Chapter IX: The Order Ungulata (2)

_Auchenia._[189]—Dentition of adults normally: _i_ ¹⁄₃, _c_ ¹⁄₁, _p_ ²⁄₂, _m_ ³⁄₃; total 32—one of the lower premolars may, however, be wanting. In the upper jaw there is a compressed, sharp, pointed laniariform incisor near the hinder edge of the premaxilla, followed, in the male at least, by a moderate-sized, pointed, curved true canine in the anterior part of the maxilla. The isolated canine-like premolar which follows in the Camels is not present. The teeth of the molar series, which are in contact with each other, consist of two very small premolars (the first almost rudimentary) and three broad molars, constructed generally like those of _Camelus_. In the lower jaw the three incisors are long, spatulate, and procumbent; the outer ones being the smallest. Next to these is a curved, suberect canine, followed after an interval by an isolated, minute, and often deciduous simple conical premolar; then a contiguous series of one premolar and three molars, which differ from those of existing species of _Camelus_ in having a small accessory column at the anterior outer edge. The skull generally resembles that of _Camelus_, the relatively larger brain-cavity and orbits and less developed cranial ridges being due to its smaller size. The nasal bones are shorter and broader, and are joined by the premaxillæ. Vertebræ: C 7, D 12, L 7, S 4, C 15-20. Ears rather long and pointed. No dorsal hump. Feet narrow, the toes being more separated than in the camels, each having a distinct plantar pad. Tail short. Hairy covering long and woolly. Size (in existing forms) smaller, and general form lighter than in the Camels. At present and within historic times the genus is entirely confined to the western side and southernmost parts of South America, but fossil remains have been found in the caves of Brazil, in the pampas of the Argentine republic, and in Central and North America.

The word Llama, sometimes spelt Lama, is the name by which the Peruvians designated one of a small group of closely allied animals, which, before the Spanish conquest of America, were the only domesticated hoofed mammals of the country, being kept, not only for their value as beasts of burden, but also for their flesh, hides, and wool,—in fact, supplying in the domestic economy of the people the place of the horse, the ox, the goat, and the sheep of the Old World. The word is now sometimes restricted to one particular species or variety of the group, and sometimes used in a generic sense to cover the whole. Although they were often compared by early writers to sheep, and spoken of as such, their affinity to the camel was very soon perceived, and they were included in the genus _Camelus_ in the _Systema Naturæ_ of Linnæus. They were, however, separated by Cuvier in 1800 under the name of _Lama_, changed by Illiger in 1811 to _Auchenia_ (in allusion to the great length of neck, αὐχήν), a term afterwards adopted by Cuvier, and almost universally accepted by systematic zoologists, although there has been of late a disposition to revive the earlier name.

In essential structural characters, as well as in general appearance and habits, all the animals of this genus very closely resemble each other, so that the question as to whether they should be considered as belonging to one, two, or more species has been one which has led to a large amount of controversy among naturalists. The question has been much complicated by the circumstances of the great majority of individuals which have come under observation being either in a completely or partially domesticated state, and descended from ancestors which from time immemorial have been in like condition, one which always tends to produce a certain amount of variation from the original type. It has, however, lost much of its importance since the doctrine of the distinct origin of species has been generally abandoned.

The four forms commonly distinguished by the inhabitants of South America are recognised by some naturalists as distinct species, and have had specific designations attached to them, though usually with expressions of doubt, and with great difficulties in defining their distinctive characteristics. These are (1) the Llama, _Auchenia glama_ (Linn.), or _Lama peruana_ (Tiedemann); (2) the Alpaca, _A. pacos_ (Linn.); (3) the Guanaco or Huanaco, _A. huanacus_ (Molina); and (4) the Vicugna, _A. vicugna_ (Molina), or _A. vicunna_, (Cuv.) The first and second are only known in the domestic state, and are variable in size and colour, being often white, black, or piebald. The third and fourth are wild, and of a nearly uniform light-brown colour, passing into white below. They certainly differ from each other, the Vicugna being smaller, more slender in its proportions, and having a shorter head (Fig. 116) than the Guanaco (Fig. 117). It may therefore, according to the usual view of species, be considered distinct. It lives in herds on the bleak and elevated parts of the mountain range bordering the region of perpetual snow, amidst rocks and precipices, occurring in various suitable localities throughout Peru, in the southern part of Ecuador, and as far south as the middle of Bolivia. Its manners very much resemble those of the Chamois of the European Alps; and it is as vigilant, wild, and timid. The wool is extremely delicate and soft, and highly valued for the purposes of weaving, but the quantity which each animal produces is not great.

