While there is little difference of opinion as to the propriety of including in the family †Hypertragulidæ the four phyla described in the foregoing pages, the systematic position and the relationships of that family as a whole are matters of debate and likely long to remain so. Dr. Matthew refers the entire group to the suborder Tragulina and regards †Leptomeryx as being closely related to the direct ancestry of the American deer, a view which is accepted by Professor Osborn, but in which I am unable to concur. My own belief is that the family was an early offshoot from the cameline stock and therefore referable to the Tylopoda, in which suborder they are here included. It would be out of place to enter upon a discussion of this perplexing problem, which can hardly receive a definitive solution until the artiodactyls of the Uinta stage are thoroughly understood. As in so many other series, the key of the mystery lies hidden in the Uinta fauna, which is still so inadequately known.

Suborder Pecora. True Ruminants

This is the most advanced, specialized and diversified group of the artiodactyls, though the ruminating habit is shared by both Tylopoda and Tragulina. In this multitude of forms, giraffes, deer, antelopes, sheep, goats, oxen, buffaloes, bisons, etc., it is difficult to find a clue to a natural arrangement or classification. As a whole, the suborder is a well-defined group, and many structural characters, not all of which is it needful to enumerate here, are common to all of its members. The upper incisors are invariably absent, and, save in a few of the deer, the upper canine also, while the lower canine has become incisiform; the premolars are always three in number in each jaw and the molar-pattern is selenodont throughout. The odontoid process of the axis is spout-shaped. Except in a few deer, the Pecora all have bony outgrowths of the skull in the form of antlers or horns, at least in the males, many females being hornless. The ulna is coössified with the radius and the fibula is lost, except the lower end, which is a separate malleolar bone. There is always, in both fore and hind feet, a cannon-bone, the lower ends of which are parallel, not divergent, as they are in the Tylopoda, and each articular surface is encircled all around by a prominent median keel, as in the horses, which in the other suborders, as in mammals generally, is confined to the posterior side and not visible from the front. (Cf. Figs. [220] and [214, p. 401].) In no existing member of the Pecora are there complete lateral digits, and in several modern genera they have been completely suppressed; but in most there is, behind the functional pair of digits, a pair of “dew-claws,” the bones of which are more or less completely reduced, often to mere nodules. The stomach, which in the Tylopoda and Tragulina is three-chambered, is in the Pecora divided into four distinct parts.

Fig. 219.—Left manus of Patagonian Deer (Hippocamelus bisulcus). S., scaphoid. L., lunar. Py., pyramidal. Td., M., coössified trapezoid and magnum. Un., unciform. Mc. II and V, rudimentary second and fifth metacarpals. Mc. III and IV, cannon-bone. Ph. 1, 2, first and second phalanges. Ung., ungual phalanx.

Fig. 220.—Left pes of Patagonian Deer. Cal., calcaneum. As., astragalus. N., Cb., coössified navicular and cuboid. Mt. III, IV, cannon-bone. Other letters as in [Fig. 219].

As already intimated, the subdivision of the Pecora into smaller groups is far from easy. “The great difficulty which all zoölogists have felt in subdividing them into natural minor groups arises from the fact that the changes in different organs (feet, skull, frontal appendages, teeth, cutaneous glands, etc.) have proceeded with such apparent irregularity and absence of correlation that the different modifications of these parts are most variously combined in different members of the group.”[9] Two main sections of the suborder are, however, sufficiently well defined, (1) the Cervicornia and (2) the Cavicornia.

SECTION CERVICORNIA. DEER AND GIRAFFES

This section includes two families, the giraffes and the deer. Inasmuch as the former have not now and never did have any representatives in the western hemisphere, for the purposes of this book the section becomes identical with the deer family.