Chapter XI: UNGULATA (continued)--ARTIODACTYLA (EVEN-TOED UNGULATES)--SIRENIA (2)
FAM. 7. GIRAFFIDAE.--Undoubtedly the type of a distinct family, Giraffidae, is the genus _Giraffa_. It is characterised by {302} 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 {303} 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[204] 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 (2-1/2 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.[205] 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 {304} 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[206] 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 {305} 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.[207] 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 {306} 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[208]) 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 {307} 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.[209] 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[210] 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 {308} 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[211] 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 {309} 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._ x 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.[212] 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 {310} 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 {311} 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 {312} 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._ x 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. {313}
_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._ x 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 {314} 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.[213]
FIG. 163.--Beatrix Antelope. _Oryx beatrix._ x 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 {315} 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_ ([female]). x 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 {316} 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._ x 1/25.
The last genus of this section or sub-family is the African Eland, genus _Oreas_[214] (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. {317}
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.[215]
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 {318} slightly through domestication. It is, however, said not to cross in a state of nature with the Gaur.[216]
FIG. 166.--Gayal. _Bos frontalis._ x 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 {319} 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,[217] 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._ x 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. {320}
FIG. 168.--Yak. _Bos grunniens._ x 1/15.
FIG. 169.--British Wild Ox. _Bos taurus._ From Vaynol Park, Bangor. x 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 {321} 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._ x 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 {322} 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._ x 1/12. (From _Nature_.)
FIG. 172.--Blanford's Sheep. _Ovis blanfordi._ x 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 {324} 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._ x 1/10.
FIG. 174.--Thar. _Capra jemlaica._ x 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, {325} 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._ x 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 {326} 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 L200.
"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._ x 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, {327} which has the peculiarity of having the shortest cannon bones of any Ruminant. Its name denotes its range.
FIG. 177.--Goral. _Nemorrhaedus goral._ x 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. Loennberg.[218] 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 {328} has a slight naked strip above the nostrils, as in the Sheep, but there is no fissure of the upper lip.
EXTINCT FAMILIES OF ARTIODACTYLA.
The origin of the Artiodactyla is placed by Cope in the family PANTOLESTIDAE,[219] allied to the genus _Protogonodon_ of the Condylarthra. As, however, this family is represented by but a few back teeth and a fragment of the hind-foot, it seems premature to regard it as the necessary starting-point of the Bunodont and Ruminant groups.
FAM. ANTHRACOTHERIIDAE.--This well-known and ancient family consists of creatures of for the most part a Pig-like form, with teeth approaching the selenodont shape, and a complete dentition. The carpals, tarsals, metacarpals, and metatarsals are all free. The toes are four (or five) to each foot, with the outermost beginning to be reduced. These of course are all generalised and primitive characters, pointing nowhere in particular, except, of course, to an Artiodactyle stock, on account of the teeth and the two predominating toes.
The type genus of the family, _Anthracotherium_, is not, as its name might seem to denote, a relic of the Carboniferous period; its remains were found in lignite, which may also show that it was at least semi-aquatic in habit. Its form, however, must have been Pig-like, so at least one would presume from the elongated skull and shortish legs. There were species as great as a Rhinoceros, and smaller forms. The genus began in the Oligocene and continued down to the Pliocene. It is known from Europe, Asia, and America.
The skull is long with a prominent sagittal crest. The facial part is also very long, and the orbits are not closed by a bony ring. The premolars are simple teeth; the molars distinctly bunodont with a tendency in one or two to the selenodont condition. The canines are powerful, as are also the incisors. The scapula has been specially compared with that of the Camel. It has no acromion, which is usually though not always absent in Ungulates. An ally of the present animal, for instance, the Hippopotamus, has the acromion developed. The radius and ulna, the tibia and fibula, are all fully developed. {329}
_Ancodus_ (or _Hyopotamus_, as it has been called) is also Oligocene in range, and its remains have been found in the same countries as have those of _Anthracotherium_. Both genera are indeed closely allied. _Ancodus_ seems to be a more slightly-built creature. The skull looks weaker, but presents much the same features of organisation. In _A. velaunus_, a species found in French rocks, a metacarpal of digit I. was present in the manus, while _A. brachyrhynchus_ had a completely five-fingered manus.
The Miocene genus _Merycopotamus_ (from the lower layers of the Siwalik formation in India) is more distinctly selenodont than the forms already discussed. On this ground it has been placed in a separate sub-family. As, however, in other respects it does not depart from the Anthracotherian type of structure, this proceeding seems to be hardly necessary. There are two species known, of which one, _M. nanus_, is, as its name denotes, a dwarf form.
FAM. CAENOTHERIIDAE.--While the last family consisted of animals rather more akin to the Pigs, the present is more Pecorine in its characters. The molars are selenodont; but as in the Tragulidae the premolars are more of the nature of cutting teeth. The dentition, like that of so many of these early Ungulates, is complete, and the canines are not prominent. The feet are four-toed, the lateral toes not reaching the ground.
