so nearly in the shape and disposition of their upper disks, that the true scales, also osseous, of the Acanthodes sulcatus, (fig. 3, a,) a Ganoid of the Coal Measures, can scarce be distinguished from them, even when microscopically examined. It is only when seen in section that the distinctive difference appears. The true scale of the Acanthodes, though considerably elevated in the centre, seems to have been planted on the skin; whereas the scale-like shagreen of the dog-fish is elevated over it on an osseous pedicle or footstalk (fig. 5, a) as a mushroom is elevated over the sward on its stem; and the base of the stalk is found to resemble in its stellate character that of a shagreen point of the prickly type. The apparent scale is, we find, a bony prickle bent at right angles a little over its base, and flattened into a rhomboidal disk atop.

In small fragments of shagreen, (fig. 2 b) which have been detected in the bone-bed of the Upper Ludlow Rocks, (Upper Silurian,) and constitute the most ancient portions of this substance known to the palæontologist, the osseous tubercles are, as in the minuter spikes of the ray, of the upright thorn-like type; they merely serve to show that the placoids of the first period possessed, like those of the existing seas, an ability of secreting solid bone on their cuticular surfaces; and that, though at least such of them as have bequeathed to us specimens of their dermal armature possessed it in the form farthest removed from that of their immediate successors the ganoid fishes, they resembled them not less in the substance of which their dermoskeletal, than in that of which their endoskeletal, parts were composed. For the internal skeleton in both orders, during these early ages, seems to have been equally cartilaginous, and the cuticular skeleton equally osseous. In the ichthyolitic formation immediately over the Silurians,—that of the Lower Old Red Sandstone,—the Ganoids first appear; and the members of at least one of the families of the deposit, the Acanths,—a family rich in genera and species,—seem to have formed connecting links between this second order and their placoid predecessors. They were covered with true scales (fig. 4, a,) and their free gills were protected by gill-covers; and so they must be regarded as real Ganoids but as the shagreen of the spotted dog-fish nearly approaches, in form and character, to ganoid scales, without being really such, the scales of this family, on the other hand, approached equally near, without changing their nature, to the shagreen of the Placoids, especially to that of the spiked dogfish, (Spinax Acanthias.) (Fig. 4, b.) We even find on their under surfaces what seems to be an approximation to the characteristic footstalk. They so considerably thicken in the middle from their edges inwards, (fig. 5, c,) as to terminate in their centres in obtuse points. With these shagreen-like scales, the heads, bodies, and fins of all the species of at least two of the Acanth genera,—Cheiracanthus and Diplacanthus,—were as thickly covered as the heads, bodies, and fins of the sharks are with their shagreen; and so slight was the degree of imbrication, that the portion of each scale overlaid by the two scales in immediate advance of it did not exceed the one twelfth part of its entire area. In the scale of the Cheiracanthus we find the covered portion indicated by a smooth, narrow band, that ran along its anterior edges, and which the furrows that fretted the exposed surface did not traverse. It may be added, that both genera had the anterior edge of their fins armed with strong spines,—a characteristic of several of the Placoid families.

In the Dipterian genera Osteolepis and Diplopterus the scales were more unequivocally such than in the Acanths, and more removed from shagreen. The under surface of each was traversed longitudinally by a raised bar, which attached it to the skin, and which, in the transverse section, serves to remind one of the shagreen footstalk. They are, besides, of a rhomboidal form; and, when seen in the finer specimens, lying in their proper places on what had been once the creature’s body, they seem merely laid down side by side in line, like those rows of glazed tiles that pave a cathedral floor; but on more careful examination, we find that each little tile was deeply grooved on its higher side and end, (for it lay diagonally in relation to the head,) like the flags of a stone roof, (fig. 6, a,)—that its lateral and anterior neighbors impinged upon it along these grooves to the extent of about one third its area,—and that it impinged, in turn, to the same extent on the scales that bordered on it posteriorly and latero-posteriorly. Now, in the Cœlacanth family, (and on this special point the foregoing remarks are intended to bear,) the scales, which were generally of a round or irregularly oval form, (fig. 6, b,) overlapped each other to as great an extent as in any of the existing fishes of the Cycloid or Ctenoid orders,—to as great an extent, for instance, as in the carp, salmon, or herring. In a slated roof there is no part on which the slates do not lie double, and along the lower edge of each tier they lie triple;—there is more of slate covered than of slate seen: whereas in a tile-roof, the covered portion is restricted to a small strip running along the top and one of the edges of each tile, and the tiles do not lie double in more than the same degree in which the slates lie triple. The scaly cover of the two genera of Dipterians to which I have referred was a cover on the tile-roof principle; and this is an exceedingly common characteristic of the scales of the Ganoids. The scaly cover of the Cœlacanths, on the other hand, was a cover on the slate-roof principle;—there was in some of their genera about one third more of each scale covered than exposed; and this is so rare a ganoidal mode of arrangement, that, with the exception of the Dipterus,—a genus which, though it gives its name to the Dipterian sept, differed greatly from every other Dipterian,—I know not, beyond the limits of the ancient Cœlacanth family, a single Ganoid that possessed it. The bony covering of the Cœlacanths was farthest removed in character from shagreen, as that of their contemporaries the Acanths approximated to it most nearly; they were, in this respect, the two extremes of their order; and did we find the Cœlacanths in but the later geological formations, while the Acanths were restricted to the earlier, it might be argued by assertors of the development hypothesis, that the amply imbricated, slate-like scale of the latter had been developed in the lapse of ages from the shagreen tubercle, by passing in its downward course—broadening and expanding as it descended—through the minute, scarcely imbricated disks of the Acanths, and the more amply imbricated tile-like rhombs of the Dipterians and Palæonisci, until it had reached its full extent of imbrication in the familiar modern type exemplified in both the Cœlacanths and the ordinary fishes. But such is not the order which nature has observed;—the two extremes of the ganoid scale appear together in the same early formation: both become extinct at a period geologically remote; and the ganoid scales of the existing state of things which most nearly resemble those of ancient time are scales formed on the intermediate or tile-roof principle.

