If the Devonian witnessed the culmination of the Palæozoic corals, its later stages saw the final decadence of the great dynasty of the Trilobites. Of these creatures there are in the Devonian some large and ornate species, remarkable for their spines and tubercles; as if in this, the latter day of their dominion, they had fallen into habits of luxurious decoration unknown to their sterner predecessors, and at the same time had found it necessary to surround their now disputed privileges with new safeguards of defensive armour. Not improbably the decadence of the Trilobites may have been connected with the introduction of the numerous and formidable fishes of the period.
But while the venerable race of the Trilobites was preparing to fight its last and unsuccessful battle, another and scarcely less ancient tribe of crustaceans, the Eurypterids, already strong in the Silurian, was armed with new and formidable powers. The Pterygotus anglicus, which should have been named scoticus, since its head-quarters are in Scotland, was in point of size the greatest of known crustaceans, recent or fossil. According to Mr. Henry Woodward, who has published an admirable description and figures of the creature in the Palæontographical Society’s Memoirs, it must have been six feet in length, and nearly two feet in breadth. Its antennæ were, unlike the harmless feelers of modern Crustacea, armed with powerful claws. Two great eyes stood in the front of the head, and two smaller ones on the top. It had four pairs of great serrated jaws, the largest as wide as a man’s hand. At the sides were a pair of powerful paddles, capable of urging it swiftly through the water as it pursued its prey; and when attacked by any predaceous fish, it could strike the water with its broad tail, terminated by a great flat “telson,” and retreat backward with the rapidity of an arrow. Woodward says it must have been the “shark of the Devonian seas;” rather, it was the great champion of the more ancient family of the lobsters, set to arrest, if possible, the encroachments of the coming sharks.
The Trilobites and Eurypterids constitute a hard case for the derivationists. Unlike those Melchisedeks, the fishes of the Silurian, which are without father or mother, the Devonian crustaceans may boast of their descent, but they have no descendants. No distinct link connects them with any modern crustaceans except the Limuli, or horse-shoe crabs; and here the connection is most puzzling, for while there seems some intelligible resemblance between the adult Eurypterids and the horse-shoe, or king-crabs, the latter, in their younger state, rather resemble Trilobites, as Dr. Packard has recently shown. Thus the two great tribes of Eurypterids and Trilobites have united in the small modern group of king-crabs, while on the other hand, there are points of resemblance, as already stated, between Trilobites and Isopods, and the king-crabs had already begun to exist, since one species is now known in the Upper Silurian. So puzzling are these various relationships, that one naturalist of the derivationist school has recently attempted to solve the difficulty by suggesting that the Trilobites are allied to the spiders! Thus nature sports with our theories, showing us in some cases, as in the corals and fishes, partnerships split up into individuals, and in others distinct lines of being converging and becoming lost in one slender thread. Barrande, the great palæontologist of Bohemia, has recently, in an elaborate memoir on the Trilobites, traced these and other points through all their structures and their whole succession in geological time thereby elaborating a most powerful inductive argument against the theory of evolution, and concluding that, so far from the history of these creatures favouring such a theory, it seems as if expressly contrived to exclude its possibility.
But, while the gigantic Eurypterids and ornate Trilobites of the Devonian were rapidly approaching their end, a few despised little crustaceans,—represented by the Amphipeltis of New Brunswick and Kampecaris of Scotland,—were obscurely laying the foundation of a new line of beings, that of the Stomapods, destined to culminate in the Squillas and their allies, which, however different in structure, are practically the Eurypterids of the modern ocean. So change the dynasties of men and animals.
"Thou takest away their breath, they die, They return to their dust; Thou sendest forth Thy Spirit, They are created; Thou renewest the form of the earth."
