American Big Game in Its Haunts Part 2

In the opinion of many palaeontologists, the ancestors of the present hoofed beasts, or ungulates, were contained among these _Condylarthra_, as they were named by Prof. Cope.

Of course, these early mammals are known to us only by their fossil and mostly fragmentary skeletons, but it may be said that at least in the ungulate line, the successive geological periods show steady structural progression in certain directions. Of great importance are a decrease in the number of functional digits; a gradual elevation of the heel, so that their modern descendants walk on the tips of their toes, instead of on the whole sole; a constant tendency to the development of deeply grooved and interlocked joints in place of shallow bearing surfaces; and to a complex pattern of the molar crowns instead of the simple type mentioned. To this may be added as the most important factor of all in survival, that these changes have progressed together with an increase in the size of the brain and in the convolutions of its outer layer.

The _Condylarthra_ seem to have gone out of existence before the time of the middle Eocene, but before this they had become separated into the two great divisions of odd-toed and even-toed ungulates, into which all truly hoofed beasts now living fall.

The first group (_Perissodactyla_) has always one or three toes functionally developed, either the third, or third, second and fourth, the two others having entirely disappeared, except for a remnant of the fifth in the forefoot of tapirs. They have retained some at least of the upper incisor teeth, and, except in some rhinoceroses, the canines are also left; the molars and premolars are practically alike in all recent species, and in all of which we know the soft parts, the stomach has but one compartment, and there is an enormous caec.u.m. It is probable that they took rise earlier than their split-footed relations, and their Tertiary remains are far more numerous, but their tendency is toward disappearance, and among existing mammals they are represented only by horses, a.s.ses, rhinoceroses, and tapirs.

Contrasted with these, _Artiodactyla_ have always an even number of functional digits, the third and fourth reaching the ground symmetrically, bearing the weight and forming the "split hoof;" the second and fifth remain, in most cases, as mere vestiges, showing externally as the accessory hoofs or dewclaws; in the hippopotamus alone they are fully developed and the animal has a four-toed foot. In deer and bovine animals the incisors and frequently the canines have disappeared from the upper jaw, and the molars are unlike the premolars in having two lobes instead of one. The stomach is always more or less complex; at its extreme reaching the ruminant type with four compartments, in a.s.sociation with which is a caec.u.m reduced in size and simple in form. Nearly all have horns or antlers, at least in one s.e.x.

Most split-hoofed animals are ruminants, but there is a small remnant, probably of early types, which are not. The present ungulates may be summed up in this way:

Odd-toed: _(Perissodactyla)_-- Horse, a.s.s, Rhinoceros, Tapir.

Even-toed: _(Artiodactyla)_--

Non-ruminants-- Hippopotamus, Swine, Peccaries.

Ruminants-- Camels, Llamas, Chevrotains, Giraffe, Antelopes, Sheep, Goats, Musk-ox, Oxen, Deer.

The non-ruminant artiodactyls need not detain us long. Hippopotamuses are little more than large pigs with four toes; they were never American, though many species, some very small, are found in the European Tertiary. The two existing species are African.

In the western hemisphere swine are represented by the peccaries, differing from them chiefly in having six less teeth, one less accessory toe on the hind foot, and in a stomach of more complex character.

Peccaries also have the metapodial bones supporting the two functional digits fused together at their upper ends, forming an imperfect "cannon bone," which is a characteristic of practically all the ruminants, but of no other hoofed beasts. One species only enters the United States along the Mexican border.

All non-ruminant ungulates have from four to six incisors in the upper jaw; the canines are present, and sometimes, as in the wart hogs, reach an extraordinary size.

Coming now to the ruminants, all digits except the third and fourth have disappeared from camels and llamas, and the nails on these are limited to their upper surface without forming a hoof, the under side being a broad pad, upon which they tread. No camel-like beasts have inhabited North America since the Pliocene age. Chevrotains, or muis deer (_Tragulidae_), are not deer in any true sense, as they have but three compartments to the stomach; antlers are absent and in their place large and protruding canine teeth are developed in the upper jaw, and the lateral metacarpal bones are complete throughout their length, instead of being represented by a mere remnant. They are the smallest of ungulates, and inhabit only portions of the Indo-Malayan region. Camels also have upper canines, and the outer, upper incisors as well.

