Chapter 24 of 28 · 43594 words · ~218 min read

CHAPTER XI

THE ORDER CARNIVORA

Though the existing Carnivora as at present restricted[422] form a very natural and well-defined order among the Mammalia, it is difficult to find any important common diagnostic characters by which they can be absolutely separated; so that, as in the case of so many other natural groups, it is by the possession of a combination of various characters that they must be distinguished. Thus they are all unguiculate, and never have less than four well-developed toes on each foot, with nails more or less pointed, rarely rudimentary or absent. The pollex and hallux are never opposable to the other digits. They are regularly diphyodont and heterodont, and their teeth are always rooted.[423] Their dentition consists of small pointed incisors, usually three in number, on either side of each jaw, of which the first is always the smallest and the third the largest, the difference being most marked in the upper jaw; strong conical, pointed, recurved canines; cheek-teeth variable, but generally, especially in the anterior part of the series, more or less compressed, pointed, and trenchant; if the crowns are flat and tuberculated they are never complex or divided into lobes by deep inflexions of enamel. The condyle of the lower jaw is a transversely placed half-cylinder working in a deep glenoid fossa of corresponding form. The brain varies much in relative size and form, but the hemispheres are never destitute of well-marked convolutions (Fig. 23, p. 71). The stomach (Fig. 234) is always simple and pyriform. The cæcum is either absent or short and simple (Fig. 235), and the colon is not sacculated, or greatly wider than the small intestine. Vesiculæ seminales are never present. Cowper’s glands are present in some, absent in other groups. The uterus is bicornuate. The mammæ are abdominal, and very variable in number. The placenta is deciduate, and almost always zonary. The clavicle is often entirely absent, and when present is never complete. The humerus often has an entepicondylar foramen. The radius and ulna are distinct. The scaphoid and lunar bones are united into one, and there is never a distinct os centrale in the adult. The fibula is always a distinct slender bone.

Several of these characters are, however, not applicable to all the members of the extinct group of Carnivores for which the name Creodonta has been proposed, as will be noticed in the sequel.

The large majority of the species composing this order subsist chiefly upon some variety of animal food, though many are omnivorous, and some few chiefly, though not entirely, vegetable eaters. The more typical forms live altogether on recently killed warm-blooded animals, and their whole organisation is thoroughly adapted to a predaceous mode of life. In conformity with this manner of obtaining their subsistence they are generally bold and savage in disposition, though some species are capable of being domesticated, and when placed under favourable circumstances for the development of such qualities exhibit a very high degree of intelligence and fidelity. The existing representatives of the order are naturally divided into two suborders, the members of the one being the more typical, and mainly terrestrial in their mode of life; while those of the other are aberrant, having the whole of their organisation specially modified for living habitually in water. These are called respectively the True, or Fissiped, and the Pinniped Carnivora.

_Suborder_ CARNIVORA VERA.

[Illustration: FIG. 220.—Left upper carnassial teeth of Carnivora. I, _Felis_; II, _Canis_; III, _Ursus_. 1, Anterior, 2, middle (paracone), and 3, posterior (metacone) cusp of blade; 4, inner tubercle (protocone) supported on distinct root; 5, inner cusp posterior in position, and without distinct root, characteristic of the _Ursidæ_.]

Generally adapted for terrestrial progression and mode of life, though some may be partially aquatic in their habits. The fore limbs never have the first digit, or the hind limbs the first and fifth digits, longer than the others. Incisors ³⁄₃ on each side, with very rare exceptions. Cerebral hemispheres more or less elongated; always with three or four gyri on the outer surface forming arches above each other, the lowest surrounding the Sylvian fissure. The molar series of teeth have not the uniform characters of those of the Pinnipedia. There is always one tooth in each jaw which is specially modified, and to which the name of “sectorial” or “carnassial” tooth has been applied. The teeth in front of this are more or less sharp pointed and compressed; while those behind it are broad and tuberculated. The characters of the carnassial teeth deserve special attention, as, though fundamentally the same throughout the suborder, they are greatly modified in different genera. The upper carnassial is the most posterior of the teeth which have predecessors, and is therefore reckoned as the last premolar (_p_ ⁴⁄ of the typical dentition). It consists essentially of a more or less compressed blade supported on two roots and an inner tubercle supported by a distinct root (see Fig. 220). The blade when fully developed has three cusps or lobes (1, 2, and 3), but the anterior is always small, and often absent. The middle lobe is conical, high, and pointed; the posterior lobe has a compressed straight knife-like edge. The inner tubercle (4) varies very much in extent, but is generally placed near the anterior end of the blade, though sometimes it is median in position. In the _Ursidæ_ alone both the inner tubercle and root are wanting, and there is often a small internal and posterior cusp (5) without root. In this aberrant family also the carnassial is relatively to the other teeth much smaller than in the rest of the Carnivora. The lower carnassial (see Fig. 221) is the most anterior of the teeth without predecessors in the milk-series; it is therefore reckoned the first true molar (_m_ ¹⁄). It has two roots supporting a crown, consisting when fully developed of a compressed bilobed blade (1 and 2), a heel, or talon (4), and an inner cusp (3). The lobes of the blade, of which the hinder (2) is the larger, are separated by a notch, generally prolonged into a linear fissure. In the most specialised Carnivora, as the _Felidæ_ (I), the blade alone is developed, both talon and inner cusp being absent or rudimentary. In others, as _Meles_ (V) and _Ursus_ (VI), the heel is greatly developed, broad, and tuberculated. The blade in these cases is generally placed obliquely, its flat or convex (outer) side looking forwards, so that the two lobes are almost side by side, instead of anterior and posterior. The inner cusp (3) is generally conical, pointed, and placed to the inner side of the hinder lobe of the blade. The special characters of these teeth are more disguised in the Sea Otter (_Latax_) than in any other form, but even in it they can be traced.

[Illustration: FIG. 221.—Left lower carnassial teeth of Carnivora. I, _Felis_; II, _Canis_; III, _Herpestes_; IV, _Lutra_; V, _Meles_; VI, _Ursus_. 1, Anterior lobe (paraconid) of blade; 2, posterior (protoconid) lobe of blade; 3, inner cusp (metaconid); 4, talon (hypoconid). It will be seen that the relative size of the two roots varies according to the development of the portion of the crown they have respectively to support.]

The homology of the various parts of the Carnivorous carnassial with the primitive tritubercular type (p. 30) is indicated in the figures. It may be observed, however, that the anterior lobe of the three-lobed upper carnassial is an element added on to the more primitive two-lobed type. When the talon of the lower carnassial, as in _Canis_, consists of a large outer and small inner cusp, the latter (not seen in the figure) is the entoconid.

The toes are nearly always armed with large, strong, curved, and tolerably sharp claws, ensheathing the ungual phalanges, and held more firmly in their places by broad laminæ of bone reflected over their attached ends from the bases of the phalanges. In some forms, most notably the _Felidæ_, these claws are retractile; that is to say, the ungual phalanx, with the claw attached, folds back in the fore foot into a sheath by the outer or ulnar side of the middle phalanx of the digit, being retained in this position when the animal is at rest by a strong elastic ligament. In the hind foot the ungual phalanx is retracted on to the top, and not the side of the middle phalanx. By the action of the deep flexor muscles, the ungual phalanges are straightened out, the claws protruded from their sheath, and the soft “velvety” paw becomes suddenly converted into a most formidable weapon of offence. The habitual retraction of the claws preserves their points from wear in ordinary progression.

The skeleton of the Lion represented in Fig. 15 (p. 45) illustrates the digitigrade mode of progression of the _Felidæ_, as well as the essential characters of the bony framework of a typical Carnivore.

The Fissipedal Carnivora were divided by Cuvier into two groups, according to the position of the feet in walking,—the Plantigrada, or those that place the whole of the soles to the ground, and the Digitigrada, or those that walk only on the toes; and the difference between these groups was considered of equal importance to that which separated the Pinnigrada or Seals from both of them. The distinction is, however, quite an artificial one, since every intermediate condition exists between the extreme typical plantigrade gait of the Bears and the truly digitigrade walk of the Cats and Dogs; in fact, the greater number of the Carnivora belong to neither one form nor the other, but may be called “subplantigrade”; often when at rest applying the whole of the sole to the ground, but keeping the heel raised to a greater or less extent when walking.

An amended classification of the existing forms is into three distinct sections, of which the Cats, the Dogs, and the Bears may be respectively taken as representatives, and which are hence called Æluroidea, Cynoidea, and Arctoidea. This division is founded mainly on characters exhibited by the base of the skull, but is corroborated by the structure of other parts.[424] The presence or absence of a bridge of bone, covering the external carotid artery in a part of its course by the side of the alisphenoid bone, and enclosing the “alisphenoid canal” (see Fig. 8, p. 38), a character to which the late Mr. H. N. Turner first drew attention, might seem unimportant at first sight, but it is curiously constant in certain groups, which we have other reasons, derived often from a combination of less easily definable characters, to regard as natural. It is therefore generally mentioned in the following family definitions.

It must, however, be stated that while the arrangement is a convenient one as regards the existing Carnivores, it will not hold good when the fossil forms are included. Thus there is ample evidence to show that the Dogs and Bears were formerly so intimately connected that in a palæontological classification the _Canidæ_ cannot be satisfactorily separated from the _Ursidæ_; while in another direction the _Canidæ_ were closely allied to the ancestral _Viverridæ_. The most important objection against this classification is, however, the apparent intimate connection exhibited by fossil forms between the _Viverridæ_ and the _Mustelidæ_, which, so far as the present evidence goes, tends to show that the latter are derived from the former. If this be eventually fully proved, it would seem to indicate that the Arctoidea are not a natural group; and that the resemblances between the _Ursidæ_ and _Mustelidæ_ have been independently acquired, in the course of the descent of the one family from a Canoid, and of the other from a Viverroid stock.

_Section_ ÆLUROIDEA.

[Illustration: FIG. 222.—Left side of the palatal aspect of the cranium and mandible of the Suricate (_Suricata tetradactyla_). _c_, Carotid foramen; _f_, fissure in floor of auditory meatus. From Mivart, _Proc. Zool. Soc._ 1882, p. 184.]

The Æluroidea or Cat-like Carnivores include the _Felidæ_, _Viverridæ_, _Proteleidæ_, and _Hyænidæ_. The existing representatives of this section present the following common features. Auditory bulla (Fig. 222) much dilated, rounded smooth, thin-walled, and (except in the _Hyænidæ_) divided into two chambers, by a septum. Bony auditory meatus short. Paroccipital process applied to, and spread over the hinder part of the bulla (Fig. 222). Mastoid process never very salient, and often obsolete. Carotid canal (Fig. 8, p. 38, _car_) small, sometimes very inconspicuous. Condyloid and glenoid foramina concealed or wanting. Cæcum small, rarely absent. Os penis generally small and irregular (large in _Cryptoprocta_). Cowper’s glands present; prostate distinctly lobed. Some details of the anatomy of the soft parts will be found under the head of _Genetta_.

_Family_ FELIDÆ.

In all the forms, both recent and fossil, which can be included in this family the canines are strongly developed, there are never more than one upper and two lower molars, and the three lower incisors are placed in the same horizontal line. With one exception, the humerus has an entepicondylar foramen.

The following characters are common to all the existing members. True molars reduced to one above and below, that of the upper jaw very small and transversely extended. Only two inferior premolars. Upper carnassial with three lobes to the blade; lower without talon or inner cusp. Auditory bulla not externally constricted. No alisphenoid canal. Carotid canal very minute. Digits 5-4. Dorsal vertebræ 13.

[Illustration: FIG. 223.—Front view of skull of Lion (_Felis leo_).]

_Felis._[425]—The whole structure of the animals of this genus exhibits the Carnivorous type in its fullest perfection. Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ³⁄₂, _m_ ¹⁄₁; total 30. A distinctly cusped inner tubercle to the upper carnassial. Claws completely retractile. The upper anterior premolar (_p._ 2), always small, and may be absent without any other modification in the dental or other structures. Such a variation should not therefore be considered as of generic importance. Incisors very small. Canines large, strong, slightly recurved, with trenchant edges and sharp points, and placed wide apart (Fig. 223). Premolars compressed and sharp pointed. The most posterior in the upper jaw (the carnassial), a very large tooth, consisting of a subcompressed blade, divided into three unequal lobes supported by two roots, with a very small inner tubercle placed near the front end of the tooth and supported by a distinct root (Fig. 220). The upper true molar a very small tubercular tooth placed more or less transversely at the inner side of the hinder end of the last. In the lower jaw the true molar (carnassial) reduced to the blade alone, which is very large, trenchant, and much compressed, divided into two subequal lobes. Occasionally it has a rudimentary talon, but never an inner cusp. The skull is generally short and rounded, though proportionally more elongated in the larger forms. The facial portion is especially short and broad, and the zygomatic arches are very wide and strong. The auditory bullæ are large, rounded, and smooth. Vertebræ: C 7, D 13, L 7, S 3, C 13-29. Clavicles better developed than in other Carnivora, but not articulating with either the scapulæ or sternum. Limbs digitigrade. Anterior feet with five toes, the third and fourth nearly equal and longest, the second slightly and the fifth considerably shorter; the pollex still shorter, not reaching as far as the metacarpo-phalangeal articulation of the second. Hind feet with only four toes. The third and fourth the longest, the second and fifth somewhat shorter and nearly equal; the hallux represented only by the rudimentary metatarsal bone. The claws all very large, strongly curved, compressed, very sharp, and exhibiting the retractile condition in the highest degree. The tail varies greatly in length, being in some a mere stump, in others nearly as long as the body. Ears of moderate size, more or less triangular and pointed. Eyes rather large. Iris very mobile, and with a pupillary aperture which contracts under the influence of light in some species to a narrow vertical slit, in others to an oval, and in some to a circular aperture. Tongue thickly covered with sharp-pointed, recurved horny papillæ. Cæcum small and simple.

As in structure so in habits, the Cats may be considered the most specialised of all the Carnivora. All the known members of the genus feed, in the natural state, almost exclusively on warm-blooded animals which they have themselves killed. One Indian species (_F. viverrina_) preys on fish and even (it is said) on freshwater molluscs. Unlike the Dogs, they never associate in packs, and rarely hunt their prey in open ground, but from some place of concealment wait until the unsuspecting victim comes within reach, or with noiseless and stealthy tread, crouching close to the ground for concealment, approach near enough to make the fatal spring. In this manner they frequently attack and kill animals considerably exceeding their own size. They are mostly nocturnal, and the greater number, especially the smaller species, more or less arboreal. None are aquatic, and all take to the water with reluctance, though some may habitually haunt the banks of rivers or pools, because they more easily obtain their prey in such situations.

The numerous species of the genus are very widely diffused over the greater part of the habitable world, though most abundant in the warm latitudes of both hemispheres. No species are, however, found in the Australian region, or in Madagascar. Although the Old-World and New-World Cats (except perhaps the Northern Lynx) are all specifically distinct, no common structural character has been pointed out by which the former can be separated from the latter. On the contrary, most of the minor groups into which the genus has been divided have representatives in both hemispheres.

Notwithstanding the considerable diversity in external appearance and size between different members of this extensive genus, the structural differences are but slight, and so variously combined in different species that the numerous attempts hitherto made to subdivide it are all unsatisfactory and artificial. The principal differences are to be found in the form of the cranium, especially of the nasal and adjoining bones, the completeness of the bony orbit posteriorly, the development of the first upper premolar and of the inner tubercle of the upper carnassial, the length of the tail, the form of the pupil, and the condition and coloration of the fur, especially the presence or absence of tufts or pencils of hair on the external ears. Writing in 1881 Professor Mivart[426] gave the number of existing species of _Felis_ as 48, but by Mr. Blanford’s reduction of the number of Indian species[427] the list may now be diminished to some 41. The following account is chiefly devoted to some of the more important and better known species.

