The Evolution of Man, vol 2, Ernst Haeckel [online e reader txt] 📗

impossible. It is claimed that several (three to six) traces of provertebrae have been discovered in the anterior (pre-chordal) part of the Selachii-skull; this would bring up the number of cranial somites to twelve or sixteen, or even more.

(FIGURES 2.343 TO 2.345. Arm and hand of three anthropoids.

FIGURE 2.343. Chimpanzee (Anthropithecus niger).

FIGURE 2.344. Veddah of Ceylon (Homo veddalis).

FIGURE 2.345. European (Homo mediterraneus). (From Paul and Fritz Sarasin.))

In the primitive skull of man (Figure 2.323) and the higher Vertebrates, which has been evolved from that of the Selachii, five consecutive sections are discoverable at a certain early period of development, and one might be induced to trace these to five primitive vertebrae; but these sections are due entirely to adaptation to the five primitive cerebral vesicles, and correspond, like these, to a large number of metamera. That we have in the primitive skull of the mammals a greatly modified and transformed organ, and not at all a primitive formation, is clear from the circumstance that its original soft membranous form only assumes the cartilaginous character for the most part at the base and the sides, and remains membranous at the roof. At this part the bones of the subsequent osseous skull develop as external coverings over the membranous structure, without an intermediate cartilaginous stage, as there is at the base of the skull. Thus a large part of the cranial bones develop originally as covering bones from the corium, and only secondarily come into close touch with the primitive skull (Figure 2.333). We have previously seen how this very rudimentary beginning of the skull in man is formed ontogenetically from the “head-plates,” and thus the fore end of the chorda is enclosed in the base of the skull. (Cf. Figs 1.145 and Chapters 1.13 and 1.14.)

The phylogeny of the skull has made great progress during the last three decades through the joint attainments of comparative anatomy, ontogeny, and paleontology. By the judicious and comprehensive application of the phylogenetic method (in the sense of Gegenbaur) we have found the key to the great and important problems that arise from the thorough comparative study of the skull. Another school of research, the school of what is called “exact craniology” (in the sense of Virchow), has, meantime, made fruitless efforts to obtain this result. We may gratefully acknowledge all that this descriptive school has done in the way of accurately describing the various forms and measurements of the human skull, as compared with those of other mammals. But the vast empirical material that it has accumulated in its extensive literature is mere dead and sterile erudition until it is vivified and illumined by phylogenetic speculation.

Virchow confined himself to the most careful analysis of large numbers of human skulls and those of anthropoid mammals. He saw only the differences between them, and sought to express these in figures.

Without adducing a single solid reason, or offering any alternative explanation, he rejected evolution as an unproved hypothesis. He played a most unfortunate part in the controversy as to the significance of the fossil human skulls of Spy and Neanderthal, and the comparison of them with the skull of the Pithecanthropus (Figure 2.283). All the interesting features of these skulls that clearly indicated the transition from the anthropoid to the man were declared by Virchow to be chance pathological variations. He said that the roof of the skull of Pithecanthropus (Figure 2.335, 3) must have belonged to an ape, because so pronounced an orbital stricture (the horizontal constriction between the outer edge of the eye-orbit and the temples) is not found in any human being. Immediately afterwards Nehring showed in the skull of a Brazilian Indian (Figure 2.335, 2), found in the Sambaquis of Santos, that this stricture can be even deeper in man than in many of the apes. It is very instructive in this connection to compare the roofs of the skulls (seen from above) of different primates. I have, therefore, arranged nine such skulls in Figure 2.335, and reduced them to a common size.

(FIGURE 2.346. Transverse section of a fish’s tail (from the tunny). (From Johannes Muller.) a upper (dorsal) lateral muscles, a apostrophe, b apostrophe lower (ventral) lateral muscles, d vertebral bodies, b sections of incomplete conical mantle, B attachment lines of the inter-muscular ligaments (from the side).)

We turn now to the branchial arches, which were regarded even by the earlier natural philosophers as “head-ribs.” (Cf. Figures 1.167 to 1.170). Of the four original gill-arches of the mammals the first lies between the primitive mouth and the first gill-cleft. From the base of this arch is formed the upper-jaw process, which joins with the inner and outer nasal processes on each side, in the manner we have previously explained, and forms the chief parts of the skeleton of the upper jaw (palate bone, pterygoid bone, etc.) (Cf. Chapter 2.25.) The remainder of the first branchial arch, which is now called, by way of contrast, the “upper-jaw process,” forms from its base two of the ear-ossicles (hammer and anvil), and as to the rest is converted into a long strip of cartilage that is known, after its discoverer, as “Meckel’s cartilage,” or the promandibula. At the outer surface of the latter is formed from the cellular matter of the corium, as covering or accessory bone, the permanent bony lower jaw. From the first part or base of the second branchial arch we get, in the mammals, the third ossicle of the ear, the stirrup; and from the succeeding parts we get (in this order) the muscle of the stirrup, the styloid process of the temporal bone, the styloid-hyoid ligament, and the little horn of the hyoid bone. The third branchial arch is only cartilaginous at the foremost part, and here the body of the hyoid bone and its larger horn are formed at each side by the junction of its two halves. The fourth branchial arch is only found transitorily in the mammal embryo as a rudimentary organ, and does not develop special parts; and there is no trace in the embryo of the higher Vertebrates of the posterior branchial arches (fifth and sixth pair), which are permanent in the Selachii. They have been lost long ago. Moreover, the four gill-clefts of the human embryo are only interesting as rudimentary organs, and they soon close up and disappear. The first alone (between the first and second branchial arches) has any permanent significance; from it are developed the tympanic cavity and the Eustachian tube. (Cf. Figures 1.169 and 2.320.)

