This genus (which has also received the name of _Lophotragus_) is

represented by a small Deer (Fig. 127) from China of about the same

size as the Indian Muntjac. The male has minute simple antlers and very

large canine teeth. There are no supraorbital glands, nor is there a

tufted gland on the metatarsus. The limbs have the same peculiarities

as in _Cervulus_, but the mesocuneiform may also ankylose with the

ectocuneiform, and traces of the metacarpals may remain. The hair is

coarse and somewhat quill-like.

[Illustration: FIG. 127.—Male of _Elaphodus michianus_. From Sclater

_Proc. Zool. Soc._ 1876, p. 273.]

_Cervus._[199]—The great majority of the Deer of the Old World may be

included in this large genus, which is one not easy of definition. The

antlers of the male are, however, large, and two or three times the

length of the head, and may be either rounded or palmate; the canines are

never large; the ectocuneiform of the tarsus remains distinct from the

naviculo-cuboid; the lateral digits are represented by their phalanges;

and the skull does not carry prominent frontal ridges. Vertebræ: C 7, D

13, L 6, S 4, C 11-14. The size of the lachrymal fossa and vacuity, and

the degree of inflation of the auditory bulla, are subject to variation

in the different groups into which the genus may be divided.

The _Rusine_ group is characteristic of the Oriental region, where it is

typically represented by the Sambur (_C. aristotelis_) of India, Burma,

and China. The antlers are rounded, and often strongly grooved, without a

bez tine, and with the beam simply forked at the extremity, upright, and

but slightly curved; the angle formed by the brow tine, which rises close

to the burr, being acute. The molars are markedly hypsodont, with small

accessory columns. The lachrymal fossa is deep and the vacuity large; the

auditory bulla is slightly inflated and rugose. Tail moderate; neck maned.

The Sambur, which is abundant in hilly districts, is a fine animal,

standing nearly 5 feet in height, and of massive build; the general

colour being deep brown. _C. equinus_, of Borneo, Sumatra, and Singapore,

_C. swinhoei_, of Formosa, _C. philippinus_, and _C. alfredi_ of the

Philippines, are closely allied species, of which the two latter are

of smaller dimensions. The Indian Hog Deer (_C. porcinus_) is a still

smaller form, not larger than the Roe. _C. hippelaphus_ of Java, _C.

timoriensis_, and _C. moluccensis_ are distinguished by the posterior

branch of the beam of the antler being considerably larger than the

anterior.

The _Rucervine_ group is another strictly Oriental one, and is

represented by the Swamp Deer (_C. duvaucelli_) of India, the closely

allied _C. schomburgki_ of Siam, of which the antlers are shown in Fig.

119 (p. 309), and _C. eldi_ of Burma and Hainan. The beam of the antler

is somewhat flattened, and more curved than in the Rusine group; the

large brow tine is given off from the beam at an obtuse angle and curves

upwards; the beam bifurcates into two branches, which again divide. Skull

as in the Rusine group, but relatively narrower. Tail short; neck maned.

The Swamp Deer is somewhat smaller than the Sambur, and of a full

yellowish colour. Fossil representatives of this group occur in the

Pliocene of India.

The _Elaphurine_ group is represented only by the very aberrant _C.

davidianus_ of Northern China. In size and proportions this species

approximates to the Swamp Deer, but the antlers are peculiar in rising

straight from the brow and then giving off a long and straight back tine

(correlated by Sir V. Brooke with the posterior branch of the Rusine

antler); the summit of the beam is forked, and in old individuals the

two tines of the fork may again branch. Nasals long, and much expanded

between the lachrymal vacuities, of which they form the inner border;

lachrymal fossa large and deep. Tail long; neck maned.

The _Axine_ group includes only the well-known Axis of India, readily

distinguished by the white spots with which the body is marked. Antlers

of a Rusine type, the beam being much curved, and the brow tine usually

given off at an acute or right angle. Molars very hypsodont. The

coloration of the Axis is more brilliant than that of any other member of

the family.

Here may be noticed a group of Deer mainly characteristic of the

eastern Palæarctic region, frequently known as the _Pseudaxine_ group,

which appears to connect the Axine with the Elaphine type. Well-known

representatives of this group are _C. sika_ (Fig. 128) of Japan, _C.

mantchuricus_ of China, and _C. taëvanus_ of Formosa. The antlers have a

brow and tres tine, and then a forked beam, of which the posterior tine

is the smaller. The lachrymal vacuity and fossa are of moderate size;

and the auditory bulla is only moderately inflated, and quite smooth

externally. Tail moderate; neck maned. In summer the coat is spotted, but

is plain in winter. A herd of _C. sika_ have been acclimatised in Ireland

by Viscount Powerscourt, at Powerscourt, County Wicklow. A number of Deer

from the Pliocene of Europe, such as _C. perrieri_ and _C. etueriarum_,

appear to be allied both to the Pseudaxine and Axine groups.

[Illustration: FIG. 128.—The Japanese Deer (_Cervus sika_). From Lord

Powerscourt, _Proc. Zool. Soc._ 1884, p. 209.]

The _Elaphine_ or typical group is at once characterised by the presence

of a bez tine to the antlers (Fig. 129), in which the beam is rounded,

and splits up near the summit into a larger or smaller number of snags,

often arranged in a cup-like manner. Skull as in the preceding group. All

the species large. The Red Deer, _C. elaphus_, which is dark brown in

colour, with a light patch on the rump, inhabits Europe, Western Asia,

and Northern Africa—the so-called Barbary Deer not being specifically

distinct. A full-grown Scotch Stag is fully 4 feet in height at the

withers. The antlers are shed between the end of February and the early

part of April; old animals shedding earlier than younger ones. The young,

which (as in all the members of the genus except some of the Rusine

species) are spotted, are born at the end of May or the beginning of

June. The points on the antlers increase in number with the age of the

creature, and when twelve are present it is known in Scotland as a “royal

stag.” This number, however, is sometimes exceeded, as in the case of a

pair of antlers, weighing 74 lbs., from a stag killed in Transylvania,

which had forty-five points. The antlers during the second year consist

of a simple unbranched stem, to which a tine or branch is added in each

succeeding year, until the normal development is attained, after which

their growth is somewhat irregular. Many of the antlers dug up in British

peat-beds (as Fig. 118) are larger than those of living individuals,

and in the cave-deposits of England and the Continent antlers are met

with rivalling those of the Wapiti in size; these large fossil antlers

probably indicating the ancestral form from which the Red Deer and

several of the allied species are descended.

[Illustration: FIG. 129.—Head of the Wapiti (_Cervus canadensis_).]

The North American Wapiti (_Cervus canadensis_, Fig. 129), the Persian

Maral (_C. maral_), the Kashmir Stag (_C. cashmeerianus_), as well as

_C. affinis_ of Tibet, are all closely allied to the Red Deer, but are

of larger size, this being especially the case with the first two. A

fine example of the antlers of the Wapiti is shown in the accompanying

woodcut, and exhibits the absence of a cup at the surroyals, by which

this species is distinguished from the Red Deer.

The last, or _Damine_ group of existing Deer includes the Common and the

Persian Fallow Deer. These are readily characterised by the palmation

of the antlers in the region of the surroyals and the spotted coat.

The Common Fallow Deer (_C. dama_) stands about three feet in height.

The Persian Fallow Deer (_C. mesopotamicus_) is very closely allied,

differing only in its slightly larger size and the form of the antlers,

the two breeding together. The common species, although now kept in

English parks, does not appear to be a native of this country, having

probably been introduced from the regions bordering the Mediterranean.

The fur is of a yellowish-brown colour (whence the name “fallow”), marked

with white spots; there is, however, a uniformly dark brown variety found

in Britain. The bucks and does live apart, except during the pairing

season; and the doe produces one or two, and sometimes three fawns at

a birth. The Fallow Deer from the Pleistocene and Pliocene deposits

of the East Coast described under the names of _C. browni_ and _C.

falconeri_ appear to have been closely allied to the existing species.

The remarkable _C. verticornis_, of the Norfolk Forest-bed, is regarded

as an aberrant member of this group, in which the antlers are very short

and thick, with the brow tine cylindrical and downwardly curved, and the

beam expanded above the tres tine into a crown with two points.

The extinct Irish Deer (_Cervus giganteus_), of which the skeleton is

shown in the woodcut (Fig. 130), is the only representative of the

_Megacerotine_ group. The antlers, which may have a span of over 11

feet, are enormously palmated, and have a bifurcated brow tine, a small

bez tine, and a third posterior tine. The skeleton measures upwards of

6 feet at the withers. Remains of this species are especially common in

the peat-bogs of Ireland, but are also met with in Pleistocene deposits

over a large part of Europe. In addition to the forms already mentioned

there are many other fossil species of _Cervus_, some of which, like the

English Pleistocene _C. sedgewicki_, cannot be included in any of the

existing groups. There is no conclusive evidence of the existence of any

species of _Cervus_ before the Lower Pliocene period.

=Telemetacarpalia.=—This section includes all the Deer of the New World,

together with some Old World forms, and is characterised by retaining the

distal extremities of the lateral (second and fifth) metacarpals. With

the exception of _Alces_, _Capreolus_, and _Hydropotes_ (which are either

partly or entirely Old World types), the vomer is so much ossified as to

divide the posterior bony nares into two distinct orifices (Fig. 132).

[Illustration: FIG. 130.—Skeleton of the Gigantic Irish Deer (_Cervus

giganteus_). After Owen.]

_Rangifer._[200]—The Reindeer, or Caribou as it is termed in North

America, is the sole representative of the genus _Rangifer_, which is

sufficiently distinguished from all its allies by the presence of antlers

in both sexes. The lachrymal vacuity is small. This animal is distributed

over the northern parts of Europe, Asia, and America; the differences

which may be observable in specimens from different regions not being

sufficient to allow of specific distinction. The Reindeer is a heavily

built animal, with short limbs, in which the lateral hoofs are well

developed, and the cleft between the two main hoofs is very deep, so that

these hoofs spread out as the animal traverses the snow-clad regions in

which it dwells. The antlers (Fig. 131) are of very large relative size.

There is a bez as well as a brow tine, which are peculiar in being either

branched or palmated. In the American race (Caribou), as well as in some

of the specimens found fossil in the English Pleistocene (Fig. 131), one

of the brow tines is generally aborted to allow of the great development

of the other. The dentition of the Reindeer is frequently remarkable

for the very small size of the posterior lobe of the last lower molar.

Vertebræ: C 7, D 14, L 5, S 5, C 11.

[Illustration: FIG. 131.—Skull and antlers of the Reindeer (_Rangifer

tarandus_), from an English Pleistocene deposit. _br_, Brow tine; _bz_,

bez tine. (After Owen.)]

The Reindeer has long been domesticated in Scandinavia, and is of

especial value to the Laplanders, whom it serves as a substitute for

the Horse, Cow, Sheep, and Goat. It is capable of drawing a weight of

300 lbs., and its fleetness and endurance are remarkable. Harnessed to

a sledge it will travel without difficulty 100 miles a day over the

frozen snow, on which its broad and deeply cleft hoofs are admirably

adapted for travelling. During the summer the Lapland Reindeer feeds

chiefly on the young shoots of the willow and birch; and since at this

season migration to the coast seems necessary to the well-being of this

animal, the Laplander, with his herds, sojourns for several months in

the neighbourhood of the sea. In winter its food consists chiefly of the

so-called reindeer-moss and other lichens which the animal makes use

of its hoofs in seeking for beneath the snow. The wild Reindeer grows

to a much greater size than the tame breed; but in Northern Europe the

former are being gradually reduced through the natives entrapping and

domesticating them. The tame breed found in Northern Asia is much larger

than the Lapland form, and is there used to ride on. Remains referable

to the existing species are found in the cavern and other Pleistocene

deposits of Europe.

[Illustration: FIG. 132.—Hinder part of the base of the cranium of the

Virginian Deer (_Cariacus virginianus_). From Garrod, _Proc. Zool. Soc._

1877, p. 13.]

_Alces._[201]—The Elk or Moose (_Alces machlis_) has the same general

distribution as the Reindeer, and is likewise the single existing

representative of its genus. It is the largest existing member of the

family, attaining sometimes a height of 8 feet at the withers. The

antlers (Fig. 133) have neither brow nor bez tine, but form an enormous

basin-shaped palmation, primarily composed of an anterior and a posterior

branch; their weight may be as much as 60 lbs. The nasal bones are very

short, and the narial aperture of great size. The Elk is covered with a

thick coarse fur of a brownish colour, longest on the neck and throat.

Its legs are long and its neck short, and as it is thus unable to feed

close to the ground, it browses on the tops of low plants, the leaves of

trees, and the tender shoots of the willow and birch. Its antlers attain

their full length by the fifth year, but in after years they increase in

breadth and in the number of snags, until fourteen of these are produced.

Although spending a large part of their lives in forests, Elks do not

suffer much inconvenience from the great expanse of their antlers, as in

making their way among trees they are carried horizontally to prevent

entanglement with the branches. Their usual pace is a shambling trot, but

when frightened they break into a gallop. The natural timidity of the

Elk forsakes the male at the rutting season, and he will then attack

whatever animal comes in his way. The antlers and hoofs are his principal

weapons, and with a single blow from the latter he has been known to kill

a wolf. The female often gives birth to two fawns, and with these she

retires into the deepest recesses of the forest, the young remaining with

her till their third year. The Elk ranges, but in scanty numbers, over

the whole of Northern Europe and Asia, as far south as East Prussia, the

Caucasus, and North China, and over North America from the New England

States westward to British Columbia. Fossil species are found in the

Pleistocene deposits of Europe.

_Cervalces._[202]—A remarkable extinct Deer from the Pleistocene of North

America, described as _Cervalces_, appears in some respects (although

a true Telemetacarpalian) to connect _Alces_ with _Cervus_. Thus the

palmated antlers are divided into anterior and posterior branches, but

below this division there are two tines apparently corresponding to the

bez and posterior tines of _Cervus giganteus_ (Fig. 130).

[Illustration: FIG. 133.—Head of Elk (_Alces machlis_).]

