The Cambridge natural history, Vol. 10 (of 10) Papyras Classics

← The Cambridge natural history, Vol. 10 (of 10)

Chapter 33

C. Pecora

The Pecora are a group which possess so many characters in common that it is not an easy task further to subdivide them.

In all there are but two functional digits on the feet, and the metacarpals and metatarsals of these are fused. There are no upper incisors, and canines in the upper jaw are not universal, and generally small. Horns are confined to this group of the Selenodontia. The premolar teeth are of a simpler form than the molars. The stomach has four chambers, of which two may be regarded as belonging to its cardiac half and two to the pyloric. The former are, in the first place, a large paunch or rumen, followed by a smaller reticulum, so called on account of the network arrangement of the folds of its lining membrane. Connected with the latter, and constituting the first part of the pyloric half of the stomach, is the psalterium or "manyplies," so called on account of the longitudinal folds, like the leaves of a book, into which its lining membrane is raised. Finally there is the abomasum, out of which proceeds the small intestine. Garrod has observed that the chamber of the stomach which varies most among the Pecora is the psalterium. This chamber is specially large in Bos, and particularly small in the Antelopes Nannotragus and Cephalophus. But its variation relates more especially to the folds of its mucous membrane. These folds are of varying lengths and have a definite arrangement There may be as many as five sets of laminae of regular depths. The most simple psalterium is that of Cephalophus, where there are only two sets of laminae of different sizes, a deeper set and a very much shallower set; this form is termed by Garrod "duplicate." Most common is the "quadruplicate" arrangement, with four sets of laminae of differing depths. In all Pecora the liver is but little divided by fissures.

Fam. 6. Cervidae.—The Deer tribe is a very extensive one, and, with the exception of Africa and Australia, world-wide in distribution.

The Deer are absolutely distinguished from all other Ruminant animals by the existence of antlers, which are invariably present in the male sex, save in the aberrant genera Moschus and Hydropotes; in the Reindeer alone are antlers present in both sexes. The general characters of these appendages have been dealt with on a former page (p. ), where they are compared to, or rather contrasted with, the horns of the Bovidae. These antlers, so characteristic of the Cervidae, are very variously developed among the members of the family. Thus in Elaphodus the antlers are very small and entirely unbranched. In the Muntjacs, Cervulus, the antlers are hardly larger, but they have a small anterior branch arising from near the pedicel, the "brow tine." In Cariacus antisiensis only one branch, the brow tine, is present, but it is nearly as long as the main stem of the antler, the "beam." In Capreolus capraea the beam bears two tines; in Cervus sika three; in C. duvauceli two of the three tines present bear secondary branches. There are other complications (some of which are illustrated in Figs. 152-157) of the simple antler which culminate in the complex antlers with their expanded "palms" of the Elk and the Fallow Deer.

Another highly-interesting fact concerning these same antlers is their gradual increase in complexity of tines and palm from the Miocene Cervus matheroni to the great Irish Elk of post-Tertiary times.

Beyond the antlers there seems to be no character of universal applicability which distinguishes the Cervidae from the nearly-related Antelopes. There are, however, a number of structural features which are nearly universally characteristic. Excepting Moschus (which Professor Garrod would not allow to be a "Deer"), no Cervine has a gall-bladder to its liver. All Bovidae (including Antelopes) have, with the exception of Cephalophus.

A small but constant character of the Deer is the existence of two orifices to the lachrymal duct. The genus Tragelaphus alone among Antelopes shows this character.

So far as is known the placenta of the Deer has but few cotyledons, that of the Bovidae many. But not many types are known.

The navicular, cuboid and ectocuneiform are often united. This is never the case in the Bovidae.

The first and second phalanges of the lateral (imperfectly developed) digits are always present except in the Muntjacs; they are never found in Bovidae. The Deer always present a light brown to a darker brown coloration. Elaphodus michianus is almost black. There is commonly white on the under parts and beneath the short tail. Some Deer, such as the Fallow Deer, are spotted; and the young of others that are uniformly coloured when adult are spotted. In some cases a winter coat, darker than the summer coat, is developed.

Altogether some sixty species of Deer are known, of which the preponderance are Old-World forms. The Deer of the Old World are distributed among the genera Cervus (all Europe and Asia); Cervulus, the Muntjacs (India, Burmah, China, etc.); Hydropotes (Eastern China); Capreolus (Europe and Central Asia); Elaphodus (Eastern China); there is one American Cervus, the Wapiti. The American genera are Cariacus and Pudua. The Elk (Alces) and the Reindeer (Rangifer) are circumpolar. The principal structural modification which occurs within the family Cervidae concerns the rudimentary fifth and second toes. In Capreolus, Hydropotes, Moschus, Alces, Rangifer, and Pudua there are considerable remains of the lower parts of metacarpals II. and V.; in the other genera smaller traces of the upper ends of the same bones.

The two most abnormal genera are Moschus and Hydropotes, more particularly the former, which neither Sir V. Brooke nor Professor Garrod allow to be members of the family at all. Moschus is usually placed in a special sub-family by itself, Moschinae, the remaining Deer being referred to another sub-family, Cervinae.

Sub-Fam. 1. Cervinae.—The genus Cervus comprises rather over twenty existing species, which, except the Wapiti (C. canadensis), are exclusively Old World in distribution. The principal features of variation in the genus, in accordance with which it has been divided up into sub-genera, are (1) palmated (Fallow Deer, Dama) or non-palmated antlers; (2) adults spotted with white at all ages and seasons (Axis), or in summer only (Pseudaxis), or not at all; (3) spotted or unspotted young; (4) existence or absence of rudimentary canines in the upper jaw.

Among the members of this genus, Cervus (Elaphurus) davidianus is interesting as having been first observed by the missionary Père David in a park belonging to the Emperor of China near Pekin. Its horns are remarkable for dividing early into two branches of equal length, of which the anterior again branches into two. Specimens of this Deer were ultimately obtained for the Zoological Society's Gardens.

