It is not yet possible to trace this phylum below the level of the uppermost White River beds, yet that will very probably be accomplished by future exploration.

The second phylum of the family was represented in the lowest Miocene by †Hypertragulus, a genus of much smaller animals than those of the preceding series, which went back to White River times without essential change, and was abundant in the John Day stage. Despite this fact, the structure of the genus is still incompletely known and much remains to be learned, but enough has already been ascertained to justify the association of this phylum with the †Protoceras-†Syndyoceras series in one family as reasonable. The number of upper incisors in †Hypertragulus has not been ascertained, but the canines were enlarged and tusk-like, the lower one not having gone over to the incisors, as it had in the preceding group. The skull had much resemblance to that of the contemporary camels, the sudden narrowing of the facial region giving it a very llama-like appearance; the orbit was open and on the face in front of it was a conspicuous vacuity. The ulna and radius were coössified and there were four digits in the manus, two in the pes, but no cannon-bone was formed.

The third phylum, that of †Leptomeryx, had about the same range in time as the preceding one, though it has not yet been found in the John Day, and the genus is assuredly known only from the White River beds, in which it is not uncommon. †Leptomeryx comprised a number of species, all very small animals, and none larger than a jack-rabbit. (See Fig. 277, p. 563.) In size, proportions and appearance, these dainty little creatures must have been very like the existing chevrotains or “mouse-deer” of Asia and the Malay islands, and by many writers they have been classed in the same suborder, the Tragulina. The upper incisors had been suppressed and the upper canine reduced to very small size, while the lower canine had become functionally one of the incisors. The skull had a very long and slender facial region, but had a less llama-like appearance than in †Hypertragulus. The neck was short and the fore limbs much shorter than the hind, so that the back sloped downward from the rump to the shoulders, as in the chevrotains. There was a remarkable, indeed quite unparalleled, difference between the fore and hind limbs and feet, the hinder extremity being not only much longer, but also much more specialized, while the anterior one retained in very large degree its primitive characteristics. Thus, in the fore-arm the ulna was complete and separate from the radius, but in the lower leg the fibula was reduced to its minimum. In the manus there were four entire and functional digits, in the pes only two, which were joined in a cannon-bone.

Finally, there was a fourth phylum, that of †Hypisodus, which was confined to the White River stage and is still incompletely known. This was a tiny creature, much smaller than any of the preceding ones, and is the only known White River ungulate with fully hypsodont grinding teeth. Another very exceptional peculiarity of its dentition was that in the lower jaw it had ten incisor-like teeth; not only the canine, but the first premolar as well, had assumed the character of the incisors. This same peculiarity is found in the lower Miocene †gazelle-camel, †Stenomylus (see p. 394), but in no other mammal.

A considerable assemblage of genera belonging to this family occurs in the upper Eocene, but the material yet obtained is too fragmentary to permit the assignment of these forms to the different phyla, though it is very probable that among them are to be found ancestors of all the White River and subsequent genera.

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.

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.

8. Cervidæ. Deer

In most of the deer now existing the male has antlers. The antler is a bony outgrowth from the frontal bone of the skull and is annually shed and replaced, increasing, as a rule, in size and in the number of branches with each renewal. During the period of growth the antler is richly supplied with blood-vessels and covered with skin and is then said to be “in the velvet,” which dries and peels off when growth is complete; after the rutting season a layer of bone at the base of the antler is resorbed, loosening the antler, which is then shed. There is, however, a permanent, cylindrical process, of greater or less length, from each frontal, the “pedicle,” from which the antler is annually reproduced, and around the base of the antler and shed with it is a roughened ring, the “burr.” Among the different genera of deer there is great variety in the form and size of the antler, from a single spike to the immense and complicated appendages of the Wapiti (Cervus canadensis). As a rule, the “beam” or main stem of the antler and its branches or “tines” are cylindrical and tapering; but in some cases, as in the Moose (Alce) and the Fallow Deer (Dama), the antler is very broad and flat and is then said to be “palmated.” Except in the Reindeer and Caribou (Rangifer) the female is without antlers.

In the skeleton there is little difference between the deer and the Cavicornia, but there are some differences in the teeth. In the males of those deer which have no antlers, such as the Musk-Deer (Moschus moschiferus) and the Chinese Water-Deer (Hydropotes inermis), as well as in certain forms with very small antlers, like the muntjacs of Asia (Cervulus and Elaphodus), the upper canine is a long, thin, recurved and sabre-like tusk, a very effective weapon. Speaking of the Indian Muntjac or “Barking Deer” (Cervulus muntjac), Flower and Lydekker say, “When attacked by dogs the males use their sharp canine teeth with great vigour, inflicting upon their opponents deep and even dangerous wounds.” In other forms of deer the upper canines are small or absent. The grinding teeth are brachyodont, but in the existing genera they have higher crowns than in the Tertiary progenitors of the family, and in the Axis and Hog Deer of India (Axis axis and A. porcinus) the molars are quite hypsodont.