It was Carl Gegenbaur again who solved the difficult problem of tracing the skeleton of the limbs of the Vertebrates to a common type. Few parts of the vertebrate body have undergone such infinitely varied modifications in regard to size, shape, and adaptation of structure as the limbs or extremities; yet we are in a position to reduce them all to the same hereditary standard. We may generally distinguish three groups among the Vertebrates in relation to the formation of their limbs. The lowest and earliest Vertebrates, the Acrania and Cyclostomes, had, like their invertebrate ancestors, no pairs of limbs, as we see in the Amphioxus and the Cyclostomes to-day (Figures 2.210 and 2.247). The second group is formed of the two classes of the true fishes and the Dipneusts; here there are always two pairs of limbs at first, in the shape of many-toed fins—one pair of breast-fins or fore legs, and one pair of belly-fins or hind legs (Figures 2.248 to 2.259). The third group comprises the four higher classes of Vertebrates—the amphibia, reptiles, birds, and mammals; in these quadrupeds there are at first the same two pairs of limbs, but in the shape of five-toed feet. Frequently we find less than five toes, and sometimes the feet are wholly atrophied (as in the serpents). But the original stem-form of the group had five toes or fingers before and behind (Figures 2.263 to 2.265).

The true primitive form of the pairs of limbs, such as they were found in the primitive fishes of the Silurian period, is preserved for us in the Australian dipneust, the remarkable Ceratodus (Figure 2.257). Both the breast-fin and the belly-fin are flat oval paddles, in which we find a biserial cartilaginous skeleton (Figure 2.336). This consists, firstly, of a much segmented fin-rod or “stem” (A, B), which runs through the fin from base to tip; and secondly of a double row of thin articulated fin-radii (r, r), which are attached to both sides of the fin-rod, like the feathers of a feathered leaf. This primitive fin, which Gegenbaur first recognised, is attached to the vertebral column by a simple zone in the shape of a cartilaginous arch. It has probably originated from the branchial arches. ( While Gegenbaur derives the fins from two pairs of posterior separated branchial arches, Balfour holds that they have been developed from segments of a pair of originally continuous lateral fins or folds of the skin.)

We find the same biserial primitive fin more or less preserved in the fossilised remains of the earliest Selachii (Figure 2.248), Ganoids (Figure 2.253), and Dipneusts (Figure 2.256). It is also found in modified form in some of the actual sharks and pikes. But in the majority of the Selachii it has already degenerated to the extent that the radii on one side of the fin-rod have been partly or entirely lost, and are retained only on the other (Figure 2.337). We thus get the uniserial fin, which has been transmitted from the Selachii to the rest of the fishes (Figure 2.338).

(FIGURE 2.347. Human skeleton. (Cf. Figure 2.326.)

FIGURE 2.348. Skeleton of the giant gorilla. (Cf. Figure 1.209.))

Gegenbaur has shown how the five-toed leg of the Amphibia, that has been inherited by the three classes of Amniotes, was evolved from the uniserial fish-fin. ( The limb of the four higher classes of Vertebrates is now explained in the sense that the original fin-rod passes along its outer (ulnar or fibular) side, and ends in the fifth toe. It was formerly believed to go along the inner (radial or tibial) side, and end in the first toe, as Figure 2.339 shows.) In the dipneust ancestors of the Amphibia the radii gradually atrophy, and are lost, for the most part, on the other side of the fin-rod as well (the lighter cartilages in Figure 2.338). Only the four lowest radii (shaded in the illustration) are preserved; and these are the four inner toes of the foot (first to fourth). The little or fifth toe is developed from the lower end of the fin-rod. From the middle and upper part of the fin-rod was developed the long stem of the limb—the important radius and ulna (Figure 2.339 r and u) and humerus (h) of the higher Vertebrates.

In this way the five-toed foot of the Amphibia, which we first meet in the Carboniferous Stegocephala (Figure 2.260), and which was inherited from them by the reptiles on one side and the mammals on the other, was formed by gradual degeneration and differentiation from the many-toed fish-fin (Figure 2.341). The reduction of the radii to four was accompanied by a further differentiation of the fin-rod, its transverse segmentation into upper and lower halves, and the formation of the zone of the limb, which is composed originally of three limbs before and behind in the higher Vertebrates. The simple arch of the original shoulder-zone divides on each side into an upper (dorsal) piece, the shoulder-blade (scapula), and a lower (ventral) piece; the anterior part of the latter forms the primitive clavicle (procoracoideum), and the posterior part the coracoideum. In the same way the simple arch of the pelvic zone breaks up into an upper (dorsal) piece, the iliac-bone (os ilium), and a lower (ventral) piece; the anterior part of the latter forms the pubic bone (os pubis), and the posterior the ischial bone (os ischii).

There is also a complete agreement