Fig. 149.
Fig.149.—Lama.Lama huanacos.×1⁄12.
Extinct Camels.—The earliest cameloid type is the genusProtylopus,[186]of which we are acquainted with an imperfect skulland the greater part of a radius and ulna belonging to one individual, and most portions of the hind-limbs in other specimens. The one species,P. petersoni, was about the size of a "jack rabbit," and is late Eocene (Uinta formation) and American in range. The teeth of this mammal are the typical forty-four, and the canines are not pronounced, being incisiform in shape. In the skull the nasals overhang, as in the genusPoebrotherium. The orbit is not closed by bone. There is in this ancient Camel a trace of the supra-orbital notch so characteristic of the Camel tribe. "The vertebrae resemble those of the modern Lamas closely in their general proportions." The lumbars have the usually Cameloid formula of 7. This genus has but two functional toes on the hind-feet, the second and fifth being reduced to vestiges. It is interesting to note that the radius and ulna appear to remain distinct, except in very old animals, in which they come to be co-ossified in the middle only, thus foreshadowing their complete union in the next genus,Poebrotherium. The present genus, moreover, as well asPoebrotherium, was distinctly unguligrade; it has not acquired the characteristic phalangigrade mode of progression of the modern types of Camels.
Fig. 150.
Fig.150.—Skull ofPoebrotherium wilsoni.i1,i2,i3, Incisors 1-3. × ½. (After Wortman.)
The American and OligocenePoebrotheriumhas been recently and exhaustively studied by Professor Scott.[187]It was considerably smaller than a Lama. Its neck was long as compared with other Artiodactyles, but still shorter than that of the Lama. It was a lightly-built, graceful creature, with apparently some external likeness to a Lama. It is an important fact to notice that at thisperiod, and for a long time after, there were no types referable to the Camelidae in the Old World. Though a Camel in many features of its organisation,Poebrotheriumwas "generalised" in many ways. Thus the metacarpals and metatarsals were not fused to form a cannon bone, and the two lateral digits were represented by splint rudiments of metacarpals and metatarsals. The dentition was complete. The skull though distinctly Tylopodan, also shows more generalised characters. Thus the orbit is not quite, though nearly, completed by bone. In the Camel it is quite closed. The nasal bones are much longer, reaching nearly to the end of the snout. The odontoid process of the axis vertebra is not spout-like as in existing forms, but cylindrical, though slightly flattened upon the upper surface. The scapula is described as being more like that of the Lama than of the Camel, though variations occur which approximate to the Camel. The brain, judging of course from casts, has those sulci "which are common to the whole series of Ungulates, and closely resemble those of a foetal Sheep."
Fig. 151.
Fig.151.—Anterior surface of axis of Red Deer, × ⅔.o, Odontoid process;pz, posterior zygapophysis;sn, foramen for second spinal nerve. (From Flower'sOsteology.)
Later in historical sequence thanPoebrotherium, and structurally intermediate between it andProtolabis, is the Miocene genusGomphotherium. It shows an advance in structure uponPoebrotherium, in that the orbit is completely encircled by bone, though the posterior wall is thin; the lower canines instead of being incisiform are curved back as in later Camels, and separated by a wide diastema from the preceding and the succeeding teeth.
Later in age thanPoebrotheriumisProtolabis, aTylopodin which the full number of teeth is still retained; its skull presents no particular changes from the Poebrotherine type; the nasals, however, are somewhat shortened.