The Guanaco has an extensive geographical range, from the highlands of the Andean region of Ecuador and Peru to the open plains of Patagonia, and even the wooded islands of Tierra del Fuego. It constitutes the principal food of the Patagonian Indians, and its skin is invaluable to them, as furnishing the material out of which their long robes are constructed. It is about the size of a European Red Deer, and is an elegant animal, being possessed of a long, slender, gracefully curved neck and fine legs. Dr. Cunningham,[190] speaking from observation on wild animals, says:—

“It is not easy to describe its general appearance, which combines some of the characters of a camel, a deer, and a goat. The body, deep at the breast but very small at the loins, is covered with long, soft, very fine hair, which on the upper parts is of a kind of fawn-colour, and beneath varies from a very pale yellow to the most beautiful snow-white. The head is provided with large ears, in general carried well back, and is covered with short grayish hair, which is darkest on the forehead. Occasionally the face is nearly black. As a rule it lives in flocks of from half a dozen to several hundreds, but solitary individuals are now and then to be met with. They are very difficult to approach sufficiently near to admit of an easy shot, as they are extremely wary, but, on being disturbed, canter off at a pace which soon puts a safe distance between them and the sportsman, even though he should be mounted. Despite their timidity, however, they are possessed of great curiosity, and will sometimes advance within a comparatively short distance of an unknown object, at which they will gaze fixedly till they take alarm, when they effect a speedy retreat. Their cry is very peculiar, being something between the belling of a deer and the neigh of a horse. It would be difficult to overestimate their numbers upon the Patagonian plains; for in whatever direction we walked we always came upon numbers of portions of their skeletons and detached bones.”

Darwin, who has given an interesting account of the habits of the Guanaco in his _Naturalist’s Voyage_, says that they readily take to the water, and were seen several times at Port Valdes swimming from island to island.

The Llama is only known as a domestic animal, and is chiefly met with in the southern part of Peru. Burmeister, a very competent writer on the subject, says that he is perfectly satisfied that it is the descendant of the wild Guanaco, an opinion opposed to that of Tschudi. It generally attains a larger size than the Guanaco, and is usually white or spotted with brown or black, and sometimes altogether black. The earliest and often-quoted account of this animal by Agustin de Zarate, treasurer-general of Peru in 1544, will bear repeating as an excellent summary of the general character and uses to which it was put by the Peruvians at the time of the Spanish conquest. He speaks of the Llama as a sheep, observing, however, that it is camel-like in shape though destitute of a hump:—

“In places where there is no snow the natives want water, and to supply this they fill the skins of sheep with water and make other living sheep carry them; for, it must be remarked, these sheep of Peru are large enough to serve as beasts of burden. They can carry about one hundred pounds or more, and the Spaniards used to ride them, and they would go four or five leagues a day. When they are weary they lie down upon the ground; and as there are no means of making them get up, either by beating or assisting them, the load must of necessity be taken off. When there is a man on one of them, if the beast is tired and urged to go on, he turns his head round and discharges his saliva, which has an unpleasant odour, into the rider’s face. These animals are of great use and profit to their masters, for their wool is very good and fine, particularly that of the species called Pacas, which have very long fleeces; and the expense of their food is trifling, as a handful of maize suffices them, and they can go four or five days without water. Their flesh is as good as that of the fat sheep of Castile. There are now public shambles for the sale of their flesh in all parts of Peru, which was not the case when the Spaniards came first; for when one Indian had killed a sheep his neighbours came and took what they wanted, and then another Indian killed a sheep in his turn.”

The disagreeable habit here noticed of spitting in the face of persons whose presence is obnoxious is common to all the group, as may be daily witnessed in specimens in confinement in the menageries of Europe. One of the principal labours to which the Llamas were subjected at the time of the Spanish conquest was that of bringing down ore from the mines in the mountains. Gregory de Bolivar estimated that in his day as many as three hundred thousand were employed in the transport of the produce of the mines of Potosi alone; but since the introduction of horses, mules, and donkeys the importance of the Llama as a beast of burden has greatly diminished.

The Alpaca, though believed by many naturalists to be a variety of the Vicugna, is more probably, like the Llama, derived from the Guanaco, having the naked callosities on the hind limbs, and the relatively large skull of the latter. It is usually found in a domesticated or semi-domesticated state, being kept in large flocks which graze on the level heights of the Andes of southern Peru and northern Bolivia at an elevation of from 14,000 to 16,000 feet above the sea-level, throughout the year. It is smaller than the Llama, and, unlike that animal, is not used as a beast of burden, but is valued only for its wool, of which the Indian blankets and ponchas are made. Its colour is usually dark brown or black.