The principal genus is the Eocene and Miocene _Caenotherium_. Of this genus there were a considerable number of species all European in range, and of small size--not more than a foot or so in length. Their small size is suggestive of the Chevrotains. In the skull the orbital cavity is nearly or quite surrounded by bone, and the tympanic bulla is large and inflated. A common feature of Artiodactyles, a failure of the nasals and maxillae to meet at the side of the face, is to be seen in this ancient forerunner of the Pecora.
_Plesiomeryx_, also European, and from the same geological horizon, is a very closely allied form.
FAM. XIPHODONTIDAE.--This family consists of slender, small Artiodactyles which are, like the Caenotheriidae, related to the Pecora. They are confined in their range to Europe.
The type genus _Xiphodon_ has selenodont molars and elongated, slender, cutting premolars. The dentition was complete and the canines not highly developed. Like _Caenotherium_, {330} _Xiphodon_ was a hornless creature, but with only two toes, the two lateral digits being represented by the merest rudiments of metacarpals. The other metacarpals were unusually long.
_Amphimeryx_ (also called _Xiphodontotherium_) is much more imperfectly known, but belongs to this family or to that of the Caenotheriidae. _Dichodon_ is another member of the same family.
FAM. OREODONTIDAE.--This family, consisting of numerous genera, is limited to the North American continent. Its range in time is from the Eocene to the Lower Pliocene. The family as a whole is to be distinguished by a number of primitive characters. The dentition is complete; the feet are four- or even five-toed; the orbit is sometimes open behind. The canines of the lower jaw are not more pronounced than the incisors. The characteristics of the group will be further developed by a consideration of some of the principal genera which are included in this family.
_Oreodon_ is a Miocene form about as large as a Peccary. The skull has a short face with a completely-closed orbital cavity. In front of the orbit is a deep pit, not a mere deficiency of ossification, such as occurs in many Artiodactyles. This is placed on the lachrymal bone, and is in fact a lachrymal fossa, such as occurs in other forms. The odontoid process of the axis vertebra is somewhat cheese-taster shaped, as in recent Artiodactyles. There are fourteen dorsal vertebrae and a very large number of caudals. The radius and the ulna are completely separated, as are the carpals. There are five digits to the fore-limbs. The fibula is complete and independent. The hind-foot is four-toed. Several species of the genus are known.
_Merycochoerus_ is an allied Miocene genus. It is more massive in form than the last, but otherwise does not present differences of importance.
_Mesoreodon_ is another genus of this family which presents some curious features of organisation. In the skull and teeth there is nothing very noteworthy, but the hyoid is remarkable. This appendage of the skull is by no means always preserved, and when it is, it might be denied that it belonged to any particular skull. In the present case there appears to be no doubt as to the identity of the bones, which resemble the corresponding bones of the Perissodactyla much more than they do those of other Artiodactyles. Associated with the bones an ossified {331} thyroid cartilage of the larynx was found. As the skull was that of a male, this character may be a sexual one. It is quite comparable to the ossification of the same cartilage in the American monkey _Callithrix_. "The function of the bone," observes Professor Scott,[220] "was probably similar to that performed by the enormously-inflated basihyal of the howling monkeys, and must have given to these animals most unusual powers of voice." Another important anatomical fact about _Mesoreodon_ is the apparent existence of a clavicle. It is of course conceivable that the remains of some other animal have got mixed up with that of the individuals upon which the present genus is founded; but failing that, here is a clavicle in an Ungulate. The spine of the scapula possesses a metacromion. This greater development of the spine of the scapula in Artiodactyles than in Perissodactyles is, it is suggested, to be correlated with the earlier loss of the clavicle in the latter group of Ungulates.
_Cyclopidius_ (synonymous with _Brachymeryx_) is a kind of pug form of _Oreodon_. The skull is short and broad, and the end of the snout a little turned up. The upper incisors are small and drop out early. On each side of the nasals is a large oval vacuity which is perhaps to be compared to the lateral deficiency to be found in other Artiodactyles. One species of this singular-looking form is appropriately called _C. simus_.
Other allied genera are _Merychyus_ and _Leptauchenia_. The former extends as far down as the Lower Pliocene, and is thus one of the newest forms of Oreodontidae.
_Agriochoerus_[221] (Fig. 178) is placed in a separate sub-family from the types which have just been considered. It is Miocene in range. It differs from _Oreodon_ and its closer relatives by the fact that the orbit is open behind and not closed. The most remarkable fact about this creature is that the terminal phalanges of the digits (five in the fore- and four in the hind-feet) being pointed, seem to suggest their encasement with claws rather than hoofs. The pollex, though small, seems to have been opposable. As with other Oreodonts, the molars are selenodont. The premaxillae are toothless--at least in adults, for two teeth are present in the {332} young. There are several species. _Agriochoerus_, like _Oreodon_ and primitive Ungulates in general, had a long tail. The genus thus shows a mixture of ancient and specialised characters.