The scales of the Cœlacanths were, in almost all the genera which compose the family, of great size; in some species, of the greatest size to which this kind of integument ever attained. Of a Cœlacanth of the Coal Measures, the Holoptychius Hibberti, the scales in the larger specimens were occasionally from five to six inches in diameter. Even in the Holoptychius nobilissimus, in an individual scarcely exceeding two and a half feet in length, they measured from an inch and a half to an inch and three quarters each way. In the splendid specimen of this last species, in the British Museum, there occur but fourteen scales between the ventrals, though these lie low on the creature’s body, and the head; and in a specimen of a smaller species,—the Holoptychius Andersoni,—but about seventeen. The exposed portion of the scale was in most species of the family curiously fretted by intermingled ridges and furrows, pits and tubercles, which were either boldly relieved, as in the Holoptychius, or existed, as in the Glyptolepis, as slim, delicately chiselled threads, lines, and dots. The head was covered by strong plates, which were roughened with tubercles either confluent or detached, or hollowed, as in the Bothriolepis, into shallow pits. The jaws were thickly set with an outer range of true fish teeth, and more thinly with an inner range of what seem reptile teeth, that stood up, tall and bulky, behind the others, like officers on horseback seen over the heads of their foot-soldiers in front. The double fins,—pectorals and ventrals,—were characterized each by a thick, angular, scale-covered centre, fringed by the rays; and they must have borne externally somewhat the form of the sweeping paddles of the Ichthyosaurian genus,—a peculiarity shared also by the double fins of the Dipterus. The single fins, in all the members of the family of which specimens have been found sufficiently entire to indicate the fact, were four in number,—an anal, a caudal, and two dorsal fins; and, with the exception of the anterior dorsal, which was comparatively small, and bent downwards along the back, as if its rays had been distorted when young,[11] they were all of large size. They crowded thickly on the posterior portion of the body,—the anterior dorsal opposite the ventrals, and the posterior dorsal opposite the anal fin. The fin-rays of the various members of the family, and such of their spinous processes as have been detected, were hollow tubular bones; or rather, like the larger pieces in the framework of the Placoids, they were cartilaginous within, and covered externally by a thin osseous crust or shell, which alone survives; and to this peculiarity they owe their family name, Cœlacanth, or “hollow-spine.” The internal hollow, i. e. cartilaginous centre, was, however, equally a characteristic of the spinous processes of the Coccosteus. In their general proportions, the Cœlacanths, if we perhaps except one species,—the Glyptolepis microlepidotus,—were all squat, robust, strongly-built fishes, of the Dirk Hatterick or Balfour-of-Burley type; and not only in the larger specimens gigantic in their proportions, but remarkable for the strength and weight of their armor, even when of but moderate stature. The specimen of Holoptychius nobilissimus in the British Museum could have measured little more than three feet from snout to tail when most entire; but it must have been nearly a foot in breadth, and a bullet would have rebounded flattened from its scales. And such was that ancient Cœlacanth family, of which the oldest of our Scotch Ganoids,—the Asterolepis of Stromness,—formed one of the members, and which for untold ages has had no living representative.

Let us now enter on our proposed inquiry regarding the cerebral development of the earlier vertebrata, and see whether we cannot ascertain after what manner the first true brains were lodged, and what those modifications were which their protecting box, the cranium, received in the subsequent periods. Independently of its own special interest, the inquiry will be found to have a direct bearing on our general subject.