The species of Cervus are fairly distributed between the Palaearctic and the Indian regions. The Palaearctic species, such as Lühdorff's Deer (Fig. 152), are mainly Asiatic. Cervus elaphus and Cervus dama alone are European and British. The former of course is the Red Deer, the latter the Fallow Deer. The Red Deer is reddish-brown in summer and greyish-brown in winter, with the white patch on the rump so common in the Deer tribe. The Red Deer is genuinely wild in Scotland, in certain parts of Devonshire and Westmoreland, and in the New Forest. At the beginning of the last century, according to Gilbert White, there were 500 head of deer in Wolmer Forest, which were inspected by Queen Anne. The antlers may have as many as forty-eight points; and a stag with more than the three anterior tines is termed a "Royal Hart." The Fallow Deer has palmated antlers, and is usually spotted. It seems to be an introduced species, common report pointing to the Romans as the introducers. It would be more correct to say "re-introduced," for fossil remains of this Deer have been met with.

Fig. 152.—Lühdorff's Deer. Cervus luehdorffi. × 1⁄15. (From Nature.)

Elaphodus contains probably two species, E. cephalophus of Milne-Edwards and E michianus of Swinhoe, both from China. The antlers are small and unbranched; the canines in the male are massive; it differs from Cervulus, to which it is closely allied, principally in the absence of frontal glands. The second species has a dark iron-grey pelage, and the late Mr. Consul Swinhoe described it as very Goat-like in aspect.

Capreolus.—The Roe Deer has fairly complex antlers. It is a small Deer and has spotted young. The common Roe Deer, C. capraea, is a native of this country. It is the smallest of our Deer, and its antlers only have three tines in stags of the third year. It is a singular fact about this Deer that though the pairing season is in July and August, the young are not born until the following May or June, a period which does not represent that of gestation. The germ remains dormant for some time before developing.

Fig. 153.—Mule Deer. Cariacus macrotis. × 1⁄15. (From Nature.)

The Muntjacs, Cervulus, form a distinct generic type confined to the Indian and the South-Eastern Palaearctic region. They are small Deer with spotted young, and short one-branched antlers placed upon pedicels as long as themselves. The canines are strongly developed in the males. There are about half-a-dozen species.

Cariacus is exclusively American in range, and contains about twenty species. There are or are not upper canines. The young are spotted. The antlers are occasionally very simple; in C. rufus and a few allies (placed in a special sub-genus Coassus) they are simple spikes without branches. In this genus, and in the nearly allied and also New-World Pudua, the vomer is prolonged backwards and divides the posterior nares into two. The bulk of the species are South American.

Fig. 154.—Chilian Deer. Cariacus chilensis. × 1⁄12. (From Nature.)

Pudua, just mentioned, comes from the Chilian Andes. It is a small Deer without canines and with minute antlers. Other generic names have been proposed for various species of American deer.

Hydropotes inermis is a small perfectly hornless Deer, living on the islands of the Yang-tse-kiang. The male has tusks; the young are spotted. Though, like other deer, Hydropotes has no gall-bladder, both Mr. Garrod and Mr. Forbes found the rudiments of one in the shape of a white ligamentous cord. Mr. Forbes has especially dwelt upon the likeness of the brain to that of Capreolus. The female has four teats, and produces three to six young at a time.

Fig. 155.—Water Deer. Hydropotes inermis. × 1⁄10. (From Nature.)

Alces machlis, the Elk or Moose, is a circumpolar species with palmated antlers and is of large size. The young are unspotted. This animal is the largest of the Deer tribe. The aspect of this creature is by no means that of a Deer, the long, thick, and rather prehensile upper lip not by any means suggesting the family to which it belongs; the legs, too, are ungainly through their unusual length. The Moose has a curious method of protecting himself from Wolves. Instead of moving about during heavy snowstorms, and being thus on the heavy ground an easy prey for these agile enemies, the animal forms what is known as a "Moose yard." An area of ground is kept well trampled down, and the animal contents itself with browsing upon the adjacent stems. The well-trampled ground gives an easy footing, and by his powerful horns the great stag is able to keep his enemies at bay.

Fig. 156.—Moose. Alces machlis. × 1⁄20.

Rangifer tarandus, the Reindeer, is unique among Deer by reason of the fact that both sexes wear antlers. These antlers are palmated. The brow tine and the next or bez tine are also palmated and are directed forwards and a little downwards. The young are unspotted. The pelage alters in winter. Like the Moose, the Reindeer is circumpolar. As is well known, during the Pleistocene period the Reindeer extended its range as far as the South of France. Even in the historic period it is said to have been hunted in Caithness.

Reindeer, like so many other particularly Arctic animals, have regular migrations. In Spitzbergen, for instance, the animal migrates in the summer to the inland region of the island, and in the autumn back again to the sea coast to browse upon the seaweed. These migrating herds have been stated to be led by a large female.

Fig. 157.—Reindeer. Rangifer tarandus. × 1⁄15.

Sub-Fam. 2. Moschinae.—Moschus moschiferus is a native of the Asiatic Highlands. It is 3 feet or so high, perfectly hornless, and with very large canines in the male. It is noteworthy that in Hydropotes, where the canines are also very large, horns are absent. These are examples, perhaps, of correlation. The musk sac (whence the name) is present on the abdomen of the male only. There is no crumen or suborbital gland, which is so generally (though by no means universally) present in Cervidae. But the male has, in addition to the musk glands, glands near the tail and on the outside of the thigh. Unlike other Deer, the lachrymal bone of Moschus bears but one orifice. The feet, so far as concerns the preservation of the outer rudimentary metacarpals, are of the more ancient type represented in Alces, Hydropotes, etc. A gall-bladder is present. The young, as in so many Cervidae are spotted; but the adult is of a greyish-brown colour.

Fig. 158.—Musk Deer. Moschus moschiferus. × 1⁄6. (From Nature.)