Mention has already been made of the occurrence of fossil Llamas in America, but some diversity of view obtains as to the generic position of some of these forms, owing to variations in their dental formula. Remains apparently referable to the existing species occur in the cavern-deposits of Brazil. In the Pleistocene of Mexico we meet with _A. (Palauchenia) magna_, which attained the size of a Camel, and had always two, and occasionally three, lower premolars; while in one South American Pleistocene species, which has been generically separated as _Hemiauchenia_, there were invariably three premolars in each jaw. In _A. (Holomeniscus) hesterna_, from the Pleistocene of North America, which was equal in size to _A. magna_, the premolars were reduced to one in each jaw; and the same condition obtains in _A. (Eschatius) vitakeriana_, where, however, the upper one is of simpler structure.

_Extinct Cameloids._—Until within the last few years the existence of two genera having so very much in common as the Camels and the Llamas, and yet so completely isolated geographically, had not received any satisfactory explanation; for the old idea that they in some way “represented” each other in the two hemispheres of the world was a mere fancy without philosophical basis. The discoveries made mostly within the past twenty years of a vast and previously unsuspected extinct fauna in the American continent of the Tertiary period, as interpreted by Leidy, Cope, Marsh, and others, has thrown a flood of light upon the early history of this family, and upon its relations to other mammals.

There have been found in these regions many Camel-like animals exhibiting different generic modifications; and, what is more interesting, a gradual series of changes, coinciding with the antiquity of the deposits in which they are found, have been traced from the thoroughly differentiated species of the modern epoch down through the Pliocene to the early Miocene beds, where, their characters having become by degrees more generalised, they have lost all that specially distinguishes them as _Camelidæ_, and are merged into forms common to the ancestral type of all the other sections of the Artiodactyles. Hitherto none of these annectant forms have been found in any of the fossiliferous strata of the Old World; and it may therefore be fairly surmised (according to the evidence at present before us) that America was the original home of the Tylopoda, and that the true Camels have passed over into the Old World, probably by way of the north of Asia, where we have every reason to believe there was formerly a free communication between the continents, and then, gradually driven southward, perhaps by changes of climate, having become isolated, have undergone some further special modifications; while those members of the family that remained in their original birthplace have become, through causes not clearly understood, restricted solely to the southern or most distant part of the continent. The occurrence in the dentition of the fossil Siwalik Camels of a feature now found only in _Auchenia_ is especially interesting from this point of view.

Briefly referring to some of these fossil types, we may note that _Pliauchenia_, of the Loup Fork beds (Lower Pliocene) of the United States, has three lower premolars, while in _Procamelus_ there were four of these teeth. In _Protolabis_ of the Miocene we have a more generalised form, in which the dental formula is _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ³⁄₃; and from this type a transition may be traced to _Poëbrotherium_, which, while having the same dental formula, was no larger than a Fox, and had the third and fourth metacarpals separate, with rudiments of the fourth and fifth. The earliest undoubted representative of the group is _Leptotragulus_, of the Uinta Eocene, which appears to have been closely allied to _Poëbrotherium_. It is, however, probable that the first lower premolar was wanting; while the other premolars of the mandible were much shorter antero-posteriorly than in the last-named genus. The manus, moreover, appears to have been less reduced, the second metacarpal retaining its connection with the magnum. It is suggested that _Leptotragulus_ may have been derived from the Bunodont genus _Homacodon_ of the Bridger Eocene, mentioned among the _Cænotheriidæ_.

TRAGULINA.

_Family_ TRAGULIDÆ.

No teeth in premaxillæ. Upper canines well developed, especially in the males; narrow and pointed. Lower canines incisiform. No caniniform premolars in either jaw, all the premolars except the last in the upper jaw being secant. Molariform teeth in a continuous series, consisting of _p_ ³⁄₃, _m_ ³⁄₃. Odontoid process of axis vertebra conical. Fibula complete. Four complete toes on each foot. The middle metapodials generally confluent, the outer ones (second and fifth) very slender but complete, _i.e._ extending from the carpus or tarsus to the digit. Navicular, cuboid, and ectocuneiform bones of tarsus united. Tympanic bullæ of skull filled with cancellar tissue. No frontal appendages. Ruminating, but the stomach with only three distinct compartments, the manyplies or third cavity of the stomach of the Pecora being rudimentary. Placenta diffused.

