The Descent of Man and Selection in Relation to Sex, Vol. I

by Charles Darwin

The Marsupials stand in many important characters below the placental mammals. They appeared at an earlier geological period, and their range was formerly much more extensive than what it now is. Hence the Placentata are generally supposed to have been derived from the Implacentata or Marsupials; not, however, from forms closely like the existing Marsupials, but from their early progenitors. The Monotremata are plainly allied to the Marsupials; forming a third and still lower division in the great mammalian series. They are represented at the present day solely by the Ornithorhynchus and Echidna; and these two forms may be safely considered as relics of a much larger group which have been preserved in Australia through some favourable concurrence of circumstances. The Monotremata are eminently interesting, as in several important points of structure they lead towards the class of reptiles.

In attempting to trace the genealogy of the Mammalia, and therefore of man, lower down in the series, we become involved in greater and greater obscurity. He who wishes to see what ingenuity and knowledge can effect, may consult Prof. Häckel’s works.[273] I will content myself with a few general remarks. Every evolutionist will admit that the five great vertebrate classes, namely, mammals, birds, reptiles, amphibians, and fishes, are all descended from some one prototype; for they have much in common, especially during their embryonic state. As the class of fishes is the most lowly organised and appeared before the others, we may conclude that all the members of the vertebrate kingdom are derived from some fish-like animal, less highly organised than any as yet found in the lowest known formations. The belief that animals so distinct as a monkey or elephant and a humming-bird, a snake, frog, and fish, &c., could all have sprung from the same parents, will appear monstrous to those who have not attended to the recent progress of natural history. For this belief implies the former existence of links closely binding together all these forms, now so utterly unlike.

Nevertheless it is certain that groups of animals have existed, or do now exist, which serve to connect more or less closely the several great vertebrate classes. We have seen that the Ornithorhynchus graduates towards reptiles; and Prof. Huxley has made the remarkable discovery, confirmed by Mr. Cope and others, that the old Dinosaurians are intermediate in many important respects between certain reptiles and certain birds—the latter consisting of the ostrich-tribe (itself evidently a widely-diffused remnant of a larger group) and of the Archeopteryx, that strange Secondary bird having a long tail like that of the lizard. Again, according to Prof. Owen,[274] the Ichthyosaurians—great sea-lizards furnished with paddles—present many affinities with fishes, or rather, according to Huxley, with amphibians. This latter class (including in its highest division frogs and toads) is plainly allied to the Ganoid fishes. These latter fishes swarmed during the earlier geological periods, and were constructed on what is called a highly generalised type, that is they presented diversified affinities with other groups of organisms. The amphibians and fishes are also so closely united by the Lepidosiren, that naturalists long disputed in which of these two classes it ought to be placed. The Lepidosiren and some few Ganoid fishes have been preserved from utter extinction by inhabiting our rivers, which are harbours of refuge, bearing the same relation to the great waters of the ocean that islands bear to continents.

Lastly, one single member of the immense and diversified class of fishes, namely the lancelet or amphioxus, is so different from all other fishes, that Häckel maintains that it ought to form a distinct class in the vertebrate kingdom. This fish is remarkable for its negative characters; it can hardly be said to possess a brain, vertebral column, or heart, &c.; so that it was classed by the older naturalists amongst the worms. Many years ago Prof. Goodsir perceived that the lancelet presented some affinities with the Ascidians, which are invertebrate, hermaphrodite, marine creatures permanently attached to a support. They hardly appear like animals, and consist of a simple, tough, leathery sack, with two small projecting orifices. They belong to the Molluscoida of Huxley—a lower division of the great kingdom of the Mollusca; but they have recently been placed by some naturalists amongst the Vermes or worms. Their larvæ somewhat resemble tadpoles in shape,[275] and have the power of swimming freely about. Some observations lately made by M. Kowalevsky,[276] since confirmed by Prof. Kuppfer, will form a discovery of extraordinary interest, if still further extended, as I hear from M. Kowalevsky in Naples he has now effected. The discovery is that the larvæ of Ascidians are related to the Vertebrata, in their manner of development, in the relative position of the nervous system, and in possessing a structure closely like the chorda dorsalis of vertebrate animals. It thus appears, if we may rely on embryology, which has always proved the safest guide in classification, that we have at last gained a clue to the source whence the Vertebrata have been derived. We should thus be justified in believing that at an extremely remote period a group of animals existed, resembling in many respects the larvæ of our present Ascidians, which diverged into two great branches—the one retrograding in development and producing the present class of Ascidians, the other rising to the crown and summit of the animal kingdom by giving birth to the Vertebrata.

We have thus far endeavoured rudely to trace the genealogy of the Vertebrata by the aid of their mutual affinities. We will now look to man as he exists; and we shall, I think, be able partially to restore during successive periods, but not in due order of time, the structure of our early progenitors. This can be effected by means of the rudiments which man still retains, by the characters which occasionally make their appearance in him through reversion, and by the aid of the principles of morphology and embryology. The various facts, to which I shall here allude, have been given in the previous chapters. The early progenitors of man were no doubt once covered with hair, both sexes having beards; their ears were pointed and capable of movement; and their bodies were provided with a tail, having the proper muscles. Their limbs and bodies were also acted on by many muscles which now only occasionally reappear, but are normally present in the Quadrumana. The great artery and nerve of the humerus ran through a supra-condyloid foramen. At this or some earlier period, the intestine gave forth a much larger diverticulum or cæcum than that now existing. The foot, judging from the condition of the great toe in the fœtus, was then prehensile; and our progenitors, no doubt, were arboreal in their habits, frequenting some warm, forest-clad land. The males were provided with great canine teeth, which served them as formidable weapons.

