<h3>FROM QUADRUPED TO BIPED</h3>
<p>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.</p>
<p>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<span class="pagenum"><SPAN name="Page_40" id="Page_40">[40]</SPAN></span> nature, he is chiefly emulated by such
lowly-organized creatures as the ants and the bees.</p>
<p>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<span class="pagenum"><SPAN name="Page_41" id="Page_41">[41]</SPAN></span> 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.</p>
<p>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."</p>
<p><span class="pagenum"><SPAN name="Page_42" id="Page_42">[42]</SPAN></span>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.</p>
<p>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.</p>
<p>The most distinguishing characteristic of the<span class="pagenum"><SPAN name="Page_43" id="Page_43">[43]</SPAN></span> 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.</p>
<p>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<span class="pagenum"><SPAN name="Page_44" id="Page_44">[44]</SPAN></span> 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.</p>
<p>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 spread<span class="pagenum"><SPAN name="Page_45" id="Page_45">[45]</SPAN></span>ing its toes for
this purpose may well have been the opposable great toe.</p>
<p>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.</p>
<p>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 grasp<span class="pagenum"><SPAN name="Page_46" id="Page_46">[46]</SPAN></span>ing
power in the hands and feet of the ape being seemingly that given above.</p>
<p>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.</p>
<p>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.</p>
<p>The power of walking on a lower limb and grasping an upper one once
attained, a succeeding step in evolution quickly appeared, and one<span class="pagenum"><SPAN name="Page_47" id="Page_47">[47]</SPAN></span> 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.</p>
<p>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<span class="pagenum"><SPAN name="Page_48" id="Page_48">[48]</SPAN></span>.
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.</p>
<p>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<span class="pagenum"><SPAN name="Page_49" id="Page_49">[49]</SPAN></span> aid in this,
but the arms are the principal organs of motion, and seem to have
developed in length accordingly.</p>
<p>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.</p>
<p>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.</p>
<p>The natural presumption from this interesting<span class="pagenum"><SPAN name="Page_50" id="Page_50">[50]</SPAN></span> 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.</p>
<p>It is of interest to find that all the existing large<span class="pagenum"><SPAN name="Page_51" id="Page_51">[51]</SPAN></span> 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.</p>
<p>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.</p>
<p>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.<span class="pagenum"><SPAN name="Page_52" id="Page_52">[52]</SPAN></span> 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.</p>
<p>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.</p>
<p><span class="pagenum"><SPAN name="Page_53" id="Page_53">[53]</SPAN></span>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.</p>
<hr style="width: 65%;" />
<p><span class="pagenum"><SPAN name="Page_54" id="Page_54">[54]</SPAN></span></p>
<h2><SPAN name="V" id="V"></SPAN>V</h2>
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