Deer

Deer (O. E. déor, díor, a common Teutonic word, meaning a wild animal, cf. Ger. Tier, Du. dier, &c., probably from a root dhus-, to breathe), originally the name of one of two British species, the red-deer or the fallow-deer, but now extended to all the members of the family Cervidae, in the section Pecora of the suborder Artiodactyla of the order Ungulata. (See Pecora; Artiodactyla and Ungulata.) Briefly, deer may be defined as Pecora presenting the following characteristics:—either antlers present in the male, or when these are absent, the upper canines large and sabre-like, and the lateral metacarpal bones represented only by their lower extremities. This definition will include the living and also most of the extinct forms, although in some of the latter the lateral metacarpal bones not only retain their lower ends, but are complete in their entire length.

The leading characters of antlers are described under Pecora, but these structures may be defined somewhat more fully in the following passage from the present writer’s Deer of all Lands:—

“Antlers are supported on a pair of solid bony processes, or pedicles, arising from the frontal bones of the skull, of which they form an inseparable portion; and if in a fully adult deer these pedicles be sawn through, they will generally be found to consist of solid, ivory-like bone, devoid of perceptible channels for the passage of blood-vessels. The pedicles are always covered with skin well supplied with blood-vessels; and in young deer, or those in which the antlers have been comparatively recently shed, the covering of skin extends over their summits, when they appear as longer or shorter projections on the forehead, according to the species. When the first or a new antler is about to be formed, the summits of these pedicles become tender, and bear small velvet-like knobs, which have a high temperature, and are supplied by an extra quantity of blood, which commences to deposit bony matter. This deposition of bony matter progresses very rapidly, and although in young deer and the adults of some species the resulting antler merely forms a simple spike, or a single fork, in full-grown individuals of the majority it assumes a more or less complexly branched structure. All this time the growing antler is invested with a skin clothed with exceedingly fine short hairs, and is most liberally supplied with blood-vessels; this sensitive skin being called the velvet. Towards the completion of its growth a more or less prominent ring of bone, termed the burr or coronet, is deposited at its base just above the junction with the pedicle; this ring tending to constrict the blood-vessels, and thus cut off the supply of blood from the antlers....

“When the antlers are freed from the velvet—a process usually assisted by the animal rubbing them against tree stems or boughs—they have a more or less rugose surface, owing to the grooves formed in them by the nutrient blood-vessels. Although a few living species have the antlers in the form of simple spikes in the adult male, in the great majority of species they are more or less branched; while in some, like the elk and fallow-deer, they expand into broad palmated plates, with tines, or snags, on one or both margins. In the antlers of the red-deer group, which form the type of the whole series, the following names have been applied to their different component parts and branches. The main shaft is termed the beam; the first or lowest tine the brow-tine; the second the bez-tine; the third the trez-tine, or royal; and the branched portion forming the summit the crown, or surroyals. But the antlers of all deer by no means conform to this type; and in certain groups other names have to be adopted for the branches.

“The antlers of young deer are in the form of simple spikes; and this form is retained in the South American brockets, although the simple antlers of these deer appear due to degeneration, and are not primitive types. Indeed, no living deer shows such primitive spike-like antlers in the adult, and it is doubtful whether such a type is displayed by any known extinct form, although many have a simple fork. In the deer of the sambar group, where the antlers never advance beyond a three-tined type, the shedding is frequently, if not invariably, very irregular; but in the majority at least of the species with complex antlers the replacement is annual, the new appendages attaining their full development immediately before the pairing-season. In such species there is a more or less regular annual increase in the complexity of the antlers up to a certain period of life, after which they begin to degenerate.”

The Cervidae are distributed all over Europe, Asia, Northern Africa and America, but are unknown in Africa south of the Sahara. They are undoubtedly a group of European or Asiatic origin, and obtained an entrance into America at a time when that continent was connected with Asia by way of Bering Strait.

The existing members of the family are classified in the writer’s Deer of all Lands as follows:—

A. Subfamily Cervinae.—Antlers, with one exception, present in the male; liver without a gall-bladder; a face-gland, and a gland-pit in the skull.

I. Reindeer, Genus Rangifer.—Lateral metacarpal bones represented only by their lower extremities; antlers present in both sexes, complex. Northern part of both hemispheres.

II. Elk, Genus Alces.—Lateral metacarpals as in preceding; antlers (as in the following genera) present only in the male, arising at right angles to the median longitudinal line of the skull, and extending at first in the plane of the forehead, after which, when in their fullest development, they expand into a broad palmation margined with snags. Northern portion of both hemispheres.