_Capreolus._[203]—Antlers (in the existing species) less than twice the

length of the head, usually with three tines on each. Brow tine developed

from the anterior surface of the upper half of the antler, and directed

upwards. Lachrymal vacuity small. Premaxillæ not always articulating with

nasals. Auditory bullæ slightly inflated, rugose externally. Vertebræ: C

7, D 13, L 6, S 6, C 8. Tail very short. Glands in fore feet rudimentary;

large in hind feet.

The Roe, or Roe Deer (_Capreolus caprea_), is a small form distributed

over Europe and Western Asia, being one of the species found in the

British Isles. The male is somewhat over two feet in height at the

withers, of a dark reddish-brown colour in summer, with a white patch

on the rump. The small antlers are approximated at their bases, and

consist of a rugged beam rising vertically for some distance, then

bifurcating, and the posterior branch again dividing. The Roe dates from

the Pleistocene period. Extinct Deer from the Continental Pliocene have

been provisionally referred to _Capreolus_.

_Hydropotes._[204]—No antlers in either sex. Lachrymal fossa deep and

short (Fig. 134); lachrymal vacuity of moderate size. Orbits small and

but slightly prominent. Auditory bulla much inflated. Angle of mandible

much produced backwardly (Fig. 134); alveolar margins of mandible in

diastema sharp and everted. Canines of male very large, and slightly

convergent. Vertebræ: C 7, D 12, L 6, S 4, C 10. No tufts on metatarsals.

Foot glands small in fore feet, deep in hind ones.

[Illustration: FIG. 134.—The left lateral view of the skull of a male

Chinese Water Deer (_Hydropotes inermis_), with the wall of the maxilla

cut away to show the root of the canine. ½ natural size. (From Sir V.

Brooke, _Proc. Zool. Soc._ 1872, p. 524.)]

[Illustration: FIG. 135.—Upper surface of the brain of _Hydropotes

inermis_. (From Garrod, _Proc. Zool. Soc._ 1877, p. 792.)]

The Chinese Water Deer (_H. inermis_) is the sole representative of this

genus. In the absence of antlers and the large canines of the male it

resembles _Moschus_, although very different in other respects. Thus

the brain (Fig. 135) has the hemispheres much convoluted, as in other

_Cervinæ_, and approximates to that of _Pudua_; while the placenta

and viscera likewise agree with those of the true Deer. In the total

absence of any ossification of the vomer to divide the posterior nares

_Hydropotes_ resembles _Capreolus_ and differs from all the following

genera. The Chinese Water-Deer is nearly of the same size as the Indian

Muntjac. It has short legs and a long body, the hair covering the latter

being of a light reddish-brown. It is a remarkably prolific animal,

differing from all other Deer in producing five or six young at a time.

The mandible of a ruminant from the Middle Miocene of Gers in France,

described under the name of _Platyprosopus_, presents such a marked

resemblance to _Hydropotes_ in the form of the angle as to suggest a more

or less intimate affinity.

_Cariacus._[205]—Skull (Fig. 132) with the vomer dividing the posterior

nares into two distinct chambers; premaxillæ not reaching nasals. Antlers

never greatly exceeding the length of the head. Lachrymal vacuity very

large, and lachrymal fossa small. Auditory bullæ slightly inflated.

Vertebræ: C 7, D 13, L 6, S 4, C 13. Tail long or short. Colour uniform

in adult.

This genus, which agrees with the Reindeer in the division of the

posterior nares by the ossified vomer, comprises a number of species

confined to the New World, none of which attain very large dimensions,

and the antlers of which are relatively smaller than in the existing

species of _Cervus_. The genus may be divided into groups.

The typical _Cariacine_ group, as represented by _C. virginianus_, has

well-developed antlers, with a short brow tine rising from the inner side

of the beam, and directed upwards, and several branches; a long tail; and

no upper canines. In this species, as well as in _C. mexicanus_ and other

forms, the antlers do not divide dichotomously, and the lachrymal fossa

is of moderate depth. The Mule Deer (_C. macrotis_) of North America is

distinguished by the dichotomous branching of the antlers and the deeper

lachrymal fossa. The Virginian Deer is somewhat smaller than the Fallow

Deer, and of a uniform reddish-yellow colour in summer, and light gray in

winter.

The _Blastocerine_ group of South America is represented by _C.

paludosus_ and _C. campestris_, and has dichotomous antlers, with no

brow tine, and the posterior branch the larger, a short tail, and no

upper canines. The _Furciferine_ group includes _C. chilensis_ and

_C. antisiensis_, confined to western South America. The antlers are

not longer than the head, with a large anterior tine curving forwards

at right angles to the simple posterior one. Auditory bullæ slightly

inflated, and rugose. Upper canines may be present. The species are of

medium size. _C. clavatus_, of Central America, while resembling this

group in the characters of the skull and the arrangement of the hair on

the face, agrees with the next one in having simple spike-like antlers.

The South American _Coassine_ group comprises the small forms known as

Brockets, in which the antlers form simple spikes not exceeding half the

length of the head. Some six species are known.

Remains of _Cariacus_, mostly or entirely referable to existing species,

are of common occurrence in the Brazilian cave-deposits. _Blastomeryx_,

of the Pliocene of North America, is believed to be an allied type.

_Pudua._[206]—Antlers in the form of minute simple spikes. Distinguished

from the Coassine group of _Cariacus_ by the articulation of the

premaxillæ with the nasals (as in the _Furciferine_ group), and the

coalescence of the ectocuneiform with the naviculo-cuboid, as well as by

various external characters. No upper canines. Represented only by the

very small _P. humilis_ of the Chilian Andes.

_Extinct Genera._—In the European and other Tertiary deposits several

genera of extinct _Cervidæ_ occur, of which the more important may

be briefly mentioned. _Amphitragulus_, of the Lower Miocene of the

Continent, has four lower premolars, brachydont molars, and no antlers;

the largest species being somewhat bigger than the Musk-Deer. The closely

allied _Palæomeryx_ (_Dremotherium_ or _Micromeryx_) generally has but

three lower premolars, and the brachydont upper molars (Fig. 122), like

those of _Amphitragulus_, want the small accessory inner column[207]

found in modern Deer. In _P. feignouxi_, of the Lower Miocene, the

lateral metacarpals, although slender, were complete, and the males had

large canines, but no antlers. _P. furcatus_, of the Middle Miocene, had

small antlers, and the canines appear to have been reduced in size. This

genus, besides being represented in the European Miocene, also occurs in

the Pliocene of India and China; some of the species being as large as

the Red Deer.

_Family_ GIRAFFIDÆ.

In the existing genus the frontal appendages consist of a pair of

short, erect, permanent bony processes placed over the union of the

frontal and the parietal bones, ossified from distinct centres, though

afterwards ankylosed to the skull, covered externally with a hairy

skin, present in both sexes, and even in the new-born animal. Anterior

to these is a median protuberance on the frontal and contiguous parts

of the nasal bones, which increases with age, and is sometimes spoken

of as a third horn. Skull with a lachrymal vacuity. No upper canines.

Molars brachydont, with rugose enamel; the upper ones having no inner

accessory column. Lateral digits entirely absent on both fore and hind

feet, even the hoofs not developed. Humerus with double bicipital groove.

Vertebræ: C 7, D 14, L 5, S 3, C 20. Gall-bladder generally absent. Male

reproductive organs and placenta of a Bovine type. Dentition: _i_ ⁰⁄₃,

_c_ ⁰⁄₁, _p_ ³⁄₃, _m_ ³⁄₃.

[Illustration: FIG. 136.—The Giraffe (_Giraffa camelopardalis_).]

_Giraffa._[208]—The Giraffe (_G. camelopardalis_) is the sole existing

representative of the genus, now confined to the Ethiopian region.

In addition to the characters noticed above, the Giraffe is characterised

by its great size and peculiar proportions; the neck and limbs being

of great length, and the back inclining upwards from the loins to the

withers.

To produce the extremely elongated neck the seven cervical vertebræ are

proportionately long, which gives a somewhat stiff and awkward motion

to the neck. The ears are large, the lips long and thin, the nostrils

closable at the will of the animal, the tongue very long and extensile,

and the tail of considerable length, with a large terminal tuft. An adult

male may have a total height of 16 feet. The coloration consists of large

blotches of darker or lighter chestnut-brown on a paler ground, the

lower limbs and under parts being of a uniform pale colour. The Giraffe

feeds almost exclusively on the foliage of trees, showing a preference

for certain varieties of mimosa, and for the young shoots of the prickly

acacia, for browsing on which its prehensile tongue and large free lips

are specially adapted. It is gregarious in its habits, living in small

herds of about twenty individuals, although Sir S. Baker, who hunted it

in Abyssinia, states that he has seen as many as a hundred together.

Fossil species of _Giraffa_ occur in Pliocene deposits over Greece,

Persia, India, and China, thus affording one of many striking instances

of the former wide distribution of the generic types now confined to the

Ethiopian region.

_Allied Extinct Types._—The Pliocene deposits of many parts of the Old

World yield remains of a number of large Ruminants which show such

evident signs of affinity with the Giraffe that it is difficult to draw

up a definition by which they can be separated in characters of family

value from that genus. On the other hand, some of these forms approximate

in the characters of the skull to some of the brachydont members of the

_Bovidæ_, although it is quite clear from the nature of the cranial

appendages that they cannot be included in that family. All these forms

have brachydont molars, with rugose enamel, like those of the Giraffe;

while several of them have limb-bones approximating to those of the

latter—the humerus, when known, having a double bicipital groove. The

nature of the cranial appendages (when present) is not fully understood,

but it appears that in some cases these approximated more to the type of

an antler than to that of a horn; although, from the absence of a “burr,”

they appear never to have been shed. A gradual diminution in the length

of the limbs and neck can be traced from the more Giraffoid to the more

Bovoid forms of this extinct group; and it is manifest that if these

animals be included in the _Giraffidæ_ the definition of that family as

given above must be somewhat modified. Only brief mention can be made of

the more important genera.

The imperfectly known _Vishnutherium_, of the Pliocene of India and

Burma, seems to make the nearest approach to the Giraffe, but the limbs

and cervical vertebræ were decidedly shorter, although of a similar

slender type. _Helladotherium_, of the Pliocene of Greece and India, is

represented by a species of considerably larger size than the Giraffe,

with no appendages or lachrymal vacuity to the skull, and with shorter

and stouter limbs and neck.

_Hydaspitherium_, _Bramatherium_, and _Sivatherium_ are Indian genera,

characterised by the presence of large palmated and antler-like cranial

appendages, varying considerably in arrangement. The former genus has a

large lachrymal vacuity which is absent in the two latter. In the first

and second genera all the appendages rise from a common base; but in

_Sivatherium_ there is a pair of simple horn-like projections on the

orbits in addition to the posterior palmated antlers. _Sivatherium_ was

an animal of huge bulk, being the largest known representative of the

Pecora.

Another apparently allied type is _Samotherium_, of the Pliocene of

the Isle of Samos, which appears also to have some affinity with the

Antelopes. The skull is nearly as large as that of the Giraffe, and is

of the same elongated shape, although depressed between the conical

horn-cores, which rise vertically above the orbits, and without a median

bony prominence on the frontals. The horn-cores form mere processes of

the frontals. The diastema and the mandibular symphysis are shorter than

in the Giraffe, and the latter is less deflected. The teeth, although

larger, are almost indistinguishable from those of the Giraffe, the only

well-marked difference being that the last lower premolar has a double in

place of a single postero-internal column.

_Family_ ANTILOCAPRIDÆ.

Closely allied to the _Bovidæ_, but the horns deciduous and branched.

_Antilocapra._[209]—The Prong-buck, or Prong-horned Antelope

(_Antilocapra americana_), as the single existing member of this family

is called, is an animal of nearly the same size as the Fallow Deer, but

of a lighter and more graceful build. It is an inhabitant of the prairies

of North America, where it is one of the few representatives of the

Cavicorn Pecora. The bony horn-cores are unbranched, and form vertical,

blade-like projections immediately above the orbit. The horns themselves

are compressed, and nearly one foot in length, having a gentle backward

curvature, the short branch arising somewhat above the middle of its

height, and inclining forwards. When the horn is about to be cast off

it becomes loosened, and a new one is formed upon the bony core beneath

it. The ears are long and pointed, and the tail is short. The neck has a

thick mane of long chestnut-coloured hair, and there is a white patch on

the rump.

_Family_ BOVIDÆ.

Frontal appendages, when present, in the form of non-deciduous horns.

Molars frequently hypsodont. Usually only one orifice to the lachrymal

canal, situated inside the rim of the orbit. Lachrymal bone almost always

articulating with the nasal. Canines absent in both sexes. The lateral

toes may be completely absent, but more often they are represented by

the hoofs alone, supported sometimes by a very rudimentary skeleton,

consisting of mere irregular nodules of bone. Distal ends of the lateral

metapodials never present. Gall-bladder almost always present. The number

of cotyledons in the placenta generally varies from 60 to 100; whereas

in the _Cervidæ_ the number is usually from 5 to 12, _Capreolus_ and

_Hydropotes_ having the fewest. In _Giraffa_ the number is upwards of

180. The nature of the horns and horn-cores has been already explained;

in the majority of genera these appendages are present in both sexes,

although much larger in the male (see p. 310).

The _Bovidæ_, or hollow-horned Ruminants (Cavicornia), form a most

extensive family, with members widely distributed throughout the Old

World, with the exception of the Australian region; but in America they

are less numerous, and confined to the Arctic and northern temperate

regions, no species being indigenous either to South or Central America.

There is scarcely any natural and well-defined group in the whole

class which presents greater difficulties of subdivision than this;

consequently zoologists are as yet very little agreed as to the extent

and boundaries of the genera into which it should be divided. For the

present the genera provisionally adopted may be arranged under a number

of sections or groups, which some writers regard as subfamilies. The

series may be commenced with the Antelopes, the greater number of which

are now characteristic of the Ethiopian region.

_Alcelaphine Section._—Includes large African Antelopes, of which the

type genus ranges into Syria; generally characterised by their great

height at the withers as compared with the rump. Skull with large frontal

sinuses, extending into the horn-cores, and the horns lyre-shaped or

recurved, and more or less approximated at the base. No large pits at

apertures of supraorbital foramina in frontals; upper molars hypsodont

and narrow. Horns in both sexes. General colour mostly uniform.

