That a land-connection with the Old World existed during the Paleocene epoch, is indicated by the similarity of the faunas of North America and Europe.
CHAPTER VIII
HISTORY OF THE PERISSODACTYLA
In attempting to trace the evolutionary history of the various mammalian groups, it is necessary to bear in mind the inevitable limitations of work of this kind. Speaking of plants, Dr. D. H. Scott says: “Our ideas of the course of descent must of necessity be diagrammatic; the process, as it actually went on, during ages of inconceivable duration, was doubtless infinitely too complex for the mind to grasp, even were the whole evidence lying open before us. We see an illustration, on a small scale, of the complexity of the problem, in the case of domesticated forms, evolved under the influence of man. Though we know that our cultivated plants, for instance, have been developed from wild species within the human period, and often within quite recent years, yet nothing is more difficult than to trace, in any given instance, the true history of a field-crop or garden plant, or even, in many cases, to fix its origin with certainty.”[5] With some mammalian groups the task, though difficult enough, is not so hopeless, because of more complete records, yet in dealing with mammals a very troublesome complication is introduced by the existence within the families, and even within the genera, of two or more parallel phyla, or genetic series. Without complete and perfect material it is impossible to make sure that we are not confusing the different phyla with one another and placing in one series species and genera that properly belong in a different one. Thus, Osborn distinguishes no less than seven such phyla among the true rhinoceroses of the Old and New Worlds, which long followed parallel, but quite independent, courses of development, and five phyla among the American horses. While these phyla add so much to the difficulty of working out the genealogical series, it is possible to simplify the problem and treat it in a broad and comprehensive manner that will sufficiently establish the essential steps of change.
Fig. 145.—Left manus of Tapir (Tapirus terrestris). S., scaphoid. L., lunar. Py., pyramidal. Pis., pisiform. Td., trapezoid. M., magnum. Un., unciform. The metacarpals are erroneously numbered. Mc. I., second metacarpal. Mc. II., third do. Mc. III., fourth do. Mc. IV., fifth do. Ph. 1, first phalanx. Ph. 2, second do. Ung., ungual phalanx.
In external appearance and general proportions the different families of existing perissodactyls have very little in common; that tapirs and rhinoceroses should be related is not surprising, but the horses would seem to be as far removed from both of the former as possible. Why, then, should they be included in the same order? A study of the skeleton, however, reveals the community of structure which obtains between the three families, a community which removes them widely from all other hoofed mammals. In all existing perissodactyls, though not in most of the Eocene genera, all the premolars, except the first, have the size and pattern of the molars. The foramina of the skull, or perforations by which blood-vessels and nerves enter and leave the cranium, are arranged in a way characteristic of the order and different from that seen in other hoofed mammals. The femur always has the third trochanter. The number of digits in each foot is usually odd, 1, 3 or 5, but four-toed forms occur, as the tapirs, which have four toes in the front foot, three in the hind; the important character is that the median plane of the foot bisects the third digit, which is symmetrical. The third and fourth, each asymmetrical, together form a symmetrical pair. Especially characteristic is the form of the astragalus and calcaneum (ankle and heel bones); the astragalus has but a single, deeply grooved and pulley-like surface, that for the tibia, the lower end is nearly flat and rests almost entirely upon the navicular, covering but little of the cuboid (see Figs. [146], [148]). The calcaneum does not articulate with the fibula and its lower end is broad and covers most of the cuboid.
While the foregoing list includes the most important of the structural features which are common to all perissodactyls and differentiate them from other hoofed animals, there are many others which it is needless to enumerate.
Fig. 146.—Left pes of Tapir. Cal., calcaneum. As., astragalus. N., navicular. Cn. 1, Cn. 2, Cn. 3, first, second and third cuneiforms. Mr. II, III, IV, second, third and fourth metatarsals.