There is no doubt that Moschus is more nearly related to the Cervidae than to any other Ruminants. It is regarded by Sir W. Flower as "an undeveloped deer—an animal which in most points (absence of horns, smooth brain, retention of gall-bladder, etc.) has ceased to progress with the rest of the group, while in some few (musk gland, mobile feet) it has taken a special line of advance of its own."

The musk itself, which gives its name to the creature, is found in a gland on the belly, about the size of a hen's egg. The whole gland is cut out and sold in this condition. Such quantities of musk deer have been and are killed for this purpose that the rarity of the animal is increasing. In the seventeenth century it was so common that the traveller Tavernier purchased 7673 musk "pods" in one journey, or, according to Buffon, 1663. The tusks, which recall those of Hydropotes, to which Moschus is not nearly allied, and of Tragulus, with which it has of course still less connexion, are said to be used for the digging up of roots. Its feet, in relation to its mountain-ranging habits, are very mobile.

Extinct Species of Deer.—It has been already mentioned that the most primitive kinds of Deer had no horns at all, resembling in this the modern Moschus and Hydropotes, and that with lapse of time went hand in hand an increasing complexity of antler; the facts of palaeontology harmonising in the most striking manner with the facts of individual development from year to year. The oldest forms seem to be more nearly akin to the living Muntjacs, and their remains occur in the lowest Miocene beds of both Europe and America. At present the group is confined to the warmer parts of Asia and some of the islands belonging to that continent.

One of the oldest types is Amphitragulus. This genus, which consists of several species, inhabited Europe, and differed from living Muntjacs in being totally hornless in both sexes; the skull had no lachrymal fossa or deficient lateral ossification.

Nearly allied is Dremotherium of similar age and range.

The Middle Miocene has furnished the remains of the genus Dicroceras. This is the earliest Deer in which horns have been found. The horns are, as the name of the genus implies, bifid, and have, like those of the living Muntjac, a very long pedicel. This is also a European genus like the last. From this period we come across true Deer, which commence in the Upper Miocene and have branched horns. Moreover they belong, at least for the most part, to the existing genera. One of the most remarkable forms is Cervus sedgwicki (sometimes placed in a separate genus, Polycladus) from the Forest Bed of Norfolk and from the Upper Pliocene of the Val d'Arno. This creature was remarkable for its multitudinously-branched antlers. These end in no less than twelve points. No Deer exists or has existed in which the horns are so completely branched. They are like those of a Red Deer exaggerated.

Fig. 159.—Giraffe. Giraffa camelopardalis. × 1⁄40.

Fam. 7. Giraffidae.—Undoubtedly the type of a distinct family, Giraffidae, is the genus Giraffa. It is characterised by the long neck, which, nevertheless, consists of only the normal seven vertebrae, and by the "horns" which differ from those of all other Ruminants; they are small bony prominences of the frontal bones, which become fused with the skull, and which are covered with unmodified skin. They are not shed. Between them is a median prominence. This cranial armature is present in the female as well as in the male, and is well developed even in the new-born young. The orbits are completely encircled by bone, and there is no lachrymal fossa, so common in Deer and Antelopes. There are no canines above; but these are present in the lower jaw. The rudimentary digits of other Ruminants have disappeared in this genus. There are fourteen pairs of ribs as in many other Artiodactyla. The liver of the Giraffe is, as in many, but not all, Ruminants, devoid of a gall-bladder; neither has it a caudate or a Spigelian lobe. The caecum is actually largish (2½ feet in length), but is relatively very small, as the small and the large intestines measure 196 and 75 feet in length respectively. The Giraffe has a well-marked "ileo-caecal" gland, found in many Ruminants; its appearance in Giraffa is especially compared by Garrod with its appearance in Alces.

Considered by itself, Giraffa forms a very isolated type of Ruminant. But after we have dealt with certain facts concerning extinct forms clearly allied to Giraffa, the isolation of the family will be found to be less marked.

The Giraffe ("one who walks swiftly," the word means in Arabic) is, as every one knows, limited in its range to the African continent. It is not, however, so familiar a fact that there are two quite distinct species of Giraffe, one a northern form from Somaliland, and the other South African. The distinctness of these two, G. camelopardalis and G. australis, has been lately worked out in some detail by Mr. de Winton. The principal point of difference between them consists in the large size of the median horn in the Cape species, which is represented by the merest excrescence in the other species. The Giraffe of West Africa is held to differ from the northern and southern species, coming nearer to the former. It appears in the first place to be a larger animal, and slight differences in the skull have been pointed out. This series of peculiarities may be expressed, for those who do not object to trinomial nomenclature, by calling this novel western form Giraffa camelopardalis peralta. The existence of the three horns covered with unaltered skin is the main characteristic of this Ungulate. But the Giraffe also differs from other Artiodactyles by its enormously long neck, which enables it to browse upon trees inaccessible to the common herd of Ruminants. The neck is often supposed to have some relation to this method of feeding. But a more ingenious explanation of its inordinate length is that it serves as a watch-tower. The long grass of the districts inhabited by the animal swarms with Lions and Leopards, which must be foes. The long neck allows of a wide look out being kept, and it is noteworthy that the Ostrich, living under similar conditions, is also renowned for its length of neck. It is the spots upon the Giraffe which have given it its name of Cameleopard; these spots present in the southern form a series of chocolate-coloured areas, sharply marked off by white spaces. Of these spots it is asserted that they serve as a means of concealing their possessor. Sir Samuel Baker wrote of it in the following words: "The red-barked mimosa, which is its favourite food, seldom grows higher than 14 or 15 feet. Many woods are almost entirely composed of these trees, upon the flat heads of which the giraffe can feed when looking downwards. I have frequently been mistaken when remarking some particular dead tree-stem at a distance that appeared like a decayed relic of the forest, until upon nearer approach I have been struck by the peculiar inclination of the trunk; suddenly it has started into movement and disappeared."

