The Origin of Species

more curious than that the hand of a man, formed for grasping, that of a mole for digging, the leg of the horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern, and should include similar bones, in the same relative positions? How curious it is, to give a subordinate though striking instance, that the hind feet of the kangaroo, which are so well fitted for bounding over the open plains⁠—those of the climbing, leaf-eating koala, equally well fitted for grasping the branches of trees⁠—those of the ground-dwelling, insect or root-eating, bandicoots⁠—and those of some other Australian marsupials⁠—should all be constructed on the same extraordinary type, namely with the bones of the second and third digits extremely slender and enveloped within the same skin, so that they appear like a single toe furnished with two claws. Notwithstanding this similarity of pattern, it is obvious that the hind feet of these several animals are used for as widely different purposes as it is possible to conceive. The case is rendered all the more striking by the American opossums, which follow nearly the same habits of life as some of their Australian relatives, having feet constructed on the ordinary plan. Professor Flower, from whom these statements are taken, remarks in conclusion: “We may call this conformity to type, without getting much nearer to an explanation of the phenomenon;” and he then adds “but is it not powerfully suggestive of true relationship, of inheritance from a common ancestor?”

Geoffroy St. Hilaire has strongly insisted on the high importance of relative position or connection in homologous parts; they may differ to almost any extent in form and size, and yet remain connected together in the same invariable order. We never find, for instance, the bones of the arm and forearm, or of the thigh and leg, transposed. Hence the same names can be given to the homologous bones in widely different animals. We see the same great law in the construction of the mouths of insects: what can be more different than the immensely long spiral proboscis of a sphinx-moth, the curious folded one of a bee or bug, and the great jaws of a beetle? Yet all these organs, serving for such widely different purposes, are formed by infinitely numerous modifications of an upper lip, mandibles, and two pairs of maxillae. The same law governs the construction of the mouths and limbs of crustaceans. So it is with the flowers of plants.

Nothing can be more hopeless than to attempt to explain this similarity of pattern in members of the same class, by utility or by the doctrine of final causes. The hopelessness of the attempt has been expressly admitted by Owen in his most interesting work on the Nature of Limbs. On the ordinary view of the independent creation of each being, we can only say that so it is; that it has pleased the Creator to construct all the animals and plants in each great class on a uniform plan; but this is not a scientific explanation.

The explanation is to a large extent simple, on the theory of the selection of successive slight modifications, each being profitable in some way to the modified form, but often affecting by correlation other parts of the organisation. In changes of this nature, there will be little or no tendency to alter the original pattern, or to transpose the parts. The bones of a limb might be shortened and flattened to any extent, becoming at the same time enveloped in thick membrane, so as to serve as a fin; or a webbed hand might have all its bones, or certain bones, lengthened to any extent, with the membrane connecting them increased, so as to serve as a wing; yet all these modifications would not tend to alter the framework of the bones or the relative connection of the parts. If we suppose that an early progenitor⁠—the archetype, as it may be called⁠—of all mammals, birds and reptiles, had its limbs constructed on the existing general pattern, for whatever purpose they served, we can at once perceive the plain signification of the homologous construction of the limbs throughout the class. So with the mouths of insects, we have only to suppose that their common progenitor had an upper lip, mandibles, and two pairs of maxillae, these parts being perhaps very simple in form; and then natural selection will account for the infinite diversity in structure and function of the mouths of insects. Nevertheless, it is conceivable that the general pattern of an organ might become so much obscured as to be finally lost, by the reduction and ultimately by the complete abortion of certain parts, by the fusion of other parts, and by the doubling or multiplication of others, variations which we know to be within the limits of possibility. In the paddles of the gigantic extinct sea-lizards, and in the mouths of certain suctorial crustaceans, the general pattern seems thus to have become partially obscured.

There is another and equally curious branch of our subject; namely, serial homologies, or the comparison of the different parts or organs in the same individual, and not of the same parts or organs in different members of the same class. Most physiologists believe that the bones of the skull are homologous⁠—that is, correspond in number and in relative connection⁠—with the elemental parts of a certain number of vertebrae. The anterior and posterior limbs in all the higher vertebrate classes are plainly homologous. So it is with the wonderfully complex jaws and legs of crustaceans. It is familiar to almost everyone, that in a flower the relative position of the sepals, petals, stamens, and pistils, as well as their intimate structure, are intelligible on the view that they consist of metamorphosed leaves, arranged in a spire. In monstrous plants, we often get direct evidence of the possibility of one organ being transformed into another; and we can actually see, during the early or