The Descent of Man, and Selection in Relation to Sex. Charles Darwin
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on the other, in the same manner as we do. They can thus also lift rather large objects, such as the neck of a bottle, to their mouths. Baboons turn over stones, and scratch up roots with their hands. They seize nuts, insects, or other small objects with the thumb in opposition to the fingers, and no doubt they thus extract eggs and young from the nests of birds. American monkeys beat the wild oranges on the branches until the rind is cracked, and then tear it off with the fingers of the two hands. In a wild state they break open hard fruits with stones. Other monkeys open mussel-shells with the two thumbs. With their fingers they pull out thorns and burs, and hunt for each other's parasites. They roll down stones, or throw them at their enemies: nevertheless, they are clumsy in these various actions, and, as I have myself seen, are quite unable to throw a stone with precision.
It seems to me far from true that because "objects are grasped clumsily" by monkeys, "a much less specialised organ of prehension" would have served them (70. 'Quarterly Review,' April 1869, p. 392.) equally well with their present hands. On the contrary, I see no reason to doubt that more perfectly constructed hands would have been an advantage to them, provided that they were not thus rendered less fitted for climbing trees. We may suspect that a hand as perfect as that of man would have been disadvantageous for climbing; for the most arboreal monkeys in the world, namely, Ateles in America, Colobus in Africa, and Hylobates in Asia, are either thumbless, or their toes partially cohere, so that their limbs are converted into mere grasping hooks. (71. In Hylobates syndactylus, as the name expresses, two of the toes regularly cohere; and this, as Mr. Blyth informs me, is occasionally the case with the toes of H. agilis, lar, and leuciscus. Colobus is strictly arboreal and extraordinarily active (Brehm, 'Thierleben,' B. i. s. 50), but whether a better climber than the species of the allied genera, I do not know. It deserves notice that the feet of the sloths, the most arboreal animals in the world, are wonderfully hook-like.
As soon as some ancient member in the great series of the Primates came to be less arboreal, owing to a change in its manner of procuring subsistence, or to some change in the surrounding conditions, its habitual manner of progression would have been modified: and thus it would have been rendered more strictly quadrupedal or bipedal. Baboons frequent hilly and rocky districts, and only from necessity climb high trees (72. Brehm, 'Thierleben,' B. i. s. 80.); and they have acquired almost the gait of a dog. Man alone has become a biped; and we can, I think, partly see how he has come to assume his erect attitude, which forms one of his most conspicuous characters. Man could not have attained his present dominant position in the world without the use of his hands, which are so admirably adapted to act in obedience to his will. Sir C. Bell (73. 'The Hand,' etc., 'Bridgewater Treatise,' 1833, p. 38.) insists that "the hand supplies all instruments, and by its correspondence with the intellect gives him universal dominion." But the hands and arms could hardly have become perfect enough to have manufactured weapons, or to have hurled stones and spears with a true aim, as long as they were habitually used for locomotion and for supporting the whole weight of the body, or, as before remarked, so long as they were especially fitted for climbing trees. Such rough treatment would also have blunted the sense of touch, on which their delicate use largely depends. From these causes alone it would have been an advantage to man to become a biped; but for many actions it is indispensable that the arms and whole upper part of the body should be free; and he must for this end stand firmly on his feet. To gain this great advantage, the feet have been rendered flat; and the great toe has been peculiarly modified, though this has entailed the almost complete loss of its power of prehension. It accords with the principle of the division of physiological labour, prevailing throughout the animal kingdom, that as the hands became perfected for prehension, the feet should have become perfected for support and locomotion. With some savages, however, the foot has not altogether lost its prehensile power, as shewn by their manner of climbing trees, and of using them in other ways. (74. Haeckel has an excellent discussion on the steps by which man became a biped: 'Natürliche Schöpfungsgeschichte,' 1868, s. 507. Dr. Buchner ('Conférences sur la Théorie Darwinienne,' 1869, p. 135) has given good cases of the use of the foot as a prehensile organ by man; and has also written on the manner of progression of the higher apes, to which I allude in the following paragraph: see also Owen ('Anatomy of Vertebrates,' vol. iii. p. 71) on this latter subject.)