The reign of fishes began in the Upper Silurian, for in the rocks of this age, more especially in England, several species have been found. They occur, however, only in the newer beds of this formation, and are not of large size, nor very abundant. It is to be observed that, in so far as the fragments discovered can be interpreted, they indicate the existence already of two distinct types of fishes, the Ganoids, or gar-fishes, protected with bony plates and scales, and the Placoids, or shark-like fishes; and that in the existing world these fishes are regarded as occupying a high place in their class. Further, these two groups of fishes are those which throughout a large portion of geological time continue to prevail to the exclusion of other types, the ordinary bony fishes having been introduced only in comparatively recent periods. With the Devonian, however, there comes a vast increase to the finny armies; and so characteristic are these that the Devonian has been called the age of fishes par excellence, and we must try, with the help of our illustration, to paint these old inhabitants of the waters as distinctly as we can. Among the most ancient and curious of these fishes are those singular forms covered with broad plates, of which the Pteraspis of the Upper Silurian is the herald, and which are represented in the Lower Devonian by several distinct genera. Of these, one of the most curious is the Cephalaspis, or buckler-head, distinguished by its broad flat head, rounded in front and prolonged at the sides into two great spines, which project far beyond the sides of the comparatively slender body. This fish, it may be mentioned, is the type of a family highly characteristic of the Lower Devonian, as well as of the Upper Silurian, and all of which are provided with large plate-like cephalic coverings, sometimes with a long snout in front, and, in so far as is known, a comparatively weak body and tail. They were all probably ground-living creatures, feeding on worms and shell-fishes, and “rooting” for these in the mud, or burrowing therein for their safety. In these respects they have a most curious analogy to the Trilobites, which in habits they must have greatly resembled, though belonging by their structure to an entirely different and much higher class. So close is this resemblance, that their head-shields used to be mistaken for those of Trilobites. The case is one of those curious analogies which often occur in nature, and which must always be distinguished from the true affinities which rest on structural resemblances. Another group of small fishes, likewise cuirassed in bony armour of plates, may be represented by the Pterichthys, with its two strong bony fins at the sides, which may have served for swimming, but probably also for defence, and for creeping on or shovelling up the mud at the bottom of the sea. But, besides the Ganoids which were armed in plated cuirasses, there were others, active and voracious, clad in shining enamelled scales, like the bony pikes of the American rivers and the Polypterus of the Nile. Some of these, like the Diplacanthus, or “double-spine” were of small size, and chiefly remarkable for their sharp defensive bony spines. Others, like Holoptychius (wrinkled-scale) and Osteolepis (bone-scale), were strongly built, and sometimes of great size. One Russian species of Asterolepis (star-scale) is supposed to have been twenty feet in length, and furnished with strong and trenchant teeth in two rows. These great fishes afford a good reason for the spines and armour-plates of the contemporary trilobites and smaller fishes. Just as man has been endeavouring to invent armour impenetrable to shot, for soldiers and for ships, and, on the other hand, shot and shells that can penetrate any armoury so nature has always presented the spectacle of the most perfect defensive apparatus matched with the most perfect weapons for destruction. In the class of fishes, no age of the world is more eminent in these respects than the Devonian.[M] In addition to these fishes, there were others, represented principally by their strong bony spines, which must have been allied to some of the families of modern sharks, most of them, however, probably to that comparatively harmless tribe which, furnished with flat teeth, prey upon shell-fishes. There are other fishes difficult to place in our systems of classification; and among these an eminent example is the huge Dinichthys of Newberry, from the Hamilton group of Ohio. The head of this creature is more than three feet long and eighteen inches broad, with the bones extraordinarily strong and massive. In the upper jaw, in addition to strong teeth, there were in front two huge sabre-shaped tusks or incisors, each nearly a foot long; and corresponding to these in the massive lower jaw were two closely joined conical tusks, fitting between those of the upper jaw. No other fish presents so frightful an apparatus for destruction; and if, as is probable, this was attached to a powerful body, perhaps thirty feet in length, and capable of rapid motion through the water, we cannot imagine any creature so strong or so well armed as to cope with the mighty Dinichthys.
[M] Many of these were discovered and successfully displayed and described by Hugh Miller, and are graphically portrayed in his celebrated work on the “Old Red Sandstone,” published in 1841.