III. True Deer, Genus Cervus.—Lateral metacarpals represented only by their upper ends. Antlers arising at acute angles to the median line of the skull (as in the following genera), at first projecting from the plane of the forehead, and then continued upwards nearly in that plane, supported on short pedicles, and furnished with a brow-tine, never regularly forked at first division, but generally of large size, and with not less than three tines; the skull without ridges on the frontals forming the bases of the pedicles of the antlers. Upper canine teeth small, or wanting. Europe, Asia and N. America.

1\. Red-deer Group, Subgenus Cervus.—Antlers rounded, usually with five or more tines, generally including a bez (second), and always a trez (third); coat of adult generally unspotted, with a large light-coloured disk surrounding the tail; young, spotted. Europe, Northern and Central Asia and North America.

2\. Sika Deer, Subgenus Pseudaxis.—Antlers smaller and simpler, four-tined, with a trez (third), but no bez (second); coat of adult spotted, at least in summer, with a white area bordered by black in the region of the tail, which is also black and white. North-Eastern Asia.

3\. Fallow-deer, Subgenus Dama.—Antlers without a bez, but with a trez-tine, above which the beam is more or less palmated, and generally furnished with numerous snags; coat of adult spotted in summer, uniform in winter, with black and white markings in the region of the tail similar to those of Pseudaxis; young, spotted. Mediterranean region, but more widely spread in Europe during the Pleistocene epoch, and also introduced into many European countries.

4\. Sambar Group, Subgenus Rusa.—Antlers rounded, three-tined, with the bez- and trez-tines wanting, and the beam simply forked at the summit; coat either uniform or spotted at all seasons. Indo-Malay countries and part of China.

5\. Barasingha Group, Subgenus Rucervus.—Antlers flattened or rounded, without bez- or trez-tine, the beam dichotomously forking, and one or both branches again forked, so that the number of tines is at least four; brow-tine forming a right angle or a continuous curve with the beam; coat of adult generally more or less uniform, of young spotted. Indo-Malay countries.

IV. Muntjacs, Genus Cervulus.—Lateral metacarpals as in Cervus; antlers small, with a brow-tine and an unbranched beam, supported on long bony pedicles, continued downwards as convergent ridges on the forehead; upper canines of male large and tusk-like. Indo-Malay countries and China.

V. Tufted Muntjacs, Genus Elaphodus.—Nearly related to the last, but the antlers still smaller, with shorter pedicles and divergent frontal ridges; upper canines of male not everted at the tips. Tibet and China.

VI. Water-deer, Genus Hydrelaphus.—Lateral metacarpals as in Rangifer; antlers wanting; upper canines of males tusk-like and growing from semi-persistent pulps; cheek-teeth tall-crowned (hypsodont); tail moderate. China.

VII. Roe-deer, Genus Capreolus.—Lateral metacarpals as in Rangifer; antlers rather small, without a brow-tine or sub-basal snag, dichotomously forked, with the upper or posterior prong again forking; tail rudimentary; vomer not dividing posterior nasal aperture of skull. Europe and Northern Asia.

VIII. Père David’s Deer, Genus Elaphurus.—Lateral metacarpals as in Cervus; antlers large, without a brow-tine or sub-basal snag, dichotomously forked, with the upper prong of the fork curving forwards and dividing, and the lower prong long, simple, and projected backwards, the beam making a very marked angle with the plane of the face; tail very long; vomer as in Capreolus. North-East Asia.

IX. American Deer, Genus Mazama.—Lateral metacarpals as in Rangifer; antlers very variable in size, forming a marked angle with the plane of the face, without a brow-tine; when consisting of more than a simple prong, dichotomously forked, frequently with a sub-basal snag, and always with the lower prong of the fork projected from the front edge of the beam, in some cases the lower, in others the upper, and in others both prongs again dividing; tail long; tarsal gland generally present; metatarsal gland very variable, both as regards presence and position; vomer dividing the inner aperture of the nostrils in the skull into two distinct chambers. America.

1\. White-tailed Group, Subgenus Dorcelaphus or Odocoileus.—Antlers large and complex, with a sub-basal snag, and the lower prong more or less developed at the expense of the upper one; metatarsal gland usually present; tail long or moderate, and hairy below; face very long and narrow; the face-gland small, and the gland-pit in the skull of moderate extent; no upper canines; size generally large. North America to Northern South America.

2\. Marsh-deer Group, Subgenus Blastoceros.—Antlers large and complex, without a sub-basal snag, and the upper prong more developed than the lower one; metatarsal gland absent; tail short; face moderately long; face-gland and gland-pit well developed; upper canines usually present in male. Size large or rather small. South America.

3\. Guemals, Subgenus Xenelaphus.—Antlers small and simple, forming a single dichotomous fork; metatarsal gland absent; tail short; face moderately long; face-gland and gland-pit well developed; upper canines present in both sexes. Size medium. South America.