A. _Old World Species._—The Lion (_F. leo_, Fig. 224) has been well known to man from the earliest historic times. Its geographical habitat made it familiar to all the races among whom human civilisation took its origin, and its strongly marked physical and moral characteristics have rendered it proverbial, perhaps to an exaggerated degree, and have in all ages afforded favourite types for poetry, art, and heraldry. The literature of the ancient Hebrews abounds in allusions to the Lion; and the almost incredible numbers that are stated to have been provided for exhibition and destruction in the Roman amphitheatres (as many as six hundred on a single occasion by Pompey, for example) show how abundant these animals must have been within accessible distance of the capital of the world.

The geographical range of the Lion was once far more extensive than at present, even within the historic period covering the whole of Africa, the south of Asia, including Syria, Arabia, Asia Minor, Persia, and the greater part of Northern and Central India, and also the south-eastern portion of Europe, as shown by the well-known story told by Herodotus of the attacks by Lions on the Camels which carried the baggage of the army of Xerxes on its march through the country of the Pæonians in Macedonia. The very circumstantial account of that historian shows that the animal in his time ranged through the country south of the Balkans, through Roumania to the west of the River Carasu, and through Thessaly as far south as the Gulf of Lepanto and the Isthmus of Corinth, having as its western boundary the River Potamo and the Pindus mountains. The whole of the evidence relating to the existence of Lions in Europe, and to their retreat from that continent shortly before the commencement of the Christian era, has been collected in the article on “_Felis spelæa_” in Boyd Dawkins and Sandford’s _British Pleistocene Mammalia_ (1868). Fossil remains attest a still wider range, as it is shown in the same work that there is absolutely no osteological or dental character by which the well-known Cave Lion (_F. spelæa_), so abundantly found in cave-deposits of the Pleistocene age in Western Europe, can be distinguished from the existing _F. leo_.

[Illustration: FIG. 224.—Lion and Lioness, after a drawing by Wolf in Elliot’s Monograph of the _Felidæ_.]

At the present day the Lion is found in localities suitable to its habits, and where not exterminated (as it probably was in Europe) by the encroachments of man, throughout Africa from Algeria to the Cape Colony, and in Mesopotamia, Persia, and some parts of the north-west of India. According to Blanford,[428] Lions are still very numerous in the reedy swamps bordering the Tigris and Euphrates, and also occur on the west flanks of the Zagros mountains and the oak-clad ranges near Shiraz, to which they are attracted by the immense herds of swine which feed on the acorns. The Lion nowhere exists in the table-land of Persia, nor is it found in Baluchistan. In India, where it is verging on extinction, it appears now to be confined to parts of Kattywar and Rajputana, though within the present century its range extended through the north-west part of India, from Bahawalpur and Sind to at least the Jumna (about Delhi), southward as far as Khandesh, and in Central India through the Saugor and Narbada territories, Bundelkund, and as far east as Palamau. It was extirpated in Harriana about 1824. One was killed at Rhyli, in the Dumaoh district, Saugor and Narbada territories, so late as in the cold season of 1847-48; and one was shot in 1810 near Kot-Deji, Sind.[429]

The great variations in external characters which different Lions present, especially in the colour and the amount of mane, has given rise to the idea that there are several species, or at all events distinct varieties peculiar to different localities. It was at one time supposed, on the authority of Captain Walter Smee,[430] that the Lion of Gujerat differed essentially from that of Africa in the absence of a mane, but subsequent evidence has not supported this view, which was probably founded upon young specimens having been mistaken for adults. Lions from that district as well as from Babylonia, which have lived in the gardens of the London Zoological Society, have had as fully developed manes as any other of the species. Mr. F. C. Selous[431] has shown that in South Africa the so-called Black-maned Lion and others with yellow scanty manes are found, not only in the same locality, but even among individuals of the same parentage.

The Lion belongs to a well-defined group, containing the largest members of the genus, and differing from the others in the well-marked character that the anterior cornu of the hyoid arch is but little ossified, so that this arch is connected with the cranium by a long ligament, instead of by a continuous chain of bones, and by the less important one that the pupil of the eye, when contracted, is a circular hole, instead of a vertical slit as in the cat. The Lion agrees with the Tiger and the Leopard in these respects, but differs from them in its uniform style of colouring, and from all the other _Felidæ_ in the arrangement of its hairy covering; thus the hair of the top of the head, chin, and neck, as far back as the shoulder, is not only very much longer, but also differently disposed from the hair elsewhere, being erect or directed forwards, and so constituting the characteristic ornament called the mane. There is also a tuft of elongated hairs at the end of the tail, one upon each elbow, and in most lions a copious fringe along the middle line of the under surface of the body, wanting, however, in some examples.[432] It must, however, be observed that these characters are peculiar to the adults of the male sex only, and that young lions show indications of the darker stripes and mottlings so characteristic of the greater number of the members of the genus.

The usual colour of the adult is yellowish-brown, but it may vary from a deep red or chestnut brown to an almost silver gray. The mane, as well as the long hair of the other parts of the body, sometimes scarcely differs from the general colour, but it is usually darker and not unfrequently nearly black. The mane begins to grow when the animal is about three years old, and is fully developed at five or six.

In size the Lion is only equalled or exceeded by the Tiger among the existing _Felidæ_; though both species present great variations, the largest specimens of the latter appear to surpass the largest Lions. A full-sized South African Lion, according to Selous, measures slightly less than 10 feet from nose to tip of tail, following the curves of the body. Harris gives 10 feet 6 inches, of which the tail occupies 3 feet. The Lioness is about a foot less. The tongue, like that of the other species of the genus, is long and flat, and remarkable for the development of the papillæ of the anterior part of the dorsal surface, which (except near the edge) are modified so as to resemble long, compressed, recurved, horny spines or claws; these, near the middle line, attaining the length of one-fifth of an inch. They give the part of the tongue on which they occur the appearance and feel of a coarse rasp, and serve the purpose of such an instrument in cleaning the flesh from the bones of the animals on which the Lions feed.

The habits of the Lion in a state of nature are fairly well known from the united observations of numerous travellers and sportsmen who have explored those districts of the African continent in which it is still common. It lives chiefly in sandy plains and rocky places interspersed with dense thorn-thickets, or frequents the low bushes and tall rank grass and reeds that grow along the sides of streams and near the springs where it lies in wait for the larger herbivorous animals on which it feeds. Although it is occasionally seen abroad during the day, especially in wild and desolate regions, where it is subject to but little molestation, the night is, as in the case of so many other predaceous animals, the period of its greatest activity. It is then that its characteristic roar is chiefly heard, as thus graphically described by Gordon Cumming:—

“One of the most striking things connected with the Lion is his voice, which is extremely grand and peculiarly striking. It consists at times of a low, deep moaning, repeated five or six times, ending in faintly audible sighs; at other times he startles the forest with loud, deep-toned, solemn roars, repeated in quick succession, each increasing in loudness to the third or fourth, when his voice dies away in five or six low muffled sounds very much resembling distant thunder. At times, and not unfrequently, a troop may be heard roaring in concert, one assuming the lead, and two, three, or four more regularly taking up their parts, like persons singing a catch. Like our Scottish stags at the rutting season, they roar loudest in cold frosty nights; but on no occasions are their voices to be heard in such perfection, or so intensely powerful, as when two or three troops of strange Lions approach a fountain to drink at the same time. When this occurs, every member of each troop sounds a bold roar of defiance at the opposite parties; and when one roars, all roar together, and each seems to vie with his comrades in the intensity and power of his voice. The power and grandeur of these nocturnal concerts are inconceivably striking and pleasing to the hunter’s ear.”

“The usual pace of a Lion,” C. J. Andersson[433] says, “is a walk, and, though apparently rather slow, yet, from the great length of his body, he is able to get over a good deal of ground in a short time. Occasionally he trots, when his speed is not inconsiderable. His gallop—or rather succession of bounds—is, for a short distance, very fast—nearly or quite equal to that of a horse. Indeed, unless the steed has somewhat the start when the beast charges, it will be puzzled to escape. Many instances are on record of horsemen who have incautiously approached too near to the Lion, prior to firing, who have been pulled down by him before they could get out of harm’s way. Happily, however, the beast soon tires of the exertion of galloping, and unless his first rush succeeds he, for the most part, soon halts and beats a retreat.” “The Lion, as with other members of the feline family,” the same writer tells us, “seldom attacks his prey openly, unless compelled by extreme hunger. For the most part he steals upon it in the manner of a cat, or ambushes himself near to the water or a pathway frequented by game. At such times he lies crouched upon his belly in a thicket until the animal approaches sufficiently near, when, with one prodigious bound, he pounces upon it. In most cases he is successful, but should his intended victim escape, as at times happens, from his having miscalculated the distance, he may make a second or even a third bound, which, however, usually prove fruitless, or he returns disconcerted to his hiding-place, there to wait for another opportunity.” His food consists of all the larger herbivorous animals of the country in which he resides—buffaloes, antelopes, zebras, giraffes, or even young elephants or rhinoceroses, though the adults of these latter he dare not attack. In cultivated districts the cattle, sheep, and even human inhabitants are never safe from his nocturnal ravages. He appears, however, as a general rule, only to kill when hungry or attacked, and not for the mere pleasure of killing, as with some other carnivorous animals. Moreover, he by no means limits himself to animals of his own killing, but, according to Selous, often prefers eating game that has been killed by man, even when not very fresh, to taking the trouble to catch an animal himself. All books of African travel and sport abound with stories, many of which are apparently well authenticated, of the lion’s prodigious strength, as, exemplified by his being able to drag off a whole ox in his mouth to a long distance, even leaping fences and dykes with it.

The Lion appears to be monogamous, a single male and female continuing attached to each other irrespectively of the pairing season. At all events the Lion remains with the Lioness while the cubs are young and helpless, and assists in providing her and them with food, and in educating them in the art of providing for themselves. The number of cubs at a birth is from two to four, usually three. They are said to remain with their parents till they are about three years old. The following account by an eye-witness gives a good idea of Lion family life[434]:—

“I once had the pleasure of, unobserved myself, watching a lion family feeding. I was encamped on the Black Umfolosi in Zululand, and towards evening, walking out, about half a mile from camp, I saw a herd of zebra galloping across me, and when they were nearly 200 yards off, I saw a yellow body flash towards the leader, and saw him fall beneath the lion’s weight. There was a tall tree about 60 yards from the place, and anxious to see what went on, I stalked up to it, while the lion was still too much occupied to look about him, and climbed up. He had by this time quite killed the beautifully striped animal, but instead of proceeding to eat it, he got up and roared vigorously, until there was an answer, and in a few minutes a lioness, accompanied by four whelps, came trotting up from the same direction as the zebra, which no doubt she had been to drive towards her husband. They formed a fine picture as they all stood round the carcase, the whelps tearing it and biting it, but unable to get through the tough skin. Then the lion lay down, and the lioness driving her offspring before her did the same four or five yards off, upon which he got up, and, commencing to eat, had soon finished a hind leg, retiring a few yards on one side as soon as he had done so. The lioness came up next and tore the carcase to shreds, bolting huge mouthfuls, but not objecting to the whelps eating us much as they could find. There was a good deal of snarling and quarrelling among these young lions, and occasionally a stand-up fight for a minute, but their mother did not take any notice of them, except to give them a smart blow with her paw if they got in her way.... There was now little left of the zebra but a few bones, which hundreds of vultures were circling round waiting to pick, while almost an equal number hopped awkwardly about on the ground within 50 or 60 yards of it, and the whole lion family walked quietly away, the lioness leading, and the lion, often turning his head to see that they were not followed, bringing up the rear.”

Though not strictly gregarious, Lions appear to be sociable towards their own species, and often are found in small troops, sometimes consisting of a pair of old Lions, with their nearly full-grown cubs, but occasionally of adults of the same sex; and there seems to be good evidence that several Lions will associate together for the purpose of hunting upon a preconcerted plan. As might be supposed, their natural ferocity and powerful armature are sometimes turned upon one another; combats, often mortal, occur among male Lions under the influence of jealousy; and Andersson relates an instance of a quarrel between a hungry Lion and Lioness over the carcase of an Antelope which they had just killed, and which did not seem sufficient for the appetite of both, ending in the Lion not only killing, but even devouring his mate. Old Lions, whose teeth have become injured with constant wear, often become “man-eaters,” finding their easiest means of obtaining a subsistence in lurking in the neighbourhood of villages, and dashing into the tents at night and carrying off one of the sleeping inmates. Lions differ from most of the smaller _Felidæ_ in never climbing trees; indeed, as mentioned before, they are rarely found in forests.

With regard to the character of the Lion, those who have had opportunities of observing it in its native haunts differ greatly. The exaggerated accounts of early writers as to its courage, nobility, and magnanimity have led to a reaction, which causes some modern authors to speak of it in language quite the reverse, and to accuse it of positive cowardice and all kinds of meanness. Livingstone goes so far as to say, “Nothing that I ever learned of the lion could lead me to attribute to it either the ferocious or noble character ascribed to it elsewhere,” and he adds that its roar is not distinguishable from that of the ostrich. Of course these different estimates depend to a great extent upon the particular standard of the writer, and also upon the circumstance that Lions, like other animals, undoubtedly show considerable individual differences in character, and behave differently under varying circumstances. They are certainly not so reckless as to be entirely devoid of the instinct of self-preservation, and if one, perhaps satiated with a good meal the night before, unexpectedly disturbed in the daytime, will occasionally retreat when confronted, even by an unarmed man, that is scarcely a reason for assigning cowardice as one of the characteristics of the species. The latest authority, Selous, while never denying the daring courage of the Lion when hungry or provoked, and vindicating the awe-inspiring character of the roar of several Lions in unison, when heard at close quarters, as the grandest sound in nature, says with regard to its outward aspect:—

“It has always appeared to me that the word ‘majestic’ is singularly inapplicable to the lion in its wild state, as when seen by daylight he always has a stealthy furtive look that entirely does away with the idea of majesty. To look majestic a lion should hold his head high. This he seldom does. When walking he holds it low, lower than the line of his back, and it is only when he first becomes aware of the presence of man that he sometimes raises his head and takes a look at the intruder, usually lowering it immediately, and trotting away with a growl. When at bay, standing with open mouth and glaring eyes, holding his head low between his shoulders, and keeping up a continuous low growling, twitching his tail the while from side to side, no animal can look more unpleasant than a lion; but there is then nothing majestic or noble in his appearance.”

Notwithstanding this evidently truthful description of the animal when seen under what may be called unfavourable circumstances, no one with an eye for beauty can contemplate the form of a fine specimen of a Lion, at all events in a state of repose, even though in the confinement of a menagerie, without being impressed with the feeling that it is a grand and noble-looking animal.

The Tiger (_F. tigris_) is so closely related to the Lion that it is chiefly by external characters that the two species are distinguished. There are, however, slight distinctions in the proportionate size of the lower teeth, the general form of the cranium, and the relative length of the nasal bones and ascending processes of the maxillaries by which the skull of the Lion and Tiger can be easily discriminated by the practised observer.