This section is represented only by the single family _Tragulidæ_, containing a few animals of small size, commonly known as Chevrotains, intermediate in their structure between the Deer, the Camels, and the Pigs. The large size of the canines of the male and the absence of horns caused them to be associated formerly with _Moschus_, one of the _Cervidæ_; hence they are often spoken of as “Pigmy Musk-Deer,” although they have no musk-secreting gland, or, except in the above-named trivial external characters, no special affinities with the true Musk-Deer. There has scarcely been a more troublesome and obdurate error in zoology than in this association of animals so really distinct. It has been troublesome, not only in preventing a just conception of the relations of existing Artiodactyles, but also in causing great confusion and hindrance in palæontological researches among allied forms; and most obdurate, inasmuch as all that has been recently done in advancing our knowledge of both groups has not succeeded in eradicating it, not only from nearly every one of our zoological text-books, whether British or Continental, but even from works of the highest scientific pretensions.

The family is now generally divided into two genera.

_Tragulus_,[191] containing the smallest of the existing Ungulates, animals having more of the general aspects and habits of some Rodents, as the Agoutis, than of the rest of their own order. The best-known species are _T. javanicus_, _T. napu_, _T. stanleyanus_, and _T. memmina_. The first three are from the Malay Peninsula, or the islands of the Indo-Malayan Archipelago, the last from Ceylon and India. A fossil species occurs in the Pliocene of the latter country.

_Dorcatherium_[192] is distinguished chiefly by the feet being stouter and shorter, the outer toes better developed, and the two middle metacarpals not ankylosed together. Its dental formula (as that of _Tragulus_) is usually _i_ ⁰⁄₃, _c_ ¹⁄₁, _p_ ³⁄₃, _m_ ³⁄₃ = 34. Vertebræ: C 7, D 13, L 6, S 5, C 12-13. The only existing species, _D. aquaticum_ (Fig. 118), from the west coast of Africa, is rather larger than any of the Asiatic Chevrotains, which it otherwise much resembles, but it is said to frequent the banks of streams, and have much the habits of Pigs. It is of a rich brown colour, with back and sides spotted and striped with white. It is evidently the survivor of a very ancient form, as remains of the type species (_D. naui_), only differing in size, occur in the lower Pliocene and Miocene of Europe; fossil species are also found in the Indian Pliocene. In _D. naui_ there are, at least frequently, four lower premolars, while the existing species has but three of these teeth.

_Extinct Traguloids._—A number of small selenodont Artiodactyles from various Miocene and Pliocene deposits appear to connect the modern Tragulina so closely with _Gelocus_ (p. 294), and thus with the ancestral _Cervidæ_, that their classification is almost an impossibility. Thus _Leptomeryx_, from the Miocene of the United States, is regarded as a Traguloid, having four premolars in each jaw and with the metatarsals fused into a cannon-bone. _Prodremotherium_, of the Upper Eocene Phosphorites of France, differs in that the metacarpals also form a cannon-bone; while in the American _Hypertragulus_, both metacarpals and metatarsals remain separate. _Bachitherium_, of the French Phosphorites, apparently presents affinity with _Gelocus_, _Prodremotherium_, and _Dorcatherium_. In this genus the first of the four lower premolars assumes the character and function of a canine, the true canine being incisor-like, and there are traces of minute upper incisors.

PECORA, OR COTYLOPHORA.

No premaxillary teeth or caniniform premolars. Upper canines generally absent, though sometimes largely developed. Inferior incisors, three on each side with an incisiform canine in contact with them. Molariform teeth consisting of _p_ ³⁄₃, _m_ ³⁄₃, in continuous series. Auditory bullæ simple and hollow within. Odontoid process in the form of a crescent, hollow above. Distal extremity of the fibula represented by a distinct malleolar bone of peculiar shape, articulating with the outer surface of the lower end of the tibia. Third and fourth metacarpals and metatarsals confluent. Outer or lateral toes small and rudimentary, or in some cases entirely suppressed; their metapodial bones never complete in existing forms. Navicular and cuboid bones of tarsus united. Horns or antlers usually present, at least in the male sex. Left brachial artery arising from a common innominate trunk, instead of coming off separately from the aortic arch as in the preceding sections. Stomach with four complete cavities. Placenta cotyledonous.[193]

The Pecora or true Ruminants form at the present time an extremely homogeneous group, one of the best-defined and most closely united of any of the Mammalia. But, though the original or common type has never been departed from in essentials, variation has been very active among them within certain limits; and the great difficulty which all zoologists have felt in subdividing them into natural minor groups arises from the fact that the changes in different organs (feet, skull, frontal appendages, teeth, cutaneous glands, etc.) have proceeded with such apparent irregularity and absence of correlation that the different modifications of these parts are most variously combined in different members of the group. It appears, however, extremely probable that they soon branched into two main types, represented in the present day by the _Cervidæ_ and the _Bovidæ_,—otherwise the antlered and horned Ruminants. Intermediate smaller branches produced the existing Musk-Deer and Giraffe, as well as the extinct _Helladotherium_ inclining to the first-named group, and the extinct _Sivatherium_, _Brahmatherium_, _Hydaspitherium_, and others more allied to the latter, although upon the true relationship of these forms there is a difference of opinion.