The difference between the fishes of the Devonian and those of the modern seas is well marked by the fact that, while the ordinary bony fishes now amount to probably 9,000 species, and the ganoid fishes to less than thirty, the finny tribes of the Devonian are predominantly ganoids, and none of the ordinary type are known. To what is this related, with reference to conditions of existence? Two explanations, different yet mutually connected, may be suggested. One is that armour was especially useful in the Devonian as a means of defence from the larger predaceous species, and the gigantic crustaceans of the period. That this was the case may be inferred from the conditions of existence of some modern ganoids. The common bony pike of Canada (Lepidosteus), frequenting shallow and stagnant waters, seems to be especially exposed to injury from its enemies. Consequently, while it is rare to find an ordinary fish showing any traces of wounds, a large proportion of the specimens of the bony pike which I have examined have scars on their scales, indicating injuries which they have experienced, and which possibly, to fishes not so well armed, might have proved fatal. Again, in the modern Amia, or mud-fish, in the bony pike and Polypterus, there is an extremely large air-bladder, amply supplied with blood-vessels, and even divided into cells or chambers, and communicating with the mouth by an “air-duct.” This organ is unquestionably in function a lung, and enables the animal to dispense in some degree with the use of its gills, which of course depend for their supply of vital air on the small quantity of oxygen dissolved in the water. Hence, by the power of partially breathing air, these fishes can live in stagnant and badly aerated waters, where other fishes would perish. In the case of the Amia, the grunting noises which it utters, its habit of frequenting the muddy creeks of swamps, and its possession of gill-cleaners, correspond with this view. It is possible that the Devonian fishes possessed this semi-reptilian respiration; and if so, they would be better adapted than other fishes to live in water contaminated with organic matter in a state of decay, or in waters rich in carbonic acid or deficient in oxygen. Possibly the palæozoic waters, as well as the palæozoic atmosphere, were less rich in pure oxygen than those of the present world; and it is certain that, in many of the beds in which the smaller Devonian fishes abound, there was so much decaying vegetable matter as to make it probable that the water was unfit for the ordinary fishes. Thus, though at first sight the possession of external armour and means to respire air, in the case of these peculiar fishes, may seem to have no direct connection with each other, their obvious correlation in some modern ganoids may have had its parallel on a more extensive scale among their ancient relatives. Just as the modern gar-fish, by virtue of its lungs, can live in stagnant shallows and hunt frogs, but on that account needs strong armour to defend it against the foes that assail it in such places; so in the Devonian the capacity to inhabit unaërated water and defensive plates and scales may have been alike necessary, especially to the feebler tribes of fishes. We shall find that in the succeeding carboniferous period there is equally good evidence of this.
We have reserved little space for the Devonian plants and insects; but we may notice both in a walk through a Devonian forest, in which we may include the vegetation of the several subordinate periods into which this great era was divisible. The Devonian woods were probably, like those of the succeeding carboniferous period, dense and dark, composed of but few species of plants, and these somewhat monotonous in appearance, and spreading out into broad swampy jungles, encroaching on the shallow bays and estuaries. Landing on one of these flats, we may first cast our eyes over a wide expanse, covered with what at a distance we might regard as reeds or rushes. But on a near approach they appear very different; rising in slender, graceful stems, they fork again and again, and their thin branches are sparsely covered with minute needle-like leaves, while the young shoots curl over in graceful tresses, and the older are covered with little oval fruits, or spore-cases; for these plants are cryptogamous, or flowerless. This singular vegetation stretches for miles along the muddy flats, and rises to a height of two or three feet from a knotted mass of cylindrical roots or root-stocks, twining like snakes through and over the soil. This plant may, according as we are influenced by its fruit or structure, be regarded as allied to the modern club-mosses or the modern pill-worts. It is Psilophyton, in every country one of the most characteristic plants of the period, though, when imperfectly preserved, often relegated by careless and unskilled observers to the all-engulfing group of fucoids. A little further inland we see a grove of graceful trees, forking like Psilophyton, but of grander dimensions, and with the branches covered with linear leaves, and sometimes terminated by cones. These are Lepidodendra, gigantic club-mosses, which were developed to still greater dimensions in the coal period. Near these we may see a still more curious tree, more erect in its growth, with rounded and somewhat rigid leaves and cones of different form, and with huge cable-like roots, penetrating the mud, and pitted with the marks of long rootlets. This is Cyclostigma, a plant near to the Lepidodendron, but distinct, and peculiar to the Devonian. Some of its species attain to the dimensions of considerable trees; others are small and shrubby. Another small tree, somewhat like the others, but with very long shaggy leaves, and its bark curiously marked with regular diamond-shaped scars, is the Leptophleum. All these plants are probably allied to our modern club-mosses, which are, however, also represented by some low and creeping species cleaving to the ground. A little further, and we reach a dense clump of Sigillariæ, with tall sparsely forking stems, and ribbed with ridges holding rows of leaf-scars a group of plants which we shall have further occasion to notice in the coal formation; and here is an extensive jungle of Calamites, gigantic and overgrown mares'-tails, allies of the modern equisetums.