As was shown in Chapter V, the existing deer of North America are of two kinds: (1) the northern, which are plainly of Old World origin and so closely similar to Old World species that many naturalists deny the necessity of making distinct species for the American forms. The best known of these are the Wapiti (Cervus canadensis), the Caribou (Rangifer caribou) and the Moose (Alce americanus). (2) The southern deer, of which the common Virginia Deer (Odocoileus virginianus) is a familiar example, though overlapping in their range that of the northern genera, are peculiar to the Americas, and, though not exactly autochthonous, they must have had a long American ancestry. In the Pleistocene we find the same genera and mostly the same species, their distribution over the continent shifting in accordance with the many climatic changes of that epoch. There was, however, at least one Pleistocene genus (†Cervalces) different from any now living and different from any known in the eastern hemisphere. The most complete specimen of this animal is a skeleton in the museum of Princeton University, found beneath a bog in northern New Jersey, though other bones, collected in Kentucky and elsewhere, are very probably referable to it. †Cervalces was very nearly related to the Moose, the neck, body, limbs and feet being almost identical in the two genera, but the skull and antlers were notably different; the nasal bones were not nearly so much shortened as in the Moose, indicating that the proboscis-like snout was not so large or inflated as in the latter. The antlers were quite unique; though in general like those of the Moose, they were much less palmated and they had, in addition, a great trumpet-like plate of bone on the lower side of each antler (see Fig. 117, p. 209), such as occurs in no other known member of the family. Although †Cervalces has not been found in the Old World, it was almost certainly an immigrant from eastern Asia.

The Moose, Caribou and Wapiti were unquestionably immigrants and came in not earlier than the Pleistocene. Nothing is known in the Pliocene or more ancient Tertiary epochs of North America which could be twisted into forms ancestral to these typically Old World genera. With the southern deer (Odocoileus, etc.) the matter stands differently, for these have a probable American ancestry extending back to the lower Miocene and possibly much farther. On the other hand, it is not altogether certain that these may not have been Pliocene immigrants, for their genealogy is still in an extremely fragmentary and unsatisfactory condition. The North American genus, Odocoileus, extended back to the Pliocene with very little change. The annoying, unrecorded gap of the upper Pliocene and the meagre representation of the middle Pliocene mammals given by the Blanco leave us without information regarding the deer of that time. In the lower Pliocene and through the whole Miocene we meet with frequent remains of a genus (†Blastomeryx) which was quite probably the ancestor of the American types of deer. It was considerably smaller than any of the existing North American species and had no antlers, but possessed the sabre-like, upper canine tusks, which characterize the muntjacs and hornless deer of Asia. The limb-bones had already attained nearly their present state of development, as regards the reduction of ulna and radius, formation of cannon-bones, etc. †Blastomeryx probably entered North America in the lower Miocene, but, as was mentioned previously (p. 409), Dr. Matthew and Professor Osborn regard the genus as autochthonous and descended from the †Hypertragulidæ.

In the middle Miocene †Blastomeryx gave rise to an aberrant genus (†Merycodus) which has been made the type of a distinct family (†Merycodontidæ, see table, p. 362), but this is perhaps unnecessary. †Merycodus had deer-like antlers, but completely hypsodont teeth such as no known member of the Cervidæ possesses. The middle Miocene species (†M. osborni) was a little creature, not more than eighteen or twenty inches high at the shoulder, and had a branched antler of three tines, which was considerably longer than the skull, while in the species of the upper Miocene (†M. furcatus) the antler was shorter and simply forked. From the number of specimens of these animals found in which the burr is incomplete or absent, it may be inferred that the antler was not always deciduous. The legs were long and very slender, and apparently there was no trace of the lateral digits, even in the fore foot. These peculiar hypsodont deer persisted even in the older Pleistocene.

Deer are the only members of the Pecora which inhabit South America, where there are several genera of them, all much more nearly allied to North American than to Old World forms. No record of the presence of the family in the southern continent has been found in beds older than the Pleistocene, but in view of the degree of specialization which they have there undergone, it is probable that the immigration took place in the Pliocene.