Although examples of both species present considerable variations in size, and reliance cannot always be placed upon alleged dimensions, especially when taken from skins stripped from the body, it seems well ascertained that the length of the largest-sized Bengal Tiger may exceed that of any Lion. According to Mr. W. T. Blanford,[435] adult males measure from 5½ to 6½ feet from the nose to the root of the tail; the tail itself measuring some 3 feet in length. Measured along the curves of the head and back to the tip of the tail, males usually give a length of from 9 to 10 feet, but some specimens reach to 12 feet. The female is somewhat smaller, and has a lighter and narrower head. The Tiger has no mane, but in old males the hair of the cheeks is rather long and spreading. The ground colour of the upper and outer parts of the head, body, limbs, and tail is a bright rufous fawn, and these parts are beautifully marked with transverse stripes of a dark, almost black colour. The markings vary much in different individuals, and even on the two sides of the same individual. The under parts of the body, the inside of the limbs, the cheeks, and a large spot over each eye are nearly white. The Tigers which inhabit hotter regions, as Bengal and the south Asiatic islands, have shorter and smoother hair, and are more richly coloured and distinctly striped than those of Northern China and Siberia, in which the fur is longer, softer, and lighter coloured.

[Illustration: FIG. 225.—The Tiger (_Felis tigris_).]

The Tiger is exclusively Asiatic, but has a very wide range in that continent, having been found in almost all suitable localities south of a line drawn from the river Euphrates, passing along the southern shores of the Caspian and Sea of Aral by Lake Baikal to the Sea of Okhotsk. Its most northern range is the territory of the Amur, its most southern the islands of Sumatra, Java, and Bali. Westward it reaches to Turkish Georgia and eastward to the island of Saghalin. It is absent, however, from the great elevated plateau of Central Asia, nor does it inhabit Ceylon, Borneo, or the other islands of the Indo-Malayan Archipelago, except those above mentioned. Its absence from Ceylon leads Mr. Blanford to conclude that the Tiger has only recently migrated into Southern India.

The principal food of the Tiger in India is cattle, deer, wild hog, and pea-fowl, and occasionally human beings. The regular “man-eater” is generally an old Tiger whose vigour is passed, and whose teeth are worn and defective; it takes up its abode in the neighbourhood of a village, the population of which it finds an easier prey than the larger or wilder animals named above. Though chiefly affecting grassy plains or swamps, it is also found in forests, and seems to be fond of haunting the neighbourhood of old ruins. As a rule, Tigers do not climb trees; but when pressed by fear, as during an inundation, they have been known to do so. They take to the water readily and are good swimmers. The Tigers of the Sundarbans (Ganges delta) continually swim from one island to the other to change their hunting-grounds for deer. The following extract on the habits of the Tiger is taken from Sir J. Fayrer’s _Royal Tiger of Bengal_ (1875):—

“The tigress gives birth to from two to five, even six cubs; but three is a frequent number. She is a most affectionate and attached mother, and generally guards and trains her young with the most watchful solicitude. They remain with her until nearly full grown, or about the second year, when they are able to kill for themselves and begin life on their own account. Whilst they remain with her she is peculiarly vicious and aggressive, defending them with the greatest courage and energy, and when robbed of them is terrible in her rage; but she has been known to desert them when pressed, and even to eat them when starved. As soon as they begin to require other food than her milk, she kills for them, teaching them to do so for themselves by practising on small animals, such as deer and young calves or pigs. At these times she is wanton and extravagant in her cruelty, killing apparently for the gratification of her ferocious and bloodthirsty nature, and perhaps to excite and instruct the young ones, and it is not until they are thoroughly capable of killing their own food that she separates from them. The young tigers are far more destructive than the old. They will kill three or four cows at a time, whilst the older and more experienced rarely kill more than one, and this at intervals of from three or four days to a week. For this purpose the tiger will leave its retreat in the dense jungle, proceed to the neighbourhood of a village or gowrie, where cattle feed, and during the night will steal on and strike down a bullock, drag it into a secluded place, and then remain near the ‘marrie,’ or ‘kill,’ for several days, until it has eaten it, when it will proceed in search of a further supply, and, having found good hunting ground in the vicinity of a village or gowrie, continue its ravages, destroying one or two cows or buffaloes a week. It is very fond of the ordinary domestic cattle, which in the plains of India are generally weak, half-starved, under-sized creatures. One of these is easily struck down and carried or dragged off. The smaller buffaloes are also easily disposed of; but the buffalo bulls, and especially the wild ones, are formidable antagonists, and have often been known to beat the Tiger off, and even to wound him seriously.”

In many districts of India the number of Tigers has been very considerably diminished of late years. In some other countries they appear, however, to be on the increase; thus according to one of the administration reports of Java laid before the Dutch Chambers, portions of that island are being depopulated through Tigers. In 1882 the population of a village in the south-west of the Bantam province was removed and transferred to an island off the coast in consequence of the trouble caused to the people by Tigers. These animals have now become an intolerable pest in parts of the same province. The total population is about 600,000, and, in 1887, sixty-one were killed by Tigers, and in consequence of the dread existing among the people, it has been proposed to deport the inhabitants of the villages most threatened to other parts of the country where Tigers are not so common, and where they can pursue their agricultural occupations with a greater degree of security. At present they fear to go anywhere near the borders of the forest. The people seem disinclined, or they lack the means and courage, to attack and destroy their enemy, although considerable rewards are offered by Government for the destruction of beasts of prey. In 1888 the reward for killing a Royal Tiger was raised to two hundred florins. It appears also that the immunity of the Tiger is in part due to superstition, for it is considered wrong to kill one unless he attacks first or otherwise does injury.

The Leopard (_F. pardus_, Fig. 226), although belonging to the same restricted group as the two preceding species, is distinguished from both by its inferior size, and its coloration. The animal now commonly known as the Leopard was called Pard (πάρδος and πάρδαλις) or Panther (πάνθηρ) by the ancients. Leopard (_leo-pardus_) is a later term, originally applied, it is believed, to the Cheeta or Hunting Leopard, upon the supposition that it was a creature intermediate between the Lion and the true Pard. If so it has been completely transferred to the more common species, and though in this sense a perfectly unnecessary and unmeaning term, has gradually superseded those by which this was originally known. Pard, so commonly used by Elizabethan authors, is now nearly obsolete in the English language, and Panther has either become synonymous with Leopard, or is used vaguely for any similar large feline animal, even the Puma of America.

[Illustration: FIG. 226.—The Leopard (_Felis pardus_).]

Owing to their extensive geographical range, and the great variations, both in size, form, and coloration to which Leopards are subject, zoologists have scarcely decided whether all the forms popularly referred to this animal should be regarded as specifically alike, or whether they should constitute several distinct species, but the prevailing opinion is in favour of the former view. The attempts to separate a larger and more robust variety, under the name of Panther, from a smaller and more graceful form, to which the term Leopard might properly be restricted, have failed, owing to the existence of intermediate conditions which cannot be assigned definitely to either one or the other form.[436] The most marked anatomical difference yet noted in different varieties of leopard is in the length of the tail as compared with that of the body, even the number of the caudal vertebræ showing variation, though within what limits, and whether correlated with other characters, has not yet been clearly ascertained. The fur of those specimens which inhabit the most northern confines of its range of distribution, as North China, is longer and softer, and the markings are consequently less distinct than on those from more congenial climates, and the well-marked variation thus produced has given rise to the idea of specific distinction.

The size of different individuals, as before said, varies greatly, the head and body usually measuring from 3½ to 4½ feet in length, and the tail from 2½ to 3 feet, but specimens have been met with which fall short of or exceed these limits. The ground colour of the fur varies from a pale fawn to a rufous buff, graduating into a pure white on the under parts and inside of the limbs. This is spotted over with dark brown or black; the spots on the back and sides being arranged in rosettes or broken rings, which vary greatly in size and distinctness in different individuals, but are without the central spot seen in those of the Jaguar. The spots on the under parts and limbs are simple and blacker than those on the other parts of the body. The bases of the ears behind are black, the tips buff. The upper side of the tail is buff, spotted with broken rings like the back, its under surface white with simple spots. The hair of the cubs is longer than that of the adults, its ground colour less bright, and its spots less distinct. Perfectly black Leopards, which, however, in certain lights show the characteristic markings on the fur, are not uncommon. These appear to be examples of melanism, occurring as individual variations, sometimes in one cub out of a litter of which the rest are normally coloured, and therefore not indicating a distinct race, much less a species. These are met with chiefly in Southern Asia. We are not aware of any recorded case from Africa, though there seems no reason why they should not occur.

In habits the Leopard resembles the other large Cat-like animals, yielding to none in the ferocity and bloodthirstiness of its disposition. It is exceedingly quick and active in its movements, but seizes its prey by waiting in ambush or stealthily approaching to within springing distance, when it suddenly rushes upon it and tears it to the ground with its powerful claws and teeth. It preys upon almost any animal it can overcome, such as antelopes, deer, sheep, goats, monkeys, peafowls, and is said to have a special liking for dogs. It not unfrequently attacks human beings in India, chiefly children and old women, but instances have been known of a Leopard becoming a regular “man-eater.” When favourable opportunities occur, it often kills many more victims than it can devour at once, apparently to gratify its propensity for killing, or only for the sake of their fresh blood. It generally inhabits woody districts, and can climb high trees with facility if necessary for its safety when hunted, but usually lives on or near the ground, among rocks, bushes, and roots and low branches of large trees.

The present geographical range of the Leopard is very extensive, as it is met with in various suitable localities, where not too much interfered with by human cultivation, throughout the greater part of Africa from Algeria to the Cape Colony, and through the whole of the South of Asia from Palestine to China, including all India south of the Himalaya, and the islands of Ceylon, Java, Sumatra, and Borneo. Fossil bones and teeth, indistinguishable from those of existing Leopards, have been found in cave-deposits of Pleistocene age in Spain, France, Germany, and England. The evidence of the former existence of the Leopard in England is described at length by Boyd Dawkins and Sanford in their _British Pleistocene Mammalia_.[437]

The Ounce, or Snow Leopard (_F. uncia_), inhabits the highlands of Central Asia, from the lofty mountains of Tibet to the southern parts of Siberia, at altitudes of from 9000 to 18,000 feet above the sea. It is about the size of the common Leopard, but lighter in colour, with longer fur, less distinct spots, and a long thick tail. Its skull differs in shape from that of all the other _Felidæ_; the facial portion being very broad, the nasal bones especially being wide and depressed, and the zygomatic arches very strong and deep. The Clouded Tiger (_F. nebulosa_[438]) is a beautifully marked species, with elongated head and body, long tail, and rather short limbs. The canine teeth are proportionally longer than in any existing member of the genus. It is thoroughly arboreal, and is found in the forests of South-East Asia and the islands of Sumatra, Java, Borneo, and Formosa. _F. serval_, the Serval, from South Africa, is yellow with black spots, and has a short tail and large ears. Numerous smaller species called Tiger Cats and Wild Cats, of which the Oriental _F. marmorata_ (Fig. 227) is a good example, are found throughout the warmer parts of Asia and Africa. The Wild Cat of Europe, _F. catus_, still inhabits the mountainous and wooded parts of Great Britain.

[Illustration: FIG. 227.—The Marbled Cat (_Felis marmorata_). From Blanford, _Mammalia of British India_, p. 74, after Elliot.]

The Caffre Cat (_F. caffra_[439]), of Africa and Southern Asia, was the species held in veneration by the ancient Egyptians, and immense numbers of its mummified remains have recently been found in Egypt, whence they have been imported in large quantities to this country for manure. This species is generally regarded as the main ancestral stock from which the European Domestic Cat has been derived; one of the arguments in support of this opinion being that the whole of the sole of the hind foot of _F. caffra_ is black, and that the same feature obtains in the darker varieties of the Domestic Cat; while in _F. catus_ there are only spots of black upon this portion of the limb. Remains of the Caffre Cat occur in the Pleistocene cave-deposits at Gibraltar. The Indian _F. rubiginosa_ is the smallest species of Cat.

The Caracal or Persian Lynx (_F. caracal_) is an animal about the size of a fox, of slender build, with a moderately long tail, reaching down to the heels. It is of a uniform vinous or bright fulvous brown colour above, and is paler, sometimes almost white, beneath. It is quite or almost entirely unspotted. The tail has a black tip, and the ears are black externally, long, upright, pointed, and surmounted by a pencil of fine black hairs. It inhabits Central and North-West India, Persia, Arabia, Syria, and the greater part of Africa.

The true Lynxes comprise various species or varieties found in the northern and temperate regions of both the Old and New World, all larger than the true Wild Cats, with long limbs, short stumpy tail, ears tufted at the tip, and pupil of the eye linear when contracted. Their fur is generally long and soft, varying, however, according to season and locality, and always longish upon the cheeks. Their colour is always light brown or gray, and generally more or less spotted with a darker shade. The naked pads of the feet are more or less covered by the hair that grows between them. The skull and skeleton do not differ markedly from those of the other cats, but the small anterior upper premolar tooth found in many other species is usually wanting; and the lower carnassial has a rudimental talon. Their habits are exactly those of the other Wild Cats, and they are exceeded by none in the untameable savageness of their disposition. They capture their prey in the same manner, either lying in wait, or noiselessly stealing within reach, and then making a sudden rush or spring upon it. Their food consists of any mammals or birds which they can overpower. In inhabited countries they commit extensive ravages upon sheep, lambs, and poultry. Lynxes generally frequent rocky places and forests, being active climbers, and passing much of their time among the branches of the trees. Their skins are of considerable commercial value.

Zoologists are by no means agreed at present as to the specific distinctions, if any really exist, between the various modifications of this group. As many as eight species are sometimes recognised, four belonging to the Old and four to the New World. The former are _F. lynx_, of Scandinavia, Russia, Northern Asia, and till lately the forest regions of Central Europe; though not an inhabitant of Britain during the historic period, its remains have been found in cave-deposits of Pleistocene age; _F. cervaria_, Siberia; _F. pardina_, Turkey, Greece, Sicily, Sardinia, and Spain; and _F. isabellina_, Tibet. The American varieties are _F. canadensis_, the most northern species, and _F. rufa_, the American Wild Cat or Bay Lynx, extensively distributed from the Atlantic to the Pacific throughout nearly the whole latitude of the United States, but replaced in Texas and southern California by _F. maculata_, and in northern Oregon and Washington territory by _F. fasciata_.

[Illustration: FIG. 228.—European Lynx (_Felis lynx_). From a drawing by Wolf in Elliot’s _Monograph of the Felidæ_.]

In both cases, as might be supposed, specimens obtained from the more southern climates are shorter in their fur, more brightly coloured, and more distinctly spotted than those from colder regions. When only a few individuals of each most markedly different form are examined the distinctions are sufficiently evident. The occurrence, however, of transitional or intermediate forms makes it extremely difficult to draw the line between the different varieties or species, or to assign definite characters by which they can be separated. Wherefore it is best at present to accept the so-called species as only provisional, and wait until more abundant materials, with fuller knowledge of the localities from which they are derived, and of the variations due to age, sex, season, and climate, have been more carefully studied. We shall then probably come to the conclusion that all or nearly all the existing forms of northern Lynxes, whether American or Eurasian, belong to what may fairly be called a species, which is becoming by degrees differentiated into several more or less strongly marked local varieties. Mr. W. T. Blanford has indeed shown that the Tibetan Lynx (_F. isabellina_) is inseparable from _F. lynx_; the specimens from Gilgit being intermediate in colour between the typical forms of the two races. On the other hand, from the evidence of cranial characters, Professor Mivart is disposed to regard _F. pardina_ as a valid species.

[Illustration: FIG. 229.—The Puma (_Felis concolor_).]