Later still in point of time isProcamelus. In this form we have apparently an ancestral stock, whence both Camels and Lamas were derived. The upper incisors are as in existing forms, but the first and second persist for a somewhat longer time. The skull shows two well-marked types of structure; inP. occidentalisthere are more points of likeness to the Lama, inP. angustidensto the Camel. In both, the orbits are completely encircled by bone. The nasals are much shortened. The odontoid process of the axis is still more concave than inPoebrotherium, but not spout-like as in existing forms. This fact shows that the spout-like character of the Camels' odontoid process is not a point of affinity to other Artiodactyles—in fact the occurrence of the same form of odontoid process in Perissodactyles is enough proof of this. We must come to the conclusion that the form is adaptive in all cases. If we were not obliged on palaeontological evidence to come to this conclusion, the structure in question is just one which would be fastened upon as evidence of genetic affinity; for it is a resemblance in a small though distinctive point of structure having no obvious relation to utility. The metacarpals and metatarsals have coalesced to form the cannon bones, though a rudiment of one metacarpal seems to remain. The genera referred to appear to be on the direct line of descent of the modern representatives of the family. But there are other forms which are offshoots of the main stem. Such areHomocamelus,Eschatia, andHolomeniscus. The last two are Pliocene and American; the teeth are much reduced.
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.[188]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 abook, 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 inBos, and particularly small in the AntelopesNannotragusandCephalophus. 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 ofCephalophus, 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.[189]
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 generaMoschusandHydropotes; 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.200), 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 inElaphodusthe 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." InCariacus antisiensisonly one branch, the brow tine, is present, but it is nearly as long as the main stem of the antler, the "beam." InCapreolus capraeathe beam bears two tines; inCervus sikathree; inC. duvaucelitwo 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 MioceneCervus matheronito 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 arenearlyuniversally characteristic. ExceptingMoschus(which Professor Garrod would not allow to be a "Deer"), no Cervine has a gall-bladder[190]to its liver. All Bovidae (including Antelopes) have, with the exception ofCephalophus.
A small but constant character of the Deer is the existence of two orifices to the lachrymal duct. The genusTragelaphusalone 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 michianusis 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[191]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 AmericanCervus, the Wapiti. The American genera areCariacusandPudua. The Elk (Alces) and the Reindeer (Rangifer) are circumpolar. The principal structural modification which occurs withinthe family Cervidae concerns the rudimentary fifth and second toes. InCapreolus,Hydropotes,Moschus,Alces,Rangifer, andPuduathere 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 areMoschusandHydropotes, more particularly the former, which neither Sir V. Brooke nor Professor Garrod allow to be members of the family at all.Moschusis 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 genusCervuscomprises 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) davidianusis 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 ofCervusare fairly distributed between the Palaearctic and the Indian regions. The Palaearctic species, such as Lühdorff's Deer (Fig. 152), are mainly Asiatic.Cervus elaphusandCervus damaalone 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 haspalmated 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.
Fig.152.—Lühdorff's Deer.Cervus luehdorffi.×1⁄15. (FromNature.)
Elaphodus[192]contains probably two species,E. cephalophusof Milne-Edwards andE michianusof Swinhoe, both from China. The antlers are small and unbranched; the canines in the male are massive; it differs fromCervulus, to which it is closely allied, principally in the absence of frontal glands. The secondspecies 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.
Fig.153.—Mule Deer.Cariacus macrotis.×1⁄15. (FromNature.)
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.
Cariacusis exclusively American in range, and contains about twenty species. There are or are not upper canines. The youngare spotted. The antlers are occasionally very simple; inC. rufusand a few allies (placed in a special sub-genusCoassus) they are simple spikes without branches. In this genus, and in the nearly allied and also New-WorldPudua, the vomer is prolonged backwards and divides the posterior nares into two. The bulk of the species are South American.
Fig. 154.
Fig.154.—Chilian Deer.Cariacus chilensis.×1⁄12. (FromNature.)
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 inermisis 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,Hydropoteshas no gall-bladder, both Mr. Garrod[193]and Mr. Forbes[194]found therudiments of one in the shape of a white ligamentous cord. Mr. Forbes has especially dwelt upon the likeness of the brain to that ofCapreolus. The female has four teats, and produces three to six young at a time.
Fig. 155.
Fig.155.—Water Deer.Hydropotes inermis.×1⁄10. (FromNature.)
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 welltrampled 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.
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 inthe 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.
Fig.157.—Reindeer.Rangifer tarandus.×1⁄15.