_Alcelaphus._[210]—If _Damalis_ be included, this genus is represented

by some nine or ten living species. Head more or less long and narrow,

with the muffle moderately broad and naked. Nostrils approximated, edged

with stiff hairs. Horns compressed and ringed at the base, more or less

lyrate, and bent back at the tips. Hoofs small. Tail of moderate length,

and heavy. Two mammæ.

[Illustration: FIG. 137.—The Harte-beest (_Alcelaphus caama_).]

In the typical forms, such as the Bubaline Antelope (_A. bubalinus_), the

Harte-beest (_A. caama_, Fig. 137), and the Tora Antelope (_A. tora_,

Fig. 138), the horns, which present the peculiar curvature shown in the

figures, are situated on a crest at the vertex of the skull, and the

facial portion of the cranium is greatly elongated. The Harte-beest,

which is found throughout Central and Southern Africa, stands nearly 5

feet high at the withers, and is a somewhat ungainly looking animal, with

short hair, which is grayish-brown above and nearly white beneath. In

the Pliocene of the Siwalik Hills in Northern India there occur remains

of an _Alcelaphus_ (_A. palæindicus_) in which the skull had the long

facial portion characteristic of the typical group, while the horns

approximate to those of the Bontebok. The Blessbok (_A. albifrons_)

and Bontebok (_A. pygargus_), belonging to the genus _Damalis_ of many

authors, have the facial portion of the skull shorter, the horns situated

more in advance of the plane of the occiput, and inclining regularly

backwards. Of the Blessbok Mr. C. J. Anderson observes that “it is of a

beautiful violet colour, and is found in company with black Wildebeests

and Springboks in countless thousands on the vast green plains of short

crisp, sour grass occupying a central position in South Africa. Cattle

and horses refuse to pasture on the grassy products of these plains,

which afford sustenance to myriads of this Antelope, whose skin emits a

most delicious and powerful perfume of flowers and sweet-smelling herbs.”

Since the time this was written these Antelopes have been greatly reduced

in number. _A. (Damalis) hunteri_, from East Africa, appears to be allied

to _A. senegalensis_, but in the more elongated facial portion of the

skull approximates to the Harte-beest, and thus confirms the view that

_Damalis_ should not form a distinct genus.

[Illustration: FIG. 138.—Head of _Alcelaphus tora_. From Sclater, _Proc.

Zool. Soc._ 1873, p. 762.]

_Connochætes._[211]—Head short and massive, with the muffle very broad

and bristly. Nostrils widely separated, hairy within. Horns on the

vertex of the skull, immediately over the occiput, approximated at base,

cylindrical, bent outwards, and recurving upwards at the tip. Extremities

of premaxillæ much expanded laterally, and firmly ankylosed. Vertebræ:

C 7, D 14, L 6, S 4, C 16. Hoofs very narrow. Tail very long, covered

throughout with long hairs. Four mammæ. Two species, _C. taurina_ and

_C. gnu_ (Fig. 139), both from South Africa. The former, or Brindled

Gnu, is distinguished by the absence of long hair on the face, the black

(instead of white) tail, and the presence of dark vertical streaks on the

shoulders; it is never found to the south of the Orange River.

The White-tailed Gnu stands about 4 feet 6 inches at the withers. These

animals were formerly found in large herds, and are remarkable not only

on account of their peculiar form, but also for their grotesque actions

when alarmed. Some interesting observations have recently been published

upon the mode of development of the horns of the Gnu,[212] from which

it appears that in very young individuals the horns are straight and

divergent, situated some distance below the vertex of the head, and

separated by a wide hairy interval. These young horns form the straight

tips of those of the adult, the basal downwardly curved portion being

subsequently developed. In the fully adult animal the base of the horns

forms a helmet-like mass on the forehead which completely obliterates the

hairy frontal space of the young.

[Illustration: FIG. 139.—The White-tailed Gnu (_Connochætes gnu_).]

_Cephalophine Section._—Small or medium-sized African and Indian

Antelopes, with simple horns present only in the males, a more or less

elongated suborbital gland, a lachrymal depression in the skull, and

square-crowned upper molars (Fig. 140). Lateral hoofs well developed.

_Cephalophus._[213]—One pair of horns, arising far back on the frontals,

conical, short, angulated at the base, and erect or recurved. Suborbital

gland opening in the form of a slit, or as a row of pores. Auditory bulla

divided by a distinct septum. Muffle large and moist. Tail very short.

Head tufted. Upper molars of larger species with an accessory internal

column. Dorsal vertebræ fourteen in number. Some sixteen species,

confined to southern and tropical Africa.

The Duikerboks, as the members of this genus are called, are among

the most graceful of the African Antelopes, the smallest species not

being larger than a rabbit. The West African _C. sylvicultor_ and _C.

longiceps_ are the largest species.

_Tetraceros._[214]—Two pairs of conical horns, of which the anterior are

much the smaller. Suborbital gland elongated, and lachrymal fossa very

large. Upper molars (Fig. 140) without accessory internal column. One

existing Indian species (_T. quadricornis_).

[Illustration: FIG. 140.—Palatal and outer aspects of the three right

upper premolars and first molar of the Four-horned Antelope (_Tetraceros

quadricornis_). From the _Palæontologia Indica_.]

The Four-horned Antelope is found throughout the peninsula of India in

jungle. The general colour is brown, lighter beneath and on the inside of

the limbs. Remains of this species are found fossil in the cave-deposits

of Madras, and a small Ruminant from the Pliocene of the Siwalik Hills

has been provisionally referred to this genus.

_Cervicaprine Section._—Small or large Antelopes now confined to the

Ethiopian region, with horns present only in the males, lachrymal vacuity

generally large, more or less distinct pits at the apertures of the

supraorbital foramina in the frontals, and narrow upper molars in which

there is no accessory internal column.

_Neotragus._[215]—Distinguished from the next genus by having the crown

of the head tufted, muzzle hairy, premaxillæ long and reaching the

lachrymals, nasals very short, mesethmoid much ossified, third lobe of

last lower molar either absent or very small, and the hinder lobe of the

corresponding upper molar much reduced.

Three species, Salt’s Antelope (_N. saltianus_), from Abyssinia, and

also _N. kirki_ and _N. damarensis_; the two latter having a small

third lobe to the last molar. Writing of the first-named species, Mr.

W. T. Blanford[216] observes that “the _Beni-Israel_, or _Om-dig-dig_,

one of the smallest Antelopes known, abounds on the shores of the Red

Sea and throughout the tropical and subtropical regions of Abyssinia.

It is occasionally, but rarely, found at higher elevations; I heard of

instances of its being shot both at Serafie and Dildi, but it is not

often seen above about 6000 feet. It inhabits bushes, keeping much to

heavy jungle on the banks of water-courses, and is usually single, or

in pairs, either a male and female or a female and young being found

together; less often the female is accompanied by two young ones, which

remain with her until full grown.”

_Nanotragus._[217]—Horns small, parallel with frontals, and rising

immediately above postorbital process of frontals, in front of the

fronto-parietal suture. Lachrymal fossa very large, suddenly descending

in front of the orbit, and extending on to the maxilla; lachrymal vacuity

small. Auditory bulla large and smooth, without internal septum. Nasals

of moderate length. Crown of the head smooth; naked part of muffle small;

aperture of suborbital gland small. Lateral hoofs small or absent. Nine

species.[218]

The typical species is the Royal Antelope (_N. pygmæus_) of Guinea, the

smallest existing representative of the Pecora. This species, together

with _N. moschatus_ and _N. tragulus_ have no lateral hoofs, or tufts on

the knees. In the _Scopophorine_ group, comprising _N. scoparia_, _N.

montanus_, and _N. hastatus_, both these appendages are present; while in

the _Oreotragine_ group (_N. melanotis_ and _N. oreotragus_) the former

are present and the latter absent.

_Pelea._[219]—Horns rather small, compressed, upright, scarcely

diverging, and placed immediately over the orbits. No suborbital gland,

nor lachrymal fossa; premaxillæ not reaching nasals. Tail short and

bushy. Colour uniform. One species—the Rehbok (_P. capreola_), South

Africa, is nearly of the size of a Fallow Deer, although more resembling

a Chamois in build and habits. The colour is of a uniform light gray.

This animal inhabits bare rocky districts, and thus differs widely from

the Water-buck and its allies.

_Cobus._[220]—Large Antelopes, with the horns large, elongate, sublyrate,

and ringed at the base, and with rudimentary suborbital glands. Skull

with a deep frontal hollow, no lachrymal depression, large lachrymal

vacuity, and the premaxillæ reaching the very long nasals. Tail long,

with a ridge of hair above, and slightly tufted at the end. Colour

uniform. Six species, African.

The Antelopes of this genus are water-loving animals, the Water-buck

(_C. ellipsiprymnus_) and the Singsing (_C. defassus_) being well-known

examples. Both these species are much alike, standing as much as 4 feet

6 inches at the withers. The Water-buck of South and Eastern Africa is

characterised by the coarseness of its long hair; while in the Singsing

of West and Central Africa the hair is remarkably fine and soft.

Fossil Antelopes from the Pliocene of India are referred to _Cobus_.

_Helicophora_, from the Lower Pliocene of Attica, is regarded as allied

to _Cobus_, but it has no distinct supraorbital pits.

_Cervicapra._[221]—An allied South African genus in which the tail is

short and bushy and the premaxillæ do not reach the nasals. Three species.

The Reitbok (_C. arundineum_) is of a grizzly ochre colour; it stands

nearly 3 feet in height, and has horns about 1 foot in length. The Nagor

(_C. redunca_) is about 6 inches shorter, with horns of half the length,

and fulvous brown above and white below; the West African _C. bohor_

being rather larger.

_Antilopine Section._—A large group of moderate-sized or small Antelopes,

most abundant in the deserts bordering the Palæarctic, Oriental, and

Ethiopian regions. Horns generally compressed and lyrate, or recurved, or

cylindrical and spiral, ringed at base, sometimes present in both sexes.

Skull with large pits at apertures of supraorbital foramina of frontals,

and generally a distinct lachrymal fossa. Molars of upper jaw narrow,

without inner accessory column, and resembling those of the Sheep and

Goats. Tail moderate, compressed, hairy above.

_Antilope._[222]—Horns, present only in the male, long, cylindrical,

subspiral, and diverging. Suborbital gland large, with a somewhat linear

opening; lachrymal depression of skull very large, and a small lachrymal

fissure. Glands in the feet; lateral hoofs present. One species, India.

The well-known Black-buck (_A. cervicapra_) is found on open plains

all over India, except in lower Bengal and Malabar. Old males are deep

blackish-brown in colour on the back and sides and the outer surfaces of

the limbs, the under parts and inner surfaces of the limbs white, and

the back of the head, nape, and neck yellowish. Young males and females

are fawn-coloured above. Very large herds are seen in the plains about

Delhi and Mattra, which are said in some instances to reach to thousands.

Horn-cores are found in the Pleistocene deposits of the valley of the

Jumna which cannot be distinguished from those of the existing species.

_Æpyceros._[223]—Horns compressed, lyrate, and wide-spreading; present

only in male. No suborbital gland, or lachrymal depression in the skull.

No lateral hoofs. Two species; one from South and the other from West

Africa.

The Palla (_Æ. melampus_) is a large Antelope standing over 3 feet

high at the withers, and readily distinguished by its dark red colour,

gradually shading to white below. It is usually found on or near hills in

herds of from twenty to thirty. _Æ. petersi_ is from the Congo.

_Saiga._[224]—Nose very large, convex, and inflated. Supraorbital gland

present. Lachrymal fossa of skull small, and fissure absent; narial

aperture very large; nasals extremely short; supraorbital pits rather

small. Horns yellow, lyrate, of moderate length; present only in male.

Vertebræ: C 7, D 13, L 6, S 4, C 10. One species, Eastern Europe and

Western Asia.

The Saiga (_S. tatarica_) is a clumsily built and somewhat sheep-like

Antelope inhabiting the steppes; it occurs fossil in the Pleistocene of

France and England.

_Pantholops._[225]—Allied in the characters of the head and skull to

_Saiga_, but the nose less convex, the nostrils of the male more swollen,

and the horns of that sex black, very long, compressed, and lyrate; those

of female very short. One species, Central Asia.

The Chiru (_P. hodgsoni_) inhabits the highlands of Western Tibet and

Turkestan. In the former area it generally goes in small herds of from

three to six, and in the summer may be found grazing in early morning on

the level spaces frequently found in the river valleys at elevations of

about 15,000 feet. It is excessively shy and difficult to approach. The

large size of the narial aperture in the skull of Chiru is suggestive of

a connection with respiration at a high altitude, but this appears to be

negatived by the occurrence of the same feature in the Saiga.

_Gazella._[226]—Delicately built and sandy-coloured Antelopes, with

lyrate or recurved horns, which may be absent in the female, and are

always smaller and simpler in that sex than in the male. Skull with

moderate lachrymal fossa, and a distinct lachrymal fissure. Vertebræ: C

7, D 13, L 6, S 4, C 14. Suborbital gland frequently small, and covered

with hair. Face with a white streak running from the outer side of the

base of each horn nearly down to the upper end of each nostril, cutting

off a dark triangular central patch, and bordered externally by a

diffused dark line (see Fig. 121, p. 310). The Gazelles, of which there

are some twenty-four existing species, are typically Palæarctic desert

forms, the Springbok (_G. euchore_) being an outlying South African

species. _G. picticaudata_ and _G. gutturosa_ are respectively found in

Western Tibet and Mongolia, the former at great elevations. The majority

of the Gazelles do not exceed 30 inches in height, although _G. mohr_ is

36. Sir Victor Brooke classifies[227] the Gazelles as follows:—

A. No stripe on back; three lower premolars.

_a._ White of rump not encroaching on the fawn of the haunches.

I. Female with horns.

1. Horns lyrate or sublyrate—_G. dorcas_, _G. isabella_,

_G. rufifrons_, _G. lævipes_, _G. tilonura_, _G. naso_.

