Sir R. Murchison groups with this upper division of the Old Red of Scotland certain light-red and yellow sandstones and grits which occur in the northernmost part of the mainland, and extend also into the Orkney and Shetland Islands. They contain Calamites and other plants which agree generically with Carboniferous forms.
Middle Old Red Sandstone.—In the northern part of Scotland there occur a great series of bituminous schists and flagstones, to the fossil fish of which attention was first called by the late Hugh Miller. They were afterwards described by Agassiz, and the rocks containing them were examined by Sir R. Murchison and Professor Sedgwick, in Caithness, Cromarty, Moray, Nairn, Gamrie in Banff, and the Orkneys and Shetlands, in which great numbers of fossil fish have been found. These were at first supposed to be the oldest known vertebrate animals, as in Cromarty the beds in which they occur seem to form the base of the Old Red system resting almost immediately on the crystalline or metamorphic rocks. But in fact these fish-bearing beds, when they are traced from north to south, or to the central parts of Scotland, thin out, so that their relative age to the Lower Old Red Sandstone, presently to be mentioned, was not at first detected, the two formations not appearing in superposition in the same district. In Caithness, however, many hundred feet below the fish-zone of the middle division, remains of Pteraspis were found by Mr. Peach in 1861. This genus has never yet been found in either of the two higher divisions of the Old Red Sandstone, and confirms Sir R. Murchison’s previous suspicion that the rocks in which it occurs belong to the Lower “Old Red,” or agree in age with the Arbroath paving-stone.[[2]]
Fossil Fish of the Middle Old Red Sandstone.—The Devonian fish were referred by Agassiz to two of his great orders, namely, the Placoids and Ganoids. Of the first of these, which in the Recent period comprise the shark, the dog-fish, and the ray, no entire skeletons are preserved, but fin-spines, called ichthyodorulites, and teeth occur. On such remains the genera Onchus, Odontacanthus, and Ctenodus, a supposed cestraciont, and some others, have been established.
By far the greater number of the Old Red Sandstone fishes belong to a sub-order of Ganoids instituted by Huxley in 1861, and for which he has proposed the name of Crossopterygidæ,[[3]] or the fringe-finned, in consideration of the peculiar manner in which the fin-rays of the paired fins are arranged so as to form a fringe round a central lobe, as in the Polypterus (see a, Fig. 499), a genus of which there are several species now inhabiting the Nile and other African rivers. The reader will at once recognise in Osteolepis (Fig. 500), one of the common fishes of the Old Red Sandstone, many points of analogy with Polypterus. They not only agree in the structure of the fin, at first pointed out by Huxley, but also in the position of the pectoral, ventral, and anal fins, and in having an elongated body and rhomboidal scales. On the other hand, the tail is more symmetrical in the recent fish, which has also an apparatus of dorsal finlets of a very abnormal character, both as to number and structure. As to the dorsals of Osteolepis, they are regular in structure and position, having nothing remarkable about them, except that there are two of them, which is comparatively unusual in living fish.
Among the “fringe-finned” Ganoids we find some with rhomboidal scales, such as Osteolepis, Fig. 500; others with cycloidal scales, as Holoptychius, before mentioned (see Fig. 498). In the genera Dipterus and Diplopterus, as Hugh Miller pointed out, and in several other of the fringe-finned genera, as in Gyroptychius and Glyptolepis, the two dorsals are placed far backward, or directly over the ventral and anal fins. The Asterolepis was a ganoid fish of gigantic dimensions. A. Asmusii, Eichwald, a species characteristic of the Old Red Sandstone of Russia, as well as that of Scotland, attained the length of between twenty and thirty feet. It was clothed with strong bony armour, embossed with star-like tubercles, but it had only a cartilaginous skeleton. The mouth was furnished with two rows of teeth, the outer ones small and fish-like, the inner larger and with a reptilian character. The Asterolepis occurs also in the Devonian rocks of North America.
If we except the Placoids already alluded to, and a few other families of doubtful affinities, all the Old Red Sandstone fishes are Ganoids, an order so named by Agassiz from the shining outer surface of their scales; but Professor Huxley has also called our attention to the fact that, while a few of the primary and the great majority of the secondary Ganoids resemble the living bony pike, Lepidosteus, or the Amia, genera now found in North American rivers, and one of them, Lepidosteus, extending as far south as Guatemala, the Crossopterygii, or fringe-finned Ichthyolites, of the Old Red are closely related to the African Polypterus, which is represented by five or six species now inhabiting the Nile and the rivers of Senegal. These North American and African Ganoids are quite exceptional in the living creation; they are entirely confined to the northern hemisphere, unless some species of Polypterus range to the south of the line in Africa; and, out of about 9000 living species of fish known to M. Günther, and of which more than 6000 are now preserved in the British Museum, they probably constitute no more than nine.
If many circumstances favour the theory of the fresh-water origin of the Old Red Sandstone, this view of its nature is not a little confirmed by our finding that it is in Llake Superior and the other inland Canadian seas of fresh water, and in the Mississippi and African rivers, that we at present find those fish which have the nearest affinity to the fossil forms of this ancient formation.