B. _New World Species._—The Puma or Couguar (_F. concolor_, Fig. 229), commonly called “Panther” in the United States, is about the size of a Leopard, but of an uniform brown colour. It usually measures from nose to root of tail about 40 inches, the tail being rather more than half that length. The head is rather small compared with that of other Cats and has no mane. The ears are large and rounded. The tail is cylindrical, with some bushy elongation of the hairs near the end, but not forming a distinct tuft as in the Lion. The general colour of all the upper parts and sides of the adult is a tawny yellowish-brown, sometimes having a gray or silvery shade, but in some individuals dark or inclining to red. The lower parts of the body, inner surface of the limbs, the throat, chin, and upper lip are dirty white; the outside of the ears, particularly at their base, and a patch on each side of the muzzle black; the end of the tail dusky. The young are, when born, spotted with dusky brown and the tail ringed; these markings gradually fading, and quite disappearing before the animal becomes full-grown.

The Puma has an exceedingly wide range of geographical distribution, extending over a hundred degrees of latitude, from Canada in the north to Patagonia in the south, and was formerly pretty generally diffused in suitable localities from the Atlantic to the Pacific Ocean, but the advances of civilisation have in recent years considerably curtailed the extent of the districts which it inhabits. In Central America it is still common in the dense forests which clothe the mountain ranges as high as 8000 or 9000 feet above the sea-level, where the hideous sound of its howling is said to be almost continuously heard at night during the breeding season. Though an expert climber, it is by no means confined to wooded districts, being frequently found in scrub and reeds along the banks of rivers, and even in the open pampas and prairies. Its habits much resemble those of the rest of the group to which it belongs; and, like the Leopard, when it happens to come within reach of an abundant and easy prey, as the sheep or calves of an outlying farming station, it kills far more than it can eat, either for the sake of the blood only or to gratify its propensity for destruction. It rarely attacks man, and, when pursued, escapes if possible by ascending lofty trees. Several instances have occurred of Pumas becoming tame in captivity. Edmund Kean, the celebrated actor, had one which followed him about like a dog. When caressed they express their pleasure by purring like a domestic cat.

_F. onca_, the Jaguar, is a larger and more powerful animal than the last, and more resembles the Leopard in its colours. It also is found in both North and South America, but with less extensive range, reaching northwards only as far as Texas, and southwards nearly to Patagonia. It climbs as well as the Puma, and preys to a great extent upon monkeys. Several allied smaller elegantly spotted forms inhabiting the intratropical regions of America are commonly included under the name of Ocelot or Tiger Cat, though zoologists are still undecided whether under this designation several distinct species have not been confused, or whether all the Ocelots are to be referred to a single species (_F. pardalis_) showing great individual or racial variation. Their fur has always a tawny yellow or reddish-gray ground colour, and is marked with black spots, aggregated in streaks and blotches, or in elongated rings enclosing an area which is rather darker than the general ground colour. They range through the wooded parts of tropical America, from Arkansas in the north as far south as Paraguay, and in their habits resemble the other smaller members of the Cat tribe, being ready climbers and exceedingly bloodthirsty.

_F. yaguarundi_, rather larger than the Domestic Cat, with an elongated head and body, and of a uniform brownish-gray colour, ranges from Matamoras to Paraguay. _F. eyra_ is a small Cat, very Musteline in form, having an elongated head, body, and tail, and short limbs, and is also of a uniform light reddish-brown colour. It is a native of South America and Mexico. _F. pajeros_ is the Pampas Cat. The American Lynxes have been already noticed with those of the Old World.

[Illustration: FIG. 230.—The Ocelot (_Felis pardalis_).]

C. _Fossil Species._—It has been already incidentally mentioned that several of the existing species of _Felis_, such as the Lion, Leopard and Caffre Cat, are met with in a fossil condition in the European Pleistocene deposits, and it may be added that the Pardine Lynx has left its remains in the cavern-deposits of Gibraltar. The caves of Brazil have yielded remains of the Jaguar and Ocelot; while the Puma is found in the Pleistocene of the United States. Existing species now inhabiting India are met with in cavern-deposits in Madras. In the Pliocene Siwaliks of Northern India the huge extinct _F. cristata_ shows characters connecting it both with the Tiger and the Jaguar; and the same deposit also contains the remains of a small species of the size of _F. bengalensis_. In Europe numerous species occur in the Upper and Lower Pliocene, some of which were as large as a Leopard. _F. atrox_ and _F. augusta_, of the Pliocene of the United States, were of the dimensions of the Lion.

_Cynælurus._[440]—The Cheeta or Hunting Leopard (_C. jubatus_) is distinguished from the other _Felidæ_ by the inner tubercle of the upper carnassial, though supported by a distinct root, having no salient cusp upon it; by the tubercular molar being more in a line with the other teeth; and by the claws being smaller, less curved, and less completely retractile, owing to the feebler development of the elastic ligaments. The skull is short and high, with the frontal region broad and elevated in consequence of the large development of the frontal air-sinuses. The head is small and round, the body light, the limbs and tail long. Its colour is pale yellowish-brown with small black spots. The Cheeta is less savage and more easily tamed than most of the Cats. In Asia it has been trained for the chase of the Antelope. It has rather an extensive geographical range from the Cape of Good Hope, throughout Africa and the south-western parts of Asia, as far as Southern India.

_Extinct Genera._—A number of forms are gradually becoming known, especially through the researches of American palæontologists, which, though evidently animals of the same general type, and therefore to be placed in or near the family _Felidæ_, depart so much in various details of structure that they must be referred to different genera. As one of the points in which _Felis_ manifests its specialisation is the reduction of the number of the molar series of teeth, with concomitant shortening of the jaws, it might be supposed that in the earlier and perhaps ancestral forms these teeth would be more numerous and approach more nearly to the primitive or typical number of the heterodont mammals, viz. seven on each side. This is actually the case. Similarly we find that many of these forms exhibit a less specialised structure of the teeth themselves, as is shown by the absence of the anterior lobe of the upper carnassial, and the retention of the hind talon in the corresponding lower tooth. Again, some of them have an alisphenoid canal in the skull; while the femur may have a third trochanter, and the claws be very imperfectly retractile.

An extremely generalised form is the small _Proælurus_, from the Upper Eocene and Lower Miocene, with _p_ ⁴⁄₄, _m_ ¹⁄₂, an alisphenoid canal, and a third trochanter to the femur. _Dinictis_, of the North American Miocene, is a larger allied form, with _p_ ³⁄₃, _m_ ¹⁄₂; the upper carnassial having no anterior lobe, and the ungual phalanges being devoid of bony sheaths. The characters of the base of the skull, and the form and relations of the astragalus, differ very considerably from _Felis_. _Pseudælurus_, from the French Miocene, is another very generalised Feline, in which there may be either three or four premolars, and the lower carnassial may retain its inner cusp. _Ælurictis_, of the French Phosphorites, with _p_ ³⁄₃₋₄, _m_ ¹⁄₁₋₂, together with several American Miocene genera, such as _Nimravus_ (_p_ ³⁄₂, _m_ ¹⁄₂), _Archælurus_ (_p_ ³⁄₃₋₄, _m_ ¹⁄₂), _Pogonodon_ (_p_ ³⁄₃, _m_ ¹⁄₁), and _Hoplophoneus_ (_p_ ²⁻³⁄₂, _m_ ¹⁄₁), approach more closely to the modern Cats, although many or all of them retain the alisphenoid canal, and have not yet developed the anterior lobe to the upper carnassial, or lost the talon to the lower one. _Hoplophoneus_ has a descending flange to the mandible; and its scapholunar bone has a line indicating its dual origin; while the femur still retains the third trochanter, of which all traces are lost in the modern Cats.

On the other hand, some of the extinct _Felidæ_ show a most remarkable tendency towards a specialisation not occurring in any of the surviving members of the family, viz. an enormous development of the upper canines, with which is usually associated an expansion downwards and flattening of the anterior part of the ramus of the lower jaw, on the outer side of which the canine lies, when the mouth is closed. In _Machærodus næogeus_, the Sabre-toothed Tiger, from the caves of Brazil and also from Pleistocene deposits near Buenos Ayres, an animal about the size of a Tiger, these teeth are 7 inches in length, greatly compressed, and finely serrated on the trenchant anterior edges. Similar serrations are seen on a much fainter scale in the unworn teeth of modern Tigers. Many modifications of this commonly-called “machærodont” type have been met with both in the Old and New World. In _M. cultridens_, of the Upper Pliocene of Italy and France, the upper canine is long and narrow, with smooth cutting edges; the smaller form described as _M. meganthereon_ being apparently the female of this species. _M. crenatidens_, of the same deposits, is distinguished by the shorter and broader upper canine, in which both edges are strongly serrated; the same feature occurring in the closely allied or identical _M. latidens_ of the English cavern-deposits. The Italian Pliocene form described as _M. nestianus_ has serrations only on the hinder edge of the upper canine, and the third lower premolar is separated by a long interval from the fourth. _M. necator_, of the Pleistocene of South America, is remarkable as being the only member of the family in which the humerus has no entepicondylar foramen. A very remarkable form, _Eusmilus_, from the Upper Eocene Phosphorites of Central France, differs from all other known Felines in having only two pairs of incisors in the lower jaw, and a small canine separated by a very long diastema from the cheek-teeth, which consist only of one premolar and one sectorial true molar. The lower jaw is enormously expanded towards the symphysis to protect the large upper canines. This animal then, although of Eocene age, appears to form the culminating development of the sabre-toothed or machærodont dentition, the most specially carnivorous type of structure known.

Other species of _Machærodus_ are found in the Pliocene deposits of Europe and Asia. The accompanying woodcut exhibits the last two upper teeth of the Indian _M. sivalensis_, from which it will be seen that the inner tubercle of the carnassial is much reduced in size, while the molar is very minute.

_Family_ VIVERRIDÆ.

[Illustration: FIG. 231.—Oral surface of the left upper carnassial and molar of _Machærodus sivalensis_.]

Premolars ³⁄₃ or ⁴⁄₄. Molars ¹⁄₁ or ²⁄₂. Upper carnassial usually without an anterior lobe, and the lower one with a well-developed talon; second lower incisor (as in all the following families) raised above the level of the first and third. Auditory bulla externally constricted, and divided by a septum. An alisphenoid canal (with very rare exceptions). Carotid canal distinct as a groove on the side of the bulla. Humerus usually with an entepicondylar foramen. Digits usually 5-5, but sometimes the pollex or hallux or both may be wanting. Dorsal vertebræ 13 or 14. Limited in distribution to the Old World.

The subfamily =Cryptoproctinæ= contains the single genus _Cryptoprocta_.[441] Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ¹⁄₁; total 36. The teeth generally closely resemble those of the _Felidæ_. The first premolar of both jaws is very minute and early deciduous. The upper carnassial has a very small inner tubercle, quite at the anterior part of the tooth. The true molar is very small and placed transversely. The lower carnassial has a large trenchant bilobed blade, and a very minute talon, but no inner cusp. Skull generally like that of _Felis_, but proportionately longer and narrower. Orbit widely open behind. Vertebræ: C 7, D 13, L 7, S 3, C 29. Body elongated. Limbs moderate in size. Feet subplantigrade; five well-developed toes on each, with sharp, compressed, retractile claws. Ears moderate. Tail long and cylindrical.

The only known species, _C. ferox_, the “Foussa” of the Malagasy, is peculiar to Madagascar, being the largest carnivorous animal in the island. It is about twice the size of the common Cat (5 feet from nose to end of tail), with short close fur of nearly uniform pale brown. Little is as yet known of its habits, except that it is nocturnal, frequently attacks and carries off goats, and especially kids, and shows great ferocity when wounded, on which account it is much dreaded by the natives.

The remaining numerous specific and generic modifications found in the existing animals belonging to this family seem to arrange themselves mainly into two tolerably distinct groups, distinguishable by the characters of the auditory bulla and neighbouring parts of the base of the skull, and by the structure of the feet. The one form has the genus _Viverra_ or Civet Cats for its most typical representative, and the other _Herpestes_ or the Ichneumons.

Subfamily =Viverrinæ=.—Auditory bulla oval, or rather conical, broad and truncated and not everted behind, narrow in front and more or less compressed at the sides. The outer or anterior chamber very small and flat. The meatus with scarcely any inferior lip, its orifice being close to the tympanic ring. Paroccipital process triangular, its apex projecting slightly beyond the bulla. Claws strongly curved and more or less retractile. Perineal scent-glands generally present.

This subfamily includes both Ethiopian and Oriental forms, but the former are the more numerous.

The typical section, which includes five genera, has the following characters. Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₂ (¹⁄₂ in _Prionodon_); total 40. Skull elongated; facial portion small and compressed. Orbits well-defined but incomplete behind. Teeth always sectorial, never very small. Vertebræ: C 7, D 13, L 7 (or D 14, L 6), S 3, C 22-30. Body elongated and compressed. Head pointed in front; ears rather small. Extremities short. Feet small and rounded. Toes short, five on each foot. First toe both on fore and hind feet much shorter than the others. Palms and soles covered with hair, except the pads of the feet and toes, and in some species a narrow central line on the under side of the sole, extending backwards nearly to the heel. Tail moderate or long; usually marked with dark and light rings. A pair of large glandular follicles situated on the perineum (in both sexes), and secreting in most species an oily substance of a peculiarly penetrating odour.

The numerous species of this section form a large series, the two extremes of which differ considerably, but the several genera into which they may be divided blend so into one another that it is difficult to differentiate them sharply.

All the animals of this section are, for their size, extremely active, fierce, and rapacious. They feed chiefly on small mammals and birds.

_Viverra._[442]—This includes the largest species. The teeth (Fig. 232) are stouter and less compressed than in the other genera; the second upper molar being especially larger. The auditory bulla smaller and more pointed in front. Body shorter and stouter; limbs longer; tail shorter, tapering. Under side of tarsus completely covered with hair. Claws longer and less retractile. Fur rather long and loose, and in the middle line of the neck and back usually elongated so as to form a sort of crest or mane; neck with a black gorget. Pupil circular when contracted. Perineal glands greatly developed. These characters apply especially to _V. civetta_, the African Civet, or “Civet-Cat” as it is commonly called, an animal rather larger than a common Fox, and an inhabitant of intratropical Africa. _V. zibetha_, the Indian Civet, of about equal size, inhabits Bengal, China, the Malay Peninsula, and adjoining islands. _V. tangalunga_, from Java, Sumatra, Borneo, and the Philippines, and _V. megaspila_, from Burma, are smaller but nearly allied animals; the latter being more distinctly spotted than either of the others. From these species and the next the civet of commerce, once so much admired as a perfume in England, and still largely used in the East, is obtained. The animals are kept in cages, and the odoriferous secretion collected from the interior of the perineal follicles with a spoon or spatula.

[Illustration: FIG. 232.—The left upper dentition of the Indian Civet (_Viverra zibetha_). From the _Palæontologia Indica_.]

The Rasse or Lesser Indian Civet (_V. malaccensis_) may be regarded as the representative of a distinct group of _Viverra_, although often referred to a separate genus (_Viverricula_). The size of this animal is smaller than in the typical group, the build is slighter, the muzzle finer, the claws sharper and more curved, and there is no erectile mane along the back. Generally there is an alisphenoid canal in the skull; and the anterior chamber of the auditory bulla is much more inflated than the hinder one, so that the apparent length of the whole bulla is increased. This species is found over the greater part of India, and extends to the Malay Peninsula and Southern China.

Large species of _Viverra_ occur in the Pleistocene and Pliocene of India, and also in the Pliocene of France, which approximate in some characters of the dentition to the extinct genus _Ictitherium_, mentioned at the end of the family. Species of this genus have also been described from the Miocene and Upper Eocene of Europe. The Lower Miocene _V. antiqua_ has an alisphenoid canal, and all the other cranial characters of the typical forms.