4\. Brockets, Subgenus Mazama.—Antlers in the form of simple unbranched spikes; metatarsal, and in one case also the tarsal gland absent; tail very short; face elongated; face-gland small and gland-pit deep and triangular; hair of face radiating from two whorls: upper canines sometimes present in old males. Size small. Central and South America.

X. Genus Pudua.—Skull and metacarpals generally as in Mazama; size very small; hair coarse and brittle; antlers in the form of short, simple spikes; cannon-bones very short; tail very short or wanting; no whorls in the hair of the face; face-gland moderately large, and gland-pit deep and oval; tarsal and metatarsal glands wanting; ectocuneiform bone of tarsus united with the naviculocuboid. South America.

B. Subfamily Moschinae.—Antlers wanting in both sexes; liver furnished with a gall-bladder; no face-gland or gland-pit.

XI. Musk-deer, Genus Moschus.—Hair coarse and brittle; upper canines of male very long; no tarsal or metatarsal glands or tufts; lateral metacarpals represented by their lower extremities; lateral hoofs very large; tail very short; naked portion of muzzle extensive; male with a large abdominal gland. Central Asia.

Of the above, Reindeer and Elk are dealt with in separate articles (qq.v.).

The first or typical group of the genus Cervus includes the red-deer (Cervus elaphus) of Europe and western Asia, of which there are several local races, such as the large C. elaphus maral of eastern Europe and Persia, which is often partially spotted above and dark-coloured below, the smaller C. e. barbarus of Tunisia and Morocco, and the still smaller C. e. corsicanus of Corsica. The Scandinavian red-deer is the typical form of the species. In all red-deer the antlers are rounded, and show a more or less marked tendency to form a cup at the summit. Wapiti, on the other hand, show a marked tendency to the flattening of the antlers, with a great development of the fourth tine, which is larger than all the others, and the whole of the tines above this in the same plane, or nearly so, this plane being the same as the long axis of the animal. Normally no cup is developed at the summit of the antler. The tail, too, is shorter than in the red-deer; while in winter the under parts become very dark, and the upper surface often bleaches almost white. The cry of the stags in the breeding season is also different. The typical representative of the group is the North American wapiti C. canadensis, but there are several closely allied races in Central Asia, such as C. canadensis songaricus and C. c. bactrianus, while in Manchuria the subgroup is represented by C. c. xanthopygus, in which the summer coat is reddish instead of grey. The hangul (C. cashmirianus) of Kashmir is a distinct dark-coloured species, in which the antlers tend to turn in at the summit; while C. yarcandensis, of the Tarim Valley, Turkestan, is a redder animal, with a wholly rufous tail, and antlers usually terminating in a simple fork placed in a transverse plane. Another Asiatic species is the great shou (C. affinis) of the Chumbi Valley, in which the antlers curve forwards in a remarkable manner. Lastly C. albirostris, of Tibet, is easily recognized by its white muzzle, and smooth, whitish, flattened antlers, which have fewer tines than those of the other members of the group, all placed in one plane.

The second group of the genus Cervus, forming the subgenus Pseudaxis, is typified by the handsome little Japanese deer, or sika, C. (P.) sika, in which the antlers are four-tined, and covered with red “velvet” when first grown, while the coat is fully spotted in summer, but more or less uniformly brown in winter. The most distinctive feature of the deer of this group is, however, the patch of long erectile white hairs on the buttocks, which, although inconspicuous when the animals are quiescent, is expanded into a large chrysanthemum-like bunch when they start to run or are otherwise excited. The patch then forms a guiding signal for the members of the herd when in flight. On the mainland of Manchuria both the typical sika, and a larger race (C. sika manchuricus), occur. A still larger and finer animal is the Pekin sika (C. hortulorum), of northern Manchuria, which is as large as a small red-deer; it is represented in the Yang-tse valley by a local race, C. h. kopschi. Formosa possesses a species of its own (C. taëvanus), which, in correlation with the perpetual verdure of that island, is spotted at all seasons.

For the fallow-deer, Cervus [Dama] dama, see Fallow-deer.