2. Horns non-lyrate—_G. cuvieri_, _G. leptoceros_, _G.

spekei_, _G. arabica_, _G. bennetti_, _G. fuscifrons_,

_G. muscatensis_.

II. Female without horns.

_G. subgutturosa_, _G. gutturosa_, _G. picticaudata_.

_b._ White of rump projecting forwards in an angle into the

fawn colour of the haunches. Horns in both sexes.

_G. dama_, _G. mohr_, _G. soemmerringi_, _G. granti_

(Fig. 121), _G. thomsoni_.

B. A white stripe down the back, two lower premolars. Horns in

both sexes.—_G. euchore._

The East African _G. walleri_ is an aberrant species, in which the

females are hornless, which has been made the type of the genus

_Lithocranius_. It is characterised by the extreme density of the horns

and skull, the slenderness of the mandible, and the small size of the

cheek-teeth, the upper molars being relatively broader and lower than

usual. The cranium is remarkable for the shortness of its facial portion,

the large size and production backwards of the supraoccipital, and for

the circumstance that the long basicranial axis is nearly parallel with

the plane of the palate.

Fossil species of _Gazella_ are found in the Pliocene and Pleistocene

deposits of Europe and India. _G. deperdita_ (_brevicornis_), of the

Lower Pliocene of France and Greece, appears to be a generalised species

in which the lower molars frequently have accessory columns, traces of

which are found in some of the existing forms.

_Hippotragine Section._—Includes very large African Antelopes, with long

horns, present in both sexes, which are placed over or behind the orbit,

and are either recurved, straight, or subspiral. Skull with no distinct

pits at apertures of supraorbital foramina in frontals, no lachrymal

fossa, and only a small lachrymal fissure. No suborbital gland. Tail

long, cylindrical, and tufted at the end. Upper molars extremely

hypsodont, very broad, and with large accessory columns, thus closely

resembling those of the Oxen. Some authorities divide this section into

two. In the Pliocene it occurs in India and Europe.

_Hippotragus._[228]—Horns stout, rising vertically from a crest over

the orbit at an obtuse angle to the plane of the nasals, then recurved;

lachrymal fissure in some instances almost obliterated. Neck with an

erect recurved mane. Tail very distinctly tufted. Four species, tropical

Africa and south to the Cape.

The Sable Antelope (_H. niger_) is one of the best-known examples of this

genus, occurring in South and East Africa. It stands upwards of 4½ feet

in height at the withers, and, except for some white streaks on the face

and the whole of the under surface of the body, is of a black colour.

The Blaubok (_H. leucophæus_) is distinguished by the glaucous hue of

the hair. The other species are the Equine Antelope (_H. equinus_) and

Baker’s Antelope (_H. bakeri_) from the Sudan, both closely allied, but

the latter distinguished by its pale fulvous colour, pencilled ears, and

black stripes on the shoulder.

Skulls of fossil Antelopes from the Pliocene of India have been referred

to _Hippotragus_ (_H. sivalensis_), and Sir V. Brooke suggests that the

European Pliocene _Antilope recticornis_ is not generically separable.

_Oryx._[229]—Horns long, slender, nearly straight or somewhat recurved,

rising behind the orbit, and inclining backwards in the plane of the

nasals; lachrymal fossa distinct. Nape maned; tail long, and more haired

than in _Hippotragus_. Four species, ranging over all the African deserts

to Arabia and Syria.

The Gemsbok (_O. gazella_, Fig. 141), is a South African species

characterised by its straight horns, the presence of a tuft of hair on

the throat, as well as by the large patches and stripes of black on the

head, back, limbs, and flanks. It stands nearly 4 feet in height at the

shoulder, and the horns are 2 feet 9 inches in length. The colour of the

upper part of the body is a rusty gray, and of the under part white,

while these are separated from each other by a well-defined black band

on either side. These bands unite on the breast, and are continued as a

single black band until reaching the lower jaw, where they again divide

and form two transverse bands on the head, terminating at the base of

the horns. The head otherwise is white, as also are the limbs, with the

exception of the thighs, which are black. The Gemsbok generally goes

in pairs, or in small herds of three or four. The Beisa (_O. beisa_)

of Abyssinia is distinguished by the absence of the tuft of hair on

the throat. Writing of this species in his _Geology and Zoology of

Abyssinia_, Mr. W. T. Blanford observes that “the appearance of a herd of

Oryx is very imposing. They are some of the most elegant and symmetrical

of animals, the motions being those of a wild Horse rather than of an

Antelope. Their favourite pace appears to be either a steady quick walk

or a trot; they rarely break into a gallop unless greatly alarmed. When

frightened they dash off, sometimes snorting and putting their heads down

as if charging, raising their long tails, and looking very formidable.

They are wary animals, though far less so than some other Antelopes. It

is said that they frequently attack when wounded, and their long straight

horns are most deadly weapons.” The Arabian Beatrix Antelope (_O.

beatrix_) is a much smaller animal, with the black markings confined to

the head, fore limbs, and flanks. Finally, the Leucoryx (_O. leucoryx_)

of North Africa, while agreeing in size with the Beatrix, differs by its

curved horns and uniform coloration.

[Illustration: FIG. 141.—The Gemsbok (_Oryx gazella_).]

The extinct _Palæoryx_, of the Lower Pliocene of Europe and the Isle of

Samos, appears to have been an ancestral form of _Oryx_, said to show

some signs of affinity with _Hippotragus._

_Addax._[230]—Horns with the same inclination as in _Oryx_, but with a

slight spiral twist. No mane on nape, but a slight one on the throat.

Hoofs rounded. One species (_A. nasomaculatus_), from North Africa and

Arabia, the colour of which is nearly white.

_Tragelaphine Section._—Includes large, so-called Bovine, Antelopes now

mainly characteristic of the Ethiopian region, but with one Oriental

genus. Horns usually present in the male only (if developed in the female

smaller), with a more or less distinct ridge in front, and usually

twisted spirally, the front ridge twisting outwards from the base of the

horn. Skull without lachrymal fossa, but with a large or small lachrymal

fissure; usually large pits at the apertures of the supraorbital foramina

on the frontals; premaxillæ reaching nasals. Muffle large and moist;

nostrils approximated. Molars hypsodont or brachydont. Vertical white

stripes frequently present on the body.

_a._ _Hind limbs much shorter than the fore. Horns behind the orbit,

short, conical, faintly angulated. Nose bovine. Body without vertical

stripes. Molars_ (Fig. 123, p. 311) _hypsodont, with a large accessory

column in those of the upper jaw. One Oriental genus._

_Boselaphus._[231]—The one genus of this subsection is represented only

by the well-known Nilghai (_B. tragocamelus_) of India. The male stands

over 4 feet in height at the shoulder, with horns about 8 inches in

length; the hornless female being about one third smaller. Both sexes

have a short erect mane, and the male has also a tuft of hair upon the

throat. When adult the sexes are very different in colour, the male being

of a dark iron gray or slate colour, approaching black on the head and

legs, while the female and young are of a bright light brown or fawn

colour. In both male and female at all ages the lips, chin, and under

parts, as well as two transverse stripes on the inner sides of the ears

and rings on the fetlocks, are white, and the mane and tip of the tail

black. The Nilghai is one of the few Antelopes occurring in India, where

it is found from near the foot of the Himalaya to the south of Mysore,

though rare to the north of the Ganges and also in the extreme south. It

is most abundant in Central India, and does not occur in Assam or the

countries to the east of the Bay of Bengal. It frequents forests and low

jungles, though often found in tolerably open plains, associating in

small herds. One, or very often two, young produced at a birth. Fossil

remains of species of this genus occur in the Pleistocene and Pliocene

deposits of India.

_b._ _Fore and hind limbs equal. Horns long, and spirally twisted. Nose

cervine, and aperture of suborbital gland very small. Body generally

striped. Molars brachydont, those of the upper jaw in existing forms with

a smaller inner accessory column. Three existing Ethiopian genera._

_Tragelaphus._[232]—Female hornless. Horns of males (Fig. 142) over

orbit, with one or two spiral turns, obscurely ridged, the posterior

ridge being more developed than the anterior. Skull with small

supraorbital pits, very small lachrymal fissure, and no deep intercornual

depression in the frontals. Neck maned or smooth. Hoofs short or long.

Coloration usually brilliant, differing markedly in the two sexes, and

the white bands on the body, when present, numerous and distinct. Seven

species.

[Illustration: FIG. 142.—Head of _Tragelaphus gratus_. From Sclater,

_Proc. Zool. Soc._ 1883, p. 36.]

The Harnessed Antelopes are among the handsomest of the whole group. The

small Guib (_T. scriptus_) is not larger than a Goat, but _T. angasi_ is

3 feet 4 inches in height at the shoulder. In _T. scriptus_, _T. angasi_,

and _T. euryceros_, the two sexes differ in colour, the body is marked by

white stripes descending from a white dorsal streak, and the hoofs are

short; the third species differing from the others by the absence of a

mane on the neck, back, and belly. _T. gratus_ agrees with this group in

coloration (the mane being absent), but differs in the extreme elongation

of its hoofs. The Nakong, _T. spekei_, while having the long hoofs of

_T. gratus_, has a perfectly plain body coloration, with a mane on the

neck. The two species with elongated hoofs inhabit swampy districts, for

which this peculiar structure is admirably adapted; and the Nakong, when

frightened, will rush into the water and leave only its nostrils and the

tips of the horns above the surface. The small Bushbuck (_T. sylvaticus_)

of South Africa has no stripes, and short hoofs.

[Illustration: FIG. 143. The Kudu (_Strepsiceros kudu_). From Sclater,

_List of Animals in Zoological Society’s Gardens_, 1883, p. 136.]

_Strepsiceros._[233]—Females hornless. Horns (Fig. 143) more twisted

than in _Tragelaphus_, forming an open spiral, with the anterior ridge

very strongly developed, and rising at an obtuse angle to the plane of

the nasals. Skull with large supraorbital pits, large lachrymal fissure,

and deep intercornual depression. Hoofs short. Body with white vertical

stripes descending from a longitudinal dorsal streak. Two existing

species.

The Kudu (_S. kudu_, Fig. 143) extends from South Africa to Abyssinia,

and is only inferior in size to the Eland. The horns are about 4 feet

in length, and form a very open spiral, and there is a fringe of long

hair down the front of the neck. The Lesser Kudu (_S. imberbis_), of

Somali-land is a much smaller form, without the fringe of hair on the

neck, and with a much smaller axis formed by the spiral of the horns.

An imperfect skull from the Pliocene of Northern India has been referred

to _Strepsiceros_.

_Oreas._[234]—Females horned. Horns twisted on their own axis, with very

strong ridges, inclining upwards and outwards in the plane of the nasals.

General characters of skull as in preceding genus. Stripes on body, if

present, very faintly marked. One existing species.

The Eland (_O. canna_) is the largest of all the Antelopes, the males

standing nearly 6 feet at the withers. One variety from South Africa

is of a uniform pale fawn colour, while the Central African form is of

a bright tan colour, marked by a number of thin pale vertical stripes

descending from a dark dorsal ridge—these markings fading more or less in

the adults. The males have a large dewlap, a tuft of brown hair on the

forehead, and a small mane on the neck. The straight black horns of the

male are usually about 18 inches long. Elands were formerly extremely

abundant in Southern and Eastern Africa, but their destruction has been

so relentless that they have totally disappeared from extensive areas,

and are daily becoming scarcer.

Portions of upper jaws from the Pliocene deposits of India appear to

indicate the former existence in that area of large Antelopes closely

allied to the Eland, but distinguished from the living species by the

greater size of the inner accessory column in the upper molars.

_Allied Extinct Types._—Large Antelopes with spirally twisted horns

appear to have been common over Southern Europe in Pliocene times, but

their exact affinity is in many cases difficult to determine. Of these,

_Palæoreas_, which occurs in the Lower Pliocene of Europe and Algeria,

appears to present affinities both to _Oreas_ and _Strepsiceros_, and may

have been the ancestral type from which these two genera are derived; the

upper molars have well-developed accessory columns.

The so-called _Antilope torticornis_, of the French Pliocene, resembles

_Tragelaphus_ in the greater development of the posterior as compared

with the anterior ridge of the horn-cores, and has accordingly been

referred to that genus. _Protragelaphus_, of the Lower Pliocene of

Attica, differs from all the other types in the absence of the anterior

ridge on the horn-cores and of the supraorbital pits, while it has a

distinct lachrymal fossa.

In this place it will be convenient to notice certain fossil forms which

do not accord with any of the existing sections of the family, and for

the reception of which the _Palæotragine_ section has been formed. In

these types the horn-cores are laterally compressed like those of the

modern Goats, but the upper molars resemble those of the brachydont

Antelopes. The earliest of these genera, and the first representative

of the Antelopes yet known, is _Protragoceros_, of the Middle Miocene

of France, first described as _Antilope clavata_; _Palæotragoceros_ and

_Tragoceros_, of the Lower Pliocene, are distinguished by their larger

horns and wider molars.

A remarkable large Antelope from the Lower Pliocene of the Isle of Samos,

in the Turkish Archipelago, proposed to be described as _Criotherium_,

appears to be unlike any other form. The horns, which are placed on the

extreme vertex of the skull, are very short, tightly twisted, and project

in front of the forehead. The upper molars have short and broad crowns,

with no accessory column on the inner side.

_Rupicaprine Section._—The Caprine Antelopes, as the typical members of

this section may be termed, appear to connect the true Antelopes with the

Goats. They are mostly small or medium-sized forms, inhabiting portions

of the Palæarctic and Oriental regions, with one outlying North American

genus. The typical forms present the following features. Horns present,

and of nearly equal size in both sexes, rising behind the orbits, short,

ringed at the base, conical or somewhat compressed, and recurved.

Suborbital gland generally present, in some cases small. Build clumsy;

hoofs large; tail short, tapering, hairy above. Skull with lachrymal

fossa, but no fissure. Molars as in the Caprine section.

_Rupicapra._[235]—Horns short and cylindrical, rising perpendicularly

from the forehead for some distance, then bending sharply backwards and

downwards, forming hooks with pointed tips. Premaxillæ not reaching the

nasals. One species, Palæarctic.

