Chapter 4. From Quadruped To Biped

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_ CHAPTER IV. FROM QUADRUPED TO BIPED

In the question which now confronts us, that of the evolution of man from the lower world of animals, it is necessary first to state in what particulars he has evolved, what are the conditions which distinguish him from the lower animals. Four marked distinctions may be named: his erect attitude, with the freeing of the fore limbs from use as agents in locomotion; his employment of natural objects, instead of his bodily organs, as tools and weapons; his development of vocal language; and his great mental superiority, with the general use of the mind in his dealings with nature.

In none of these particulars does man stand quite alone; in all of them an affinity with the lower animals exists. Steps of progress in these directions have been made by many animals, though none of them have gained any considerable advance. In man's strikingly developed social habit and organization he has no close counterpart among the vertebrates, but several among the insects. And it is of much interest to find that in the highest field of man's progress, his employment of the mind in his dealings with nature, he is chiefly emulated by such lowly-organized creatures as the ants and the bees.

We do not need to look far among the lower animals for the species which come nearest to man in structure and which seem to have immediately preceded him in the line of descent. We find these forms in the monkeys or apes, and especially in their highest representatives, the anthropoid apes. These possess in a partial degree all the special characteristics of man. They are social in habit; some of them are semi-erect in posture, and their fore limbs partly freed from use in locomotion; they possess some imperfect means of vocal communication; they employ the mind to some extent in place of the body; in short, they seem arrested forms on the road from brute to man, signal-posts on the highway of evolution. In physical organization their approach to man is singularly close. In anatomy man and the higher apes are in most respects counterparts of each other. The principal anatomical distinction has been considered to be in the foot, which from the opposable character of the great toe was classed by Cuvier with the hand, the apes being named Quadrumana, or four-handed, and man Bimana, or two-handed. Fuller research has shown that this distinction does not exist, the foot of the ape being found to agree far more closely with the foot than with the hand of man. Estimated according to use, the hand is, in the whole order, the special prehensile organ; the foot, however prehensile it may be, is predominantly a walking organ. And the opposability of the great toe is approached in some men, who have great mobility in this organ, and can use it for grasping.

In regard to the brain, the organ of the mind, the difference between the higher apes and man is almost solely one of comparative size, the lower intelligence of the apes being indicated by the smaller size of their brains. The largest ape brain is scarcely half the size of the smallest human brain. But anatomically they are nearly identical. All the structural features of the brain are common to both, and the details are largely filled out in the anthropoid apes, the convolutions being all present and the pattern of arrangement the same. The brain of the orang may be said to be like that of man in all respects except size and the greater symmetry of its convolutions, which are less complicated with minor convolutions than in man. In truth, the difference between the brains of man and the orang is almost insignificant as compared with the difference between those of the orang and the lowest apes. Mr. E. W. Taylor, who has recently made an exhaustive study of the minute anatomy of the brain of the chimpanzee, remarks, "The similarity between the brain of the anthropoid apes and of man is one of the most singular and interesting facts of which we have knowledge."

In any attempt, then, to consider the origin of man from the point of view of evolution, we are irresistibly drawn to the ape tribe as the next lower link in the long chain of development, and are led to consider the characteristics of the apes as the intermediate stage between the quadruped and the biped, the bridge crossing this great gulf in organic development. This is by no means to suggest that some one of the existing anthropoid apes is the direct ancestor of man. Such an idea has never been entertained by scientists. These animals cannot even fairly be considered as brothers to man's ancestor, but must be looked upon as more or less distant cousins, with a physical organization less favorable to high development than that of man. Man's ancestry lies much farther back in time, and his progenitor must have been constituted differently from any of the existing large apes.

In the ape tribe we are able to trace nearly every step by which the gulf between quadruped and biped has been crossed, from the quadrupedal baboon to the nearly erect gibbon. And in seeking to follow this development through its successive stages, the first point to be considered is how the apes gained their special power of grasping, that characteristic to which they undoubtedly owe the partial freedom of their hands and their tendency to assume the erect attitude.

The most distinguishing characteristic of the apes and of the nearly related lemurs has not hitherto been definitely pointed out. This is that they form the only group of strictly arboreal animals. The tree is not alone their native habitat, but they are specially adapted to it in their organs of motion, a fact which cannot be affirmed of any other animal group. If we consider, for instance, the squirrels, one of the best-known groups of tree-living animals, we find them to be members of the great order of rodents, whose native habitat is the land surface. Though the squirrels have taken to the trees, there has been no adaptive change in the structure of their limbs and feet. The same may be said of almost all tree-dwellers except the lemurs and apes. The sloth, indeed, is specially adapted in organization to an arboreal residence, but this change is individual, not tribal, this animal being an aberrant form of the ground-dwelling edentata. In the apes and lemurs, on the contrary, the ground-dwellers are the aberrant forms, stray wanderers from the host. Nearly all the species live in trees, to which they are specially adapted by the formation of their feet. It remains to inquire how this deviation in structure arose, what were the steps of development of the grasping foot and hand, the special characteristic of this group.

