Darwin and Modern Science [134]
well known as to have become a commonplace, is the phylogeny of the horses, which, contrary to all that would have been expected, ran the greater part of its course in North America. So far as it has yet been traced, the line begins in the lower Eocene with the genus Eohippus, a little creature not much larger than a cat, which has a short neck, relatively short limbs, and in particular, short feet, with four functional digits and a splint-like rudiment in the fore-foot, three functional digits and a rudiment in the hind-foot. The forearm bones (ulna and radius) are complete and separate, as are also the bones of the lower leg (fibula and tibia). The skull has a short face, with the orbit, or eye-socket, incompletely enclosed with bone, and the brain-case is slender and of small capacity. The teeth are short- crowned, the incisors without "mark," or enamel pit, on the cutting edge; the premolars are all smaller and simpler than the molars. The pattern of the upper molars is so entirely different from that seen in the modern horses that, without the intermediate connecting steps, no one would have ventured to derive the later from the earlier plan. This pattern is quadritubercular, with four principal, conical cusps arranged in two transverse pairs, forming a square, and two minute cuspules between each transverse pair, a tooth which is much more pig-like than horse-like. In the lower molars the cusps have already united to form two crescents, one behind the other, forming a pattern which is extremely common in the early representatives of many different families, both of the Perissodactyla and the Artiodactyla. In spite of the manifold differences in all parts of the skeleton between Eohippus and the recent horses, the former has stamped upon it an equine character which is unmistakable, though it can hardly be expressed in words.
Each one of the different Eocene and Oligocene horizons has its characteristic genus of horses, showing a slow, steady progress in a definite direction, all parts of the structure participating in the advance. It is not necessary to follow each of these successive steps of change, but it should be emphasised that the changes are gradual and uninterrupted. The genus Mesohippus, of the middle Oligocene, may be selected as a kind of half-way stage in the long progression. Comparing Mesohippus with Eohippus, we observe that the former is much larger, some species attaining the size of a sheep, and has a relatively longer neck, longer limbs and much more elongate feet, which are tridactyl, and the middle toe is so enlarged that it bears most of the weight, while the lateral digits are very much more slender. The fore-arm bones have begun to co-ossify and the ulna is greatly reduced, while the fibula, though still complete, is hardly more than a thread of bone. The skull has a longer face and a nearly enclosed orbit, and the brain-case is fuller and more capacious, the internal cast of which shows that the brain was richly convoluted. The teeth are still very short-crowned, but the upper incisors plainly show the beginning of the "mark"; the premolars have assumed the molar form, and the upper molars, though plainly derived from those of Eohippus, have made a long stride toward the horse pattern, in that the separate cusps have united to form a continuous outer wall and two transverse crests.
In the lower Miocene the interesting genus Desmatippus shows a further advance in the development of the teeth, which are beginning to assume the long-crowned shape, delaying the formation of roots; a thin layer of cement covers the crowns, and the transverse crests of the upper grinding teeth display an incipient degree of their modern complexity. This tooth-pattern is strictly intermediate between the recent type and the ancient type seen in Mesohippus and its predecessors. The upper Miocene genera, Protohippus and Hipparion are, to all intents and purposes, modern in character, but their smaller size, tridactyl feet and somewhat shorter-crowned teeth are reminiscences of their ancestry.
Each one of the different Eocene and Oligocene horizons has its characteristic genus of horses, showing a slow, steady progress in a definite direction, all parts of the structure participating in the advance. It is not necessary to follow each of these successive steps of change, but it should be emphasised that the changes are gradual and uninterrupted. The genus Mesohippus, of the middle Oligocene, may be selected as a kind of half-way stage in the long progression. Comparing Mesohippus with Eohippus, we observe that the former is much larger, some species attaining the size of a sheep, and has a relatively longer neck, longer limbs and much more elongate feet, which are tridactyl, and the middle toe is so enlarged that it bears most of the weight, while the lateral digits are very much more slender. The fore-arm bones have begun to co-ossify and the ulna is greatly reduced, while the fibula, though still complete, is hardly more than a thread of bone. The skull has a longer face and a nearly enclosed orbit, and the brain-case is fuller and more capacious, the internal cast of which shows that the brain was richly convoluted. The teeth are still very short-crowned, but the upper incisors plainly show the beginning of the "mark"; the premolars have assumed the molar form, and the upper molars, though plainly derived from those of Eohippus, have made a long stride toward the horse pattern, in that the separate cusps have united to form a continuous outer wall and two transverse crests.
In the lower Miocene the interesting genus Desmatippus shows a further advance in the development of the teeth, which are beginning to assume the long-crowned shape, delaying the formation of roots; a thin layer of cement covers the crowns, and the transverse crests of the upper grinding teeth display an incipient degree of their modern complexity. This tooth-pattern is strictly intermediate between the recent type and the ancient type seen in Mesohippus and its predecessors. The upper Miocene genera, Protohippus and Hipparion are, to all intents and purposes, modern in character, but their smaller size, tridactyl feet and somewhat shorter-crowned teeth are reminiscences of their ancestry.