The giraffe is separated from all living ungulates by the primitive character of its so-called "horns," which are not horns in the usual sense, but simply bony prominences of the skull covered with hair. Some of the earliest deer-like animals seem to have had simple or slightly branched antlers which were not shed, and which there is reason to believe were also hairy, and in these, as well as in other characters, giraffes and the early deer may not have been far apart. The "okapi,"

Sir Harry Johnston's late discovery in the Uganda forests, seems to have come from the same ancestral stock, but the giraffe has no other existing relatives.

The true deer, to which we shall return, are readily enough distinguished from the ox tribe and its allies by their solid and more or less branched antlers, usually confined to males, and periodically shed.

So, through this rapid survey, we have dropped out of the hoofed beasts all but the bovines and their near allies, and are thus far advanced toward our definition of a bison, but from this point we shall not find it easy to draw sharp distinctions, for while the _Bovidae_, as a whole, are well enough distinguished from all other animals, their characteristics are so much mixed among themselves that it is hardly possible to find any one or more striking features peculiar to one group, and for most of them recourse must be had to a.s.sociations of a number of lesser characters.

Oxen, antelopes, sheep and goats agree in having hollow horns of material similar to that of which hair and nails are formed, permanently fixed upon the skull in all but one species; none of them have more than the two middle digits functionally developed, one on each side of the axis of the leg; none have the lower ends remaining of the meta-podial bones belonging to the two accessory digits; and none have either incisor or canine teeth in the upper jaw.

From animals so constructed we may first take out goats and sheep, in which the female horns are much smaller than those of males, and in some species are even absent. In nearly all of them the horns are noticeably compressed in section, either triangular or sub-triangular near the base, and are directed sometimes outwardly from the head with a circular sweep; at others with a backward curve, often spirally. The muzzle is always hairy; there is no small accessory column on the inner side of the upper molars, found always in oxen and in some antelopes; the tail is short, and scent glands are present between the digits of some or all the feet.

Now, as to the perplexing animals popularly known as antelopes. No definition could be framed which would include them all in one group, for every subordinate character seems to be present in some and absent in others, so that the most that can be done with this vast a.s.semblage is to arrange its contents in series of genera, which may or may not be called sub-families, but which probably correspond in some degree to their real affinities. We can only say of any one of them that it is an antelope because it is not a sheep, nor a goat, nor an ox. They concern us here only to be eliminated, for they are not American, our p.r.o.ng-buck having a sub-family all to itself, as we shall see later, and the so-called "white goat" being usually regarded as neither goat nor truly antelope.

Within the limits of the real bovine animals, four quite distinct types may be made out, chiefly by the position of the horns upon the skull and by the shape of the horns themselves. There are also differences in the relations of the nasal and premaxillary bones, the development of the neural spines of the vertebrae, and the hairy covering of the body.

In the genus _Bos_ the horns are placed high up on the vertex of the skull, which forms a marked transverse ridge from which the hinder portion falls sharply away. The horns are nearly circular in section and almost smooth; usually they curve outward, then upward and often inward at the tip; the premaxillaries are long and generally reach to the nasals, and the anterior dorsal vertebrae are without sharply elongated spines, so that the line of the back is nearly straight. These, the true oxen, as they are sometimes termed, now exist only in domesticated breeds of cattle.

In the gaur oxen (_Bibos_) the horns are situated as in _Bos_, high up on the vertex, but are more elliptical in section; the premaxillaries are short; the dorsal vertebrae, from the third to the eleventh, bear elongated spines which produce a hump reaching nearly to the middle of the back; the tail is shorter, and the hair is short all over the body. The three species--gaur, gayal and banteng--inhabit Indo-Malayan countries, and all of them are dark brown with white stockings.

The buffaloes (_Bubalus_) are large and clumsy animals with horns more or less compressed or flattened at their bases, set low down on the vertex, which does not show the high transverse ridge of true oxen and gaurs. In old bulls of the African species the horns meet at their base and completely cover the forehead. In the arni of India they are enormously long. The dorsal spines are not much elongated, and there is no distinct hump; the premaxillae are long enough to reach the nasals. Hair is scanty all over the body, and old animals are almost wholly bare. The small and interesting anoa of Celebes, and the tamarao of Mindoro, are nearly related in all important respects to the Indian buffalo, and the carabao, used for draught and burden in the Philippines, belongs to a long domesticated race of the same animal.