In considering this question, the first fact to appear is that the apes and lemurs are plantigrade animals. Their natural tendency is to walk on the sole of the foot, a habit which few other tribes of animals possess. Most of the larger animals walk on the knuckles or the toes, and develop claws or hoofs, but the ancestral form of the ape, ages in the past, was doubtless a sole-walking quadruped, its toes apparently provided with nails instead of claws. What the story of this very ancient quadruped was we are quite unable to say. It may, in the exigencies of existence, have come to a parting of the ways; a section of the group, drawn by a love of fruit, developing the climbing habit; the remaining section continuing on the ground and following a separate line of evolution. Perhaps only a single species took to the trees; for it is quite possible for a single form, in a new and advantageous habitat, to vary in time into a great number of species.

Of all this we can know nothing: but of one thing we may feel assured, which is that the plantigrade foot is the only one that could have developed into a grasping organ; such a development being impossible to the digitigrade or the hoofed animals. One can readily see how the habit of walking on the sole might tend to a spreading of the toes, in order to obtain a wider and firmer footing. And it is equally easy to see how a free and wide motion in the great toe would aid in this result. The animal may have been at first light in weight and able to support itself on its unchanged foot, but as it increased in size and weight it would need a firmer grasp, and the final result of spreading its toes for this purpose may well have been the opposable great toe.

It must be borne in mind, in this consideration, that the apes differ from the other tree-dwellers in being destitute of claws. The squirrels, the opossums, and other arboreal animals have sharp claws, by whose aid they can easily cling to the surface of the bark-covered boughs. The nails of the apes are incapable of affording them this service, and it is not easy to perceive how a foot like theirs could become adapted to locomotion in the trees otherwise than by the gaining of mobile action and grasping power in the toes.

The existing habits of the ape tribe lead us to the conclusion that the ancestral animal may have soon begun to seek support from upper limbs. The plantigrade foot is one capable of readily curving into an organ of support, and in the case of the forefoot the toes would tend to spread and gain flexibility of motion, and the first toe to become opposable to the others and yield a more complete grasping power. It does not seem difficult to comprehend, from this point of view, how the feet of a five-toed plantigrade animal may in time have developed into grasping organs, since there would be required only an increased flexibility of the joints, and a wider and fuller movement of the great toes. That such a change took place in this instance the facts appear to indicate, the most simple and probable explanation of the development of the grasping power in the hands and feet of the ape being seemingly that given above.

The relation of the lemurs to the apes is not clearly defined. It may be an ancestral one, or the two animals may represent distinct lines of descent. In the latter case we would have two lines of animal evolution in which the grasping power was gained and adaptation to arboreal life completed. Whatever their relationship, they both possess the opposable thumb as the hall-mark of their arboreal habitat, and whenever found walking on the ground they may be looked upon as estrays from their native place of residence.

Once the grasping power was gained, the first step of change from the quadrupedal to the semi-erect attitude was completed. The process may have begun in the effort to fit the sole of the foot to the rounded surface of boughs; or its first stage may have been in the seizing of overhead branches with the flexible hand; or both influences may have acted simultaneously. We see the result only, we cannot trace the exact process; but we have as an outcome the adoption of a method of locomotion different from that of all other tree-dwellers, the forefoot developing into the hand with its opposable thumb, and the hindfoot gaining a similar grasping power in the toes.

The power of walking on a lower limb and grasping an upper one once attained, a succeeding step in evolution quickly appeared, and one of prime importance to our inquiry. The animal had ceased to be in a full sense a quadruped, while not yet a biped, and a variation in the length of its limbs was almost sure to take place. This is an ordinary result when animals cease to walk on all fours. In the leaping kangaroo and jerboa a shortening of the arms and lengthening of the legs appear. Here the arms are relieved from duty and a double duty is laid on the legs, with the consequence stated. In the ancient dinosaurian reptiles, upright walkers, the same was the case. Those varied from quite small to very large animals, but in all known instances the fore limbs were greatly reduced in size. A similar condition may be seen in the birds, the bones of whose fore-limbs have largely aborted from lack of employment as walking organs.

In the case of the apes and lemurs, while a similar effect has taken place, an interesting difference appears, due to the difference in conditions. In these animals the fore limbs are not freed from duty as organs of locomotion. In many cases, on the contrary, they have an extra duty put upon them, with the result that they have grown longer instead of shorter. Very likely these animals differed considerably in the past, as they do to-day, in the degree of use of their legs and arms. Many of them walk in the quadruped manner, either on the ground or in trees. Others make much use of their hands and arms in grasping and swinging. Great differences in the use of the arms and legs may have arisen in different species. In some, the legs may have been mainly trusted to for support, and the hands used for steadying. In others the arms may have been the chief locomotive organs and the feet have given steadiness. Here the legs may have grown the longer, there the arms, the limbs developing in accordance with their degree of employment. In the lower monkeys and the lemurs, the bones of the pelvis are altogether quadrupedal in character. This is not the case in the higher forms, and in the highest apes the pelvic bones approach those of man.