The Gemse, or Alpine Chamois (_R. tragus_), inhabits the high mountains

of Europe from the Pyrenees to the Caucasus. It stands about 2 feet in

height at the withers. The body is covered in winter with long hair of a

chestnut-brown colour, that of the head being paler, with a dark brown

streak on each side. At other seasons the colour is somewhat lighter, in

spring approaching to gray. Underneath the external covering the body is

further protected from cold by a coat of short thick wool of a grayish

colour. The tail is black; the ears are pointed and erect; the hoofs have

the outer edges higher than the soles, and are thus admirably adapted for

laying hold of the slightest projection or roughness on the face of the

rocky precipices it frequents. The Chamois is gregarious, living in herds

of fifteen or twenty, and feeding generally in the morning or evening.

The old males, however, live alone, except in the rutting season, which

occurs in October, when they join the herds, driving off the young males,

and engaging in contests with each other that often end fatally. The

period of gestation is twenty weeks, when the female, beneath the shelter

of a projecting rock, produces one and sometimes two young. In summer the

Chamois ascends to the limits of perpetual snow, being only outstripped

in the loftiness of its haunts by the Ibex; and during that season it

shows its intolerance of heat by choosing such browsing grounds as have a

northern exposure.

[Illustration: FIG. 144.—_Nemorhædus crispus._ From Sclater, _List of

Animals in Zoological Society’s Gardens_, 1883, p. 151.]

_Nemorhædus._[236]—Horns rounded, gradually recurving, without distinct

hook at the end. Suborbital gland small or wanting; ears large; skull

with a large lachrymal depression, and the premaxillæ not quite reaching

the nasals. Some nine species, ranging from the Eastern Himalayas to

North China and Japan, and southwards to Formosa, the Malay Peninsula,

and Sumatra. The smallest species is the Himalayan Goral (_N. goral_). Of

the larger forms we may mention the Himalayan Serow (_N. bubalinus_) the

Cambing-Utan (_N. sumatrensis_) of Sumatra, and the Japanese _N. crispus_

(Fig. 144). Of the Serow, Colonel Kinloch remarks that “it is a large and

powerful beast. The body is covered with very coarse hair, which assumes

the form of a bristly mane on the head and shoulders, and gives the

beast a ferocious appearance, which does not belie its disposition. The

colour is a dull black on the back, bright red on the sides, and white

underneath, the legs also being dirty white. The ears are very large, the

muzzle is coarse. The Serow has an awkward gait, but in spite of this can

go over the worst ground; and it has perhaps no superior in going down

steep hills. It is a solitary animal, and nowhere numerous.”

_Haploceros._[237]—The Rocky-Mountain Goat (_Haploceros montanus_),

inhabiting the northern parts of California, appears to be very closely

allied to _Nemorhædus_. The horns are somewhat compressed at the base;

there is no suborbital gland; and the ears are small. The hair, which is

whitish in colour, is very long, and especially abundant in the region of

the throat, shoulders, flanks, and tail. The animal is about the size of

a large Sheep.

_Budorcas._[238]—The Takin (_B. taxicolor_) of the Mishmi Hills in Assam,

and an allied species from Eastern Tibet, are larger forms apparently

related to _Nemorhædus_, but with a much greater development of the

horns. The horns of what is considered to be the male[239] arise from

the vertex of the skull, and are nearly in contact in the middle line;

they first bend outwards and downwards, and then suddenly upwards

and backwards. Those regarded by Mr. Hume as referable to the female

are directed at first outwards, and then gradually curve upwards and

backwards, without any downward flexure or angulation. The horns of

the male may be 2 feet in length, with a basal diameter of 13 inches.

The muzzle is hairy, with a small naked muffle. There appear to be

considerable seasonal and sexual variations in colour; the body being in

some cases of a yellow dun, while in others it is a dusky, reddish-brown,

with much black intermingled. The heads of large males are blackish.

Scarcely anything is known of the habits of the Takin, which never

appears to have been seen alive by Europeans.

_Caprine Section._—Both sexes with horns, but those of the female small.

Horns usually compressed, triangular, with transverse ridges, and either

curving backwards or spiral. Muzzle hairy, without naked muffle.

Suborbital gland small or absent; lachrymal fossa of skull present or

absent. Tail short and flattened. Foot-glands frequently present. Molars

very hypsodont; those of the upper jaw being narrow, without an accessory

internal column. Mainly Palæarctic, but with some outlying forms.

This section includes the Goats and Sheep, which are so closely connected

that it is difficult to give well-marked generic characters that will

hold good for all the species. They seem to be one of the latest

developments of the _Bovidæ_, since they are unknown before the Pliocene

period; and are essentially mountain forms.

[Illustration: FIG. 145.—The Alpine Ibex (_Capra ibex_).]

_Capra._[240]—Horns flattened from side to side, and either curving

backwards (Fig. 145) or spirally twisted. No suborbital gland, and no

lachrymal fossa in the skull. Foot-glands, if present, only in the fore

feet. Chin more or less bearded. Males with a strong odour. Vertebræ:

C 7, D 13, L 6, S 4, C 9-13. Some dozen species, ranging over all the

higher mountains of Southern Europe, from Spain to the Caucasus; also

found in Abyssinia, Persia, Sind, and Baluchistan, thence through the

higher Himalaya, and so on to Tibet and Northern China. One outlying

species occurs in the Nilgherries of Southern India.

The European Ibex or Steinbok (Fig. 145), which may be taken as a typical

Goat, stands about 2½ feet in height at the shoulder. In summer the hair

is short and smooth, and of an ashy-gray colour, but a long coat is

developed in winter. The horns of the male rise in a bold backward sweep

from the forehead, and are characterised by the strong transverse ridges

on the broad and flat anterior surface. They are said to be not more

than some 2 feet in length, but these dimensions are greatly exceeded

by the horns of the Himalayan Ibex. The Alpine Ibex lives at a greater

height than the Chamois, spending the day just at the limit of perpetual

snow, and descending at night to graze at lower levels. Both this and the

Himalayan species generally live in small herds of from five to fifteen

or more; they are wary animals, although not so much so as many of the

wild Sheep. The following list, mainly taken from two papers by Mr.

Sclater,[241] gives the distribution of the various species of Goats,

with some remarks on their peculiarities:—

(1) _C. ibex_, confined to the Alps of Switzerland, Savoy, and the

Tyrol, and now nearly extinct, except where artificially preserved.

(2) _C. sibirica_, closely allied to the preceding, but with larger

horns, occurs in the Altai Mountains, and throughout the Himalaya from

Kashmir to Nipal, and northward towards Turkestan. (3) _C. sinaitica_,

of the mountains of Upper Egypt, the Sinaitic Peninsula, and Palestine,

is allied to the two preceding species, but has the horns somewhat

more compressed, with a difference in the ridges on the front. (4) _C.

caucasica_, a very distinct species, confined to the Caucasus, where

it inhabits the western part of the Great Caucasus; with thick horns

curving backwards and outwards in one plane, with the exception of their

tips, which incline inwards.[242] (5) _C. pallasi_ is an allied species

from the Eastern Caucasus, distinguished, among other features, by the

curvature of the horns, which lie flatter and twist more outward from

the forehead, with a greater terminal inward bend. (6) _C. pyrenaica_,

of the Pyrenees, and the higher ranges of Central Spain, Andalusia, and

Portugal, is another nearly related species. (7) _C. ægagrus_, formerly

abundant over the Grecian Archipelago, but now restricted in Europe to

Crete and some of the Cyclades, is found throughout the mountains of

Asia Minor and Persia, and thence to Baluchistan and Sind. The horns

are thinner and sharper in front than in the Ibexes, and this species

is generally regarded as the ancestral stock of the various breeds of

domestic Goats. (8) _C. dorcas_, a Goat from the island of Jura, near

Eubœa, has been described under this name, and is apparently nearly

allied to _C. ægagrus_. (9) _C. walie_, an apparently well-characterised

species from the highest ranges of Abyssinia. (10) _C. falconeri_; the

Markhoor differs from all the preceding species by the spiral twisting of

its horns, which attain enormous dimensions. It occurs in the Pir-Panjal

range south of Kashmir, and thence into Afghanistan and the Suleiman

range, and northwards to Astor, Gilgit, and Scardo (Baltistan). The

specimens from the Suleiman range have the spiral of the horns very

close, somewhat as in the Eland; while in those from Astor, Gilgit, and

Scardo it is very open, as in the Kudu. The Pir-Panjal race occupies a

somewhat intermediate position in this respect. (11) _C. jemlaica_, the

Thar, inhabits suitable regions along the whole range of the Himalaya

from Kashmir to Bhutan. Together with the next species, it differs from

the more typical Goats in its short, thick, and much compressed horns,

the anterior border of which is keeled, and the moist naked muffle. There

are no glands in the fore feet. It was generically separated by Gray as

_Hemitragus_. (12) _C. hylocrius_, the so-called Ibex of the Nilgherries,

Anamallays, and other adjoining ranges of Southern India, is an outlying

species, apparently allied to the preceding, but with somewhat different

horns, in which the external angle in front is much rounded off.

Of fossil Goats we have but little knowledge. Remains of _C. pyrenaica_

are found in cave-deposits at Gibraltar; and it is not improbable that

the genus is represented in the Upper Pliocene of France. Several species

occur in the Pliocene of India, _C. sivalensis_ being apparently closely

allied to _C. jemlaica_, while another has horns resembling those of _C.

falconeri_, and it is possible that a third may be more nearly related to

the Ibexes.

_Ovis._[243]—Horns curving backwards and downwards in a bold sweep,

with the tips everted, generally with more or less prominent transverse

ridges, and brownish in colour. Suborbital gland and lachrymal fossa

usually present, but generally small. Foot-glands in all the feet. Chin

not bearded;[244] males without a strong odour. Vertebræ: C 7, D 13, L

6, S 4, C 10-14. Some twelve species, mainly Palæarctic, but extending

into the adjacent portions of the Oriental region, and with one outlying

species in North America.

The more typical Sheep are closely connected with the Goats by the

Himalayan Bharal (_O. nahura_) and the Aoudad (_O. tragelaphus_) of

Northern Africa, both these species having no suborbital gland and no

lachrymal fossa, while their comparatively smooth and olive-coloured

horns show a decided approximation to those of the Goats. Both present,

however, the ovine character of glands in all the feet. In the typical

Sheep the basioccipital of the skull is wider in front than behind,

with the anterior pair of tubercles widely separated and much larger

than the posterior pair. The Bharal, however, resembles the Goats in

having an oblong basioccipital, with the posterior tubercles larger and

more prominent than the anterior ones, both being situated in the same

antero-posterior line. These transitions towards the caprine type are,

however, not sufficient to support the view that the Bharal should form

the type of a distinct genus (_Pseudois_), more especially since some of

the typical Sheep, like _O. canadensis_, have the lachrymal fossa of the

skull very much reduced in size.

The distinction of the various permanent modifications under which wild

Sheep occur is a matter of considerable difficulty. Trivial characters,

such as size, slight variations in colour, and especially the form and

curvature of the horns, are relied upon by different zoologists who have

given attention to the subject in the discrimination of species, but no

complete accord has yet been established. The most generally recognised

forms are enumerated below.

The geographical distribution of wild Sheep is interesting. The immense

mountain ranges of Central Asia, the Pamir and Thian-Shan of Turkestan,

may be looked upon as the centre of their habitat. Here, at an elevation

of 16,000 feet above the sea-level, is the home of the magnificent

_Ovis poli_, named after the celebrated Venetian traveller Marco Polo,

who met with it in his adventurous travels through this region in the

thirteenth century. It is remarkable for the great size of the horns of

the old rams and the wide open sweep of their curve, so that the points

stand boldly out on each side, far away from the animal’s head, instead

of curling round nearly in the same plane, as in most of the other

species. A Sheep from the same region, in which the horns retain their

more normal development, has received the name of _O. karelini_, but,

according to Mr. W. T. Blanford,[245] is not distinct specifically from

_O. poli_. Eastward and northward is found the Argali (_O. argali_),

with a wide and not very well determined range; it formerly occurred in

the Altai, but is now found in Northern Mongolia. Still farther north,

in the Stanovoi Mountains and Kamschatka, is _O. nivicola_, and away on

the other side of Behring’s Strait, in the Rocky Mountains and adjacent

highlands of western North America, is the “Bighorn” or Mountain Sheep

(_O. canadensis_), the only member of the genus found in that continent,

and indeed—except the Bison, Musk-Ox, Mountain Goat (_Haploceros_), and

the Prong-buck (_Antilocapra_)—the only hollow-horned Ruminant, being

like the rest obviously a straggler from the cradle of its race. The two

last-named species are nearly allied, and are characterised by the slight

development of the ridges on their horns and the very shallow lachrymal

fossa. Turning southward from the point from which we started, and

still a little to the east, in Nipal and Western Tibet, is the Himalayan

Argali (_O. hodgsoni_), having massive and strongly curved horns, with

bold ridges, like those of the true Argali. Indeed, were it not for their

isolated areas there would appear to be no grounds for distinguishing

these two closely allied forms, and it is not improbable that they

are really identical. _O. brookei_, appears to have been founded on a

hybrid between _O. hodgsoni_ and _O. vignei_. In the same districts,

and also in Southern Ladak, there occurs the Bharal (_O. nahura_), with

smaller, smoother, and more spreading horns. Passing in a south-westerly

direction we find a series of smaller forms, _O. vignei_ of Ladak, _O.

cycloceros_ of Northern India, Persia, and Baluchistan. _O. gmelini_ of

Asia Minor and Persia, _O. ophion_, confined to the elevated pine-clad

Troodos Mountains of the island of Cyprus, and said at the time of the

British occupation in 1878 to have been reduced to a flock of about

twenty-five individuals, and _O. musimon_, the Moufflon of Corsica and

Sardinia (see Fig. 146), believed to have been formerly also a native of

Spain. In the three latter species the females are hornless. Lastly, we

have the somewhat aberrant, Goat-like Aoudad (_O. tragelaphus_), of the

great mountain ranges of North Africa, in which, as already mentioned,

the skull and horns resemble those of the Bharal, although the tail is

longer, and there is a thick fringe of long hair on the throat, chest,

and fore legs.

