The toes are fastened by means of well adapted joints to the lower end of the tarsus, and form what is popularly regarded as the bird's foot. When spoken of separately, these toes are called digits, and when spoken of collectively, they are called the podium. They are composed of small bones called phalanges or internodes, which are jointed upon one another like the several parts of the human fingers. The digits can be spread out for walking purposes, or bent around so as to clasp an object. The outer bone of each digit almost always bears a nail or claw, which is sometimes very strong and hooked, as is the case with the birds of prey, while in other species it is only slightly curved and is not meant as a weapon of offense or defense, but chiefly to enable the bird to "scratch for a living."
How do the birds, in perching and roosting, retain their hold so long on a limb without becoming weary? They do not need to make a conscious effort to do this, but are held by the mechanical action of certain muscles and tendons in the leg and foot. Of course, the bird can also control these muscles by an act of its will, but a large part of their action is automatic. In some species there is a muscle called the ambiens, which has its rise in the pelvis, passes along the inner side of the thigh, whence its tendon runs over the apex of the angle of the knee joint, and down the leg till it joins the muscles that flex the toes. Now when the bird's leg is bent at the joints, as is the case in perching, the tendons of this muscle are stretched over the knee and ankle joints, thus pulling the digits together, and causing them of their own accord to grasp the perch more or less tightly. When a bird wishes to unloose its hold, it simply rises on its feet and relaxes the tendons.
All birds by no means possess this particular muscle, but all the perchers have some muscular arrangement in the legs and toes that practically answers the same purpose. If you will bend your wrist backward as far as you can, you will observe that your fingers will have a tendency to curve slightly forward. This is caused by the stretching of the tendons over the convex part of your bent wrist joints.
The typical bird has four digits, three in front and one reaching backward. The hind toe is called the hallux, and corresponds to the thumb of the human hand, so that in grasping an object it can be made to meet any of the other toes. But many birds are not provided with a quartet of digits. The ostrich has only two, the inner and hinder toes being wanting. However, this great fowl does not experience any lack, for its feet are almost solid like hoofs, and quite flat, and hence are especially adapted for traveling across the sandy desert.
No bird has ever been found with more than four toes; and four seem to be ample for all purposes. A fifth toe for a bird would be as useless as a fifth wheel on a wagon. Quite a number of species have only three toes, most of them among the walkers and waders, and none, I believe, among the true perchers. Take the plovers and sanderlings, for example, which spend most of their time, when not on the wing, in running about on the ground, especially along the seashore or the banks of streams and lakes, and seldom, if ever, sit on a perch—in their case a fourth toe would be worse than a superfluous appendage; it would be an encumbrance, dragging along in the mud and mire. In these species it is the hind toe that is lacking, their three digits all being in front, where they are of the greatest service. There is another class of birds that have hind toes, though very much reduced because their owners do not perch, but scuttle about on the beach. This class includes the little spotted sandpipers which you often see running or flying along the shores of a river or lake.
Curious to tell, several species of woodpeckers are tridactyl—that is, three-toed—and still more curious is the fact that in their case the true hind toe is lacking, while the outer front toe is bent backward, or "reversed," as it is called, and is thus made to do service for a hind toe. The other species of woodpeckers have four toes, two in front and two behind, the outer one of the latter pair being a reversed digit. Why some of the woodpeckers should have four toes and others only three is an unsolved enigma, and is especially puzzling in view of the fact that the four-toed kinds do not seem to possess any advantage over their cousins. The tridactyl species are as expert climbers as any members of the family, and are extremely hardy birds, too, some of them dwelling the year round in cold northern climates, where the food question must often be a serious one.
Here is still another conundrum for the bird student: Why do the four-toed woodpeckers have two hind digits, despite the fact that they always clamber upward when they take their promenades on the boles and branches of the trees, whereas the agile little nuthatch, which glides upward or downward, as the impulse moves him, has only one rear toe and three in front, like the true perchers? Nor is it less puzzling that the cuckoos, which are perching birds, should have two toes in front and two behind. Then, there is the little brown creeper which never perches and is forever creeping, creeping, upward, upward—save, of course, when it takes to wing—and yet its toes are arranged in the normal percher style, the hind digit having an especially long, curved claw. It is a mistake to suppose that all the problems of the bird world have been solved.
Look at the different kinds of birds' feet and see how wisely they have been planned for the various purposes to which they have been applied. In order that a bird may use his feet with the greatest dexterity in perching and flitting, his digits should be as free and movable as possible; and so we find that the toes of the perchers are usually cleft to the base, are long and slender, easily opened and closed, and possess the power to grasp an object firmly. The same is true of the raptorial birds, or birds of prey, which are strong perchers and depend largely for their food supply on clutching their victims while on the wing. In all these birds the hind toe is also well developed, and is on the same plane as the anterior digits—a wise adaptation of means to ends.