The earliest forms of true Pecora, as _Palæomeryx_, generally had no frontal appendages, and some few forms continue to the present day in a similar case. In the very large majority, however, either in both sexes or in the male only, a pair or occasionally two pairs (_Tetraceros_ and the extinct _Sivatherium_) of processes are developed from the frontal bones as weapons of offence and defence, these being almost always formed on one or other of two types.

1. “Antlers” are outgrowths of true bone, covered during their growth with vascular, sensitive integument coated with short hair. When the growth of the antler is complete, the supply of blood to it ceases, the skin dies and peels off, leaving the bone bare and insensible, and after a time, by a process of absorption near the base, it becomes detached from the skull and is “shed” (Fig. 119). A more or less elongated portion or “pedicle” always remains on the skull from the summit of which a new antler is developed. In the greater number of existing species of Deer this process is repeated with great regularity at the same period of each year. The antler may be simple, straight, subcylindrical, tapering and pointed, but more often it sends off one or more branches called “tines” or “snags” (Fig. 119). In this case the main stem is termed the “beam.” Commonly all the branches of the antler are cylindrical and gradually tapering. Sometimes they are more or less expanded and flattened, the antler being then said to be “palmated.” In young animals the antlers are always small and simple, and in those species in which they are variously branched or palmated, this condition is only gradually acquired in several successive annual growths. An interesting parallel has been observed here, as in so many other cases, between the development of the race and that of the individual. Thus the earliest known forms of Deer, those of the Lower Miocene, generally have no antlers, as in the young of the existing species. The Deer of the Middle Miocene have simple antlers, with not more than two branches, as in existing Deer of the second year; but it is not until the Pliocene and Pleistocene times that Deer occur with antlers developed with that luxuriance of growth and beauty of form characteristic of some of the existing species in a perfectly adult state. Among recent _Cervidæ_, antlers are wanting in the genera _Moschus_ and _Hydropotes_; they are present in both sexes in _Tarandus_ (the Reindeer), and in the male sex only in all others.

In those forms with the most complex antlers (Figs. 119, 120) the tine immediately over the forehead is termed the _brow tine_, the next one the _bez tine_, and the third one the _tres tine_; the mass of points at the summit of the antler being termed either the _royal_ and _surroyal tines_, or collectively the _crown_. The nodulated bony ring at the base of the antler, just above the point at which it separates from the pedicle when it is shed, is termed the _burr_.

2. The horns of the _Bovidæ_ consist of permanent, conical, usually curved bony processes, into which air-cells continued from the frontal sinuses often extend, called “horn-cores,” ensheathed in a case of true horn, an epidermic development of fibrous structure, which grows continuously, though slowly, from the base, and wears away at the apex, but is very rarely shed entire. The only existing species in which the latter process occurs regularly and periodically is the American Prong-Buck (_Antilocapra_), in which the horns also differ from those of all others in being bifurcated. Horns are not present at birth, but begin to grow very soon afterwards. The males of all existing _Bovidæ_ possess them, and they are also present (though usually not so fully developed) in the females of all except the genera _Boselaphus_, _Strepsiceros_, _Tragelaphus_, _Antilope_, _Æpyceros_, _Saiga_, _Cobus_, _Cervicapra_, _Pelea_, _Nanotragus_, _Neotragus_, _Cephalophus_, and _Tetraceros_; as well as in some species of _Gazella_, such as _G. picticandata_ and _G. walleri_.

Another character by which different members of the _Pecora_ can be distinguished among themselves is derived from the nature of the molar teeth. Although there is nothing in the general mode and arrangement of the enamel-folds, or in the accessory columns, absolutely distinctive between the two principal families, existing species may generally be distinguished, inasmuch as the true molars of the _Cervidæ_ are more or less brachydont, and those of the _Bovidæ_ generally hypsodont, _i.e._, the teeth of the former have comparatively short crowns (Fig. 122), which, as in most mammals, take their place at once with the neck (or point where the crown and root join) on a level with or a little above the alveolar border, and remain in this position throughout the animal’s life; whereas in the other forms (Fig. 123), the crown being lengthened and the root small, the neck does not come up to the alveolar level until a considerable part of the surface has worn away, and the crown of the tooth thus appears for the greater part of the animal’s life partially buried in the socket. In this form of tooth (which is almost always most developed in the posterior molars of the permanent series) the constituent columns of the crown are necessarily nearly parallel, whereas in the first-described they diverge from the neck towards the free or grinding surface of the tooth. In the completely hypsodont form the interstices of the lengthened columnar folds of enamel and dentine are filled up with cement, which gives stability to the whole organ, and is entirely or nearly wanting in the short-crowned teeth. The same modification from low to high crowns without essential alteration of pattern is seen in an even still more marked manner in some of the Perissodactyle Ungulates, the tooth of the Horse bearing to that of _Anchitherium_ the same relation as that of an Ox does to the early selenodont Artiodactyles. A parallel modification has also taken place in the molar teeth of the Proboscidea.

