Fig. 24.—Skeleton of the Ostrich. Shows the powerful legs, small feet, and rudimentary wings of the bird; the obliquity at which the bones of the legs and wings are placed, and the comparatively small angles which any two bones make at their point of junction. a Angle made by femur with ilium. b Angle made by tibia and fibula with femur. c Angle made by tarso-metatarsal bone with tibia and fibula. d Angle made by bones of foot with tarso-metatarsal bone. r Bones of wing inclined to each other at nearly right angles. Compare with fig. [4], p. 21, fig. [26], p. 55, and fig. [27], p. 59.—Adapted from Dallas.
The feet of the ostrich, like those of the horse and deer, are reduced to a minimum as regards size; so that they occasion very little friction in the act of walking and running. The foot is composed of two jointed toes,[30] which spread out when the weight of the body comes upon them, in such a manner as enables the bird to seize and let go the ground with equal facility. The advantage of such an arrangement in rapid locomotion cannot be over-estimated. The elasticity and flexibility of the foot contribute greatly to the rapidity of movement for which this celebrated bird is famous. The limb of the ostrich, with its large bones placed very obliquely to form a system of powerful levers, is the very embodiment of speed. The foot is quite worthy of the limb, it being in some respects the most admirable structure of its kind in existence. The foot of the ostrich differs considerably from that of all other birds, those of its own family excepted. Thus the under portion of the foot is flat, and specially adapted for acting on plane surfaces, particularly solids.[31] The extremities of the toes superiorly are armed with powerful short nails, the tips of which project inferiorly to protect the toes and confer elasticity when the foot is leaving the ground. The foot, like the leg, is remarkable for its great strength. The legs of the ostrich are closely set, another feature of speed.[32] The wings of the ostrich are in a very rudimentary condition as compared with the legs.[33] All the bones are present, but they are so dwarfed that they are useless as organs of flight. The angles which the bones of the wing make with each other, are still less than the angles made by the bones of the leg. This is just what we would a priori expect, as the velocity with which wings are moved greatly exceeds that with which legs are moved. The bones of the wing of the ostrich are inclined towards each other at nearly right angles. The wings of the ostrich, although useless as flying organs, form important auxiliaries in running. When the ostrich careers along the plain, he spreads out his wings in such a manner that they act as balancers, and so enable him to maintain his equilibrium (fig. 25). The wings, because of the angle of inclination which their under surfaces make with the horizon, and the great speed at which the ostrich travels, act like kites, and so elevate and carry forward by a mechanical adaptation a certain proportion of the mass of the bird already in motion. The elevating and propelling power of even diminutive inclined planes is very considerable, when carried along at a high speed in a horizontal direction. The wings, in addition to their elevating and propelling power, contribute by their short, rapid, swinging movements, to continuity of motion in the legs. No bird with large wings can run well. The albatross, for example, walks with difficulty, and the same may be said of the vulture and eagle. What, therefore, appears a defect in the ostrich, is a positive advantage when its habits and mode of locomotion are taken into account.
Fig. 25.—Ostriches pursued by a Hunter.
Professional runners in many cases at matches reduce the length of their anterior extremities by flexing their arms and carrying them on a level with their chest (fig. [28], p. 62). It would seem that in rapid running there is not time for the arms to oscillate naturally, and that under these circumstances the arms, if allowed to swing about, retard rather than increase the speed. The centre of gravity is well forward in the ostrich, and is regulated by the movements of the head and neck, and the obliquity of the body and legs. In running the neck is stretched, the body inclined forward, and the legs moved alternately and with great rapidity. When the right leg is flexed and elevated, it swings forward pendulum-fashion, and describes a curve whose convexity is directed towards the right side. When the left leg is flexed and elevated, it swings forward and describes a curve whose convexity is directed towards the left side. The curves made by the right and left legs form when united a waved line (vide figs. [18], 19, and 20, pp. 37, 39, and 41). When the right leg is flexed, elevated, and advanced, it rotates upon the iliac portion of the trunk of the bird, the trunk being supported for the time being by the left leg, which is extended, and in contact with the ground. When the left leg is flexed, elevated, and advanced, it in like manner rotates upon the trunk, supported in this instance by the extended right leg. The leg which is on the ground for the time being supplies the necessary lever, the ground the fulcrum. When the right leg is flexed and elevated, it rotates upon the iliac portion of the trunk in a forward direction, the right foot describing the arc of a circle. When the right leg and foot are extended and fixed on the ground, the trunk rotates upon the right foot in a forward direction to form the arc of a circle, which is the converse of that formed by the right foot. If the arcs alternately supplied by the right foot and trunk are placed in opposition, a more or less perfect circle is produced, and thus it is that the locomotion of animals is approximated to the wheel in mechanics. Similar remarks are to be made of the left foot and trunk. The alternate rolling of the trunk on the extremities, and the extremities on the trunk, utilizes or works up the inertia of the moving mass, and powerfully contributes to continuity and steadiness of action in the moving parts. By advancing the head, neck, and anterior parts of the body, the ostrich inaugurates the rolling movement of the trunk, which is perpetuated by the rolling movements of the legs. The trunk and legs of the ostrich are active and passive by turns. The movements of the trunk and limbs are definitely co-ordinated. But for this reciprocation the action of the several parts implicated would neither be so rapid, certain, nor continuous. The speed of the ostrich exceeds that of every other land animal, a circumstance due to its long, powerful legs and great stride. It can outstrip without difficulty the fleetest horses, and is only captured by being simultaneously assailed from various points, or run down by a succession of hunters on fresh steeds. If the speed of the ostrich, which only measures six or eight feet, is so transcending, what shall we say of the speed of the extinct Æpyornis maximus and Dinornis giganteus, which are supposed to have measured from sixteen to eighteen feet in height? Incredible as it may appear, the ostrich, with its feet reduced to a minimum as regards size, and peculiarly organized for walking and running on solids, can also swim. Mr. Darwin, that most careful of all observers, informs us that ostriches take to the water readily, and not only ford rapid rivers, but also cross from island to island. They swim leisurely, with neck extended, and the greater part of the body submerged.