_Fossa._[443]—The Fossa of Madagascar comes so close to the Rasse that its right to generic distinction seems doubtful. There is, however, no scent-pouch. The limbs are slender; and there are two small bare spots on the sole of the hind foot, above the plantar pads. There is no dark line along the back; the throat gorget of _Viverra_ is absent; and in the tail the spots only tend to form rings, which are not complete. The skull has an alisphenoid canal, and a large bulla as in the typical group of _Viverra_.

[Illustration: FIG. 233.—The Common Genet (_Genetta vulgaris_).]

_Genetta._[444]—The Genettes are smaller animals, with more elongated and slender bodies, and shorter limbs than the Civets. Skull elongated and narrow. Auditory bulla large, elongated, rounded at both ends. Teeth compressed and sharp pointed. The inner side of the third upper premolar has a tubercle not present in the previous genus, and the talon of the lower carnassial is larger. Pupil contracting to a linear aperture. Tail long, slender. Fur short and soft, spotted or cloudy. Under side of the tarso-metatarsus with a narrow longitudinal bald streak. No pouch for storing the secretion of the scent-gland. _G. vulgaris_, the common Genet (Fig. 233), is found in France south of the river Loire, Spain, South-Western Asia, and Africa from Barbary to the Cape. _G. felina_, _senegalensis_, _tigrina_, and _pardalis_ are other named species, all African in habitat.

[Illustration: FIG. 234.—Stomach of Genet cut open. _œ_, Œsophagus; _pv_, pyloric valve; _x_, sudden bend where the internal folds are interrupted. (From Mivart, _Proc. Zool. Soc._ 1882, p. 505.)]

[Illustration: FIG. 235.—Cæcum of Genet. (After Mivart, _loc. cit._ p. 508.)]

A few details (taken from Professor Mivart’s memoirs on the Æluroidea) of the anatomy of the soft parts of the Genet may be given as illustration of these parts in the Carnivora generally, and of this family and genus in particular. The salivary glands are shown in Fig. 19 (p. 56), and these conform to the general type prevalent in the Æluroidea. Thus there is a distinct zygomatic gland; the parotid with its (Steno’s) duct is well developed; and there is a small submaxillary gland. The stomach (Fig. 234), while conforming to the simple type characteristic of the Carnivora, is much larger than in the Cat; it is characterised by the presence of some strongly marked internal folds near the pyloric extremity, which stop suddenly at a point where the stomach makes an abrupt constriction and flexure. Beyond this point there are three other longitudinal folds; and the pyloric valve is small. The allied genera present modifications from this form of stomach. The cæcum (Fig. 235) is short, thick, and pointed. The liver (Fig. 236) much resembles that of the Cat, but differs in that the left lateral lobe is undivided, although having a small groove on its posterior or abdominal aspect, while the cystic fissure is less deep, and situated more to the right. The caudate lobe is relatively longer, has a deep concavity, and runs uninterruptedly into the Spigelian; the latter being relatively somewhat larger than in the Cat, with a deep groove dividing the proximal third from the distal two-thirds. In _Viverra_ the right lateral and right central lobes are nearly equal in size. The variations in the form of the liver of the allied genera are detailed in Professor Mivart’s memoir. The brain of the Genet is shown in Fig. 23 (p. 71); the small depression _d_ placed on the superior lateral gyrus appears to be the sole representative of the distinct crucial sulcus which distinguishes the brains of the _Felidæ_ from those of all other members of the Æluroidea.

[Illustration: FIG. 236.—Abdominal aspect of the liver of the Genet. _c_, Caudal lobe; _gb_, gall-bladder; _ha_, hepatic artery; _hd_, hepatic duct; _LC_, left central lobe; _LL_, left lateral lobe; _pv_, portal vein; _RC_, right central lobe; _RL_, right lateral lobe; _Sp_, Spigelian lobe; _vc_, vena cava. (From Mivart, _Proc. Zool. Soc._ 1882, p. 510.)]

[Illustration: FIG. 237.—Cæcum of _Prionodon_. (From Mivart, _Proc. Zool. Soc._ 1882, p. 508.)]

_Prionodon._[445]—This and the following genus comprise the beautiful Linsangs (Fig. 238), which are distinguished from the preceding genera by the loss of the second upper molar, which is, however, very small in some of the Genets. In the present genus the ground colour is whitish or yellowish with brown or black markings, which may either form broad continuous patches across the hinder part of the body, or may be broken up into spots. The tail is very long, the limbs comparatively short, and the fur very short and close. The pollex and hallux are well developed; the claws are almost completely retractile; and the tarsus and metatarsus are completely haired. The pupil is round. The cæcum (Fig. 237) is remarkably small. This genus is exclusively Oriental, and comprises _P. gracilis_ from Borneo, Java, and (?) Sumatra, _P. pardicolor_ from Nipal, and _P. maculosus_ from Tenasserim; the head and body of the latter measuring from 18 to 20 inches in length. Speaking of _P. pardicolor_, Mr. Hodgson observes that it is “equally at home on trees and on the ground; it dwells and breeds in the hollows of decayed trees. It is not gregarious at all, and preys chiefly upon small birds, which it is wont to pounce upon from the cover of the grass. The times of breeding are said to be February and August, and the litter to consist of two young, there being two litters each year.”

_Poiana._[446]—This African genus, represented solely by one species, _P. poënsis_ (Fig. 238), from Fernando Po, is very closely allied to the preceding, but the spots are smaller, and show no tendency to run into transverse bands or stripes, except in the region of the head and shoulder; while the sole of the foot has a narrow bald band running up towards the tarsus, as in _Genetta_. The length of the head and body is 38 inches, and that of the tail about 40 inches. It is probable that this animal should really be regarded as a slightly aberrant species of the genus _Prionodon_.

[Illustration: FIG. 238.—The African Linsang (_Poiana poënsis_). From Mivart, _Proc. Zool. Soc._ 1882, p. 160.]

The five following genera differ in several important respects from all the preceding, and collectively constitute the _Paradoxurine_ section of Professor Mivart. With the exception of one African form, they are mainly Oriental. In this section the auditory bulla is frequently in two portions, the posterior moiety in one case being unossified, and it is always much narrowed in front (Fig. 239). The palate (as in the figure) may be much produced behind the molars; and the teeth are often but slightly sectorial, and may be very small. The long tail is in most cases not ringed.

_Paradoxurus._[447]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₂; total 40. The blunt and rounded form of the cusps of the hinder premolar and the molar teeth distinguishes this genus from most of the members of the family. Vertebræ: C 7, D 13, L 7, S 3, C 29-36. Head pointed in front. Ears small, rounded. Body long. Limbs moderate. Palms and soles almost entirely naked, and joining the foot-pads without the intervention of any hairy space. Claws completely retractile. Pupil vertical. Tail long, non-prehensile; in the Indian species without rings. The Paradoxures or Palm-Civets are less strictly carnivorous than the other members of the family. They are mostly about the size of the common Cat, or rather larger, and are partly arboreal in their habits. The species are rather numerous, and present considerable variations in the details of the form and size of their molar teeth; in only a few does the bony palate extend behind the molars. They are restricted geographically to Southern Asia and the Indo-Malayan archipelago. The best known species[448] are _P. niger_, _P. hermaphroditus_, _P. jerdoni_, _P. aureus_, _P. grayi_ from India and Burma, _P. philippinensis_ of the Philippines, _P. larvatus_ of Southern China and Formosa, _P. leucomystax_ of the Malay Peninsula, Sumatra, and Borneo, and _P. musschenbroeki_ of Celebes. The name _Paradoxurus_ was applied from the mistaken notion that the tail was prehensile. Mr. Blanford[449] gives the following account of the habits of _P. niger_: “The common Palm-Civet, Tree-Cat, or Toddy-Cat, is a familiar animal in most parts of India, though, being thoroughly nocturnal in its habits, it is but rarely seen in the daytime. It is arboreal, passing the day generally in trees, either coiled up in the branches, or in a hole in the trunk, and in places where cocoa-nut palms are common it frequently selects one of them for a residence. Mango groves are also a favourite resort. It not unfrequently takes up its abode in the thatched roofs of houses; Jerdon found a large colony established in the rafters of his own house in Tellicheri. It even occurs in large towns; I have known of one being caught in the middle of Calcutta.”

[Illustration: FIG. 239.—Palatal aspect of the left side of the cranium and mandible of _Arctogale leucotis_. _a_, Anterior opening of alisphenoid canal; _o_, foramen ovale; _c_, carotid canal ¹⁄₁. (From Mivart, _Proc. Zool Soc._ 1882, p. 165.)]

_Arctogale._[450]—This genus—represented only by _A. trivirgata_ of Java, and _A. leucotis_ of Burma, Tenasserim, Sumatra, Java, etc.—is chiefly distinguished from _Paradoxurus_ by the extremely small size of the cheek-teeth (Fig. 239), which are often not in contact with one another; the upper carnassial being almost triangular in shape. Palate frequently convex longitudinally between the carnassials, and greatly produced behind the last molar, with a very narrow bony aperture of the posterior nares. The soles of the feet are still more naked than in _Paradoxurus_; and the pollex and hallux are more divergent. In _A. leucotis_ the length of the head and body is 26·5 inches, and the tail 27 inches. In many specimens the three dorsal stripes are much less distinctly marked than in others, and tend to break up into spots; while the general coloration is considerably lighter.

_Hemigale_,[451] another modification of the Paradoxure type, contains one species, _H. hardwickei_, from Borneo and Malacca, an elegant-looking animal, smaller and more slender than the Paradoxures, of light gray colour, with transverse broad dark bands across the back and loins; the proximal portion of the tail being ringed. The tarsus is hairy. The general cranial characters are those of _Paradoxurus_, but the auditory bulla is ankylosed into a single piece.

_Arctictis._[452]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₂; total 40. The posterior upper molar and the first lower premolar very often absent. Cheek-teeth generally small and rounded, with a distinct interval between them, but formed generally on the same pattern as _Paradoxurus_. Vertebræ: C 7, D 14, L 5, S 3, C 34. Body elongated. Head broad behind, with a small pointed face. Whiskers long and numerous. Ears small, rounded, but clothed with a pencil of long hairs. Eyes small. Limbs short. Soles and palms broad, entirely naked. Tail very long and prehensile; thickly covered with long hair. Fur long and harsh. Cæcum extremely small. But one species is known, _A. binturong_, the Binturong, an inhabitant of Southern Asia from Nipal through the Malay Peninsula to the islands of Sumatra and Java. Although structurally agreeing closely with the Paradoxures, its tufted ears, long, coarse, and dark hair, and prehensile tail give it a very different external appearance. It may be regarded as a very aberrant Paradoxure, connected, so far as dental characters are concerned, with _Paradoxurus_ by means of _Arctogale_. The bony palate also extends considerably behind the last molar, as in the latter. The Binturong is slow and cautious in its movements, chiefly if not entirely arboreal, and appears to feed on vegetable as well as animal substances.

_Nandinia_[453] contains one species, _N. binotata_, a somewhat aberrant Paradoxure, from West Africa. It is rather smaller than the true Paradoxures, with smaller and more pointed molar teeth, and no cæcum. The wall of the hinder chamber of the auditory bulla remains through life unossified.

The dentition appears to be of a more decidedly carnivorous type than in the other members of the section.

_Cynogale._[454]—This remarkable genus is regarded by Professor Mivart as representing a third section of the _Viverrinæ_; it contains one species, _C. bennetti_ (described by S. Müller under the name of _Potamophilus barbatus_), from Borneo, Sumatra, and the Malay Peninsula. This is a curious Otter-like modification of the Viverrine type, having semiaquatic habits, both swimming in the water and climbing trees, living upon fish, crustacea, small mammals, birds, and fruit. The number and general arrangement of its teeth are as in _Paradoxurus_, but the premolars are peculiarly elongated, compressed, pointed and recurved, somewhat as in the Seals, though the molars are tuberculated. The head is elongated, the muzzle broad and depressed. Whiskers very long and abundant. Ears small and rounded. Toes short and slightly webbed at the base. Tail short, cylindrical, covered with short hair. Fur very dense and soft, of a dark brown colour, mixed with black and gray. Humerus without entepicondylar foramen.

Subfamily =Herpestinæ=.—Auditory bulla very prominent, and somewhat pear-shaped, the posterior chamber being large, rounded, and generally with its greatest prominence to the outer side. The anterior chamber considerably dilated, and produced into a short inferior wall to the auditory meatus, in which is a depression or vacuity just below the centre of the opening of the meatus. Sometimes this vacuity is continued into the meatus, forming a narrow fissure. The paroccipital process does not project beyond the bulla, but is spread out and lost (in adult animals) on its posterior surface. Toes straight; claws lengthened, exserted, non-retractile. No perineal glands. The dentition is always of a markedly sectorial type; and the orbit may be surrounded by bone. Very generally the anus opens into a sac-like depression. The majority of the genera are Ethiopian; the type genus alone extending into the Oriental and Palæarctic regions.

_Herpestes._[455]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, sometimes ³⁄₃, _m_ ²⁄₂; total 40 or 36. Teeth of molar series generally with strongly developed, sharply-pointed cusps. Skull elongated, constricted behind the orbits. Face short and compressed. Frontal region broad and arched. Postorbital processes of frontal and jugal bones well developed, generally meeting so as to complete the circle of the orbit behind. Vertebræ: C 7, D 13, L 7, S 3, C 21-26. Head pointed in front. Ears short and rounded. Body very long and slender. Extremities short. Five toes on each foot, the first, especially that on the hind foot, very short. Toes free, or but slightly palmated. Palms generally naked. Distal portion of soles naked, under surface of tarsus and metatarsus usually clothed with hair, but considerable specific variation in this respect. Tail long or moderate, generally thick at the base, and sometimes covered with more or less elongated hair. The longer hairs covering the body and tail almost always annulated. This genus contains a very large number of animals commonly called Ichneumons, or in India Mungooses, varying in size from that of a large Cat down to a Weasel. They are widely distributed over the African continent and the southern parts of Asia, especially India and the Indo-Malayan archipelago, one species occurring also in Spain. They are mostly terrestrial in their habits, feeding on small mammals and birds, reptiles, especially snakes, eggs of birds and reptiles, and also insects. Some species are partially domesticated, being used to keep houses clear of rats, mice, and snakes. _H. ichneumon_ was a sacred animal to the ancient Egyptians. They vary considerably in appearance, some, as _H. galera_ and _H. urva_ (Fig. 240), are larger and heavier, with stouter body, longer limbs, and stronger teeth. The common Indian Mungoose (_H. mungo_) is considerably smaller than the Egyptian form; its fur is of a pale gray colour, the hairs being largely white ringed, while the cheeks and throat are more or less reddish. Like the Egyptian species, it is frequently domesticated, and put to a similar use. It is especially serviceable in India as a serpent-killer, destroying not only the eggs and young of these creatures, but attacking without hesitation and killing the most venomous adult snakes. The fact that it invariably survives those encounters has led to the belief that it either enjoys immunity from the effects of snake-poison, or that after being bitten it has recourse, as the natives maintain, to the root of a plant as an antidote. Neither of these suppositions has stood the test of scientific examination, for it has been found that when actually bitten it falls a victim to the poison as rapidly as other mammals, while there is no trustworthy evidence of its seeking a vegetable antidote. The truth seems to be that the Mungoose, by its exceeding agility and quickness of eye, avoids the fangs of the snake while fixing its own teeth in the back of the reptile’s neck. One large species, believed to be from Africa, recently described as _H. grandis_, is remarkable for the extreme complexity of the cusps on the molars, and also for the absence of an entepicondylar foramen to the humerus; the latter feature also occurring in the allied _H. albicaudatus_. The Oriental _H. urva_ (Fig. 246) is stated to be somewhat aquatic in habits, and to feed on frogs and crabs.