As the hypsodont tooth is essentially a modification of, and, as it were, an improvement upon, the brachydont, it is but natural to expect that all intermediate forms may be met with. Even among the Deer themselves, as pointed out by Lartet, the most ancient have very short molars, and the depressions on the grinding surface are so shallow that the bottom is always visible; while in the _Cervidæ_ of the more recent Tertiary periods, and especially the Pleistocene and living species, these same cavities are so deep that whatever be the state of the dentition the bottom cannot be seen. Some existing Deer, as the Axis, are far more hypsodont than the majority of the family; and, on the other hand, many of the Antelopes (as _Tragelaphus_) retain much of the brachydont character, which is, however, completely lost in the more modern and highly specialised Sheep and Oxen.

The complicated stomach of the Pecora (Fig. 124), which is necessary for the performance of the peculiar function known as “chewing the cud”—a function common also to the Tragulina and Tylopoda—is divided into four well-defined compartments, known as (1) the Rumen or Paunch, (2) the Reticulum or Honey-comb Bag, (3) the Psalterium or Manyplies, (4) the Abomasum or Reed. The paunch is a very capacious receptacle, shaped like a blunted cone bent partly upon itself. Into its broader base opens the œsophagus or gullet at a spot not far removed from its wide orifice of communication with the second stomach or honey-comb bag. Its inner walls are nearly uniformly covered with a pale mucous membrane, which is beset with innumerable close-set, short, and slender villi, resembling very much the “pile” on velvet. The honey-comb bag is very much smaller than the paunch. It is nearly globose in shape, and receives its name on account of the peculiar arrangement of its mucous membrane which forms shallow hexagonal cells all over its inner surface. Running along its upper wall there is a deep groove, coursing from the first to the third stomach. This groove plays an important part in the act of rumination. Its walls are muscular, like those of the viscus with which it is associated, which allows its calibre to be altered. Sometimes it completely closes round so as to become converted into a tube by the opposition of its edges. At others it forms an open canal. The manyplies is globular in form, and its lining membrane is raised into longitudinal folds or laminæ arranged very much like the leaves of a book, and very close together. Their surfaces are roughened by the presence of small projections or papillæ. The reed is the proper digestive stomach, corresponding with the same organ in man. Its shape is somewhat pyriform, and its walls are formed of a smooth mucous membrane, which secretes the gastric juice.

When the food is first swallowed it is conveyed into the paunch, and after undergoing a softening process there it is regurgitated into the mouth, and undergoes a further trituration by the molar teeth and mixture with the secretion of the salivary and buccal glands. It is then swallowed again, but now passes directly through the before-mentioned groove into the manyplies, and, after filtering through the numerous folds of the lining membrane of this cavity, finally reaches the fourth or digestive stomach.

The placenta of the Pecora is characterised by the fœtal villi being collected into groups or cotyledons, which may present either a convex or a concave surface to the uterus. These cotyledons are received into permanent elevations in the mucous membrane of the uterus, the surfaces of which present a curvature which is the reverse of the cotyledons.

_Family_ CERVIDÆ.

Frontal appendages, when present, in the form of antlers. First molar, at least, in both jaws brachydont. Two orifices to the lachrymal duct, situated on or inside the rim of the orbit. An antorbital or lachrymal vacuity of such dimensions as to exclude the lachrymal bone from articulation with the nasal. Upper canines usually present in both sexes and sometimes attaining a very great size in the male (see Fig. 134). Lateral digits of both fore and hind feet, almost always present, and frequently the distal ends of the metapodials. Placenta with few cotyledons. Gall-bladder absent (except in _Moschus_). This family contains numerous species, having a wide geographical distribution, ranging in the New World from the Arctic Circle as far south as Chili, and in the Old World throughout the whole of Europe and Asia, though absent in the Ethiopian and Australian regions.

It may be divided into two subfamilies.

Subfamily =Moschinæ=.—This subfamily is represented solely by the Musk-Deer, which differs so remarkably from the true Deer that it is considered by several writers as the representative of a separate family. The late Professor Garrod even suggested that it should be regarded as an extremely aberrant member of the _Bovidæ_.