The Giraffe, remarked Pliny, "is as quiet as a sheep." The Roman public, to whom the first Giraffe ever brought into Europe was exhibited, expected from its name "to find in it a combination of the size of the camel and the ferocity of a panther." As a matter of fact, Giraffes in captivity are not always sheep-like in temper. They will kick with viciousness and vigour, and will even initiate an attack upon their keeper. At the same time they are singularly nervous creatures, and have been known to die from a shock. In moving, the Giraffe uses the fore- and hind-limb of each side simultaneously; this gives to its gait a peculiar rocking motion, the singularity of which is heightened by the curving movements of the long neck, which even describes now and then a figure of eight in the air. Giraffa camelopardalis and the species (?) already referred to are the only existing Giraffes (of the genus Giraffa), and they are not found out of Africa. Sir Harry Johnston has lately given a brief account of a larger and more brilliantly coloured species from Uganda which will probably prove to belong to a distinct genus. It has five horns, the additional pair being placed above the ears.

Sir Harry Johnston has quite recently made known another genus of Giraffidae living in the Semliki forest, Belgian Congo district. The skin and two skulls, as well as the bones of the feet, are known from specimens sent by Sir Harry Johnston to the Natural History Museum, and briefly described to the Zoological Society by Professor Ray Lankester. This creature, of which the native name is "Okapi," is proposed to be called Ocapia johnstoni. The first actual specimens which reached this country were two bandoliers made from the skin of the flanks, which were striped black and white, and were not unnaturally held to be portions of the skin of a new species of Zebra. The animal is of about the size of a Sable Antelope, and the back and sides are of a rich brown colour; it is only the fore- and hind-limbs which are striped, the striping being longitudinal, i.e. parallel with the long axis of the body. The head is Giraffe-like, but there are no external horns; wisps of curled hairs seem to represent the vestiges of the horns of other Giraffes. The tail is rather short, and the neck is rather thick and short. The skull is clearly Giraffine. The basicranial axis is straight, and the fontanelle in the lachrymal region is very large. Upon the frontal bones near their parietal border is a large boss on either side, which presumably represents the horn core or "os cornu." On the mandible the great length of the diastema between the incisors and premolars is a Giraffine characteristic. The Okapi lives in pairs in the deepest recesses of the forest.

We are acquainted with a few extinct forms, belonging to Giraffa, which are extra-African in range. G. sivalensis is from the Pliocene of the Siwalik Hills in India, G. attica from Greece. These remains, however, do not include the top of the skull, so that it is doubtful whether their horns were as in G. camelopardalis.

A closely-allied genus is the extinct Samotherium. This flourished in Miocene times, and its remains have been found in the Greek island of Samos. The neck and limbs are shorter than in the Giraffe, and the horns, longer than in Giraffa, are placed just above the orbit upon the frontal bones alone, instead of upon the boundary line of frontals and parietals as in Giraffa. In several ways, therefore, the existing Giraffe is a more modified or specialised animal than its forerunner of the Miocene. In the latter, the male alone carried horns, and in neither sex does the unpaired median bony excrescence appear. The remains of this genus (probably even the same species, S. boissieri) also occur in Persia.

Helladotherium (there is but one species, H. duvernoyi) has its four limbs of nearly the same length; the skull of the only known example is hornless; the neck is shorter than in Giraffa. It is known from the Miocene deposits of Pikermi in Greece.

Palaeotragus is a genus which is not referred to the Giraffidae by all systematists. Its very name, given to it by the eminent French palaeontologist M. Gaudry, indicates his opinion as to its Antelopine affinities. The chief and indeed (according to Forsyth Major) the only reason for placing this Ruminant with the Antelopes is the large size of the horns. They undoubtedly suggest the horn cores of Antelopes. But they are placed wider apart than in those animals. It is thought that the hornless Camelopardalis parva is the female of this species, which is from Pikermi.

Rather more different from Giraffa is the extinct genus Sivatherium, from the Siwalik deposits of India. Here again there has been some discussion as to its affinities. Some place it in the neighbourhood of Antilocapra, but most palaeontologists now regard it as a Giraffe. The main peculiarity of this large beast was the existence of two pairs of horn cores; the larger are upon the parietal bones, and are of a palmated form, with a few short tines, which are highly suggestive of those of the Elk (Alces). The shorter anterior pair are upon the frontal bones. The neck is short, the limbs of equal length, and there are no additional toes upon the limbs. Sivatherium was almost as large as an Elephant, and in restorations it is depicted as having a fleshy dilated nose like the Saiga Antelope; this view is based upon the position and size of the nasal bones. Hornless skulls have been identified as the female of Sivatherium.

Vishnutherium, Hydraspotherium, and Bramatherium are allied genera.

Fam. 8. Antilocapridae.—This family contains but one genus and species, the N. American "Pronghorn," Antilocapra americana. This animal deserves a family to itself on account of the singular structure of the horns, which are intermediate in character between those of the Deer and those of the Antelopes. They are unquestionably "hollow-horned" Ruminants, in that there is an osseous horn core, upon which lies the actual horn. This, however, is softer than in Bovidae, and is semicorneous. It is, indeed, more like the velvet of the stag's horn. Moreover the horn is branched, and there are sometimes even three prongs. Furthermore, it is now certainly known that the Pronghorn sheds its horns not merely occasionally, but with definite annual periodicity. It so far resembles the Deer. But it must be borne in mind that in the Deer the horn shedding is a twofold process. There is first of all the stripping off of the velvet, and secondly the shedding of a portion of the horn core down to the burr. What happens in the Prongbuck is the shedding of the true horn only ( = the shedding of the velvet), not of the horn core. It appears, however, that occasionally (once in their lifetime?) certain undoubted Antelopes may cast their horns. Another external character of this animal is the total absence of "false hoofs," the last vestiges of the second and fifth digits. The Pronghorn is a gregarious creature running in bands of six up to hundreds.