[Illustration: FIG. 146.—The Moufflon (_Ovis musimon_). From a living

animal in the London Zoological Gardens.]

We thus find that Sheep are essentially inhabitants of high mountainous

parts of the world, for dwelling among which their wonderful powers of

climbing and leaping give them special advantages. No species frequent

by choice either level deserts, open plains, dense forests, or swamps.

By far the greater number of species are inhabitants of the continent of

Asia, one extending into North America, one into Southern Europe, and one

into North Africa. No wild Sheep exist in any other part of the world,

unless the so-called Musk-Ox of the Arctic regions, the nearest existing

ally to the true Sheep, may be considered as one. Geologically speaking,

Sheep appear to be very modern animals, or perhaps it would be safer to

say that no remains that can be with certainty referred to the genus

have been met with in the hitherto explored true Tertiary beds, which

have yielded such abundant modifications of Antelopes and Deer. They are

generally considered not to be indigenous in the British Isles, but to

have been introduced by man from the East in prehistoric times. A fossil

Sheep (_Ovis savigni_), apparently allied to the Argali, has, however,

been described from the so-called Forest-bed of the Norfolk coast.

The Sheep was a domestic animal in Asia and Europe before the dawn of

history, though quite unknown as such in the New World until after the

Spanish conquest. It has now been introduced by man into almost all

parts of the world where settled agricultural operations are carried on,

but flourishes especially in the temperate regions of both hemispheres.

Whether our well-known and useful animal is derived from any one of the

existing wild species, or from the crossing of several, or from some

now extinct species, is quite a matter of conjecture. The variations of

external characters seen in the different domestic breeds are very great.

They are chiefly manifested in the form and number of the horns, which

may be increased from the normal two to four or even eight, or may be

altogether absent in the female alone, or in both sexes; in the form and

length of the ears, which often hang pendent by the side of the head;

in the peculiar elevation or arching of the nasal bones in some Eastern

races; in the length of the tail, and the development of great masses of

fat at each side of its root, or in the tail itself; and in the colour

and quality of the fleece.

_Ovibos._[246]—This genus is generally considered to be a connecting link

between the Caprine and Bovine sections, but should rather be regarded as

an aberrant type of the former. Horns of adult male rounded, smooth, and

closely approximated at their bases, where they are depressed and rugose;

curving downwards, and then upwards and forwards. Muzzle caprine; no

suborbital gland, no lachrymal fossa or fissure in skull; orbits tubular;

a large narial aperture and very short nasals; premaxillæ not reaching

nasals. Tail short, and molar teeth caprine. One existing and two fossil

species, Palæarctic and Nearctic.

[Illustration: FIG. 147.—The Musk-Ox (_Ovibos moschatus_).]

The animal commonly known as the Musk-Ox (_Ovibos moschatus_), though

approaching in size the smaller varieties of Oxen, is in structure and

habits closely allied to the Sheep, its affinities being well expressed

by the generic name _Ovibos_ bestowed upon it by De Blainville. The

specific name, as also the common English appellatives “Musk-Ox,”

“Musk-Buffalo,” or “Musk-Sheep,” applied to it by various authors, refer

to the musky odour which the animal exhales. This does not appear to be

due to the secretion of a special gland, as in the case of the Musk-Deer;

but it must be observed that, except as regards the osteology, very

little is known of the anatomy of this species. It about equals in size

the small Welsh and Scotch cattle. The head is large and broad. The horns

in the old males have extremely broad bases, meeting in the median line,

and covering the brow and whole crown of the head. They are directed

at first downwards by the side of the face and then turn upwards and

forwards, ending in the same plane as the eye. Their basal halves are of

a dull white colour, oval in section and coarsely fibrous; their middle

part smooth, shining, and round; their tips black. In the females and

young males the horns are smaller, and their bases are separated from

each other by a space in the middle of the forehead. The ears are small,

erect, and pointed, and nearly concealed in the hair. The space between

the nostrils and the upper lip is covered with short close hair, as in

Sheep and Goats, without any trace of the bare muffle of the Oxen.

The greater part of the animal is covered with long brown hair, thick,

matted, and curly on the shoulders, so as to give the appearance of a

hump, but elsewhere straight and hanging down,—that of the sides, back,

and haunches reaching as far as the middle of the legs and entirely

concealing the very short tail. There is also a thick woolly under-fur,

shed in the summer. The hair on the lower jaw, throat, and chest is long

and straight, and hangs down like a beard or dewlap, though there is no

loose fold of skin in this situation as in Oxen. The limbs are stout

and short, terminating in unsymmetrical hoofs, the external one being

rounded, the internal pointed, and the sole partially covered with hair.

The Musk-Ox is at the present day confined to the most northern parts of

North America, where it ranges over the rocky barren grounds between the

60th parallel and the shores of the Arctic Sea. Its southern range is

gradually contracting, and it appears that it is no longer met with west

of the Mackenzie River, though formerly abundant as far as Eschscholtz

Bay. Northwards and eastwards it extends through the Parry Islands and

Grinnell Land to North Greenland, reaching on the west coast as far south

as Melville Bay; and it was also met with in abundance by the German

polar expedition of 1869-70 at Sabine Island on the east coast. No trace

of it has been found in Spitzbergen or Franz Joseph Land. As proved

by the discovery of fossil remains, it ranged during the Pleistocene

period over northern Siberia and the plains of Germany and France, its

bones occurring very generally in river deposits along with those of

the Reindeer, Mammoth, and Woolly Rhinoceros. It has also been found in

Pleistocene gravels in several parts of England, as Maidenhead, Bromley,

Freshfield near Bath, Barnwood near Gloucester, and also in the lower

brick-earth of the Thames valley at Crayford, Kent.

It is gregarious in habit, assembling in herds of twenty or thirty head,

or, according to Hearne, sometimes eighty or a hundred, in which there

are seldom more than two or three full-grown males. The Musk-Ox runs with

considerable speed, notwithstanding the shortness of its legs. Major

H. W. Feilden, naturalist to the Arctic expedition of 1875, says: “No

person watching this animal in a state of nature could fail to see how

essentially ovine are its actions. When alarmed they gather together like

a flock of sheep herded by collie dog, and the way in which they pack

closely together and follow blindly the vacillating leadership of the old

ram is unquestionably sheep-like. When thoroughly frightened they take

to the hills, ascending precipitous slopes and scaling rocks with great

agility.” They feed chiefly on grass, but also on moss, lichens, and

tender shoots of the willow and pine. The female brings forth a single

young one in the end of May or beginning of June after a gestation of

nine months. According to Sir J. Richardson, “when this animal is fat

its flesh is well tasted, and resembles that of the Caribou, but has

a coarser grain. The flesh of the bulls is highly flavoured, and both

bulls and cows when lean smell strongly of musk, their flesh at the same

time being very dark and tough, and certainly far inferior to that of

any other ruminating animal existing in North America.” The carcase of

a Musk-Ox weighs, exclusive of fat, above 3 cwt. On this subject, Major

Feilden[247] says: “The cause of the disagreeable odour which frequently

taints the flesh of these animals has received no elucidation from my

observations. It does not appear to be confined to either sex, or to any

particular season of the year; for a young unweaned animal, killed at

its mother’s side and transferred within an hour to the stew-pans, was

as rank and objectionable as any. The flesh of some of these animals of

which I have partaken was dark, tender, and as well flavoured as that of

four-year old Southdown mutton.”

Remains of two fossil species of this genus (_O. bombifrons_ and _O.

cavifrons_) have been described from Pleistocene beds in the United

States, the one from Kentucky and the other from the Arkansas River. Both

(if indeed they be valid species) appear closely allied to the living

form.

_Bovine Section._—Horns present and of nearly equal size in both sexes;

in form rounded or angulated, placed on or near the vertex of the

skull, extending more or less outwards, and curving upwards near the

extremities; external surface comparatively smooth and never marked by

prominent transverse ridges or knobs. Muzzle broad, with large naked

muffle; nostrils lateral; no suborbital gland. Skull without any trace of

lachrymal fossa or fissure. Tail long and cylindrical; generally tufted

at the extremity, rarely hairy throughout. Males usually with a dewlap on

the throat. No foot-glands. Molar teeth extremely hypsodont; those of the

upper jaw with a nearly square cross-section, and a large accessory inner

column.

The section is abundantly represented in the Palæarctic, Oriental, and

Ethiopian regions, with one Nearctic species and an outlying and aberrant

species in Celebes.

_Bos._[248]—The whole of the species of Oxen were included by Linnæus in

the single genus _Bos_, and although the species have been distributed

by modern zoologists in several genera—such as _Anoa_, _Bubalus_,

_Bison_, _Poëphagus_, _Bibos_, and _Bos_—the characters by which they are

separated are so slight that it seems, on the whole, preferable to retain

the old genus in its original wide sense. Using then the term _Bos_ in

this sense, it will include all the representatives of the section—about

a dozen in number—and may be divided into several groups.

The first group includes the Buffaloes (genus _Bubalus_), chiefly

characterised by their more or less flattened and angulated horns, which

incline upwards and backwards, with an inward curve towards their tips,

and are placed below the plane of the occiput, or vertex of the skull.

The premaxillæ reach to the nasals, and the vomer is peculiar in being

so much ossified as to join the posterior border of the palate. The back

has a distinct ridge in the region of the withers; and the forehead is

frequently convex. Oriental and Ethiopian region, and Celebes.

The most generalised representative of this group is the small Anoa

(_B. depressicornis_) of Celebes, the type of the genus _Anoa_ or

_Probubalus_, which has the same cranial structure as in the more

typical Buffaloes, to the young of which (as was pointed out by the

late Professor Garrod) it presents a striking resemblance. Its colour

is black; and the short and prismatic horns are directed upwards from

the forehead. In the Pliocene Siwaliks of India there occur the remains

of larger Buffaloes (_B. occipitalis_ and _B. acuticornis_) closely

allied to the Anoa, but with longer and more distinctly angulated horns.

The still larger _B. platyceros_ of the last-named deposits, in which

the horns are wide-spreading and much flattened, appears to be in some

respects intermediate between the preceding and following forms. The

typical Indian Buffalo (_Bos buffelus_), which has been domesticated

over South-East Asia, Egypt, and Southern Europe, is, in the wild state,

a gigantic animal with enormous horns. These horns are longer, more

slender, and more outwardly directed in the female than in the male;

and in the former sex may have a length of more than 6 feet from base

to tip. They are widely separated at their bases, the forehead is very

convex, and the ears are not excessively large, and have no distinct

fringe. These Buffaloes frequent swampy and moist districts in several

parts of India, but it is in many instances difficult to decide whether

they belong to really wild or to feral races. Very large skulls,

specifically indistinguishable from those of the existing form, occur in

the Pleistocene deposits of the Narbada valley in India; while an allied,

if not specifically identical form, occurs in the Pliocene of the same

country. There is some doubt whether _B. antiquus_ of the Pleistocene of

Algeria is most nearly related to the Indian or to the African species.

In Africa two species of Buffalo are recognised by Sir Victor

Brooke,[249] namely the large _B. caffer_, occurring typically at the

Cape, but said by this writer to range to Abyssinia, and the smaller

_B. pumilus_, which seems to have a very wide distribution. The skulls

of both these forms are shorter than in the Indian species, while the

horns are also shorter, much more curved inwardly, and more approximated

on the forehead. In the large typical form of _B. caffer_ from South

Africa the colour is black, the horns of the male are very thick, much

reflected, and closely approximated on the forehead, where they form

a helmet-like mass.[250] The large northern form described as _B.

æquinoctialis_ has the horns somewhat less thick, and thus approximates

to the so-called _B. pumilus_.

The latter occurs typically in Western Africa, where it has also been

described as _B. brachyceros_. In the typical form the horns are thinner

and less reflected than in _B. caffer_, and in some specimens they are

more widely separated on the forehead, and are marked at their bases by

distinct rugæ. The colour is ruddy brown, inclining to rufous in one

specimen. The skulls of Buffaloes from West Africa, probably referable to

the form described as _B. centralis_, appear to connect _B. pumilus_ with

_B. caffer_, as shown by their larger size and the form of their horns;

so that further observations are required to show whether the smaller

form is really entitled to rank as a distinct species, or merely as a

well-marked local race.

The second group comprises the Bisons, which are more nearly allied to

the true Oxen, having similar rounded horns, but the skull being less

massive, with a longer and more tapering frontal region, and a wider

frontal diameter. The superior part of the forehead is transversely

arched, the intercornual space elevated in the middle, the horns situated

below the plane of the occiput, and the orbits more or less prominent.

The premaxillæ do not extend upwards to reach the nasals. The Bisons

(Fig. 148) have the body covered with short, crisp, woolly hair, while on

the head and neck there is an abundance of much longer and darker hair,

which forms a mane concealing the eyes, ears, and the bases of the horns.

There is also a long beard beneath the chin; while a line of long hair

extends from the head nearly to the tail, the latter being tufted at the

extremity. The withers are much higher than the hind quarters, so that

there is a kind of hump at the shoulders.

The group is represented by two species—the European and the American

Bison. The former is the _Bos bonasus_ of Linnæus, and is also identical

with the _Bos bison_ of Ray. The German name _Wisent_ is the equivalent

of the Greek _Bison_. The American species is the _Bos americanus_ of

Gmelin. Both species are closely allied, but the American Bison is

slightly the smaller animal of the two, and is shorter and weaker in

the hind quarters, with a smaller pelvis; its body is, however, more

massive in front; and the hair on the head, neck, and fore quarters is

longer and more luxuriant. A large bull American Bison, preserved in the

Museum at Washington, stands 5 feet 8 inches in height at the withers.

The European Bison appears to have been formerly abundant over a large

portion of Europe in the Pleistocene period—the fossil race described as

_B. priscus_ not being specifically distinct; but at the present day it

exists only in the primeval forests of Lithuania, Moldavia, Wallachia,

and the Caucasus, where it is artificially preserved.

[Illustration: FIG. 148.—The American Bison (_Bos americanus_). After

Hornaday.]