Highly interesting examples of these varied results may be seen in the existing anthropoid apes. In all of them it would appear that the arm was a prominent factor in locomotion, for in each instance it is longer than the leg,--but it differs in proportional length in every instance. It is shortest in the chimpanzee, somewhat longer in the gorilla, still longer in the orang, and remarkably long in the gibbon. In all these instances the fact that the arms exceed the legs in length indicates that they must have played a large and important part in the work of locomotion, and especially so in the case of the gibbon. It is well known, in fact, that the gibbons progress very largely by the aid of their arms, swinging from limb to limb and from tree to tree with extraordinary strength and facility. The legs lend their aid in this, but the arms are the principal organs of motion, and seem to have developed in length accordingly.

As regards the other anthropoid species, Wallace's observations on the habits of the orang are of interest. This animal usually walks on all fours on the branches in a semi-erect crouching attitude, but our naturalist saw one moving by the use of its arms alone. In passing from tree to tree the arms come actively into play. The animal seizes a handful of the overlapping boughs of the two trees and swings easily across the intervening space. While seeming to move very deliberately, its actual speed was found to be about six miles an hour.

The organization of man, as he now exists, shows an interesting and important deviation from that of the manlike apes, and one which serves as strong evidence that none of these apes occupied a place in his line of descent. This is that he is a long-legged and short-armed animal, a condition the reverse of that seen in the anthropoid apes. While man's hands reach barely to the middle of the thigh, those of the chimpanzee reach below the knee, of the gorilla to the middle of the leg, of the orang to the ankle, and of the gibbon to the ground. All these apes have short legs and long arms. Man, on the contrary, has long legs and short arms.

The natural presumption from this interesting fact is that man's ancestor, which we may provisionally call the man-ape, differed essentially in its mode of progression from the other apes. The smaller forms of these usually move on all fours in the trees, though the arms are always ready for a swing or a climb. The anthropoid apes also show a tendency to a similar mode of progression, though with a difference in their mode of walking, which, as we shall see later on, is never that of the quadruped. As for the man-ape, it may have originally walked in the same manner as the related species, if we surmise that the variation in the length of the limbs was a subsequent development. Certainly after its limbs attained the proportions of those of man, its facility of swinging from tree to tree must have been diminished, while it would have found it inconvenient to move in the crouching attitude of the orang and its fellows. Its easiest attitude must then have been the erect one, and its motion a true biped walk, not the swinging and jumping movement of the other anthropoids. In short, the development of man's ancestor into a short-armed animal, however and whenever it took place, could not but have interfered seriously with its ease of motion in the trees. Though this change may have begun in the trees, it probably had its full development only after the animal made the ground its habitual place of residence.

It is of interest to find that all the existing large apes are arboreal, the gorilla being the least so, probably on account of its weight. Though they all descend at times to the ground, their awkward motion on the surface shows them to be out of their element, while they move with ease and rapidity in the trees. The organization of man renders it questionable if his primeval ancestor was arboreal to any similar extent. The indications would seem to be that it made the ground its habitual place of residence at an early period in its history, and that the result of this new habit and of its erect attitude was a change in the relative length of its limbs.

That this animal dwelt mainly in trees in the first stage of its existence, and possessed a powerful grasping power in its hands, we have corroborative evidence in recent studies of child life. The human infant, in its earliest days of life, displays a remarkable grasping power, being able to sustain its weight with its hands for a number of seconds, or a minute or more, at an age when its other muscles are flabby and powerless. It appears in this to repeat a habit normal to the ancestral infant, an instinct developed to prevent a fall from its home among the boughs.

Yet it is doubtful if the man-ape long remained a specially arboreal animal. The varied length of arm in the anthropoid apes was doubtless of early origin, and in all probability man's ancestor had originally a shorter arm than its related species. If so, this must have rendered it less agile in trees than other forms. If we could see this ancient creature in its arboreal home, we should probably find it more inclined to stand erect than the other apes, walking on a lower limb, and steadying itself by grasping an upper limb. This would be a more natural and easy mode of progression to a short-armed animal than the crouching attitude of the orang or the swinging motion of the gibbon, and its effect would be to make the erect attitude to a large extent habitual with this animal.

In short, man's ancestor may have become in considerable measure a biped while still largely a dweller in the trees, and to that degree set its arms free for other duties than that of locomotion. Like the other apes, it probably often descended to the ground, where its habit of walking erect on the boughs rendered the biped walk an easy one, or where this habit may have been originally acquired. While this is conjectural, it is supported by facts of organization and existing habit, and for the reasons given it seems highly probable that the ancestor of man took to a land residence at an early period in its history, climbing again for food or safety, but dwelling more and more habitually on the earth's surface. Even at this remote era it may have become essentially human in organization, its subsequent changes being mainly in brain development, and only to a minor extent in physical form and structure.

Fossil apes have not been found farther back than the Miocene Age of geology. It is quite probable, however, that they may yet be found in Eocene strata, since examples of their highest representatives, the anthropoid or manlike apes, have been found in Miocene rocks. The fact that these large apes are now few in number of species, is no proof that many forms of them may not have formerly existed, and among these we may class the ancestor of man. _

Read next: Chapter 5. The Freedom Of The Arms

Read previous: Chapter 3. Relics Of Ancient Man

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