Fam. 9. Bovidae.—This family, more extensive than that of the Cervidae, contains not only the Oxen, Sheep, and Goats, but also the Antelopes, save only Antilocapra, which must be placed in a family by itself. The only two points which distinguish all Bovidae from all Cervidae are the nature of the horns already described, and the polycotyledonary condition of the placenta. Moreover the horns are usually present in both sexes, though there are exceptions, such as the Sheep and Goats, and various genera of Antelopes (Tragelaphus, Tetraceros, etc.). There are never the first two phalanges belonging to the rudimentary digits II., V., as there are in all Deer excepting Cervulus. There is as a rule but one orifice to the lachrymal duct. There are never persistent upper canine teeth in either sex.

It is exceedingly difficult to separate the Antelopes from the Sheep, Oxen, and Goats. Their inclusion along with these creatures in one family, Bovidae, shows that no differences of an important character exist. The term Antelope is rather of popular than of zoological significance. As a rule there are horns in both sexes; but this rule is not without exceptions, of which one is the genus Strepsiceros, the Koodoo. Many other Bovidae are horned in the males only, e.g. Saiga, Tragelaphus. The Antelopes further differ from the true Oxen in their more graceful build, and in the fact that the horns, if they curve at all, generally curve backwards towards the neck. In the Oxen, on the other hand, the build is stouter, and the horns usually curve outwards. The same remarks apply to the Sheep. Such an Antelope, however, as the Eland (Orias) is very Ox-like in habit. Another feature which may be remarked upon, though not of absolute differential value, is that while the Antelopes are as a rule smooth and sleek in their skins, the Oxen tend to be rough and shaggy. The Zebu, however, in this, in its hump, and in general aspect, is far from being unlike an Eland. But then the Zebu is a domestic race, and we do not know what the wild stock was like. It is perhaps with the Goats that the Antelopes have the nearest affinities, and it is difficult to place such a form as Nemorrhaedus, and indeed some others. In the Antelopes as a rule the middle lower incisors are larger than the lateral ones; in the Sheep and Goats they are alike in size. The parietal bones, too, in the Antelopes are moderately large and are much shortened in the remaining Cavicornia, especially in the Oxen. As the Antelopes are the oldest, so far as we know, of all bovine animals, one would expect to find them combining the characters of the rest. But they do this so effectually that a disentanglement is really impossible. They date from the Miocene. Antelopes are now limited to Europe, Asia, and Africa; they have always had the same range, though more abundant in former times in Europe. They preponderate now in tropical Africa, and abound in genera and species. Messrs. Sclater and Thomas allow altogether thirty-five genera, of which twenty-four are exclusively Ethiopian in range.

In the following summary of the group Messrs. Sclater and Thomas's work is followed. They commence with a section or sub-family of which the type is the Hartebeest.

Bubalis, or Alcelaphus as it is sometimes called, is an African genus, ranging however into Arabia. These Antelopes are characterised by the long skull and the doubly-curved horns. There are eight species of the genus, of which B. caama is the best known; this is the animal known as the Hartebeest. The Bontebok and Blessbok belong to a closely-allied genus, Damaliscus, distinguished mainly by the fact that the bony base of the horn cores is not extended upwards, and therefore the parietal bones are visible when the skull is viewed from in front, which is not the case in Bubalis.

Fig. 160.—Brindled Gnu. Connochaetes taurinus. × 1⁄20.

The Gnus, Connochaetes, are familiar owing to their curious aspect. The hairy face, and rump and tail like those of a pony are highly characteristic. The horns are bovine in appearance, standing outwards and then curving upwards. There are three species of Gnu, all from South Africa. They are C. gnu, C. taurinus, and C. albogulatus.

Of the Cephalophine section there are two genera:—

Cephalophus is an African genus. These animals are known as Duikerboks; they are small, and have short non-curved horns in the male sex only. Their general aspect is not unlike that of certain Deer with simple horns, such as Cervulus. Messrs. Sclater and Thomas allow thirty-eight species. The smallest species do not exceed the dimensions of a Hare. None are really large.

Tetraceros is an Indian genus characterised, as its name denotes, by the fact that it possesses four horns. It is the posterior pair which correspond to the single pair of Cephalophus. The anterior pair, which are much smaller and are sometimes absent, are a new pair. The female of this Antelope is hornless. Sheep are occasionally four-horned, and there is indeed a breed of such in Kashmir. A four-horned Chamois was described by the late Mr. Alston.

The Klipspringer, Oreotragus saltator, is the first type of a third section; as its name denotes, it is an Antelope with Goat-like habits, being found particularly among rocks. The horns are short and straight. This, the only species of the genus, is African in range, of which its Dutch name gives evidence. A specimen in the Zoological Society's Gardens (as has been pointed out to me by Mr. Mercer) had the habit of depositing the secretion of the tear gland upon a mass of concrete in its enclosure, the secretion thus exuded forming a pointed heap of hardish matter. It may be that the object of this is to guide its fellows to its whereabouts.

Ourebia is a less-known genus, larger in size, but with horns of the same character, though longer.

The Grysbok and the Steinbok, genus Raphiceros, have similar horns. This as well as the last two genera have horns in the male only.

One of the smallest of Antelopes belongs to an allied genus; this is Neotragus pygmaeus. It is known as the Royal Antelope, a name apparently derived from Bosman's statement that the negroes called it "the king of the harts." Its horns are very small. The height of the animal is only 10 inches. Horns are present in the male alone. The last three genera are African.

The Cervicaprine series, which is also African, includes the Waterbucks and Reedbucks, so called on account of their water-loving propensities. As in the last series, from which they are separated by Sclater and Thomas, but with which they are united by Flower, there are horns in the male only. These horns, though not twisted, are long. The typical genus is Cobus, of which there are eleven species. The Waterbuck, C. ellipsiprymnus, and the Sing-sing, C. unctuosus, are perhaps the best-known species; the former is blackish grey, the latter browner in colour. In C. maria and one or two other species the horns are more curved backwards and again forwards than in some of the others, where their form is sublyrate.