SECTION CAVICORNIA. HOLLOW-HORNED RUMINANTS

In the animals of this second and far larger section of the Pecora there are bony outgrowths of the skull, from the frontal bones, outgrowths which are permanent and non-deciduous; these are the horn-cores, which are tapering and unbranched. The horn-core is, in turn, covered with a sheath of horn, likewise unbranched and permanent, but growing from year to year until the maximum size is attained, a process which is familiarly illustrated in the growth of a calf. Among Recent Cavicornia there is but one exception to the rule that the horny sheath is non-deciduous and unbranched and that one is the Prong Buck (Antilocapra americana). In the Cavicornia it is the very general rule that both sexes are horned, though the females commonly have smaller horns and in several genera of antelopes the does are hornless. There is almost as great variety in the shape and size of the horn as of the antler; we find small, medium-sized and enormously large horns, which may be straight, simply curved, complexly curved, spiral, lyrate or twisted. The antelopes have many types of horns, as have the sheep and goats, the oxen, buffaloes and bisons; but only a few of them are exemplified in the western hemisphere, which now, as in the preceding geological periods, is singularly poor in representatives of the Pecora.

9, 10. Antilopidæ and Antilocapridæ. Antelopes

Two very different kinds of antelopes are found in North America at the present time; one of them, the erroneously named Rocky Mountain Goat (Oreamnos montanus), is evidently a late immigrant from the Old World, and fossil remains of it have been found in the Pleistocene cave-deposits of California. This animal is a member of the true antelope family (Antilopidæ) and belongs to the chamois group of mountain-antelopes; it has no near relatives among other American mammals, living or extinct.

The Prong Buck, or Prong-horned Antelope (Antilocapra americana), occupies a very isolated position, so much so that a distinct family, the Antilocapridæ, has been created for its reception. It differs from all other Cavicornia in having a branched horn, though the bony core is simple, and in annually shedding and renewing the horny sheath; the horn is directly over the eye; there are no dew-claws and all traces of the bones of the lateral digits have completely disappeared. The grinding teeth are thoroughly hypsodont. The genus occurred in the older Pleistocene, where it was associated with the last of the †deer-antelope, or †Merycodus series (†Capromeryx), and which, so far as it is known, would seem to connect the two families, though this is doubtful. A middle Miocene genus (†Dromomeryx Fig. 128, p. 237) would be a more probable ancestor of the Prong Buck, if it were not for the long, unfilled gap of the upper Miocene and the whole Pliocene. †Dromomeryx had erect horn-cores placed directly above the eyes as in the modern genus, but low-crowned grinding teeth; it was the most ancient American cavicorn yet known. It remains to be determined by future exploration, whether this middle Miocene genus was actually the ancestor of Antilocapra, or merely an anticipation of it.

In the lower Pliocene have been found the remains, very incomplete, of several antelopes, which appear to have been immigrants from the Old World, but are too imperfectly known for any definitive reference. One resembles the flat-horned, or goat-horned, antelopes of the European Miocene and Pliocene. Others had spirally twisted horns like those of the Recent strepsicerine, or twisted-horn antelopes of Africa and Asia, but may, nevertheless, be referable to the Antilocapridæ.

Antelopes even penetrated to South America, and three genera of them have been reported from the Pleistocene of the Brazilian caverns and the Argentine pampas, but they were less successful in establishing a foothold than were the deer, and form no part of the modern Neotropical fauna.

11. Bovidæ. Sheep, Bisons, Oxen, etc.

A series of genera, of disputed systematic position, is represented to-day by the so-called Musk-Ox (Ovibos moschatus), which is now exclusively North American, but in the Pleistocene ranged over northern Asia and Europe as far west as Great Britain. The Musk-Ox, which is at present found only in the extreme north, is a heavy, short-legged animal, three and a half to four feet high, and six feet or more in length; the body is covered with a dense coat of woolly hair overlaid by a thatch of long, straight hair, which gives the animal a very shaggy appearance. The horns are broad at the base, especially so in old males, in which they meet in the middle line and cover much of the head as with a horny casque; they curve downward and then upward and forward, with the tips directed toward the front; in the females and young males the horns are very much smaller.

This series cannot be traced back of the Pleistocene, in which epoch it was not only far more widely distributed, but also very much more diversified, no less than three extinct genera, in addition to the existing one, having been found in the North American Pleistocene. One of these (†Symbos), which extended from Alaska to Arkansas, had horns which were smaller and shorter than in the modern genus, and, even when fully developed, did not meet in the middle line of the head. The other two genera, from California (†Euceratherium and †Preptoceras Fig. 116, p. 203), are of great interest as showing affinities to the Musk-Ox and also to sheep and to certain antelopes, such as the Takin (Budorcas) of northern India and Tibet. They serve to connect the musk-oxen with other Cavicornia, but the origin of all these animals is to be sought in Asia.