Finally, in the genus _Bison_ the horns are below the vertex as in buffaloes, but are set far apart at the base, which is cylindrical; they are short and their curve is forward, upward and inward; the anterior dorsal and the last cervical vertebrae have long spines which bear a distinct hump on the shoulders; the premaxillae are short and never reach the nasals; there are fourteen, or occasionally fifteen, pairs of ribs, all other oxen having but thirteen, and there is a heavy mane about the neck and shoulders. The yak of central Asia is very bison-like in some respects, but in others departs in the direction of oxen.

So at last, group by group, we have gone through the ungulates, and the bisons alone are left, and as the American animal has short, incurved horns, set low down on the skull and far apart at the base; premaxillaries falling short of the nasals; the last cervical and the anterior dorsal vertebrae with spines; fourteen pairs of ribs, and a mane covering the shoulders, we conclude that it is a bison, and as the same characteristics with minor variations are shown by the European species, often, but wrongly, called "aurochs," we say that these two alone of existing _Bovidae_ are bisons, with the yak as a somewhat questionable relative.

In all essential respects the two bisons are very similar, but minute comparison shows that the European species, _Bison bonasus_, has a wider and flatter forehead, bearing longer and more slender horns, and all the other distinctive features are less p.r.o.nounced. In the American species, _Bison bison_, the pelvis is less elevated, producing the characteristic slope of the hindquarters. It is a coincidence that the two regions originally inhabited by the bisons are those in which the white races of men have to the greatest extent thrown their restless energies into the struggle for existence, with the result that extinction to nearly the same degree has overtaken these two near cousins among oxen. A few wild members of the European species still exist in the Caucasus, as a few of the American are left in British America, but elsewhere both exist only under protection.

The carefully kept statistics of the Bielowitza herd in Grodno, western Russia, which includes nearly all but the few wild ones, shows that between 1833 and 1857 they increased in number from 768 to 1,898, but from this maximum the decrease has been constant, with trifling halts, until in 1892 less than five hundred were left; so that even if the Peace River bison are counted with the remnant of the American species, it is probable that the survivors of each race are about equal in number.

It is true that the number of our own species has lately been placed as high as a thousand, but even if these figures are correct, the seeds of decay from internal causes, such as inbreeding and the degeneration of restraint, are already sown, and the inevitable end of the race is not far off.

The Peace River, or woodland, bison has lately been separated as a sub-species _(B. bison athabascae)_, distinguished from the southern and better known form by superior size, a wider forehead, longer, more slender and incurved horns, and by a thicker and softer coat, which is also darker in color. Now, it is an interesting fact that a fossil bison skull from the lower Pliocene of India resembles the present European species, and in later geological times very similar bisons closely allied to each other, if not identical, inhabited all northern regions, including America. These were large animals with wide skulls, and there is little doubt that from this circ.u.mpolar form came both of the bisons now inhabiting Europe and America. Out of some half dozen fossil bison which have been described from America, none earlier than the latest Tertiary, _Bison latifrons_ from the Pleistocene seems likely to have been the immediate ancestor of recent American species, and as the one skull of the woodland bison which has been examined resembles both _latifrons_ and the European species more than the plains species does, it seems probable that these two more nearly represent the primitive bison, of which the former inhabitant of the prairies is a more modified descendant.

The process of elimination has at last led to this outline definition of a bison, but among the ungulates we have pa.s.sed over, there are certain others which concern us because they are American.

Sheep and goats agree together and differ from oxen in being usually of smaller size; the tail is shorter, the horns of females are much smaller than those of males, they lack the accessory column on the inner side of the upper molars, and the cannon bone is longer and more slender; but when it comes to a comparison of the one with the other, it is by no means always easy to tell the difference. It is true that the early Greeks seem to have had a rough and ready rule under which mistakes were not easy, for Aristotle tells us "Alcmaeon is mistaken when he says that goats breathe through their ears," but the severely practical methods of our own day leave us little but some very minute points of difference. One of the best of these lies in the shape of the basi-occipital bone, but naturally this can be observed only in the prepared skull. The terms often employed to denote difference in the horns can have only a general application, for they break down in certain species in which the two groups approach each other. The following table expresses some fairly definite points of separation:

SHEEP (_Ovis_). GOAT (_Capra_).

1. Muzzle hairy except between 1. Muzzle entirely hairy.

and just above the nostrils.

