The Evolution of Man

by Ernst Haeckel

Our evolutionary hypotheses can claim the same value, for the same reasons. In formulating them we are acting on the same inductive and deductive methods, and with almost equal confidence, as the geologist. We hold them to be correct, and claim the status of “biological theories” for them, because we cannot understand the nature and origin of man and the other organisms without them, and because they alone satisfy our demand for a knowledge of causes. And just as the geological hypotheses that were ridiculed as dreams at the beginning of the nineteenth century are now universally admitted, so our phylogenetic hypotheses, which are still regarded as fantastic in certain quarters, will sooner or later be generally received. It is true that, as will soon appear, our task is not so simple as that of the geologist. It is just as much more difficult and complex as man’s organisation is more elaborate than the structure of the rocks.

When we approach this task, we find an auxiliary of the utmost importance in the comparative anatomy and embryology of two lower animal-forms. One of these animals is the lancelet (Amphioxus), the other the sea-squirt (Ascidia). Both of these animals are very instructive. Both are at the border between the two chief divisions of the animal kingdom—the vertebrates and invertebrates. The vertebrates comprise the already mentioned classes, from the Amphioxus to man (acrania, lampreys, fishes, dipneusts, amphibia, reptiles, birds, and mammals). Following the example of Lamarck, it is usual to put all the other animals together under the head of invertebrates. But, as I have often mentioned already, the group is composed of a number of very different stems. Of these we have no interest just now in the echinoderms, molluscs, and articulates, as they are independent branches of the animal-tree, and have nothing to do with the vertebrates. On the other hand, we are greatly concerned with a very interesting group that has only recently been carefully studied, and that has a most important relation to the ancestral tree of the vertebrates. This is the stem of the Tunicates. One member of this group, the sea-squirt, very closely approaches the lowest vertebrate, the Amphioxus, in its essential internal structure and embryonic development. Until 1866 no one had any idea of the close connection of these apparently very different animals; it was a very fortunate accident that the embryology of these related forms was discovered just at the time when the question of the descent of the vertebrates from the invertebrates came to the front. In order to understand it properly, we must first consider these remarkable animals in their fully-developed forms and compare their anatomy.

We begin with the lancelet—after man the most important and interesting of all animals. Man is at the highest summit, the lancelet at the lowest root, of the vertebrate stem.

It lives on the flat, sandy parts of the Mediterranean coast, partly buried in the sand, and is apparently found in a number of seas.[^[28]] It has been found in the North Sea (on the British and Scandinavian coasts and in Heligoland), and at various places on the Mediterranean (for instance, at Nice, Naples, and Messina). It is also found on the coast of Brazil and in the most distant parts of the Pacific Ocean (the coast of Peru, Borneo, China, Australia, etc.). Recently eight to ten species of the amphioxus have been determined, distributed in two or three genera.

[28] See the ample monograph by Arthur Willey, Amphioxus and the Ancestry of the Vertebrates; Boston, 1894.

Johannes Müller classed the lancelet with the fishes, although he pointed out that the differences between this simple vertebrate and the lowest fishes are much greater than between the fishes and the amphibia. But this was far from expressing the real significance of the animal. We may confidently lay down the following principle: The Amphioxus differs more from the fishes than the fishes do from man and the other vertebrates. As a matter of fact, it is so different from all the other vertebrates in its whole organisation that the laws of logical classification compel us to distinguish two divisions of this stem: 1, the Acrania (Amphioxus and its extinct relatives); and 2, the Craniota (man and the other vertebrates). The first and lower division comprises the vertebrates that have no vertebræ or skull (cranium). Of these the only living representatives are the Amphioxus and Paramphioxus, though there must have been a number of different species at an early period of the earth’s history.

Fig. 210—The lancelet (Amphioxus lanceolatus), left view. The long axis is vertical; the mouth-end is above, the tail-end below; a mouth, surrounded by threads of beard; b anus, c gill-opening (porus branchialis), d gill-crate, e stomach, f liver, g small intestine, h branchial cavity, i chorda (axial rod), underneath it the aorta; k aortic arches, l trunk of the branchial artery, m swellings on its branches, n vena cava, o visceral vein.
Fig. 211—Transverse section of the head of the Amphioxus. (From Boveri.) Above the branchial gut (kd) is the chorda, above this the neural tube (in which we can distinguish the inner grey and the outer white matter); above again is the dorsal fin (fh). To the right and left above (in the episoma) are the thick muscular plates (m); below (in the hyposoma) the gonads (g). ao aorta (here double), c corium, ec endostyl, f fascie, gl glomerulus of the kidneys, k branchial vessel, ld partition between the cœloma (sc) and atrium (p), mt transverse ventral muscle, n renal canals, of upper and uf lower canals in the mantle-folds, p peribranchial cavity, (atrium), sc cœloma (subchordal body-cavity), si principal (or subintestinal) vein, sk perichorda (skeletal layer).

Opposed to the Acrania is the second division of the vertebrates, which comprises all the other members of the stem, from the fishes up to man. All these vertebrates have a head quite distinct from the trunk, with a skull (cranium) and brain; all have a centralised heart, fully-formed kidneys, etc. Hence they are called the Craniota. These Craniotes are, however, without a skull in their earlier period. As we already know from embryology, even man, like every other mammal, passes in the earlier course of his development through the important stage which we call the chordula; at this lower stage the animal has neither vertebræ nor skull nor limbs (Figs. 83–86). And even after the formation of the primitive vertebræ has begun, the segmented fœtus of the amniotes still has for a long time the simple form of a lyre-shaped disk or a sandal, without limbs or extremities. When we compare this embryonic condition, the sandal-shaped fœtus, with the developed lancelet, we may say that the amphioxus is, in a certain sense, a permanent sandal-embryo, or a permanent embryonic form of the Acrania; it never rises above a low grade of development which we have long since passed.