[Illustration: FIG. 240.—The Crab-eating Mungoose (_Herpestes urva_). From Blanford, _Mammalia of British India_, p. 130.]

Remains of the small _H. nipalensis_ occur in the cavern-deposits of Madras. Viverroids from the Miocene and Upper Eocene of Europe, which agree with _Herpestes_ in the presence of an inner tubercle to the third upper premolar and of a hinder cusp to the fourth lower premolar, have been referred to the existing genus. The species which have been separated generically under the three following names are very closely allied to _Herpestes_.

_Helogale_,[456] premolars ³⁄₃, without diastema between first and second; soles of feet completely naked. Contains two small South-African species, _H. parvula_ and _H. undulata_.

_Bdeogale_[457] contains also two small Ichneumon-like animals, _B. crassicauda_ and _puisa_, differing from _Herpestes_ proper in having only four toes on each foot, both pollex and hallux being absent. The orbit is nearly complete, the tail of moderate length and rather bushy.

_Cynictis._[458]—Pollex present, but hallux absent. Skull shorter and broader than in _Herpestes_, rather contracted behind the orbits, which are large and complete behind. Face short. Anterior chamber of the auditory bulla very large. Front claws elongated. _C. penicillata_, from South Africa. The cæcum (Fig. 241) of this genus is longer than in any other member of the family.

All the foregoing Herpestines have the nose short, with its under surface flat, bald, and with a median longitudinal groove. The remaining forms have the nose more or less produced, with its under side convex, and a space between the nostrils and the upper lip covered with close adpressed hairs, and without any median groove.

[Illustration: FIG. 241.—Cæcum of _Cynictis penicillata_. (From Mivart, _Proc. Zool. Soc._ 1882, p. 508.)]

_Rhinogale._[459]—Toes 5-5. Claws of fore feet short, compressed, acute. Under surface of tarsus hairy. Palate flat. Founded on a single specimen from East Africa, _R. melleri_.

_Crossarchus._[460]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ³⁄₃, _m_ ²⁄₂; total 36. Snout elongated. Toes 5-5. Claws on fore feet long and curved. Hallux very short. Under surface of tarsus naked. Tail shorter than the body, tapering. Palate flat. Fur harsh. Species: _C. obscurus_, the Kusimanse, a small burrowing animal from West Africa, of uniform dark brown colour; _C. fasciatus_; _C. zebra_; and _C. gambianus_.

_Suricata._[461]—A more distinct genus than any of the above. The dental formula as in the last, but the teeth of the cheek-series remarkably short in the antero-posterior direction, corresponding with the shortness of the skull generally (Fig. 222). Orbits complete behind. Vertebræ: C 7, D 15, L 6, S 3, C 20. Though the head is short and broad, the nose is pointed and rather produced and movable. Ears very short. Body shorter and limbs longer than in _Herpestes_. Toes 4-4, the pollex and hallux being absent. Claws on fore feet very long and narrow, arched, pointed, and subequal. Hind feet with much shorter claws, soles hairy. Tail rather shorter than the body. One species only is known, the Suricate, _S. tetradactyla_, a small gray-brown animal, with dark transverse stripes on the hinder part of the back, from South Africa. The cæcum is short.

[Illustration: FIG. 242.—Cæcum of _Galidea elegans_. (From Mivart, _Proc. Zool. Soc._ 1882, p. 508.)]

_Galidictis_,[462] _Galidea_,[463] and _Hemigalidea_[464] are names of three slight generic modifications of the Viverrine type, allied to the _Herpestinæ_, but placed by Mivart in a distinct subfamily, _Galidictiinæ_. They are all characterised by the absence of the alisphenoid canal in the skull, as well as of the entepicondylar foramen to the humerus; and are inhabitants of Madagascar. The best known, _Galidea elegans_, is a lively Squirrel-like little animal with soft fur and a long bushy tail, which climbs and jumps with agility. It is of a chestnut-brown colour, the tail being annulated with darker brown. The cæcum (Fig. 242) is remarkable for its comparative length and pointed termination. _Hemigalidea_ is distinguished by the absence of rings on the tail. _Galidictis vittata_ and _striata_ chiefly differ from the Ichneumons in their coloration, being gray with parallel longitudinal stripes of dark brown.

_Eupleres_[465] is another form, also from Madagascar, which has been placed in a subfamily apart. It differs remarkably from all the other _Viverridæ_ in the weak development of the jaws and the small size of the teeth (Fig. 243), in consequence of which it was, when first discovered, placed in the order Insectivora. Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₂; total 40. Vertebræ: C 7, D 13, L 7, S 3, C 20. No alisphenoid canal; an entepicondylar foramen to the humerus. But one species is known, _E. goudoti_.

[Illustration: FIG. 243.—Skull of _Eupleres goudoti_. ⅘ natural size. Mus. Roy. Coll. Surgeons.]

_Extinct Genera._—The Tertiaries of the Old World have yielded several genera allied to the existing Viverroids, some of which show decided signs of affinity with other families. Of these the Lower Miocene _Amphictis_ appears to be nearly related to _Viverra_, but is distinguished by the form of the second lower molar, which is longer and has two distinct roots. _Palæoprionodon_, of the French Phosphorites, has a dentition very like that of _Prionodon_, the molars being reduced to ¹⁄₂; the skull has an alisphenoid canal and the general basal characters of the _Viverridæ_, but resembles the _Mustelidæ_ in the presence of a glenoid foramen and in the position of the condylar foramen. In _Stenoplesictis_, of the same deposits, the dental formula is _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₂; and although the skull has a complete septum in the bulla, yet some of the cranial and dental features approximate so decidedly towards those of the extinct _Mustelidæ_, as to lead some authorities to refer the genus to that family. The most probable explanation of this resemblance is that the Musteloids have originated from generalised Viverroids allied to _Stenoplesictis_. The Lower Pliocene _Ictitherium_ differs from all other Viverroids in the presence of three distinct lobes to the upper carnassial, and thereby connects the other members of the family so closely with the _Hyænidæ_ that it is practically impossible to draw up a definition which will distinguish the two families.

The North American Eocene genera _Miacis_ and _Didymictis_ are generally regarded as representing a separate family—_Miacidæ_—with affinities both to the _Viverridæ_ and _Canidæ_.

_Family_ PROTELEIDÆ.

Skull with no alisphenoid canal; and the auditory bulla divided into two distinct chambers. Dorsal vertebræ 15. Molars ¹⁄₁. Premolar and molar teeth very small and simple in character.

_Proteles._[466]—This genus contains but a single species, _P. cristatus_, the Aard-Wolf or Earth-Wolf of the Dutch colonists of the Cape, an animal nearly allied to the Hyænas, but remarkably modified in its dentition, the molar teeth being very small, placed far apart, and almost rudimentary in character (Fig. 244). The canines are long and rather slender. The dental formula is _i_ ³⁄₃, _c_ ¹⁄₁, _p_ and _m_ ⁴⁄₃₋₄; total 30 or 32. Vertebræ: C 7, D 15, L 5, S 2, C 24. The fore feet with five toes; the pollex though short, with a distinct claw. The hind feet with four subequal toes. Claws all strong, blunt, subcompressed, and non-retractile. The general external appearance is very like that of a small Striped Hyæna, but the muzzle is more pointed and the ears larger. It has a copious mane of long hair, capable of being erected when the animal is excited, along the middle line of the neck and back. It is a native of South Africa, and is a burrowing nocturnal animal, feeding on decomposing animal substances, larvæ, and termites. Observations upon specimens in captivity indicate that it has neither inclination nor power to attack or feed upon living vertebrated animals.

[Illustration: FIG. 244.—Skull and Dentition of the Aard-Wolf (_Proteles cristatus_). ½ natural size.]

Some writers regard _Proteles_ as representing a subfamily of the _Hyænidæ_.[467]

_Family_ HYÆNIDÆ.

Skull with no alisphenoid canal; and the auditory bulla not divided by a septum into two chambers. Dorsal vertebræ 15. Molars usually ¹⁄₁, but in some fossil forms ¹⁄₂, or ²⁄₂, the second lower molar being very small; upper carnassial with three distinct lobes; lower carnassial with a large blade and small talon. No entepicondylar foramen to the humerus. This family is confined to the Old World, where it is now represented by a single genus, which, although evidently nearly related to the _Viverridæ_, is sufficiently distinct to be regarded as not referable to that family. The extinct _Ictitherium_, however, as already mentioned, connects the more generalised members of the _Hyænidæ_ very closely with the _Viverridæ_.

_Hyæna._[468]—Dentition in existing forms usually _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₃, _m_ ¹⁄₁; total 34. Teeth, especially canines and premolars, very large, strong, and conical. Upper carnassial (Fig. 245) with a very large, distinctly trilobed blade and a moderately developed inner tubercle placed at the anterior extremity of the blade. Molar very small, and placed transversely close to the hinder edge of the last, as in the _Felidæ_. Lower carnassial consisting of little more than the bilobed blade. Zygomatic arches of cranium very wide and strong. Sagittal crest high, giving attachment to very powerful biting muscles. Orbits incomplete behind. Vertebræ: C 7, D 15, L 5, S 4, C 19. Limbs rather long, especially the anterior pair, digitigrade, four subequal toes on each, with stout non-retractile claws. Pollex and hallux only represented by rudimentary metacarpal and metatarsal bones. Tail rather short. A large post-anal median glandular pouch, into which the largely developed anal scent glands pour their secretion.

[Illustration: FIG. 245.—Outer (_A_) and palatal (_B_) aspects of the right upper carnassial tooth of the Striped Hyæna (_Hyæna striata_). From the _Quart. Journ. Geol. Soc._]

The three existing species of Hyæna are divisible into two sections, to which some zoologists assign generic rank, but fossil forms show such a transition between these two types as to render any such division impracticable.

The typical or _Euhyænine_ group presents the following distinctive features. Upper molar moderately developed and three-rooted. An inner cusp and hind talon more or less developed on the lower molar. Ears large, pointed. Hair long, forming a mane on the back and shoulders. _H. striata_, the Striped Hyæna (Fig. 246) of Northern Africa and Southern Asia. _H. brunnea_, of South Africa, in some respects intermediate between this and the next group.

The Striped Hyæna is dirty gray in colour, with narrow transverse tawny or blackish stripes on the body and legs; the length of the head and body is 3½ feet, and that of the tail, with its hair, 1½ feet. It occurs throughout peninsular India, where it is most common in open hilly districts, and in North Africa. Mr. Blanford[469] gives the following account of its habits: “It is a nocturnal animal, and although an occasional individual may be met with returning to its den in the early morning, its rambles are usually commenced after sunset and ended before sunrise. During the night it roams far and wide, and no tracks of wild animals are more common in the countries where it is found than its unmistakable footprints, very like a dog’s in shape, but with the marks of the hind feet conspicuously smaller than those of the fore feet. Unlike the Spotted Hyæna, the Striped species appears to be solitary in its habits, and it is rare to meet with more than two together. The principal food of the Hyæna consists of the carcases of animals that have died of disease or been killed by beasts of prey, and very often it carries off portions of the body to its den. I once shot one that was carrying away the hind leg of a Nilghai. The powerful jaws and large teeth are admirably adapted for crushing bones, which are consumed by Hyænas, after the flesh has been picked off by vultures and jackals. Occasionally sheep or goats, and more often dogs, are carried off by Hyænas, and the latter at all events are often taken alive to the animal’s den.” The Striped Hyæna is essentially a cowardly animal, and one that is much more silent than _H. crocuta_. Remains of _H. striata_ are found in the cavern-deposits of the south of France, and also in the Upper Pliocene of the Val d’Arno in Tuscany, and in the English Red Crag.

[Illustration: FIG. 246.—The Striped Hyæna (_Hyæna striata_).]

The _Crocutine_ group presents the following characters. Upper molar extremely small, two- or one-rooted, often deciduous. Lower molar without trace of inner cusp, and with an extremely small talon. Ears moderate, rounded. Hair not elongated to form a mane. _H. crocuta_, the Spotted Hyæna (Fig. 247), from Africa south of the Sahara. In dental characters as well as in its visceral anatomy, especially as regards the reproductive organs of the female,[470] this species may be considered as by far the more specialised form. The Spotted Hyæna is a larger and bolder animal than the Striped species, hunting in packs, and uttering very frequently its unearthly cry. The coloration consists of dark brown spots on a yellowish ground. It was formerly very common at the Cape. Remains of a large race of this species are exceedingly common in the cavern-deposits of Europe, where they were first described under the name of _Hyæna spelæa_; teeth have also been met with in the Norfolk Forest-bed, and in cavern-deposits in Madras—the latter locality being exceedingly interesting from a distributional point of view.

[Illustration: FIG. 247.—The Spotted Hyæna (_Hyæna crocuta_).]

In addition to the remains of existing species, to which reference has been already made, there were numerous extinct forms of _Hyæna_ in the upper Tertiaries of Europe, from the horizon of the Lower Pliocene Pikermi beds of Greece upwards. In the Crocutine group _H. colvini_ of the Pliocene of India (Fig. 248), and _H. robusta_ of that of Italy, appear to have been ancestral forms allied to _H. crocuta_; the former being distinguished by the loss of the first upper premolar. _H. eximia_, of the Pikermi beds, is a more generalised form, in which the first lower premolar (lost in existing forms) is retained. In the typical group, _H. arvernensis_ and _H. perrieri_, of the Upper Pliocene of the Continent, approximate to _H. brunnea_; although _H. perrieri_ makes a farther step towards the Crocutine group by the loss of the inner cusp in the lower carnassial. The extinct _Hyænictine_ group, as represented by the Indian _H. sivalensis_ and the Grecian _H. græca_, connects _H. striata_ with _Palhyæna_. Both are characterised by the presence of a small second lower molar behind the carnassial; while _H. græca_ also has four lower premolars. Still more generalised is the _Lychyænine_ group; comprising _H. macrostoma_ of India and _H. chæretis_ of the Pikermi beds; in these forms the muzzle was longer, and the premolars much more compressed than in the existing species, thus making a very decided approach to the _Viverridæ_. There were four lower premolars; the lower carnassial had an inner cusp, and it is probable that there was a second lower molar; while the first upper molar was placed partially behind the carnassial. The Lower Pliocene _Palhyæna hipparionum_, in which the dental formula is _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₂, is a smaller form with long jaws and compressed premolars which approaches so closely to the Viverroid genus _Ictitherium_ as to show pretty clearly how the Hyænas have been gradually modified from that stock.

[Illustration: FIG. 248.—Outer view of part of the right ramus of the mandible of _Hyæna colvini_, showing the third and fourth premolars and the carnassial. (From the _Palæontologia Indica_.)]

_Section_ CYNOIDEA.

_Family_ CANIDÆ.

This section contains the single family of the _Canidæ_, or Dog-like animals, which appear to hold an intermediate position between the other two sections, retaining also many of the more generalised characters of the ancient members of the order. The structure of the auditory bulla and adjacent parts of the bones of the skull is intermediate between that of the Æluroid and Arctoid forms. In the number and arrangement of the teeth they more nearly approach the primitive heterodont type than any other existing Carnivora. A cæcum is always present, sometimes short and simple, but when long it is folded upon itself in a characteristic manner.

[Illustration: FIG. 249.—Right lateral aspect of the skull of the Dog (_Canis familiaris_).]