Sub-Fam. 2. Moschinae.—Moschus moschiferus[195]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 inHydropotes, 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 ofMoschusbears but one orifice. The feet, so far as concerns the preservation of the outer rudimentarymetacarpals, are of the more ancient type represented inAlces,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.
Fig.158.—Musk Deer.Moschus moschiferus.×1⁄6. (FromNature.)
There is no doubt thatMoschusis 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 seventeenthcentury 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 ofHydropotes, to whichMoschusis not nearly allied, and ofTragulus, 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 modernMoschusandHydropotes, 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 isAmphitragulus. 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 isDremotheriumof similar age and range.
The Middle Miocene has furnished the remains of the genusDicroceras. 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 isCervus 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.
Fig.159.—Giraffe.Giraffa camelopardalis.×1⁄40.
Fam. 7. Giraffidae.—Undoubtedly the type of a distinct family, Giraffidae, is the genusGiraffa. It is characterised bythe 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 thenew-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[196]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 inGiraffais especially compared by Garrod with its appearance inAlces.
Considered by itself,Giraffaforms a very isolated type of Ruminant. But after we have dealt with certain facts concerning extinct forms clearly allied toGiraffa, 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. camelopardalisandG. australis, has been lately worked out in some detail by Mr. de Winton.[197]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 formGiraffa 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 herdof 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[198]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 camelopardalisand the species (?) already referred to are the only existing Giraffes (of the genusGiraffa), 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 Ugandawhich 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.[199]This creature, of which the native name is "Okapi," is proposed to be calledOcapia 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 toGiraffa, which are extra-African in range.G. sivalensisis from the Pliocene of the Siwalik Hills in India,G. atticafrom Greece. These remains, however, do not include the top of the skull, so that it is doubtful whether their horns were as inG. camelopardalis.
A closely-allied genus is the extinctSamotherium. 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 inGiraffa, are placed just above the orbit upon the frontal bones alone, instead of upon the boundary line of frontals and parietals as inGiraffa. In several ways, therefore, the existing Giraffe is a more modified orspecialised 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 inGiraffa. It is known from the Miocene deposits of Pikermi in Greece.
Palaeotragusis 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[200]) 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 hornlessCamelopardalis parvais the female of this species, which is from Pikermi.
Rather more different fromGiraffais the extinct genusSivatherium, from the Siwalik deposits of India. Here again there has been some discussion as to its affinities. Some place it in the neighbourhood ofAntilocapra, 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.Sivatheriumwas 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 ofSivatherium.
Vishnutherium,Hydraspotherium, andBramatheriumare 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 characterbetween 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),notof the horn core. It appears, however, that occasionally (once in their lifetime?) certain undoubted Antelopes may cast their horns.[201]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 onlyAntilocapra, which must be placed in a family by itself. The only two points which distinguish all Bovidae from all Cervidae[202]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 exceptingCervulus. 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 thanof zoological significance. As a rule there are horns in both sexes; but this rule is not without exceptions, of which one is the genusStrepsiceros, 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 asNemorrhaedus, 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[203]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, orAlcelaphusas 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 whichB. caamais thebest 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 inBubalis.
Fig. 160.
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.[204]There are three species of Gnu, all from South Africa. They areC. gnu,C. taurinus, andC. albogulatus.
Of the Cephalophine section there are two genera:—
Cephalophusis 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 asCervulus. Messrs. Sclater and Thomas allow thirty-eight species. Thesmallest species do not exceed the dimensions of a Hare. None are really large.
Tetracerosis 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 ofCephalophus. 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.
Ourebiais a less-known genus, larger in size, but with horns of the same character, though longer.
The Grysbok and the Steinbok, genusRaphiceros, 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 isNeotragus 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 isCobus, of which there are eleven species. The Waterbuck,C. ellipsiprymnus, and the Sing-sing,C. unctuosus, are perhaps the best-known species; the former isblackish grey, the latter browner in colour. InC. mariaand 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 toCobus; 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 genusPelea, 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 genusAntilopeis 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, genusSaiga, 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 ofMacraucheniain 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 isS. 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 muchlegend. 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.