In Recent North America there are four or five species of sheep (Ovis) which are confined to the mountainous and broken areas of the western part of the continent and extend from Alaska to Mexico. The “Bighorn” or Rocky Mountain Sheep (Ovis canadensis) is characterized by great, spirally coiled horns in the rams, in the ewes the horns are very much smaller and nearly straight; the other species differ but slightly from this type. The species O. canadensis has been found in the Pleistocene, but nothing further is known of its history. Evidently, the sheep were late immigrants.

“The geographical distribution of wild sheep is interesting. The immense mountain ranges of Central Asia, the Pamir and Thian Shan of Turkestan, may be looked on as the centre of their habitat.” “Sheep are essentially inhabitants of the high mountainous parts of the world, for dwelling among which their wonderful powers of climbing and leaping give them special advantages. No species frequent by choice either level deserts, open plains, dense forests or swamps. By far the greater number of species are inhabitants of the continent of Asia, one extending into North America [should read, four or five] one into Southern Europe and one into North Africa.... No remains that can be with certainty referred to the genus [Ovis] have been met with in the hitherto explored true Tertiary beds, which have yielded such abundant modifications of Antelopes and Deer.”[10]

The only other division of the family which is represented in North America is that of the bisons, of which the fast vanishing remnant of a single species[11] (Bison bison) is all that is left of what was once an extensive and varied assemblage. The bisons differ from the true oxen in the form and structure of the skull, in the shoulder-hump, which is produced by the very long spines of the dorsal vertebræ and in consequence of which the back slopes downward from the shoulders to the croup. They differ further in the character of the hair, which is short and woolly on the body and hind quarters, very long and shaggy on the head and neck. In the Pleistocene of North America there were at least seven recognizable species of bisons, which ranged over the continent from Alaska to Florida, though it is not probable that they were all contemporary. One of the earliest and by far the largest of these was the gigantic B. †latifrons, a specimen of which in the American Museum of Natural History measures six feet across the horns in a straight line; this was a Mississippi Valley species and extended from Ohio to the Gulf of Mexico and westward to Kansas and Texas. Another gigantic species (B. †crassicornis) lived in Alaska in association with a second and smaller species (B. †occidentalis) which ranged as far south as Kansas. B. †occidentalis, though smaller than the preceding species, was larger than the existing one and was remarkable for the great size of the hump. The bisons were migrants from the Old World and are the only members of the great ox-tribe that ever reached America. At present the Old World has but a single species of Bison (B. bonasus), which has been saved from extermination only by the most rigid protection.

Neither sheep nor bison extended their range to South America; both are and have been essentially northern groups and seem to have been unable to cross the tropics.

From the foregoing account, confused as it unavoidably is, one thing at least stands out clearly, that North America played a very insignificant rôle in the evolution of the Pecora, and has only two peculiar groups, the Prong Buck and the American types of deer, and of these, the probable American ancestry does not extend back of the lower Miocene and perhaps not so far. Even in the Old World the story, so far as it has been deciphered, is by no means clear and consistent, which is no doubt due to the fact that the regions from which Tertiary mammals have been obtained are so small in comparison with those that have yielded nothing. Certain broad outlines of the history may, nevertheless, be discerned.

The suborder Pecora at an early date became divided into the two great branches of the Cervicornia and Cavicornia, the former giving off the giraffe series, which in the Miocene and Pliocene ramified and extended through Asia and southern Europe, though now confined to Africa. In the lower Miocene of Europe the muntjac-like deer and the antelopes, the first of the Cavicornia, were already well distinguished. From the primitive antelopes arose not only the wonderful assemblage of modern antelopes, but also the goats and sheep and the great and varied ox-tribe. From the middle Oligocene forms it may obviously be inferred that both Cervicornia and Cavicornia united in a single trunk, or, traced in the other direction, diverged from a common stock, to which also the suborder of the Tragulina goes back.

On the other hand, it is equally obvious that the camels and llamas have been separated from the Pecora at least since the middle Eocene, and, consequently, the many points of agreement between the two suborders, other than those shared with all artiodactyls, are not due to inheritance from a common ancestry, but have been independently acquired in the two series. It will be instructive to note some of the more important of these independent similarities: (1) the selenodont and more or less hypsodont character of the grinding teeth; (2) the spout-shaped odontoid process of the axis; (3) the great reduction of the ulna and its coössification with the radius; (4) the loss of the fibula, except for its lower end, which persists as a separate malleolar bone; (5) the formation of cannon-bones by the fusion of the third and fourth metapodials; (6) the development of a complex, many-chambered stomach. Other points of likeness might be cited, but those already given will suffice to show how very important this parallel mode of evolution often proves itself to be.