FIG. 178.--Skeleton of _Agriochoerus latifrons_. x 1/8. (After Wortman.)
The most ancient form of Oreodont is _Protoreodon_. This is Eocene, and became extinct during that period. It had a complete dentition, open orbit, and no lachrymal fossa. The fore-feet were five-toed, the hind four-toed.
FAM. ANOPLOTHERIIDAE.--This family is entirely Eocene in point of time, and is unknown outside Europe. The dentition of the group is complete; the molars are seleno-bunodont, like those of the Anthracotheriidae. The bones of the carpus, tarsus, metacarpus and metatarsus are all free; the toes are four to two in number on each foot. The orbit is widely open behind. The tail is long, as in _Xiphodon_, etc. {333}
These general characters only just serve to differentiate the family; but they illustrate its archaic character, in which it resembles the Xiphodontidae, and even more the Anthracotheriidae. A survey of some of the genera which have been assigned to the family will bring out other features in the organisation of these very ancient Artiodactyles.
_Anoplotherium_ is so called on account of the fact that it is, like all ancient Artiodactyles, without horns or claws. Tusks it might have, but as a matter of fact has not. There are, as in Artiodactyles generally, nineteen dorso-lumbar vertebrae; the long tail has numerous chevrons. The shoulder blade has a well-marked acromion and a distinct coracoid process; it is wide proximally. The bones of the fore-arm and fore-leg are, as is usual in primitive Artiodactyles, separate.
In the skull the chief features, in addition to that mentioned in the definition of the family, are the large size of the paroccipital processes; there is no fossa lachrymalis or deficiency in the side of the face. The animal is three-toed, both in the fore- and hind-limbs. The second toe is nearly as large as the Artiodactyle third and fourth. There are tiny rudiments of the two remaining fingers. The hind-foot is also three-toed, and there is a trace of the hallux. The fingers are so widely separated and divergent from each other that it has been suggested that the animal had webbed feet and inhabited marshes, in which it swam by the aid of its long tail. The creature was the size of a Tapir.
Closely resembling _Anoplotherium_ are a number of other genera.
_Diplobune_ ( = _Hyracodontotherium_) was much like the last, but was a more delicately-formed animal. The fingers and toes (three of each) end in such sharply-pointed phalanges that claws seem to be almost suggested. There are several species of this genus. _Dacrytherium_ differs by the presence of a lachrymal fossa.
_Dichobune_ has four-toed extremities, of which the lateral ones are more slender and shorter than the two middle ones. As in other Anoplotheriidae, the anterior premolars are furnished with a sharp cutting edge.
ORDER V. SIRENIA.
Aquatic Mammalia, with but few scattered hairs; hind-limbs absent; fore-limbs paddle-shaped; tail flattened, and either {334} Whale-like or rhomboidal to circular in form. Nostrils on upper surface of not specially-elongated snout. Clavicles are absent. The scapula has the normal mammalian form, with a well-developed and roughly median spine. The bones of the arm and hand articulate together, as in land animals; the phalanges show at most traces of increase in number above the normal. Pelvis represented by a vestige, more highly developed in some fossil than in recent forms. Stomach complex, consisting of several chambers. Lungs simple and not lobulated. Diaphragm oblique and very muscular. Brain peculiar in form and but slightly convoluted. Testes abdominal. Teats two, and pectoral in position. Placenta non-deciduous and zonary.[222]
This limited group consists of purely aquatic forms, which are both marine and fresh-water in their proclivities. They have been placed in the immediate vicinity of the Whales; but it is now believed by most zoologists that the likenesses which they undoubtedly show to the Cetacea are of an adaptive kind and related to their similar mode of life. The group is a readily-definable one. Externally they are marked by their dark coloration, somewhat Whale-like though of clumsier build, and by the total absence of external ears and hind-limbs; the latter are, however, as will be pointed out shortly, marked by certain rudimentary bones. There is a flattened tail, which in the Dugong and _Rhytina_ is precisely like that of a Whale. It is interesting to note that the former genus, whose tail is, judging it at least by the standard of the Whales, more completely modified for the aquatic life, should also show other features which indicate their longer life as marine creatures. For the flippers are more Whale-like in that the fore-arm is completely enclosed within the body, or nearly so, and the nostrils have a more decidedly superior position than in the Manatee. The fore-limbs of this group, as may be inferred from what has just been said, are flipper-like; but, contrary to what we find in Whales, the phalanges do not as a rule show any traces of multiplication, so characteristic a feature of the Cetacean hand, and the individual bones are connected by well-formed joints. Beneath the thick skin, which is sparsely provided with stout hairs in the Dugong, is a layer of blubber. Dr. Murie has called attention to the fact that this layer in the {335} Manatee[223] differs from the blubber of the Whale in that there is no free oil anywhere.[224]
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The Cambridge natural history, Vol. 10 (of 10)Chapter XI: UNGULATA (continued)--ARTIODACTYLA (EVEN-TOED UNGULATES)--SIRENIA (2)
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