If it be an advantage to man to stand firmly on his feet and to have his hands and arms free, of which, from his pre-eminent success in the battle of life there can be no doubt, then I can see no reason why it should not have been advantageous to the progenitors of man to have become more and more erect or bipedal. They would thus have been better able to defend themselves with stones or clubs, to attack their prey, or otherwise to obtain food. The best built individuals would in the long run have succeeded best, and have survived in larger numbers. If the gorilla and a few allied forms had become extinct, it might have been argued, with great force and apparent truth, that an animal could not have been gradually converted from a quadruped into a biped, as all the individuals in an intermediate condition would have been miserably ill-fitted for progression. But we know (and this is well worthy of reflection) that the anthropomorphous apes are now actually in an intermediate condition; and no one doubts that they are on the whole well adapted for their conditions of life. Thus the gorilla runs with a sidelong shambling gait, but more commonly progresses by resting on its bent hands. The long-armed apes occasionally use their arms like crutches, swinging their bodies forward between them, and some kinds of Hylobates, without having been taught, can walk or run upright with tolerable quickness; yet they move awkwardly, and much less securely than man. We see, in short, in existing monkeys a manner of progression intermediate between that of a quadruped and a biped; but, as an unprejudiced judge (75. Prof. Broca, La Constitution des Vertèbres caudales; 'La Revue d'Anthropologie,' 1872, p. 26, (separate copy).) insists, the anthropomorphous apes approach in structure more nearly to the bipedal than to the quadrupedal type.
As the progenitors of man became more and more erect, with their hands and arms more and more modified for prehension and other purposes, with their feet and legs at the same time transformed for firm support and progression, endless other changes of structure would have become necessary. The pelvis would have to be broadened, the spine peculiarly curved, and the head fixed in an altered position, all which changes have been attained by man. Prof. Schaaffhausen (76. 'On the Primitive Form of the Skull,' translated in 'Anthropological Review,' Oct. 1868, p. 428. Owen ('Anatomy of Vertebrates,' vol. ii. 1866, p. 551) on the mastoid processes in the higher apes.) maintains that "the powerful mastoid processes of the human skull are the result of his erect position;" and these processes are absent in the orang, chimpanzee, etc., and are smaller in the gorilla than in man. Various other structures, which appear connected with man's erect position, might here have been added. It is very difficult to decide how far these correlated modifications are the result of natural selection, and how far of the inherited effects of the increased use of certain parts, or of the action of one part on another. No doubt these means of change often co-operate: thus when certain muscles, and the crests of bone to which they are attached, become enlarged by habitual use, this shews that certain actions are habitually performed and must be serviceable. Hence the individuals which performed them best, would tend to survive in greater numbers.
The free use of the arms and hands, partly the cause and partly the result of man's erect position, appears to have led in an indirect manner to other modifications of structure. The early male forefathers of man were, as previously stated, probably furnished with great canine teeth; but as they gradually acquired the habit of using stones, clubs, or other weapons, for fighting with their enemies or rivals, they would use their jaws and teeth less and less. In this case, the jaws, together with the teeth, would become reduced in size, as we may feel almost sure from innumerable analogous cases. In a future chapter we shall meet with a closely parallel case, in the reduction or complete disappearance of the canine teeth in male ruminants, apparently in relation with the development of their horns; and in horses, in relation to their habit of fighting with their incisor teeth and hoofs.
In the adult male anthropomorphous apes, as Rutimeyer (77. 'Die Grenzen der Thierwelt, eine Betrachtung zu Darwin's Lehre,' 1868, s. 51.), and others, have insisted, it is the effect on the skull of the great development of the jaw-muscles that causes it to differ so greatly in many respects from that of man, and has given to these animals "a truly frightful physiognomy." Therefore, as the jaws and teeth in man's progenitors gradually become reduced in size, the adult skull would have come to resemble