_Moschus._[194]—The Musk-Deer (Fig. 125) in many respects stands by itself as an isolated zoological form, retaining characters belonging to the older and more generalised types of ruminants before they were distinctly separated into the horned and the antlered sections now dominant upon the earth. One of these characters is that both sexes are entirely devoid of any sort of frontal appendage. In this, however, it agrees with one existing genus of true Deer (_Hydropotes_); and, as in that animal, the upper canine teeth of the males are remarkably developed, long, slender, sharp pointed, and gently curved, projecting downwards out of the mouth with the ends turned somewhat backwards. Vertebræ: C 7, D 14, L 5, S 5, C 6. Among the anatomical peculiarities in which it differs from all true Deer and agrees with the _Bovidæ_ is the presence of a gall-bladder. The hemispheres of the brain are but slightly convoluted, and the cotyledons of the placenta are arranged in a peculiar linear manner.[195]

Although, owing to variations of colour presented by different individuals in different localities and seasons, several nominal species have been described, zoologists are now generally agreed that there is but one, the _Moschus moschiferus_ of Linnæus. In size it is rather less than the European Roe Deer, being about 20 inches high at the shoulder. Its limbs, especially the hinder ones, are long. The feet are remarkable for the great development of the lateral pair of hoofs, and for the freedom of motion they all present, so that they appear to have the power of grasping projecting rocky points,—a power which must be of great assistance to the animal in steadying it in its agile bounds among the crags of its native haunts. The ears are large, and the tail quite rudimentary. The hair covering the body is long, coarse, and of a peculiarly brittle and pith-like character, breaking with the application of an extremely slight force; it is generally of a grayish-brown colour, sometimes inclined to yellowish-red, and often variegated with lighter patches. The Musk-Deer has a wide distribution over the highlands of central and eastern Asia, including the greater part of southern Siberia, and extends to Kashmir on the south-west and Cochin-China on the south-east, always, however, at considerable elevations,—being rarely found in summer below 7000 feet above the sea-level, and ranging as high as the limits of the thickets of birch or pines, among which it mostly conceals itself in the daytime. It is a hardy, solitary, and retiring animal, chiefly nocturnal in its habits, and almost always found alone, rarely in pairs, and never in herds. It is exceedingly active and sure-footed, having few equals in traversing rocky and precipitous ground; and it feeds on moss, grass, and leaves of the plants which grow on the mountains among which it makes its home.

Most of the animals of the group to which the Musk-Deer belongs, in fact the large majority of mammals, have some portion of the cutaneous surface peculiarly modified and provided with glands secreting some odorous and oleaginous substance specially characteristic of the species. This, correlated with the extraordinary development of the olfactory organs, appears to offer the principal means by which animals in a state of nature become aware of the presence of other individuals of their own species, or of those inimical to them, even at very great distances, and hence it is of extreme importance both to the well-being of the individual and to the continuance of the race. The situation of this specially modified portion of skin is extremely various, sometimes between the toes, as in Sheep, sometimes on the face in front of the eyes, as in many Deer and Antelopes. Sometimes it is in the form of a simple depression or shallow recess, often very deeply involuted, and in its fullest state of development it forms a distinct pouch or sac with a narrow tubular orifice. In this sac a considerable quantity of the secretion can accumulate until discharged by the action of a compressor muscle which surrounds it. This is the form taken by the special gland of the Musk-Deer, which has made the animal so well known, and has proved the cause of an unremitting persecution to its possessor. It is found in the male only, and is a sac about the size of a very small orange, situated beneath the skin of the abdomen, the orifice being immediately in front of the preputial aperture. The secretion with which the sac is filled is of dark-brown or chocolate colour, and when fresh described as being of the consistence of “moist gingerbread,” but becoming dry and granular after keeping. It has a peculiar and very powerful scent, which when properly diluted and treated forms the basis of many of our most admired perfumes. When the animal is killed the whole gland or “pod” is cut out and dried, and in this form reaches the market of the Western World, chiefly through China.

Subfamily =Cervinæ.=—This subfamily includes all the true Deer. According to the arrangement proposed by Sir V. Brooke[196] the existing _Cervinæ_ may be divided into the sections Plesiometacarpalia and Telemetacarpalia.

=Plesiometacarpalia.=—In this section, which is mainly characteristic of the Old World, the proximal portions of the lateral (second and fifth) metacarpals persist, and the vomer is never so ossified as to divide the posterior osseous nares into two distinct passages. The premaxillæ nearly always articulate with the nasals.