At a much earlier period the uterus was double; the excreta were voided through a cloaca; and the eye was protected by a third eyelid or nictitating membrane. At a still earlier period the progenitors of man must have been aquatic in their habits; for morphology plainly tells us that our lungs consist of a modified swim-bladder, which once served as a float. The clefts on the neck in the embryo of man show where the branchiæ once existed. At about this period the true kidneys were replaced by the corpora Wolffiana. The heart existed as a simple pulsating vessel; and the chorda dorsalis took the place of a vertebral column. These early predecessors of man, thus seen in the dim recesses of time, must have been as lowly organised as the lancelet or amphioxus, or even still more lowly organised.

There is one other point deserving a fuller notice. It has long been known that in the vertebrate kingdom one sex bears rudiments of various accessory parts, appertaining to the reproductive system, which properly belong to the opposite sex; and it has now been ascertained that at a very early embryonic period both sexes possess true male and female glands. Hence some extremely remote progenitor of the whole vertebrate kingdom appears to have been hermaphrodite or androgynous.[277] But here we encounter a singular difficulty. In the mammalian class the males possess in their vesiculæ prostraticæ rudiments of a uterus with the adjacent passage; they bear also rudiments of mammæ, and some male marsupials have rudiments of a marsupial sack.[278] Other analogous facts could be added. Are we, then, to suppose that some extremely ancient mammal possessed organs proper to both sexes, that is, continued androgynous after it had acquired the chief distinctions of its proper class, and therefore after it had diverged from the lower classes of the vertebrate kingdom? This seems improbable in the highest degree; for had this been the case, we might have expected that some few members of the two lower classes, namely fishes[279] and amphibians, would still have remained androgynous. We must, on the contrary, believe that when the five vertebrate classes diverged from their common progenitor the sexes had already become separated. To account, however, for male mammals possessing rudiments of the accessory female organs, and for female mammals possessing rudiments of the masculine organs, we need not suppose that their early progenitors were still androgynous after they had assumed their chief mammalian characters. It is quite possible that as the one sex gradually acquired the accessory organs proper to it, some of the successive steps or modifications were transmitted to the opposite sex. When we treat of sexual selection, we shall meet with innumerable instances of this form of transmission,—as in the case of the spurs, plumes, and brilliant colours, acquired by male birds for battle or ornament, and transferred to the females in an imperfect or rudimentary condition.

The possession by male mammals of functionally imperfect mammary organs is, in some respects, especially curious. The Monotremata have the proper milk-secreting glands with orifices, but no nipples; and as these animals stand at the very base of the mammalian series, it is probable that the progenitors of the class possessed, in like manner, the milk-secreting glands, but no nipples. This conclusion is supported by what is known of their manner of development; for Professor Turner informs me, on the authority of Kölliker and Lauger, that in the embryo the mammary glands can be distinctly traced before the nipples are in the least visible; and it should be borne in mind that the development of successive parts in the individual generally seems to represent and accord with the development of successive beings in the same line of descent. The Marsupials differ from the Monotremata by possessing nipples; so that these organs were probably first acquired by the Marsupials after they had diverged from, and risen above, the Monotremata, and were then transmitted to the placental mammals. No one will suppose that after the Marsupials had approximately acquired their present structure, and therefore at a rather late period in the development of the mammalian series, any of its members still remained androgynous. We seem, therefore, compelled to recur to the foregoing view, and to conclude that the nipples were first developed in the females of some very early marsupial form, and were then, in accordance with a common law of inheritance, transferred in a functionally imperfect condition to the males.

Nevertheless a suspicion has sometimes crossed my mind that long after the progenitors of the whole mammalian class had ceased to be androgynous, both sexes might have yielded milk and thus nourished their young; and in the case of the Marsupials, that both sexes might have carried their young in marsupial sacks. This will not appear utterly incredible, if we reflect that the males of syngnathous fishes receive the eggs of the females in their abdominal pouches, hatch them, and afterwards, as some believe, nourish the young;[280]—that certain other male fishes hatch the eggs within their mouths or branchial cavities;—that certain male toads take the chaplets of eggs from the females and wind them round their own thighs, keeping them there until the tadpoles are born;—that certain male birds undertake the whole duty of incubation, and that male pigeons, as well as the females, feed their nestlings with a secretion from their crops. But the above suspicion first occurred to me from the mammary glands in male mammals being developed so much more perfectly than the rudiments of those other accessory reproductive parts, which are found in the one sex though proper to the other. The mammary glands and nipples, as they exist in male mammals, can indeed hardly be called rudimentary; they are simply not fully developed and not functionally active. They are sympathetically affected under the influence of certain diseases, like the same organs in the female. At birth they often secrete a few drops of milk; and they have been known occasionally in man and other mammals to become well developed, and to yield a fair supply of milk. Now if we suppose that during a former prolonged period male mammals aided the females in nursing their offspring, and that afterwards from some cause, as from a smaller number of young being produced, the males ceased giving this aid, disuse of the organs during maturity would lead to their becoming inactive; and from two well-known principles of inheritance this state of inactivity would probably be transmitted to the males at the corresponding age of maturity. But at all earlier ages these organs would be left unaffected, so that they would be equally well developed in the young of both sexes.