The Reedbucks, Cervicapra, are closely allied to Cobus; they are, however, of smaller size. Here, as in that genus, the females are hornless, and the horns of the males are of medium size. Five species are referred to the genus. They are all of a brownish fawn colour. A genus Pelea, with but one species, P. capreolus, has been separated on account of the fact that the horns are nearly straight and that there is no naked patch of skin beneath the ears. This animal has received its name on account of its resemblance to the Roebuck.

The Antilopine section includes a number of genera.

The genus Antilope is Indian in range. It includes but one species, A. cervicapra. This Antelope is of medium size, with a brown pelage getting blacker with years; it is thus known as the Black-buck. The female, which is hornless, is lighter brown. The horns are long, spirally twisted, and closely ringed.

Aepyceros, with two species, is African. The Palla (Ae. melampus) is a large Antelope, with longish lyrate horns in the male, which are half-ringed.

The Saiga Antelope, genus Saiga, is one of the most remarkable types of Antelope in its outward appearance. Its nose is very large and inflated, the two nostrils being quite widely separated, a depression indeed lying between them dorsally. The horns are lyrate in the male, absent in the female. The "ovine expression" of this bovine animal is more pronounced in the female. Corresponding with the clumsy nose are very short nostrils, the commencement of the narial aperture being therefore very far back. It is almost suggestive of Macrauchenia in this respect. The fleece is also Sheep-like. The genus occurred in this country during the Pleistocene. It is now an inhabitant of Eastern Europe and Western Asia. The only species is S. tartarica.

The Chiru, Pantholops, is allied to the Saiga. The horns of the male are long and nearly straight; they are ringed in front. The muzzle is swollen in the male; the nostrils are large, and provided with extensive sacs internally. The colour of this animal, which is exclusively Thibetan in range, is a pale fawn. The hair, in accord with its habitat, is very woolly. No living specimens have ever been brought to Europe. This creature has accumulated much legend. Its blood is believed by the Mongols to possess virtues, and by means of the rings on the horns fortunes are told. Naturally the animal is on these grounds hard to stalk and shoot.

Fig. 161.—Loder's Gazelle. Gazella loderi. × 1⁄10.

The Gazelles, genus Gazella, are fairly numerous in species, which are both Palaearctic and Ethiopian. There are altogether twenty-five of them. The genus as a whole is characterised by the small or moderate size, the sandy coloration with white belly, the presence of dark and light stripes on the face and on the flanks. These streaks, however, are not always present, and their presence or absence serves to differentiate some of the species. The horns are usually present in both sexes. The horns are of fair length, ringed, and of lyrate form.

The Springbok is separated from the rest of the Gazelles, to which genus it is clearly most nearly related, as a genus Antidorcas. This genus differs from Gazella by having only two lower premolars as in Saiga. Otherwise it resembles the Gazelles; there is but a single species, A. euchore, which is African.

Ammodorcas is closely allied to the Gazelles, but differs from them in having an elongated neck and also a long tail. A. clarkei, the only species, is limited to Somaliland.

Lithocranius, not unlike the last, has a still longer neck, which makes it almost Giraffe-like; its tail, however, is short. The scientific name is derived from the "solid stony character of the cranium." In running, this Gazelle carries the head forward in a straight line with the body. It is African.

Dorcotragus with one species, D. megalotis, is a pigmy Gazelle restricted to Somaliland. Its likeness, on account of size and in some other superficial features, to the Klipspringer, led to its original confusion with that genus (Oreotragus).

Fig. 162.—Sable Antelope. Hippotragus niger. × 1⁄20. The horns of the specimen figured have not nearly reached their full dimensions.

A sub-family Hippotraginae, or Hippotragine section, includes a number of Antelopes which agree in the possession of four mammae, and of molars more like those of the true Oxen, of horns of some length, present in both sexes, and of a longish tail. They are all African in range.

The type genus Hippotragus has its horns placed above the orbits; they are not twisted, but curved backwards. There are three species in the genus. Of these the best known is H. niger, the beautiful Sable Antelope. Its general colour is a rich, dark, glossy brown with white stripes on the face, and with a white belly. The other species are the Roan Antelope, H. equinus, and the Blaaubok, H. leucophaeus, of which the last specimen was probably killed in 1799.

Fig. 163.—Beatrix Antelope. Oryx beatrix. × 1⁄16. (From Nature.)

The genus Oryx (chiefly African, but also Arabian and Syrian) also contains a number of species, which are fairly familiar through the fact that several of them are always on view in the Zoological Society's Gardens. The genus differs from Hippotragus in that the horns, present in both sexes, are placed behind the orbits, and slant backwards in a line with the face. They are annulated. The Leucoryx (O. leucoryx) is of a pale colour, but this is not so marked as in O. beatrix, which is largely white with, however, brown legs. The Gemsbok is a handsome creature with greyish tawny colour, much darker on the legs, and with a Gazelle-like, dark, side stripe. It has received its vernacular name on account of its supposed likeness to the Chamois ("Gemse"), just as the Rehbok was so-called from its supposed likeness to the Roe Deer, and the Eland to the Elk. The Beisa (O. beisa) is of a similar tawny colour to the last, and also with darker stripes.

The Addax (Addax) of North Africa, Arabia and Syria, has but one species (A. nasomaculatus). The horns are spirally twisted.

Fig. 164.—Speke's Antelope. Tragelaphus spekii (♀). × 1⁄16.

The Tragelaphine section includes the Kudus, Elands, Nilgais, and Harnessed Antelopes. They are all long-horned (when the horns are present in both sexes), the horns being twisted; the nose is naked with a slight median groove, and all are Ethiopian or Oriental in range.