The characters of the base of the cranium are shown in Fig. 8 (p. 38), where it will be seen that the auditory bulla is inflated, although it has only a rudimental internal septum; the paroccipital process, although in contact with the bulla, is prominent, and there is a large glenoid foramen. In all the existing forms the humerus has lost the entepicondylar foramen; the crowns of the upper molars are triangular in shape (Fig. 251), and the blade of the upper carnassial consists of two lobes.

[Illustration: FIG. 250.—Cæcum of the Arctic Fox (_Canis lagopus_). _i_, Ileum; _c_, colon. In the natural position the colon is uppermost.]

In the alimentary canal the cæcum (Fig. 250) is extremely characteristic. It is a simple appendage of nearly uniform width (about equal to that of the ileum) attached to the side of the canal, just beyond the ileo-cæcal valve, and with a rounded termination. In a Dog of average size it is 5 or 6 inches long if uncoiled, but it is normally folded by its mesenteric attachments backwards and forwards several times on itself by the side of the ileum, after the manner shown in the figure.

The existing Dogs form a very compact group, with numerous species closely resembling each other in essential characters, though differing considerably externally. The most marked differences are slight variations in the number of the true molar teeth, which exceed the usual number in the Cape Long-eared Fox (_Otocyon_), and fall short of it in some other less aberrant forms to which the names of _Icticyon_ and _Cyon_ have been given, and a diminution in the number of toes in the Cape Hunting Dog (_Lycaon_), which has 4-4, instead of 5-4 as in the remainder of the family. After taking these away, there remain a great number of animals called Dogs, Wolves, Jackals, and Foxes, varying from one another only in the characters of the tail, ears, fur, form of the pupil, and some trifling peculiarities of skull and teeth, upon which some authors have divided them into many genera. These divisions are, however, extremely difficult, if not impossible, to define, on account of the numerous gradual transitions from one form to the other.

[Illustration: FIG. 251.—The last four left upper teeth of an extinct Wolf (_Canis cautleyi_). From the _Palæontologia Indica_.]

_Canis._[471]—It appears on the whole convenient to retain all the species, with the exception of _Otocyon_, _Icticyon_, and _Lycaon_, in the old genus _Canis_, the most prominent characters of which are the following. Teeth, usually _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₃; total 42. The absence of the last upper molar (_m_ ³⁄), alone distinguishes this from the generalised dentition of heterodonts, and this tooth is occasionally present in one species (_C. cancrivorus_). In certain Asiatic species (_C. primævus_ and its allies), which on this account have been separated to form the genus _Cyon_ of Hodgson, the last lower molar ⁄_m₃_ appears to be constantly absent. The milk-dentition is _di_ ³⁄₃, _dc_ ¹⁄₁, _dm_ ³⁄₃; total 28,—the first permanent premolar having no predecessor. The teeth of both permanent and milk or temporary series are figured on p. 26, Fig. 3, from the outer aspect, while the woodcut 251 shows the palatal aspect of the hinder upper teeth. The upper carnassial (_p_ ⁴⁄) consists of a stout blade, of which the anterior lobe is almost obsolete, the middle lobe large, conical, and pointed backwards, and the posterior lobe in the form of a compressed ridge; the inner tubercle is very small, and placed quite at the fore part of the tooth. The first molar is more than half the antero-posterior length of the carnassial, and considerably wider than it is long; its crown consists of two prominent conical cusps, of which the anterior is the larger, and a low broad inward prolongation, supporting two more or less distinct cusps and a raised inner border. The second molar resembles the first in general form, but is considerably smaller. The lower carnassial ⁄_m₁_ is a very large tooth, with a strong compressed bilobed blade, the hinder lobe being considerably the larger and more pointed, a small but distinct inner cusp placed at the hinder margin of the posterior lobe of the blade, and a broad, low, tuberculated talon, or heel, occupying about one-third of the whole length of the tooth. The second molar is less than half the length of the first, with a pair of cusps placed side by side anteriorly, and a less distinct posterior pair. The third is an extremely small and simple tooth, with a subcircular tuberculated crown and single root.

The cranium (Fig. 249) is more or less elongated, the facial portion tapering forwards and compressed. The jaws are elongated, and the zygomata moderately strong. The postorbital processes of the frontal short, leaving the orbit widely open posteriorly. Vertebræ: C 7, D 13, L 7, S 3, C 17-22. Clavicles present, but very rudimentary. Limbs of moderate proportions, digitigrade. Feet short; five toes on the fore foot, the pollex much shorter than the others, and not reaching to the ground. Four toes on the hind foot, the hallux being represented by a rudiment of the metatarsal.[472] All the toes are provided with exserted, non-retractile, slightly curved, and blunt claws, which, being exposed, become worn at the tips. Tail moderate, or rather long, generally somewhat bushy. The pupil of the eye, when contracted, is in some species round, in others elliptical and vertical.

This extensive genus may be considered as truly cosmopolitan. One or more species occur in every part of the American continent from Greenland to Patagonia and the Falkland Isles; and similarly, in the Old World, Europe, Africa, and Asia, with most of the large islands adjacent, and even Australia, have their wild Dogs, though in the last case they may belong to a feral race, introduced originally by man. They are generally sociable animals, hunting their prey in packs. Many species burrow in the ground; none habitually climb trees. Though mostly carnivorous, feeding chiefly on animals they have chased and killed themselves, many, especially among the smaller species, eat garbage, carrion, insects, and also fruit, berries, and other vegetable substances. The species are very numerous, and, as in most other large genera, very ill-defined, few zoologists agreeing as to which of the many slightly different modifications should be considered as local varieties and which true species. Perhaps the best cranial character by which the different members of the genus can be distinguished is that pointed out by Burmeister, viz. that in the animals generally called Dogs, Wolves, and Jackals the postorbital process of the frontal bone is regularly smooth and convex above, with its extremity bent downwards, whereas in Foxes this process is hollowed above, with its outer margin (particularly of the anterior border) somewhat raised. This modification coincides in the main with that upon which Professor Huxley[473] has based his division of the group into two parallel series, the Thooids or Lupine forms and Alopecoids or Vulpine forms, which he characterises by the presence of frontal air-sinuses in the former, which not only affect the external contour but to a still greater degree the shape of the anterior part of the cranial cavity, and the absence of such sinuses in the latter. The pupil of the eye when contracted is round in most members of the first group, and vertically elliptical in the others, but more observations are required before this character can be absolutely relied upon. The form and length of the tail is often used for the purposes of classification, but its characters do not coincide with those of the cranium, since many of the South American _Canidæ_ have the long bushy tails of Foxes and the skulls of Wolves. Taking into account various combinations of these and other minor characters, the species may be arranged in the following groups, which some authors have considered as of generic importance.

A. _Thooid or Lupine Series._—The typical group, or _Canis_ proper, contains the largest members of the genus, the true Wolves of the northern parts of both Old and New Worlds (_C. lupus_, etc.), the Jackals of Southern Asia and Africa (_C. aureus_, _mesomelas_, etc.), and the various breeds of the domestic Dog (_C. familiaris_). The true Wolves are (excluding some varieties of the domestic Dog) the largest members of the genus, and have a wide geographical range, extending over nearly the whole of Europe and Asia, and North America from Greenland to Mexico, but they are not found in South America or Africa, being replaced in both of these continents by various species of Jackals and Foxes. As might be expected from this extensive range, and the varied character of the climatic conditions of the countries they inhabit, they present great diversities of size, length and thickness of fur, and coloration, although resembling each other in all important structural characters. These differences have given rise to a supposed multiplicity of species, expressed by the names of _C. lupus_, _C. lycaon_ (Central Europe), _C. laniger_ and _C. niger_ (Tibet), _C. pallipes_ (India), _C. occidentalis_, _C. nubilis_, _C. mexicanus_, etc., of North America, but it is very doubtful whether some of these ought to be distinguished as other than local varieties. Mr. W. T. Blanford, in his recent work on the mammals of India, regards the two forms from Tibet mentioned above as inseparable from _C. lupus_. In North America there is a very distinct smaller species, called the Coyote or Prairie Wolf (_C. latrans_); and perhaps the Japanese Wolf (_C. hodophylax_) may also be distinct, although, except for its smaller size and shorter legs, it is scarcely distinguishable from the common species. Though generally distributed throughout the Indian peninsula, the Indian Wolf (_C. pallipes_), which is rather smaller and slighter than _C. lupus_, is not found in Ceylon, nor in Burma and Siam. The ordinary colour of the Common Wolf is a yellowish or fulvous gray, but specimens have been met with almost pure white and others entirely black. In northern countries the fur is longer and thicker, and the animal generally larger and more powerful than in the southern portion of its range; this being especially the case with the Tibetan races. The habits of the Wolf are similar everywhere, and it is still, and has been from time immemorial, especially known to man in all the countries it inhabits as the devastator of his flocks of sheep. They do not catch their prey by lying in ambush, or stealing up close to it and making a sudden spring as the Cat tribe do, but by fairly running it down in open chase, which their speed and remarkable endurance enable them to do; and usually, except during summer, when the young families of cubs are being separately provided for by their parents, they assemble in troops or packs, and by their combined and persevering efforts are able to overpower and kill even such great animals as the American Bison. It is singular that such closely allied species as the Domestic Dog and the Arctic Fox are among the favourite prey of Wolves, and, as is well known, children and even full-grown people are not unfrequently the objects of their attack when pressed by hunger. Notwithstanding the proverbial ferocity of the Wolf in a wild state, many instances are recorded of animals taken when quite young becoming perfectly tame and attached to the person who has brought them up, when they exhibit many of the ways of a Dog. They can, however, rarely be trusted by strangers.

The history of the Wolf in the British Isles and its gradual extirpation has been thoroughly investigated by Mr. J. E. Harting in his work on _Extinct British Animals_, from which the following account is abridged: To judge by the osteological remains which the researches of geologists have brought to light, there was perhaps scarcely a county in England or Wales in which, at one time or another, wolves did not abound, while in Scotland and Ireland they must have been still more numerous. The fossil remains which have been discovered in Britain are not larger than, nor in any way to be distinguished from, those of European wolves of the present day. Wolf-hunting was a favourite pursuit of the ancient Britons as well as of the Anglo-Saxons. In Athelstan’s reign these animals abounded to such an extent in Yorkshire that a retreat was built by one Acehorn, at Flixton, near Filey, wherein travellers might seek refuge if attacked by them. As is well known, great efforts were made by King Edgar to reduce the number of wolves in the country, but, notwithstanding the annual tribute of 300 skins paid to him during several years by the king of Wales, he was not altogether so successful as has been commonly imagined. In the reign of Henry III the number of wolves in some parts of the country was sufficient to induce the king to make grants of land to various individuals upon the express condition of their taking measures to destroy these animals wherever they could be found. In Edward II’s time the king’s forest of the Peak, in Derbyshire, is especially mentioned as infested with wolves, and it was not until the reign of Henry VII (1485-1509) that wolves appear to have become finally extinct in England. This, however, is rather a matter of inference from the cessation of all mention of them in local records than from any definite evidence of their extirpation. Their last retreat was probably in the desolate wolds of Yorkshire. In Scotland, as might be supposed from the nature of the country, the wolf maintained its hold for a much longer period. There is a well-known story of the last of the race being killed by Sir Ewen Cameron of Lochiel in 1680, but there is evidence of wolves having survived in Sutherlandshire and other parts into the following century (perhaps as late as 1743), though the date of their final extinction cannot be accurately fixed. In Ireland, in Cromwell’s time, wolves were particularly troublesome, and said to be increasing in numbers, so that special measures were taken for their destruction, such as the offering of large rewards for their heads, and the prohibition (in 1652) of the exportation of “wolf-dogs,” the large dogs used for hunting the wolves. The active measures taken then and later reduced their numbers greatly, so that towards the end of the century they became scarce, but, as in the case of the sister island, the date of their final disappearance cannot now be ascertained. It has been placed, upon the evidence of somewhat doubtful traditions, as late as 1766.

Remains of _C. lupus_ are common in the European Pleistocene; while the Indian Pliocene _C. cautleyi_, of which the upper teeth are shown in Fig. 251, was probably the ancestor of _C. pallipes_. _C. neschersensis_, of the Upper Pliocene of France, was a smaller extinct Wolf. A lower jaw from the French Pleistocene, described under the name of _Lycorus_, has only three premolars, but evidently belongs to the Wolf.

The Jackals are smaller than the Wolves, with the bushy tail about one-third the length of the head and body, and the carnassials relatively shorter as compared with the tubercular molars. The Common Jackal (_C. aureus_, Fig. 252) has a very wide distribution, ranging from South-Eastern Europe through South-Western Asia to India and Burma, and also occurring in Northern Africa; being replaced in the Ethiopian region by closely allied species. Remains indistinguishable from _C. aureus_ occur in the Pliocene Siwaliks of Northern India. Jackals hunt at night in packs, uttering the piercing cries so well known to all who have resided in countries where these animals are found.

The origin of the Domestic Dog, with its numerous breeds, has been the subject of much controversy. Some naturalists believe it to be a distinct species, descended from one that no longer exists in a wild state; others have sought to find its progenitors in some one of the wild or feral races, either of true Dogs, Wolves, or Jackals; while others again believe that it is derived from the mingling of two or more wild species or races. It was probably the earliest animal domesticated by man, and few if any other species have undergone such an extraordinary amount of variation in size, form, and proportion of limbs, ears, and tail—variations which have been perpetuated and increased by careful selective breeding. The Dingo or Australian Dog is met with wild, and also as the domestic companion of the aboriginal people. Dogs were also in the possession of the natives of New Zealand and other islands of the Pacific, where no placental mammals exist naturally, on their discovery by Europeans in the last century.

[Illustration: FIG. 252.—The Jackal (_Canis aureus_).]

The second group includes the wild Dogs of the south-east of Asia, described as _Cyon_, and distinguished by slight modifications as _C. rutilans_, _C. dukhunensis_, and _C. javanicus_, and differing from the above in wanting the small last lower tubercular molar. This difference reduces the number of the teeth to the same as in _Viverra_, and is precisely paralleled by some of the species of the extinct genus _Cynodictis_ mentioned below. The muzzle is shorter than in other species, and the facial profile is slightly convex instead of concave. The mammæ are also 12 or 14 instead of the normal 10; while there is long hair between the foot-pads. Wild Dogs inhabit not only the whole of the Oriental region, but extend into Central Asia as far north as the Altai and Amurland (_C. alpinus_). _C. dukhunensis_ ranges from the forest regions of peninsular India to Gilgit and Western Tibet, where it must inhabit open country. In their general form, and more especially the shortness of the legs, these animals come nearer to the Jackals than to the Wolves. They hunt their prey in packs. Remains of species of this group occur in the cavern-deposits of the Continent, and have been described under the name of _C. europæus_.

A group for which the name _Lycalopex_ has been proposed comprises certain South American _Canidæ_, distinguished from _Canis_ proper by their longer tails and Fox-like aspect:—_C. cancrivorus_, _C. brasiliensis_, _C. melampus_, _C. vetulus_, _C. fulvicaudus_, _C. azaræ_, _C. magellanicus_, _C. griseus_. The last three have been further separated (under the name of _Pseudalopex_) on account of slight differences in the relative size of the molar teeth, and of their pupil being elliptical when contracted. _Nyctereutes_ (one species, _C. procyonides_, from Japan and North-East Asia) has no claims to generic distinction but such as are founded upon its long loose fur, short ears, and short bushy tail, which give it some superficial resemblance to a Raccoon.