2. Interdigital glands on all 2. Interdigital glands, when the feet. present, only on fore feet.

3. Suborbital gland and pit 3. Suborbital gland and pit usually present. never present.

4. No beard nor caprine 4. Male with a beard and smell in male. caprine smell.

5. Horns with coa.r.s.e transverse 5. Horns with fine transverse wrinkles; yellowish striations, or bold k.n.o.bs or brown; sub-triangular in front; blackish; in male in male, spreading outward more compressed or angular, and forward with a sweeping backward circular sweep, points with a scythe-like curve or turned outward and forward spirally, points turned upward and backward.

These features are distinctive as between most sheep and most goats, but the Barbary wild sheep (_Ovis tragelaphus_) has no suborbital gland or pit, a goat-like peculiarity which it shares with the Himalayan bharal (_Ovis nahura_), in which the horns resemble closely those of a goat from the eastern Caucasus called tur (_Capra cylindricornis_), which for its part has the horns somewhat sheep-like and a very small beard. This same bharal has the goat-like habit of raising itself upon its hind legs before b.u.t.ting.

Both groups are a comparatively late development of the bovine stock, as they do not certainly appear before the upper Pliocene of Europe and Asia, and even at a later date their remains are not plentiful. Goats appear to have been rather the earlier, but are entirely absent from America.

The number of distinct species of sheep in our fauna is a matter of too much uncertainty to be treated with any sort of authority at this time.

Most of us grew up in the belief that there was but one, the well-known mountain sheep (_Ovis canadensis_), but seven new species and sub-species have been produced from the systematic mill within recent years, six of them since 1897. It is no part of the purpose of the present paper to dwell upon much vexed questions of specific distinctness, and it will only be pointed out here that the ultimate validity of most of these supposed forms will depend chiefly upon the exactness of the conception of species which will replace among zoologists the vague ideas of the present time. Whatever the conclusion may be, it seems probable that some degree of distinction will be accorded to, at least, one or two Alaskan forms.

As sheep probably came into America from Asia during the Pleistocene, at a time when Bering's Strait was closed by land, it might be expected that those now found here would show relationship to the Kamtschatkan species (_Ovis nivicola_); and such is indeed the case, while furthermore, in the small size of the suborbital gland and pit, and in comparative smoothness of the horns, both species approach the bharal of Thibet and India, which in these respects is goat-like.

When one considers the poverty of the new world in bovine ruminants, it seems strange that three such anomalous forms should have fallen to its share as the p.r.o.ng-horn, the white goat and the musk-ox, of none of which have we the complete history; two of the number being entirely isolated species, sometimes regarded as the types of separate families.

The p.r.o.ng-horn is a curious compound. It resembles sheep in the minute structure of its hair, in its hairy muzzle, and in having interdigital glands on all its feet. Like goats, it has no sub-orbital gland nor distinct pit. Like the chamois, it has a gland below and behind the ear, the secretion of which has a caprine odor. It has also glands on the rump. It is like the giraffe in total absence of the accessory hoofs, even to the metapodials which support them. It differs from all hollow horned ungulates in having deciduous horns with a fork or anterior branch. There is not the least similarity, however, between these horns and the bony deciduous antlers of deer, for, like those of all bovines, they are composed of agglutinated hairs, set on a bony core projecting from the frontal region of the skull.

It is well known that these horn sheaths are at times shed and reproduced, but the exact regularity with which the process takes place is by no means certain, although such direct evidence as there is goes to prove that it occurs annually in the autumn. p.r.o.ng-bucks have shed on eight occasions in the Zoological Gardens at Philadelphia, five times by the same animal, which reached the gardens in October, 1899, and has shed each year early in November, the last time on October 22, 1903,[1]

and the writer has seen one fine head killed about November 5 in a wild state, on which the horn-sheaths were loose and ready to drop off.

[Footnote 1: It is interesting to note that the first pair shed measured 7-1/4 inches, on the anterior curve; the second pair 9-1/2, and the last three 11 inches each. The largest horns ever measured by the writer were those of a buck killed late in November, 1892, near Marathon, Texas, and were 15-3/4 inches in vertical height and 21 along the curve.]

But few of these delicate animals have lived long enough in captivity to permit study of the same individual through a course of years, and the scarcity of observations made upon them in a wild state is remarkable. That irregularity in the process would not be without a.n.a.logy, is shown by the case of the Indian sambur deer, of which there is evidence from such authority as that king of sportsmen, Sir Samuel Baker, and others, that the shedding does not always occur at the same season, nor is it always annual in the same buck; and by Pore David's deer, which has been known to shed twice in one year.