The genus Tragelaphus includes the Harnessed Antelopes, so called on account of the direction of the stripes suggesting harness. The females are hornless, and the colours of the two sexes are different. The hoofs are long and the toes rather unusually separable, which state of affairs is in accord with the swampy country affected by many. T. gratus and T. spekei are larger forms; the Boschbok, T. sylvaticus, is smaller.

The Kudus, genus Strepsiceros, have more markedly twisted horns, which are absent in the female. The body is vertically striped with white. The largest species is S. kudu; a smaller form, S. imberbis, is from Somaliland.

Fig. 165.—Eland. Orias canna. × 1⁄25.

The last genus of this section or sub-family is the African Eland, genus Oreas (which it appears should be spelt Orias). The Elands are perhaps more Ox-like in appearance than the other members of this group, and in both sexes have horns, in which the spiral twisting is more close. Orias canna is the name of the common Eland. O. livingstonii has been applied to an East African variety, which has thin and faint lateral stripes like the other members of the group to which it belongs.

The genus Boselaphus includes only B. tragocamelus, the Nilgai, which is purely Indian in range. The female is hornless, and the horns of the male are smooth and not long.

The members of the Bovine section or Oxen are to be distinguished from other hollow-horned Ruminants by their stouter build and by the fact that the horns stand out from the sides of the skull and are simply curved, not twisted; and smooth, not annulate like those of other Ruminants. The muffle is naked, broad, and moist. The Oxen are widely distributed; but are entirely absent from the Australian region and from South America and Madagascar.

The true Oxen are perhaps best considered to form but a single genus, Bos. They have, however, been divided into a number of genera. Even the supposed aberrant Anoa depressicornis of Celebes hardly differs sufficiently to warrant its separation. In favour of this view, too, is the extraordinary ease with which different "genera" will cross with each other and produce fertile offspring. The following is the pedigree of an animal lately living in the Zoological Society's Gardens. The female offspring of a male Zebu and a female Gayal was mated with a male Bison. The female calf was again mated with a Bison and produced a calf, also a female, which contained therefore the three species, Bos indicus, Bibos frontalis, and Bison americanus. It is clearly unwise in view of this fact to insist too much upon generic distinctions in any of those types.

Of this genus the Oriental Gaur (Bos gaurus), the Gayal (B. frontalis), and the Banteng (B. sondaicus) form a well-marked section, characterised by their dark coloration and by the somewhat flattened horns.

The Gaur, Bos gaurus, has a more concave forehead than its allies; the horns are less curved than those of the Banteng, and less so than the horns of the Gayal (Bos frontalis). It inhabits the Indian Peninsula; and extends through Burmah to the extremity of the Malay Peninsula. The Malay name of this animal is Sakiutan, which simply means wild cattle. It chiefly frequents wooded hills and is an excellent mountain climber.

Bos frontalis, the Indian Gayal, has a white caudal disc like the last species, but the forehead is flat and the horns curve but little. It is chiefly known as a tame animal, and its occurrence in the wild state has been doubted. It has furthermore been suggested that it is merely a tame race of the Gaur altered slightly through domestication. It is, however, said not to cross in a state of nature with the Gaur.

Fig. 166.—Gayal. Bos frontalis. × 1⁄20.

The Banteng, B. sondaicus, is distributed through Chittagong, Tenasserim, and the Malay Peninsula to Java and Borneo. There are apparently two races of this animal. The species differs from the others by the fact that the horns are smaller and more curved; there is a white caudal disc; the forehead is narrower and the skull longer than in the others.

The American Bison and the European Aurochs form another section; they are indeed extremely alike, specific differences being hardly recognisable. The Bison of America, formerly present in such numbers that the prairies were black with countless herds, has now diminished to about a thousand head.

One of the largest of existing Bovidae is the Aurochs, Wisent, or European Bison, Bos bonasus (or Bison europaeus). It is exceedingly like its American relative. Formerly the animal was much more widely spread than it is now, extending its range from Europe into North America. It is now limited to certain districts on the Urals, in the Caucasus, and a herd of them are kept up through the fostering care of the Emperor of Russia in the forest of Bielovege in Lithuania. The term "Aurochs" should not really be applied to this species but to the Wild Cattle, Bos taurus. It is, however, so generally used for the Wisent (which is the German name) that it is not necessary to change it. The Sclavonic name is Zubr or Suber. It is a great beast, standing 6 feet or so in height at the shoulder. It ranged further over Europe well within the historic period. In the days of Charlemagne it was spread over Germany and was a beast of the chase. In the year 1848 the Emperor of Russia presented a pair of these Oxen to the Zoological Society of London. At the time of their presentation an interesting communication was made to the Society by M. Dolmatoff, on the method of the capture of these two examples. The creature is not easy to capture and is alarming to confront. "The eyes," says an old writer, "are red and fiery; the looks are furious and commanding." It has of course the shaggy mane and hump of the American animal. The herd in Lithuania was said to be 1900 in the year 1856. Mr. E. N. Buxton, who has lately visited the forest, quotes M. Neverli to the effect that at present the numbers are not more than 700.

Fig. 167.—Bison. Bison americanus. × 1⁄25.

Allied to this animal, and apparently still nearer to the American Bison, is the extinct B. priscus of Europe. The Pleistocene Bisons of North America, B. antiquus and B. latifrons, are not remote from the living forms. Finally, the Miocene B. sivalensis from India, and the Pliocene B. ferox and B. alleni of North America, take back this group to as remote a period as any other genus of Oxen.

Fig. 168.—Yak. Bos grunniens. × 1⁄15.

Fig. 169.—British Wild Ox. Bos taurus. From Vaynol Park, Bangor. × 1⁄20.

The Yak, Bos grunniens, is a long-haired peculiar type, confined to the Thibetan plateau. B. (Anoa) depressicornis of Celebes is characterised by its straight horns; allied to it is B. mindorensis (Philippine Islands), supposed, however, to be a hybrid between it and some other species. Africa has at least two Buffaloes. We may finally mention the Wild Ox of Europe, B. primigenius, the supposed progenitor of our domestic cattle, believed to be still surviving in the herds at Chillingham, Chartley, and elsewhere. This animal is sometimes called the Aurochs. The Romans spoke of it as the Urus, and it appears to have formerly attained to more gigantic proportions than at present. It is the small size of the present race that is the chief objection to tracing them back to the large Oxen existing near London in 1174, and found sub-fossil in the Cambridgeshire fens.