The rusine or sambar group of Cervus, of which the characteristics are given above, comprises a considerable number of long-tailed species with three-tined antlers from the Indo-Malay countries and some parts of China. The largest and handsomest is the sambar of India (Cervus [Rusa] unicolor), characterized by its massive and rugged antlers. It is represented by a number of local races, mostly of smaller size, such as the Burmese and Malay C. u. equinus, the Formosan C. u. swinhoei, and the Philippine C. u. philippinus and C. u. nigricans, of which the latter is not larger than a roe-buck, while the sambar itself is as large as a red-deer. Whether these local phases of a single variable type are best denominated races or species, must be largely a matter of individual opinion. The rusa, or Javan sambar, C. (R.) hippelaphus, is a lighter-coloured and smaller deer than the Indian sambar, with longer, slenderer and less rugged antlers. Typically from Java, this deer is also represented in the Moluccas and Timor, and has thus the most easterly range of the whole tribe. A black coat with white spots distinguishes the Philippine spotted deer, C. alfredi, which is about the size of a roe-buck; while other members of this group are the Calamianes deer of the Philippines (C. culionensis), the Bavian deer (C. kuhli) from a small island near Java, and the well-known Indian hog-deer or para (C. porcinus), all these three last being small, more or less uniformly coloured, and closely allied species. On the other hand, the larger and handsomer chital, or spotted deer (C. axis), stands apart by its white-spotted fawn-red coat and differently formed antlers.

Nearly allied to the preceding is the barasingha or rucervine group (subgenus Rucervus), in which the antlers are of a different and generally more complex character. The typical species is the Indian barasingha or swamp-deer, Cervus (Rucervus) duvauceli, a uniformly red animal, widely distributed in the forest districts of India. In Siam it is replaced by C. (R.) schomburgki, in which the antlers are of a still more complex type. Finally, we have the thamin, or Eld’s deer, C. (R.) eldi, ranging from Burma to Siam, and characterized by the continuous curve formed by the beam and the brow-tine of the antlers.

For the small eastern deer, respectively known as muntjacs (Cervulus) and tufted muntjacs or tufted deer (Elaphodus), see Muntjac; while under Water-deer will be found a notice of the Chinese representative of the genus Hydrelaphus (or Hydropotes). The roe-deer, or roe-buck (Capreolus), likewise form the subject of a separate article (see Roe-buck), as is also the case with Père David’s deer, the sole representative of the genus Elaphurus.

The American deer include such New World species as are generically distinct from Old World types. All these differ from the members of the genus Cervus in having no brow-tine to the antlers, which, in common with those of the roe-deer, belong to what is called the forked type. Including all these deer except one in the genus Mazama (of which the typical representatives are the South American brockets), the North American species constitute the subgenus Dorcelaphus (also known as Cariacus and Odocoileus). One of the best known of these is the white-tailed deer Mazama (Dorcelaphus) americana, often known as the Virginian deer. It is typically an animal of the size of a fallow-deer, reddish in summer and greyish in winter, with a long tail, which is coloured like the back above but white below, and is carried elevated when the animal is running, so as to form with the white of the inner sides of the buttocks a conspicuous “blaze.” A white fetlock-gland with a black centre is also distinctive of this species. The antlers are large and curve forwards, giving off an upright snag near the base, and several vertical tines from the upper surface of the horizontal portion. As we proceed southwards from the northern United States, deer of the white-tailed type decrease steadily in size, till in Central America, Peru and Guiana they are represented by animals not larger that a roe-buck. The most convenient plan appears to be to regard all these degenerate forms as local races of the white-tail, although here again there is room for difference of opinion, and many naturalists prefer to call them species. The large ears, brown-and-white face, short, black-tipped tail, and antlers without large basal snag serve to distinguish the mule-deer M. (D.) hemionus, of western North America; while the black tail, M. (D.) columbiana, ranging from British Columbia to California, is a smaller animal, recognizable by the larger and longer tail, which is black above and white below.

South America is the home of the marsh-deer or guazu, M. (Blastoceros) dichotoma, representing a subgenus in which the complex antlers lack a basal snag, while the hair of the back is reversed. This species is about the size of a red-deer, with a foxy red coat with black legs. The pampas-deer, M. (B.) bezoartica, of the Argentine pampas is a much smaller animal, of paler colour, with three-tined antlers. The Chilean and Peruvian Andes and Patagonia are the homes of two peculiar deer locally known as guemals (huemals), and constituting the subgenus Xenelaphus, or Hippocamelus. They are about the size of fallow-deer, and have simply forked antlers. The Chilian species is M. (B.) bisulca and the Peruvian M. (B.) antisiensis. Brockets, of which there are numerous species, such as M. rufa and M. nemorivaga, are Central and South American deer of the size of roe-bucks or smaller, with simple spike-like antlers, tufted heads and the hair of the face radiating from two whorls on the forehead so that on the nose the direction is downwards. The smallest of all deer is the Chilian pudu (Pudua pudu), a creature not much larger than a hare, with almost rudimentary antlers.

The musk-deer forms the subject of a separate article.

For deer in general, see R. Lydekker, The Deer of all Lands (London, 1898, 1908).

(R. L.*)

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