Fig. 258—Young ceratodus, shortly after issuing from the egg, magnified. k gill-cover,
l liver. (From Richard Semon.)
Fig. 259—Young ceratodus six weeks after issuing from the egg. s spiral fold of gut,
b rudimentary belly-fin. (From Richard Semon.)
Ceratodus is particularly interesting on account of the primitive build of its skeleton; the cartilaginous skeleton of its two pairs of fins, for instance, has still the original form of a bi-serial or feathered leaf, and was on that account described by Gegenbaur as a “primitive fin-skeleton.” On the other hand, the skeleton of the pairs of fins is greatly reduced in the African dipneust (Protopterus) and the American (Lepidosiren). Further, the lungs are double in these modern dipneusts, as in all the other air-breathing vertebrates; they have on that account been called “double-lunged” (Dipneumones) in contrast to the Ceratodus; the latter has only a single lung (Monopneumones). At the same time the gills also are developed as water-breathing organs in all these lung-fishes. Protopterus has external as well as internal gills.
The paleozoic Dipneusts that are in the direct line of our ancestry, and form the connecting-bridge between the Ganoids and the Amphibia, differ in many respects from their living descendants, but agree with them in the above essential features. This is confirmed by a number of interesting facts that have lately come to our knowledge in connection with the embryonic development of the Ceratodus and Lepidosiren; they give us important information as to the stem-history of the lower Vertebrates, and therefore of our early ancestors of the paleozoic age.
Chapter XXII.
OUR FIVE-TOED ANCESTORS
With the phylogenetic study of the four higher classes of Vertebrates, which must now engage our attention, we reach much firmer ground and more light in the construction of our genealogy than we have, perhaps, enjoyed up to the present. In the first place, we owe a number of very valuable data to the very interesting class of Vertebrates that come next to the Dipneusts and have been developed from them—the Amphibia. To this group belong the salamander, the frog, and the toad. In earlier days all the reptiles were, on the example of Linne, classed with the Amphibia (lizards, serpents, crocodiles, and tortoises). But the reptiles are much more advanced than the Amphibia, and are nearer to the birds in the chief points of their structure. The true Amphibia are nearer to the Dipneusta and the fishes; they are also much older than the reptiles. There were plenty of highly-developed (and sometimes large) Amphibia during the Carboniferous period; but the earliest reptiles are only found in the Permian period. It is probable that the Amphibia were evolved even earlier—during the Devonian period—from the Dipneusta. The extinct Amphibia of which we have fossil remains from that remote period (very numerous especially in the Triassic strata) were distinguished for a graceful scaly coat or a powerful bony armour on the skin (like the crocodile), whereas the living amphibia have usually a smooth and slippery skin.
The earliest of these armoured Amphibia (Phractamphibia) form the order of Stegocephala (“roof-headed”) (Fig. 260). It is among these, and not among the actual Amphibia, that we must look for the forms that are directly related to the genealogy of our race, and are the ancestors of the three higher classes of Vertebrates. But even the existing Amphibia have such important relations to us in their anatomic structure, and especially their embryonic development, that we may say: Between the Dipneusts and the Amniotes there was a series of extinct intermediate forms which we should certainly class with the Amphibia if we had them before us. In their whole organisation even the actual Amphibia seem to be an instructive transitional group. In the important respects of respiration and circulation they approach very closely to the Dipneusta, though in other respects they are far superior to them.
This is particularly true of the development of their limbs or extremities. In them we find these for the first time as five-toed feet. The thorough investigations of Gegenbaur have shown that the fish’s fins, of which very erroneous opinions were formerly held, are many-toed feet. The various cartilaginous or bony radii that are found in large numbers in each fin correspond to the fingers or toes of the higher Vertebrates. The several joints of each fin-radius correspond to the various parts of the toe. Even in the Dipneusta the fin is of the same construction as in the fishes; it was afterwards gradually evolved into the five-toed form, which we first encounter in the Amphibia. This reduction of the number of the toes to six, and then to five, probably took place in the second half of the Devonian period—at the latest, in the subsequent Carboniferous period—in those Dipneusta which we regard as the ancestors of the Amphibia. We have several fossil remains of five-toed Amphibia from this period. There are numbers of fossil impressions of them in the Triassic of Thuringia (Chirotherium).
The fact that the toes number five is of great importance, because they have clearly been transmitted from the Amphibia to all the higher Vertebrates. Man entirely resembles his amphibian ancestors in this respect, and indeed in the whole structure of the bony skeleton of his five-toed extremities. A careful comparison of the skeleton of the frog with our own is enough to show this. It is well known that this hereditary number of the toes has assumed a very great practical importance from remote times; on it our whole system of enumeration (the decimal system applied to measurement of time, mass, weight, etc.) is based. There is absolutely no reason why there should be five toes in the fore and hind feet in the lowest Amphibia, the reptiles, and the higher Vertebrates, unless we ascribe it to inheritance from a common stem-form. Heredity alone can explain it. It is true that we find less than five toes in many of the Amphibia and of the higher Vertebrates. But in all these cases we can prove that some of the toes atrophied, and were in time lost altogether.