Locomotion in Man.—The speed attained by man, although considerable, is not remarkable. It depends on a variety of circumstances, such as the height, age, sex, and muscular energy of the individual, the nature of the surface passed over, and the resistance to forward motion due to the presence of air, whether still or moving. A reference to the human skeleton, particularly its inferior extremities, will explain why the speed should be moderate.
On comparing the inferior extremities of man with the legs of birds, or the posterior extremities of quadrupeds, say the horse or deer, we find that the bones composing them are not so obliquely placed with reference to each other, neither are the angles formed by any two bones so acute. Further, we observe that in birds and quadrupeds the tarsal and metatarsal bones are so modified that there is an actual increase in the number of the angles themselves. In the extremities of birds and quadrupeds there are four angles, which may be increased or diminished in the operations of locomotion. Thus, in the quadruped and bird (fig. [4], p. 21, and fig. [24], p. 47), the femur forms with the ilium one angle (a); the tibia and fibula with the femur a second angle (b); the cannon or tarso-metatarsal bone with the tibia and fibula a third angle (c); and the bones of the foot with the cannon or tarso-metatarsal bone a fourth angle (d). In man three angles only are found, marked respectively a, b, and c (figs. [26] and [27], pp. 55 and 59). The fourth angle (d of figs. [4] and [24]) is absent. The absence of the fourth angle is due to the fact that in man the tarsal and metatarsal bones are shortened and crushed together; whereas in the quadruped and bird they are elongated and separated.
As the speed of a limb increases in proportion to the number and acuteness of the angles formed by its several bones, it is not difficult to understand why man should not be so swift as the majority of quadrupeds. The increase in the number of angles increases the power which an animal has of shortening and elongating its extremities, and the levers which the extremities form. To increase the length of a lever is to increase its power at one end, and the distance through which it moves at the other; hence the faculty of bounding or leaping possessed in such perfection by many quadrupeds.[34] If the wing be considered as a lever, a small degree of motion at its root produces an extensive sweep at its tip. It is thus that the wing is enabled to work up and utilize the thin medium of the air as a buoying medium.
Another drawback to great speed in man is his erect position. Part of the power which should move the limbs is dedicated to supporting the trunk. For the same reason the bones of the legs, instead of being obliquely inclined to each other, as in the quadruped and bird, are arranged in a nearly vertical spiral line. This arrangement increases the angle formed by any two bones, and, as a consequence, decreases the speed of the limbs, as explained. A similar disposition of the bones is found in the anterior extremities of the elephant, where the superincumbent weight is great, and the speed, comparatively speaking, not remarkable. The bones of the human leg are beautifully adapted to sustain the weight of the body and neutralize shock.[35] Thus the femur or thigh bone is furnished at its upper extremity with a ball-and-socket joint which unites it to the cup-shaped depression (acetabulum) in the ilium (hip bone). It is supplied with a neck which carries the body or shaft of the bone in an oblique direction from the ilium, the shaft being arched forward and twisted upon itself to form an elongated cylindrical screw. The lower extremity of the femur is furnished with spiral articular surfaces accurately adapted to the upper extremities of the bones of the leg, viz. the tibia and fibula, and to the patella. The bones of the leg (tibia and fibula) are spirally arranged, the screw in this instance being split up. At the ankle the bones of the leg are applied to those of the foot by spiral articular surfaces analogous to those found at the knee-joint. The weight of the trunk is thus thrown on the foot, not in straight lines, but in a series of curves. The foot itself is wonderfully adapted to receive the pressure from above. It consists of a series of small bones (the tarsal, metatarsal, and phalangeal bones), arranged in the form of a double arch; the one arch extending from the heel towards the toes, the other arch across the foot. The foot is so contrived that it is at once firm, elastic, and moveable,—qualities which enable it to sustain pressure from above, and exert pressure from beneath. In walking, the heel first reaches and first leaves the ground. When the heel is elevated the weight of the body falls more and more on the centre of the foot and toes, the latter spreading out[36] as in birds, to seize the ground and lever the trunk forward. It is in this movement that the wonderful mechanism of the foot is displayed to most advantage, the multiplicity of joints in the foot all yielding a little to confer that elasticity of step which is so agreeable to behold, and which is one of the characteristics of youth. The foot may be said to roll over the ground in a direction from behind forwards. I have stated that the angles formed by the bones of the human leg are larger than those formed by the bones of the leg of the quadruped and bird. This is especially true of the angle formed by the femur with the ilium, which, because of the upward direction given to the crest of the ilium in man, is so great that it virtually ceases to be an angle.