_Cervulus._[197]—Antlers half the length of the head, placed on pedicles nearly equal to them in length. Brow tine short, inclined inwards and upwards; terminal extremity of beam unbranched, and curved downwards and inwards. Lachrymal fossa of skull very large, and extending into facial part of jugal; lachrymal (antorbital) vacuity moderate. Ascending portion of premaxillæ at least as long as nasals. A permanent ridge extending from each pedicle over the orbit, lachrymal fossa and vacuity. Auditory bulla much inflated. Upper canines of males very large. Ectocuneiform united with naviculo-cuboid of tarsus. No traces of the phalanges of the lateral digits.

The native name Muntjac has been generally adopted in European languages for a small group of Deer indigenous to the southern and eastern parts of Asia and the adjacent islands, which are separated by very marked characters from all their allies. They are also called “Kijang” or “Kidjang,” and constitute the genus _Cervulus_ of Blainville and most zoologists;—_Styloceros_ of Hamilton-Smith, and _Prox_ of Ogilby. They are all of small size compared with the majority of Deer, and have long bodies and rather short limbs and neck. The antlers, which as in most Deer are present in the male only, are small and simple, and the main stem or beam, after giving off a very short brow tine, inclines backwards and upwards, is unbranched and pointed, and when fully developed curves inwards and somewhat downwards at the tip. These small antlers are supported upon pedicles or permanent processes of the frontal bones, longer than in any other Deer, and the front edges of which are continued downwards as strong ridges passing along the sides of the face above the orbits, and serving to protect the large supraorbital glands lying on their inner sides. The lachrymal fossa of the skull, in which is lodged the large suborbital gland or crumen, is of great depth and extent. The upper canine teeth of the males are strongly developed and sharp, curving downwards, backwards, and outwards, projecting visibly outside the mouth as tusks, and loosely implanted in their sockets. In the females they are very much smaller. The limbs exhibit several structural peculiarities not found in other Deer. The lateral digits of both fore and hind feet are very little developed, the hoofs alone being present and their bony supports (found in all other Deer) wanting. There is a tufted gland on the outer side of the metatarsus.

The Muntjacs are solitary animals, very rarely even two being seen together. They are fond of hilly ground covered with forests, in the dense thickets of which they pass most of their time, only coming to the skirts of the woods at morning and evening to graze. They carry the head and neck low and the hind-quarters high, their action in running being peculiar and not very elegant, somewhat resembling the pace of a sheep. Though with no power of sustained speed or extensive leap, they are remarkable for flexibility of body and facility of creeping through tangled underwood. They are often called by Indian sportsmen “Barking Deer,” a name given on account of their alarm cry, a kind of short shrill bark, like that of a fox but louder, which may often be heard in the jungles they frequent both by day and by night. When attacked by dogs the males use their sharp canine teeth with great vigour, inflicting upon their opponents deep and even dangerous wounds.

There is some difference of opinion among zoologists as to the number of species of the genus _Cervulus_. Sir Victor Brooke, who investigated this question in 1878 (see _Proceedings of the Zoological Society of London_ for that year, p. 898), came to the conclusion that there are certainly three which are quite well marked, viz.—

_C. muntjac_ (Fig. 126), found in British India, Burma, the Malay Peninsula, Sumatra, Java, Hainan, Banca, and Borneo. The general colour is a bright yellowish-red, darker in the upper parts of the back; the fore legs from the shoulder downwards and the lower part of the hind legs, dark bluish-brown; anterior parts of the face from the muzzle to between the eyes, brown—a blackish line running up the inside of each frontal ridge; chin, throat, inside of hind legs, and under surface of tail white. The female has a black bristly tuft of hair on the spot from which the pedicles of the antlers of the male grow. The average length of the male, according to Jerdon, is 3½ feet, tail 7 inches, height 26 to 28 inches. The female is a little smaller. The specimens from Java, Sumatra, and Borneo are of larger size than those from the mainland, and may possibly be of distinct species or race.

_C. lacrymans_ of Milne-Edwards, or Sclater’s Muntjac of Swinhoe, from Moupin, and near Hangchow, China.

_C. reevesi_, a very small species from southern China.

Subsequently the name _C. crinifrons_ has been applied to a Muntjac from Ningpo, China, readily distinguished from all other species by its bushy forehead and long tail. Another species from Tenasserim has been described as _C. feæ_.

Small Deer from the European Pliocene have been provisionally referred to _Cervulus_, but the so-called _Prox furcatus_, of the Miocene, is now included in _Palæomeryx_.

_Elaphodus._[198]—Antlers very small, unbranched, supported on long, slender, converging pedicles. Ascending rami of premaxillæ shorter than nasals. No supraorbital ridges or frontal glands. Upper canines of male long, but not everted. A distinct frontal tuft of hair. Other characters as in _Cervulus_.

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.

_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.

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.

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.