The American Bison formerly ranged over about one-third of the North

American continent. Thus, to quote from Mr. Hornaday,[251] “starting

almost at tide-water on the Atlantic coast, it extended across the

Alleghany mountain system to the prairies along the Mississippi, and

southward to the delta of that great system. Although the great plain

country of the West was the natural home of the species, where it

flourished most abundantly, it also wandered south across Texas to

the burning plains of North-Eastern Mexico, westward across the Rocky

Mountains into New Mexico, Utah, and Idaho, and northward across a

vast treeless waste to the bleak and inhospitable shores of the Great

Slave Lake itself.” In consequence of the settlement of the country by

Europeans the area inhabited by the Bison was gradually contracted, till

about 1840 one mighty herd occupied the centre of its former range.

The completion of the Union Pacific Railway in 1869 divided this great

herd into a southern and a northern division, the former comprising

a number of individuals estimated at nearly four millions, while the

latter contained about a million and a half. Before 1880 the southern

herd had, however, practically ceased to exist; while the same fate

overtook the northern one in 1883. In 1889 some twenty stragglers in

Texas represented the last of the southern herd; while there were a few

others in Colorado, Wyoming, Montana, and Dakota. A herd of some two

hundred wild individuals, derived from the northern herd, is preserved by

the United States Government in the Yellowstone National Park; and it is

believed that some five hundred of the race known as Wood-Bison exist in

British territory; but with these exceptions this magnificent species is

exterminated. The multitudes in which the American Bison formerly existed

are almost incredible; the prairies being absolutely black with them as

far as the eye could reach, and the numbers in the herds being, as we

have said, reckoned by millions. Mr. Hornaday even considers that the

whole of the game in South Africa was never equal to the number of Bison

on an equal area of the American prairies.

[Illustration: FIG. 149.—The Yak (_Bos grunniens_), domestic variety.]

An extinct Bison from the Pleistocene of Texas, known as _Bos latifrons_,

was probably the ancestor of the recent American species.

The Yak (_Bos grunniens_) appears to be allied both to the Bisons and the

true Oxen, being distinguished from the former by the different position

occupied by the long hair, which forms a fringe investing the shoulders,

flanks, and thighs, and grows over the whole of the tail. In the skull

the orbits are less tubular, the forehead flatter, and the premaxillæ

less widely separated from the nasals. There is no distinct dewlap.

Wild Yaks inhabit the higher regions of Chinese Tibet and the region of

the Karakoram, as well as the more outlying parts of Ladak, such as the

Changchemo valley. Owing, however, to incessant pursuit those now found

within the territories of the Maharaja of Kashmir are stragglers from

Chinese Tibet. The height of the Yak is somewhat lower than that of the

larger domestic cattle. The colour of the wild race is black, tending

to brown on the flanks; but many of the tame breeds which have been

crossed with ordinary cattle have more or less white (Fig. 149), and it

is the white tails of these half-breeds that are so esteemed in India as

“chowries.” Yaks are exceedingly intolerant of heat, and the wild ones

always live at very great elevations. Tame Yaks are extensively used as

beasts of burden in Tibet, where they are extremely valuable in crossing

the high and desolate wastes of that region; they have, however, the

great drawback that they refuse to eat corn, so that in districts where

there is no grass it is frequently necessary to make forced marches

with wearied beasts in order to prevent them (and thus the whole party)

perishing from starvation.

The skull of an extinct species from the Pliocene of Northern India,

described as _Bos sivalensis_, appears to indicate a species allied to

the Yak.

With the Bibovine group we come to the consideration of three Oriental

species which connect the preceding forms with the typical Oxen. The

three species are the Gaur (_B. gaurus_) the Gayal (_B. frontalis_,

Fig. 150) of India, and the Banteng (_B. sondaicus_) of Burma, Java,

Bali, and Lambok. In this group, as in the true Oxen, there are thirteen

pairs of ribs, against fourteen in the Bisons. All the three species

are characterised by the great height of the spines of the anterior

dorsal vertebræ, causing a prominent ridge down the back. The horns,

which are of a greenish colour in the Gaur, are somewhat flattened, and

after running outwards are directed upwards instead of backwards; they

occupy the vertex of the skull. The frontals are more or less concave,

the premaxillæ do not join the nasals, and the occipital aspect of

the skull is characterised by the deep incisions made by the temporal

fossæ. The lower part of the legs is white (Fig. 150), and the hoofs are

comparatively small and pointed. The Gaur (_B. gaurus_) is the largest

of the three species, and inhabits all the large forests of India from

near Cape Comorin to the foot of the Himalaya; it is commonly known

to sportsmen as the Indian Bison. It stands fully 6 feet in height

at the withers, which are much elevated; and since the whole back is

arched the line from the nose to the root of the tail forms an almost

continuous curve. The most characteristic feature of the animal is,

however, the large and convex intercornual frontal crest, which curves

forward, and thus gives a concave profile to this part of the skull. As

a rule the Gaur prefers hilly regions, although it is sometimes met with

on the flat. It is very shy and readily frightened; and it has never

been domesticated. The Gayal, or Mithan, of which a figure is given in

woodcut 150, is at once distinguished from the Gaur by the straight line

between the horns (which are black in colour), owing to the absence of

the intercornual crest of the latter. The horns are also shorter, more

rounded, and less curved. In the Indian Museum, Calcutta, there are,

however, skulls which are to a great extent intermediate between those

of typical Gaurs and those of typical Gayals, but these may belong to

hybrids. The Gayal occurs in Assam, Chittagong, and adjacent districts,

but it appears that these animals exist in a semi-domesticated condition,

no wild race being known to Europeans, although it is probable that such

may exist in the unexplored Mishmi Hills.

[Illustration: FIG. 150.—The Gayal (_Bos frontalis_). From Sclater, _List

of Animals in Zoological Society’s Gardens_, 1883.]

The Banteng (_B. sondaicus_) is a smaller and lighter built animal

than either of the preceding, with a longer and sharper head, and more

rounded and slender horns. The dorsal ridge is, moreover, but slightly

developed; while the bright dun colour of the body of the female readily

distinguishes it from the darker hue of the Gaur and Gayal.

A fossil skull from the Pleistocene deposits of the Narbada valley,

India, described as _Bos palæogaurus_, is believed to indicate a species

nearly allied to the Gaur, if indeed it be specifically distinct.

The true Oxen, or Taurine group, are now represented solely by _Bos

taurus_ and _Bos indicus_. Both of these species are now known only

by domesticated races, unless the herds of the former preserved at

Chillingham and some other British parks are the survivors of an original

wild race. The dorsal ridge of the Bibovine group is here wanting;

the horns are rounded, with their extremities directed backwards, and

are placed at the extreme vertex of the skull; while the long frontal

region is nearly flat; the temporal fossæ scarcely intrude upon the

occipital aspect of the skull; and the premaxillæ reach the nasals. The

hoofs are large and rounded. It is known that wild Oxen were abundant

in the forests of Europe at the time of Julius Cæsar, by whom they were

described as the Urus, equal to the German Aurochs; and the large skulls

found in turbary and Pleistocene deposits, and described under the name

of _Bos primigenius_, can only be regarded as having belonged to the

large original race of _B. taurus_, of which it has been thought the

Chillingham cattle are smaller descendants.[252] The subfossil skulls

described as _B. longifrons_ and _B. frontosus_ must also be looked upon

as referable to smaller races of the same species. That the domestic

cattle of Europe are descendants from the various races of the same

original species there can be no doubt, but in the case of the humped

cattle of India (_B. indicus_) it is quite probable that their origin

may be, at least in part, different. The extinct _Bos namadicus_, of the

Pleistocene deposits of India, was a species with the general characters

of the Taurine group, but with an inclination to a flattening of the

horns, and with an approximation to a Bibovine type of occiput, as well

as with the separation of the premaxillæ from the nasals.

The earliest representatives of this group occur in the Pliocene of the

Siwalik Hills in Northern India. One of these species (_B. planifrons_)

appears to be allied to _B. namadicus_; but the other (_B. acutifrons_)

was a gigantic species characterised by the sharp median angulation of

the frontal region, and the pyriform section of the enormous horn-cores.

The extinct _B. elatus_, from the Upper Pliocene of France and Italy,

is the representative of a generalised type, which may be known as the

Leptobovine group. The males had rounded horn-cores widely separated at

their bases, and placed low down on the forehead. The females (which have

been described as _Leptobos_) were often or always hornless. The limbs

were unusually slender. This group also occurs in the Pliocene of the

Siwalik Hills.

_Suborder_ PERISSODACTYLA.

This is a perfectly well-defined group of Ungulate mammals, represented

in the actual fauna of the world by only three distinct types or

families—the Tapirs, the Rhinoceroses, and the Horses—poor in genera

and species, and (except in the case of the two domesticated species of

_Equus_, which have been largely multiplied and diffused by man’s agency)

not generally numerous in individuals, though widely scattered over the

earth’s surface. Palæontological records, however, show very clearly that

these are but the surviving remnants of a very extensive and much-varied

assemblage of animals, which flourished upon the earth through the

Tertiary geological period, and which, if it could be reconstructed in

its entirety, would not only show members filling up structurally the

intervals between the existing apparently isolated forms, but would also

show several marked lines of specialisation which have become extinct

without leaving any direct successors.

[Illustration: FIG. 151.—Bones of right fore foot of existing

Perissodactyles. A, Tapir (_Tapirus indicus_), × ⅕; B, Rhinoceros

(_Rhinoceros sumatrensis_), × ⅙; C, Horse (_Equus caballus_), × ⅛. _U_,

ulna; _R_, radius; _c_, cuneiform; _l_, lunar; _s_, scaphoid; _u_,

unciform; _m_, magnum; _td_, trapezoid; _tm_, trapezium.—From Flower,

_Osteology of Mammalia_.]

The following are the principal characters distinguishing them from

the Artiodactyla. Premolar and molar teeth in continuous series, with

massive, quadrate, transversely ridged or complex crowns,—the posterior

premolars often resembling the true molars in size and structure. Crown

of the last lower molar commonly bilobed, and if a third lobe is present

in this tooth it is wanting in the last lower milk-molar. Dorso-lumbar

vertebræ never fewer than twenty-two, usually twenty-three in the

existing species. Nasal bones expanded posteriorly. An alisphenoid

canal. Femur with a third trochanter.[253] The middle or third digit on

both fore and hind feet larger than any of the others, and symmetrical

in itself, the free border of the ungual phalanx being evenly rounded

(see Fig. 151). This may be the only functional toe, or the second and

fourth may be subequally developed on each side of it. In the Tapirs

and many extinct forms, the fifth toe also remains on the fore limb,

but its presence does not interfere with the symmetrical arrangement of

the remainder of the foot around the median line of the third or middle

digit. Traces of a hallux have only been found in some extremely ancient

and primitive forms. The astragalus has a pulley-like surface above

for articulation with the tibia, but its distal surface is flattened

and unites to a much greater extent with the navicular than with the

cuboid, which bone is of comparatively less importance than in the

Artiodactyla. The calcaneum does not articulate with the lower or distal

extremity of the fibula. The stomach is always simple, the cæcum is large

and capacious, the placenta diffused, and the mammæ are inguinal. The

gall-bladder is invariably absent.

As regards the dentition, the whole of the premolar series may be

preceded by milk-teeth; and it has been demonstrated in _Rhinoceros_

that when there is no displacement of the first cheek-tooth that tooth

is a persistent milk-molar; the same condition apparently holding

good in _Palæotherium._ This feature indicates considerable dental

specialisation, the milk-molars, according to the theory generally

accepted by the leading English zoologists, being the acquired, and the

premolars the original series. Another peculiar feature of the dentition

of the Perissodactyla, very rarely met with among the Artiodactyla,

is that the premolars tend to resemble the true molars; this feature

occurring in all the existing genera, although not found in the earlier

generalised types. The cheek-teeth of all the members of the suborder

are primarily constructed on some modification of what is known as the

lophodont plan. Thus the upper molars (Fig. 155, p. 375) have an outer

antero-posterior wall from which proceed two transverse ridges, formed

by the coalescence of the primitive inner and outer columns, towards

the inner aspect of the crown; while in the lower molars there may be

either two simple transverse ridges, or these ridges may be curved into

crescents, coming into contact with one another at their extremities.

Those forms having brachydont teeth show this plan of structure in its

simplest modification; but in cases, as in the Horse, where the teeth

assume an extremely hypsodont form, the original plan is so obscured by

infoldings of the enamel that it can only be traced with difficulty.

At the present day the Perissodactyla are sharply differentiated into

Horses, Tapirs, and Rhinoceroses, but the knowledge already gained of

the extinct representatives of the suborder shows such a close alliance

between these groups that it is exceedingly difficult to make any

satisfactory classification of the whole. This is of course exactly what

might have been expected; and the same would doubtless be the case with

all other groups if we knew as much of their past history as we do of

that of the Perissodactyles.

The detailed account of the anatomy of the Horse given in the sequel will

afford much information as to the general structure of the members of the

suborder.

_Family_ TAPIRIDÆ.

Both upper and lower cheek-teeth brachydont and simply bilophodont;

hinder premolars as complex as the molars; last lower molar without third

lobe; first upper cheek-tooth with a milk-predecessor.[254] Outer columns

of upper molars conical. Four digits in the manus, and three in the pes.

_Tapirus._[255]—Dentition _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₃, _m_ ³⁄₃; total 42.

Of the upper incisors, the first and second are nearly equal, with short,

broad crowns; the third is large and conical, considerably larger than

the canine, which is separated from it by an interval. Lower incisors

diminishing in size from the first to the third; the canine, which is

in contact with the third incisor, large and conical, working against

(and behind) the canine-like third upper incisor. In both jaws there

is a diastema between the canines and the commencement of the teeth of

the cheek-series, which are all in contact. First upper premolar with a

triangular crown, narrow in front owing to the absence of the anterior

inner cusp. The other upper premolars and molars all formed on the same

plan and of nearly the same size, with four roots and quadrate crowns,

rather wider transversely than from before backwards, each having four

cusps, connected by a pair of transverse ridges, anterior and posterior.

The first lower premolar compressed in front; the others composed of a

simple pair of transverse crests, with a small anterior and posterior

circular ridge.