Fig. 170.—Punjab Wild Sheep. Ovis vignei. × 1⁄10.

Of the true sheep, genus Ovis, there are a considerable number of species. The Sheep are to be distinguished from the Goats by their rather stouter build and by the absence of the beard in the male. The horns are developed in both sexes, and are usually twisted and often of large size.

The Sheep are almost entirely Palaearctic and Nearctic. They only just get into the Oriental region. One of the finest species is the great Pamir Sheep, O. poli, whose length reaches 6 feet 7 inches, and height 3 feet 10 inches. The horns of this fine Sheep may measure more than five feet round the curves. The Rocky Mountain Bighorn (O. montana) is a Sheep ranging along the Rockies as far south as New Mexico, and also to the far north; they are not confined to the chain of mountains mentioned, but occur also on the mountains of British Columbia down to those of California. The horns are not quite as large as those of the last species, but measurements give a length (along the curve) of 32 to 40 inches.

Fig. 171.—Himalayan Burrhel Sheep. Ovis burrhel. × 1⁄12. (From Nature.)

Fig. 172.—Blanford's Sheep. Ovis blanfordi. × 1⁄10. (From Nature.)

Just as the Goats are often limited to islands and small stretches of country, so are the Sheep. Thus Cyprus has a species, O. ophion, peculiar to itself. This, which is known as the Cyprus Mouflon, is limited to a range of mountains, the Troodos, in that island. In 1878 it was believed that the animal was nearly exterminated, a flock of twenty-five members alone surviving. They have, however, since increased. Confined to the Thibetan plateau are O. hodgsoni and O. nahura. Corsica has the Mouflon, O. musimon; and the Barbary Sheep or Arui, O. tragelaphus, is found only in Northern Africa. Ovis burrhel and O. blanfordi are Indian forms.

Fig. 173.—Barbary Sheep. Ovis tragelaphus. × 1⁄10.

Fig. 174.—Thar. Capra jemlaica. × 1⁄10. (From Nature.)

Ovis nahura is chiefly responsible for the impossibility of strictly separating the Sheep and Goats. It has no suborbital glands or lachrymal fossae, which are as a rule present in the Sheep and absent from the Goats. On the other hand interdigital glands are present, which is the case with Sheep. Its habits, too, are a blending of those of the Sheep and the Goat. It lives largely on undulating ground like Sheep, and frequently lies down during the day on its feeding ground. On the other hand it is, like the Goats, a splendid climber.

The Goats, genus Capra, differ from the Sheep in their slighter build and in the fact that the horns are not spirally curved, but arched over the back. There is also the characteristic beard, and the male is odorous. The true Goats are almost exclusively Palaearctic in range. They show the limited distribution of the Sheep, a distribution which follows from their mountain-loving habits.

Fig. 175.—Sinaitic Ibex. Capra sinaitica. × 1⁄10.

Thus we have the Spanish Ibex (C. pyrenaica), limited to the Pyrenees and other mountain ranges of the peninsula; C. ibex, the Steinbok of the Alps and the Tyrol; the Markhoor, C. falconeri, of certain mountain ranges of Afghanistan; the Caucasian, Sinaitic, and Cretan Ibexes, and the Thar.

Capra aegagrus, the Persian Wild Goat, ranges from the Caucasus to Sind. It is this animal which produces the true "bezoar stone." The substance in question is a secretion apparently found in the stomach. It is still, according to Mr. Blanford, regarded as an antidote to poison in Persia. Buffon called this Goat the "Pasan," which is evidently a corruption of the word bezoar. When the substance was in repute as a medicine of the "alexipharmic" kind, the supply naturally came up to the demand. Thus the bezoar stones of the Lama in South America gained repute, and there were "Oriental bezoar, cow bezoar, hog bezoar, and monkey bezoar"! As concretions of one kind or another are not uncommon objects in the alimentary tract of mammals it was easy enough to obtain a fair amount of some substance which was sure to sell well. It is said that a stone weighing four ounces was once sold in this country (or at any rate in Europe) for £200.

"There can be no doubt," observes Mr. Blanford, "that C. aegagrus is one of the species, and probably the principal, from which tame goats are derived."

Fig. 176.—Japanese Goat Antelope. Nemorrhaedus crispus. × 1⁄12. (From Nature.)

The Chamois (Rupricapra) and the Goral (Nemorrhaedus) are best described as Goat-like Antelopes; but, as already said, it is difficult to split up the Bovidae satisfactorily. The Rocky Mountain Goat, Haploceros montanus, is a large Goat-like creature, which has the peculiarity of having the shortest cannon bones of any Ruminant. Its name denotes its range.

Fig. 177.—Goral. Nemorrhaedus goral. × 1⁄12. (From Nature.)

The Musk Ox, Ovibos moschatus, has been thought to be on the borderland between the Sheep and Oxen, as indeed expressed in its scientific name. It is a purely Arctic creature, now confined to the Nearctic region; but it formerly existed in the Arctic regions of Europe.

The anatomy of the "soft parts" of this genus has lately been investigated by Dr. Lönnberg. The animal has no foot glands such as occur in Ovis. Its kidneys, however, are non-lobate, and it has orbital glands. The cotyledons of the placenta are unusually large, and the cow has the "primary four" teats. It cannot, in fact, be definitely referred to either the Caprine or the Bovine section of the Cavicornia, and while possibly most allied to Budorcas, it may be regarded, at least for the present, as entitled to form a separate sub-family of its own. The muzzle has a slight naked strip above the nostrils, as in the Sheep, but there is no fissure of the upper lip.