Skull elevated and compressed. Orbit and temporal fossa widely

continuous, there being no true postorbital process from the frontal

bone. Anterior narial apertures very large, and extending high on the

face between the orbits; nasal bones short, elevated, triangular, and

pointed in front. Vertebræ: C 7, D 18, L 5, S 6, C about 12. Limbs short

and stout. Forefeet with four toes, having distinct hoofs: the first

is absent, the third the longest, the second and fourth nearly equal,

the fifth the shortest and scarcely reaching the ground in the ordinary

standing position. Hind feet with the typical Perissodactyle arrangement

of three toes,—the middle one being the largest, the two others nearly

equal. Nose and upper lip elongated into a flexible, mobile snout or

short proboscis, near the end of which the nostrils are situated. Eyes

rather small. Ears of moderate size, ovate, erect. Tail very short. Skin

thick and but scantily covered with hair.

The existing species of Tapir may be grouped into two sections, the

distinctive characters of which are only recognisable in the skeleton.

(A) With a great anterior prolongation of the ossification of the nasal

septum (mesethmoid), extending in the adult far beyond the nasal bones,

and supported and embraced at the base by ascending plates from the

maxillæ (genus _Elasmognathus_, Gill). Two species, both from Central

America, _Tapirus bairdi_ and _T. dowi_. The former is found in Mexico,

Honduras, Nicaragua, Costa Rica, and Panama; the latter in Guatemala,

Nicaragua, and Costa Rica. (B) With ossification of the septum not

extending farther forward than the nasal bones (_Tapirus_ proper).

Three species, _T. indicus_, the largest of the genus, from the Malay

Peninsula (as far north as Tavoy and Mergui), Sumatra, and Borneo,

distinguished by its peculiar coloration, the head, neck, fore and hind

limbs, being glossy black, and the intermediate part of the body white;

_T. americanus_ (_T. terrestris_, Linn.), the common Tapir of the forests

and lowlands of Brazil and Paraguay (Fig. 152); and _T. roulini_, the

Pinchaque Tapir of the high regions of the Andes. All the American

species are of a nearly uniform dark brown or blackish colour when adult;

but it is a curious circumstance that when young (and in this the Malay

species conforms with the others) they are conspicuously marked with

spots and longitudinal stripes of white or fawn colour on a darker ground.

The habits of all the kinds of Tapirs appear to be very similar. They are

solitary, nocturnal, shy, and inoffensive, chiefly frequenting the depths

of shady forests and the neighbourhood of water, to which they frequently

resort for the purpose of bathing, and in which they often take refuge

when pursued. They feed on various vegetable substances, as shoots of

trees and bushes, buds, and leaves. They are hunted by the natives of the

lands in which they live for the sake of their hides and flesh.

The singular fact of the existence of so closely allied animals as the

Malayan and the American Tapirs in such distant regions of the earth,

and in no intervening places, is accounted for by what is known of

the geological history of the race; for the Tapirs must once have had

a very wide distribution. There is no proof of their having lived in

the Eocene epoch, but in deposits of Miocene and Pliocene date remains

undistinguishable generically from the modern Tapirs, and described as

_T. priscus_, _T. arvernensis_, etc., have been found in France, Germany,

and in the Red Crag of Suffolk. Tapirs appear, however, to have become

extinct in Europe before the Pleistocene period, since none of their

bones or teeth have been found in any of the caverns or alluvial deposits

in which those of Elephants, Rhinoceroses, and Hippopotamuses occur in

abundance; but in other regions their distribution at this age was far

wider than at present, as they are known to have extended eastward to

China (_T. sinensis_, Owen) and westwards over the greater part of the

southern United States of America, from South Carolina to California.

Lund also distinguished two species or varieties from the caves of

Brazil, one of which appears identical with _T. americanus_. Thus we

have no difficulty in tracing the common origin in the Miocene Tapirs of

Europe of the now widely separated American and Asiatic species. It is,

moreover, interesting to observe how very slight an amount of variation

has taken place in forms isolated during such an enormous period of time.

[Illustration: FIG. 152.—The American Tapir (_Tapirus americanus_).]

The anatomy of the soft parts of the Tapirs[256] conforms to the general

Perissodactyle type, as exemplified in the Rhinoceros and the Horse,

although on the whole (as might have been expected) presenting a closer

resemblance to the former. _T. americanus_ differs from _T. indicus_

by the absence, or at any rate the less development, of the intestinal

valvulæ conniventes, the presence of a moderator band in the heart,

the shape of the glans penis, and the more elongated cæcum, which is

sacculated by four distinct longitudinal fibrous bands. The convolutions

of the hemispheres of the brain of the Tapirs are simpler than in other

Perissodactyles, thus tending to confirm the inferences which may

be drawn from the skeleton and teeth as to the comparatively low or

generalised organisation of these animals.

_Palæotapirus._—This name has been applied to an imperfectly known form

from the Upper Eocene Phosphorites of Central France, which is regarded

by Dr. Filhol as referable to this family.

_Family_ LOPHIODONTIDÆ.

Molars brachydont and bilophodont, those of the lower jaw with either

straight or imperfectly crescentoid ridges; premolars smaller and usually

simpler than the molars; last lower molar generally with a third lobe.

Outer columns of upper molars conical or flattened. Digits usually as in

the preceding family.

[Illustration: FIG. 153.—Right side of skull of _Hyracotherium

leporinum_, from the London Clay. ½ natural size. (After Owen.) 3,

Occiput; 7, sagittal crest; 11, frontals; 15, nasals; 21, maxilla; 22,

premaxilla; _d_, mandibular condyle; _a_, aperture of facial nerve; _p_

1-4, premolars; _m_ 1-3, molars.]

This family includes a number of more or less imperfectly known

forms, all of which are extinct and apparently confined to the Eocene

period, and ranging from the size of a Rabbit to that of a Rhinoceros.

Although some of these appear to have died out without giving rise to

more specialised forms, it is probable that this family contained the

ancestral types from which most or all of the modern Perissodactyles

have been derived. Only very brief mention can be made here of some

of the leading genera. _Lophiodon_, of the Middle and Upper Eocene of

Europe, with the dental formula, _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ³⁄₃, _m_ ³⁄₃,

includes the largest representatives of the family, and is generally

regarded as a stock which has died out without giving rise to later

forms. The ridges of the lower molars are straight, and the last of these

teeth has a third lobe; while the second transverse ridge of the last

upper premolar is usually incomplete; the outer columns of the upper

molars are flattened, as in the next genus. _Hyrachyus_, of the Upper

Eocene of the United States, and probably also occurring in the French

Eocenes, is an allied genus, with four premolars and no third lobe to

the last lower molar; the fourth upper premolar having the two ridges

uniting internally to form a crescent. This genus has been regarded as

the ancestor of the Rhinocerotic _Hyracodon_. The genus _Hyracotherium_

was established in 1839 by Owen for a small animal no larger than a

Hare, the skull of which was found in the London Clay at Herne Bay.

A more nearly perfect specimen, apparently of the same species, was

afterwards (in 1857) described under the name of _Pliolophus vulpiceps_,

of which the skull is figured in the accompanying woodcut. Other forms

referable to the same genus have been obtained from the Wasatch Eocene

of the United States, and were described by Professor Marsh under the

name of _Eohippus_. There were four premolars, the fourth being unlike

the molars, and in the upper jaw having only one inner cusp. The upper

molars are of the general type of those of _Lophiodon_, but have conical

outer columns, and the anterior transverse ridge imperfect, while

the ridges of the lower molars are crescentoid. _Systemodon_ differs

from _Hyracotherium_ by the absence of a diastema between the first

and second premolars; it occurs in the Wasatch Lower Eocene of the

United States. In _Pachynolophus_ (_Lophiotherium_, _Orotherium_, or

_Orohippus_), which is common to the Middle and Upper Eocene of Europe

and the Bridger Eocene of North America, the outer columns of the upper

molars are flattened, and in some cases, at least, the last premolar

resembles the molars, that of the upper jaw having two inner cusps.[257]

This genus, indeed, so closely connects _Hyracotherium_ with the genera

_Epihippus_ and _Anchilophus_ as to show that the distinction between

the _Lophiodontidæ_ and _Palæotheriidæ_ is really an arbitrary one.

_Epihippus_, of the Upper Eocene of the United States, has both the third

and fourth upper premolars as complex in the molars, and is distinguished

from _Anchilophus_ by the lower cusps and more imperfect transverse

ridges of these teeth. The so-called _Orohippus agilis_ belongs to this

genus. _Isectolophus_ is another American Eocene genus which may be

provisionally placed in this family; it is regarded by Professors Scott

and Osborn as connecting _Systemodon_ with the _Tapiridæ_; the fourth

and probably the third upper premolar approximating in structure to the

molars; the upper molars have conical outer columns. _Helaletes_ is

another closely allied form, with similar premolars, but with the outer

columns of the upper molars flattened.

[Illustration: FIG. 154.—Restoration of _Palæotherium_ (Upper Eocene).

After Cuvier.]

_Family_ PALÆOTHERIIDÆ.

[Illustration: FIG. 155.—A half-worn right upper molar of _Palæotherium

magnum_. (After Owen.) _f_, _f_, External surfaces of outer columns;

_a_, postero-external column (metacone); _b_, antero-external column

(paracone); _c_, postero-internal column (hypocone); _d_, antero-internal

column (protocone); _i_, anterior intermediate column (protoconule); _e_,

median valley; _g_, posterior valley.]

Molars (Fig. 155) brachydont, with the valleys between the ridges never

filled with cement; upper premolars either simpler than or as complex

as the molars; lower molars with crescentoid ridges, and the last of

the series with or without a third lobe. Outer columns of upper molars

flattened. Orbit (at least usually) confluent with temporal fossa. Three

digits on each foot. This family includes extinct genera ranging from

the Middle and Upper Eocene to the Miocene, and passes so gradually into

the following one that the maintenance of the two can only be supported

on the ground of convenience. The typical genus, _Palæotherium_, was

made known to science in the early part of the present century by

Cuvier, who restored the skeleton (Fig. 154) with a short neck like that

of the Tapirs, although it has been subsequently found that the neck

was considerably longer. This genus (which may be taken to include

_Paloplotherium_) ranges from the Middle to the Upper Eocene of Europe,

and usually has the full typical dentition, although the first premolar

may disappear. The last lower molar has a third lobe; and in the typical

forms the last premolar is as complex as the molars, the diastema is

short, and the canines are not large. In other forms, however, the

hinder ridge of the fourth upper premolar may be aborted. The first

upper cheek-tooth is generally a well-developed tooth, which may have

a deciduous predecessor. _Anchilophus_, of the Upper Eocene of Europe,

and _Anchitherium_, of the Miocene of Europe and North America, connect

the preceding forms with the _Equidæ_. In the latter genus there is the

full number of teeth, the last lower molar has almost completely lost

the third lobe of _Anchilophus_, and the surfaces of the two outer lobes

of the upper molars (Figs. 157, 158) lack the median vertical ridges of

that genus. In the American species of _Anchitherium_ (which have been

described as _Mesohippus_ and _Miohippus_) the lateral digits are larger

than in the European Middle Miocene _Anchitherium aurelianense_; a mere

splint represents the fifth metacarpal, and the meso- and ento-cuneiform

of the tarsus do not unite as they do in the latter.

_Family_ EQUIDÆ.

Molars hypsodont, with the outer columns of the upper ones flattened, the

valleys completely filled with cement, and the enamel thrown into folds

and plications; upper premolars as complex as molars, which they slightly

exceed in size; ridges of lower molars crescentoid, and complicated by

enamel-foldings; no distinct third lobe to last lower molar; summits of

incisors with a central infolding of enamel. Orbit completely surrounded

by bone. Digits three or one, but in the former case the median one is

alone of functional importance; ulna and fibula incomplete; meso- and

ento-cuneiform of tarsus united.

Such are the leading characters which serve to distinguish the existing

Horses and their nearest fossil allies from the _Palæotheriidæ_. The

Horse, as being the best known of the Perissodactyle Ungulates, is

selected for a somewhat detailed description; but before proceeding

to this it will be advisable to take a brief survey of the relations

of the _Equidæ_ to the extinct forms already noticed, and also of the

modifications of the family at present existing.

The earliest form which can be certainly included in this line of descent

is the American Lower Eocene genus _Phenacodus_ (noticed below under the

head of the suborder Condylarthra), in which there were five complete

digits to the feet. From this form there is but a step to _Systemodon_

and _Hyracotherium_, in which the functional digits of the manus were

reduced to four, as in _Pachynolophus_ (Fig. 156, _a_), although one

species retained a rudiment of the metacarpal of the pollex.

[Illustration: FIG. 156.—Successive stages of modification of the feet

of extinct forms of Horse-like animals (chiefly from Marsh), showing

gradual reduction of the outer and enlargement of the middle toe (III).

_a_, _Pachynolophus_ (Eocene); _b_, _Anchitherium_ (Early Miocene); _c_,

_Anchitherium_ (Late Miocene); _d_, _Hipparion_ (Pliocene); _e_, _Equus_

(Pleistocene).]

The transition from these animals of the Eocene period to the Horses of

modern times has been accompanied by a gradual increase in size. The

diminutive _Hyracotherium_ of the Lower, and _Pachynolophus_ of the

Middle and Upper Eocene were succeeded in the Miocene period by the

forms to which the name of _Anchitherium_ has been given, of the size of

sheep; these again in Pliocene times by _Hipparion_ and _Protohippus_, as

large as the modern donkeys; and it is mainly in the Pleistocene period

that _Equidæ_ occur which approach in size the existing Horse. Important

structural modifications have also taken place, with corresponding

changes in the mode of life of the animal. Thus the neck has become

elongated, the skull altered in form, the teeth greatly modified, and

the limbs have undergone remarkable changes. The last two require to be

described more in detail.

[Illustration: FIG. 157.—_a_, Grinding surface of unworn molar tooth

of _Anchitherium_; _b_, corresponding surface of unworn molar of young

Horse; _c_, the same tooth after it has been some time in use. The

uncoloured portions are the dentine or ivory, the shaded parts the

cement filling the cavities and surrounding the exterior. The black line

separating these two structures is the enamel or hardest constituent of

the tooth.]