B. _Alopecoid or Vulpine Series._—The _Vulpine_ group (_Vulpes_) includes the true Foxes, of which there are numerous varieties and species, spread over North America, Eurasia, and Africa, which have been described under the names of _C. vulpes_ (_Vulpes alopex_), the common Fox of Europe; _C. niloticus_, _adustus_, and _variegatus_, Africa; _C. flavescens_, _montanus_, _bengalensis_, _japonicus_, _corsac_, Asia; _C. fulvus_, _macrurus_, _velox_, North America. Mr. Blanford[474] concludes, however, that the Asiatic _C. flavescens_ and _C. montanus_, and very probably the North American Cross-Fox (_C. fulvus_) are merely local races of _C. vulpes_, distinguished by certain peculiarities of coloration. The English Fox measures about 2 feet in length exclusive of the tail, which is about a foot long. Its fur is of a reddish-brown colour above, and more or less white beneath; the back of the ears and the fore part of the limbs are black, and the tip of its bushy tail is white. Its long, sharp muzzle, erect pointed ears, and sharp eye, give it the well-known appearance of sagacity and cunning. The Fox is a solitary animal, inhabiting a burrow, which it either excavates for itself, or obtains by ejecting the badger or the rabbit. So averse, indeed, is the Fox to dig for itself, that when foiled in its attempts to dispossess the badger, it has been known to take up its quarters with the latter, and it can be induced to make its home in artificial burrows constructed of stone and earth for the purpose of facilitating the operation of digging out the cubs. The Fox also occurs in woods, and even in the open country without burrows, lying in its “cover” by day and stealing forth at night in search of its prey. Remains of the Common Fox occur not unfrequently in the Pleistocene deposits of Europe. The Indian _C. bengalensis_ is a very much smaller and well-marked species.

The tail of the above forms is clothed with soft fur and long hair, uniformly mixed; from them Baird distinguishes, under the name of _Urocyon_, other species which have a concealed erect mane of stiff hairs along the upper line of the tail. These have also a shorter muzzle and a wide space between the temporal crests; they are _C. virginianus_ and _C. littoralis_, both from North America. The Arctic Fox (_C. lagopus_, genus _Leucocyon_, Gray) has the tail very full and bushy and the soles of the feet densely furred below. Its colour changes according to season from bluish-gray to pure white.

Certain small elegant African Foxes (_C. zerda_, _famelicus_, and _chama_), with very large ears and corresponding large auditory bullæ, have been separated under the name of _Fennecus_, and are commonly known as Fennecs.

The earliest undoubted occurrence of the genus _Canis_ seems to be in the Upper Miocene of Switzerland, where it is represented by the Fox-like _C. œningensis_. In the Upper Pliocene of France _C. megamastoides_ is said to be allied to the Foxes and Jackals, but with some signs of affinity to the extinct _Cynodictis_. In the Pliocene Siwaliks of India there occurs _C. curvipalatus_, of the size of a small Fox, which appears to have certain resemblances to _Otocyon_.

_Lycaon._[475]—This genus resembles in most of its characters the Dogs of the Lupine series, but the teeth are rather more massive and rounded, the skull is shorter and broader, and there are but four toes on each limb, as in _Hyæna_. The one species, _L. pictus_, the Cape Hunting Dog (Fig. 253) from South and East Africa, is very distinct externally from all the other _Canidæ_. It is nearly as large as a Mastiff, with large, broadly ovate erect ears, and singularly coloured, being not only variable in different individuals, but unsymmetrically marked with large spots of white, yellow, and black. It presents some curious superficial resemblances to _Hyæna crocuta_, perhaps a case of mimetic analogy. It hunts its prey in large packs. A lower jaw from a cave-deposit in Glamorganshire, which agrees with that of the existing form in the presence of an anterior cusp to the last lower premolar, has been made the type of a distinct species (_L. anglicus_).

_Icticyon._[476]—The Bush-Dog (_I. venaticus_), from Guiana and Brazil, is a species about the size of a Fox, with close hair, and short legs and tail, distinguished from all other Dogs by the reduction of the molar teeth to ¹⁄₂, and their comparatively small size. The lower carnassial is also characterised by the loss of the inner cusp of the blade, and the secant form of its hind talon; both these features indicating a specialised type. Remains of the Bush-Dog are found in the Pleistocene cavern-deposits of Brazil, and were originally described under the name of _Speothos_.

[Illustration: FIG. 253.—The Cape Hunting Dog (_Lycaon pictus_).]

_Otocyon._[477]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ³⁻⁴⁄₄; total 46 or 48. The molar teeth are thus in excess of any other living heterodont mammal. They have the same general characters as in _Canis_, with very pointed cusps. The lower carnassial shows little of its typical characters, having five cusps on the surface; these can, however, be identified as the inner cusp, the two greatly reduced and obliquely placed lobes of the blade, and two cusps on the talon. The skull generally resembles that of the smaller Foxes, particularly the Fennecs. The auditory bullæ are very large. The hinder edge of the mandible has a very peculiar form, owing to the great development of an expanded, compressed, and somewhat inverted subangular process. Vertebræ: C 7, D 13, L 7, S 3, C 22. Ears very large. Limbs rather long. Toes 5-4. One species, _O. megalotis_, from South Africa, rather smaller than a common Fox.

Professor Huxley looks upon this as the least differentiated or most primitive existing form of the family, regarding the presence of the four molar teeth as a survival of a condition of the dentition exhibited by the common ancestors of the existing _Canidæ_ and the existing carnivorous Marsupials. There is, however, at present no palæontological proof of this, as none of the numerous fossil forms of _Canidæ_ yet discovered have more than the normal number of molars.

_Extinct Genera._—A large number of fossil Carnivora have been described from various Tertiary deposits which are more or less closely allied to the existing _Canidæ_, although, as already mentioned, connecting the latter so closely on the one hand with the _Viverridæ_ and on the other hand with the _Ursidæ_, that it is almost, if not quite impossible to say where one family begins and the other ends. A few only of the more important of these annectant types will be mentioned here. _Temnocyon_, of the Miocene of the United States, is a true Dog, which agrees with _Icticyon_ in having a secant hind talon to the lower carnassial, but preserves a generalised character in having an entepicondylar foramen to the humerus. An extremely interesting form is _Cynodictis_, of the Middle Tertiaries of Europe and the United States, which (as now restricted by Dr. Schlosser) includes a number of species mostly not larger than Foxes. The dental formula is generally the same as in _Canis_, but (as in that genus) the last lower molar may be absent. The teeth are very like those of _Viverridæ_, the lower carnassial never being greatly elongated antero-posteriorly, and its inner cusp being situated immediately on the inner side of the hinder lobe of the blade, instead of somewhat behind it, as is the case in most Dogs. In the skull the auditory bulla is inflated, but is said to have no distinct septum; while the humerus invariably has an entepicondylar foramen. It is suggested that _Cynodictis_ is not far removed from the ancestral type of many of the Viverroids and Canoids, and may itself have been derived from the under-mentioned genus _Amphicyon_. M. Boule considers, indeed, that from the resemblance of the Pliocene _Canis megamastoides_ (p. 553) to _Cynodictis_ we ought to regard the Foxes and Jackals as the descendants of _Cynodictis_, while the Wolves have been derived directly from _Amphicyon_. The last named genus, which includes some species as large as a Bear, is found in the Upper Eocene and Lower Miocene of Europe, and is represented in the Miocene of the United States by the allied _Daphœnus_. It is characterised by the presence of three upper molars—thus bringing up the dental formula to the full Eutherian number; by the five digits on all the feet, which were plantigrade; and by the presence of a third trochanter to the femur and an entepicondylar foramen to the humerus. The teeth are essentially those of a dog, and the base of the skull is also dog-like, although it is highly probable that the auditory bulla had no trace of a septum. According, however, to Dr. Filhol[478] the minute foramina described by Professor Cope[479] in the postparietal and mastoid which occur in _Ursus_, but are said to be absent in _Canis_, are present in _Amphicyon_. So far, however, as we can see, the presence or absence of those foramina cannot be regarded as diagnostic of _Ursus_ and _Canis_, although they are generally more strongly developed in the former. _Amphicyon_ may, indeed, be considered as a very generalised Dog, with affinities to the Bears in the structure of its limbs. _Dinocyon_ is a still larger form, from the Middle Miocene of France, which, so far as its teeth are concerned, connects _Amphicyon_ with the Ursoid genus _Hyænarctus_ so closely as to render it absolutely impossible to indicate any characters of family importance by which they can be distinguished. The upper carnassial of _Dinocyon_ is unknown. For other genera, see p. 562.

_Section_ ARCTOIDEA.

[Illustration: FIG. 254.—Right half of the palatal aspect of the cranium of the Raccoon (_Procyon lotor_). Letters as in Fig. 8, p. 38. (From the _Proc. Zool. Soc._ 1869, p. 10.)]

This section includes a considerable number of forms which agree in the essential characteristics of the structures of the base of the cranium and reproductive organs, and in the absence of a cæcum to the intestinal canal. They have no Cowper’s glands, but there is a rudimentary prostate and a large cylindrical penial bone; while all the members of the group have five completely developed toes on each foot. Considerable diversity is found in the characters of the base of the skull in the various forms, but the following features are common to all. The cavity of the auditory bulla is simple, and has no trace of a dividing septum; the inferior lip of the auditory meatus (_am_, Fig. 254) is considerably prolonged; the paroccipital process (_p_) of the exoccipital is more or less triangular, directed backwards, outwards, and downwards, and standing quite apart from the bulla; the mastoid process (_m_) of the periotic is always widely separated from the paroccipital, and generally very prominent; the carotid foramen (_car_) is large, and placed on the inner margin of the bulla, usually near the middle, but occasionally more posteriorly; the condyloid foramen is distinct and exposed, and never sunk into a common opening with the foramen lacerum posticum; and the glenoid foramen is always present, and usually conspicuous. The alisphenoid canal is absent except in _Ursus_, _Melursus_, and _Ælurus_.

It has been already observed (p. 501) that the evidence of fossil forms, so far as it goes, is not in favour of the Arctoidea being a natural group; so that its retention must be regarded as a somewhat provisional measure, largely based on its convenience. The group may be divided into the three families, _Ursidæ_, _Procyonidæ_, and _Mustelidæ_.[480]

_Family_ URSIDÆ.

In existing forms the true molars ²⁄₃, with broad, flat tuberculated crowns. Typically the three anterior premolars of both jaws rudimentary and often deciduous. Fourth upper premolar (carnassial) with no third or inner root. An alisphenoid canal (except in _Æluropus_). Skull with the auditory bulla depressed, and scarcely at all inflated. Feet plantigrade. No entepicondylar foramen to the humerus. Kidneys conglomerate. Geographical distribution extensive.

_Ursus._[481]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₃; total 42. The three anterior premolars above and below one-rooted, rudimentary, and frequently wanting. Usually the first (placed close to the canine) is present, and after a considerable interval the third, which is situated close to the other teeth of the molar series. The second is very rarely present in the adult state. The fourth (upper carnassial) differs essentially from the corresponding tooth of other Carnivores in wanting the inner tubercle supported by a distinct root. Its sectorial characters are very slightly marked, and it is much smaller than the first molar. The crowns of both the true molars are longer than broad, with flattened, tuberculated, grinding surfaces. The second has a large backward prolongation or heel. The lower carnassial has a small and indistinct blade and greatly developed tubercular heel. The second molar is of about the same length, but with a broader and more flattened tubercular crown. The third is smaller. The milk-teeth are comparatively small, and shed at an early age. Skull more or less elongated. Orbits small and incomplete behind. Palate prolonged considerably behind the last molar tooth. Vertebræ: C 7, D 14, L 6, S 5, C 8-10. Body heavy. Feet broad, completely plantigrade; the five toes on each foot all well developed, and armed with long compressed and moderately curved non-retractile claws. Palms and soles naked. Tail very short. Ears moderate, erect, rounded, hairy. Fur generally long, soft, and shaggy.

[Illustration: FIG. 255.—Head of the Brown Bear (_Ursus arctos_). From Sclater, _Proc. Zool. Soc._ 1867, p. 817.]

The Bears are all animals of considerable bulk, and include among them the largest members of the order. Though the species are not numerous, they are widely spread over the earth’s surface (but absent from the Ethiopian and Australian regions, and only represented by one species in the Neotropical region), and differ much among themselves in their food and manner of life. They are mostly omnivorous or vegetable feeders, and even the Polar Bear, usually purely carnivorous or piscivorous, devours grass with avidity in summer. The various species maybe arranged in the following groups:—

_Thalassarctine Group._—Head comparatively small, molar teeth small and narrow. Soles more covered with hair than in the others. This group is represented only by the well-known Polar or White Bear (_U. maritimus_) of the Arctic regions, which is one of the few mammals which are completely white at all seasons of the year.

The typical, or _Ursine_, group includes a number of species, of which the Common Brown Bear (_U. arctos_) is the best known example. This species is an exceedingly variable one, and has a very wide range in the Palæarctic region; the Syrian form described as _U. syriacus_, as well as the Hairy-eared Bear (_U. piscator_, Fig. 255) of North-Eastern Asia, and the Snow-Bear (_U. isabellinus_) of Kashmir and Nipal, not being specifically separable. The Brown Bear hibernates in cold regions, and in the Himalaya keeps to comparatively high regions, emerging from its winter lair in March, April, or May, according to the season and elevation, to feed on the numerous bulbous plants which abound in the regions it inhabits. Both the Syrian and Himalayan varieties are generally of lighter colour and smaller size than the typical European form. Bears were at one time found in the British Isles, from which, however, they have been long since exterminated. They are still found in the Pyrenees, and are comparatively abundant in parts of Norway, Hungary, and Russia. In the Kashmir Himalaya they were very abundant in some districts a few years ago, one of the present writers having in 1874 seen no less than seven examples at one time from the top of a mountain ridge; of late years their numbers have, however, been greatly diminished. The Brown Bear, although with strong powers of smelling, is very slow of sight and hearing, and in the Himalaya it is easy to approach so near that they may be shot with a smooth-bore gun. The Grizzly Bear (_U. horribilis_) of North America is so closely allied to the Brown Bear that some writers think it should only rank as a very well-marked local variety. The Black Bears of the Himalaya (_U. torquatus_), Japan (_U. japonicus_), and North America (_U. americanus_) belong to this group. The Himalayan species ranges from Persia to Assam, and thence to China and Formosa. In the greater part of this area it is essentially a forest animal, and may be found in autumn in the forests of the Kashmir valley feeding upon chestnuts and other fruits. It is also exceedingly fond of maize, mulberries, and walnuts; and a few years ago it was no very uncommon sight to see three or even five of these bears up a single mulberry or walnut tree in Kashmir. The Spectacled Bear (_U. ornatus_) of the Peruvian Andes is another member of this group.

The _Helarctine_ group is represented only by the Malay Bear or Sun Bear (_U. malayanus_), in which the head is short and broad; the molar teeth are comparatively broad (but the length still exceeding the breadth), the tongue is very long and extensile, and the fur short and smooth. This small species inhabits the Malay Peninsula, Sumatra, Java, Borneo, Tenasserim, Arakan, Chittagong, and the Garo hills of India; it inhabits forest districts, and is an expert climber.

The earliest known occurrence of the genus is in the Lower Pliocene of the Indian Siwalik Hills; where it is represented by _U. theobaldi_, which was probably the ancestor of the existing _Melursus_. The genus is represented in the Upper Pliocene of Europe by the small _U. etruscus_; and in the Pleistocene by the existing _U. arctos_, as well as by the great extinct Cave-Bear (_U. spelæus_), distinguished by the complexity of the crowns of the molars and the total loss of the three anterior premolars in the adult condition. Remains of Bears are also found in cavern-deposits in the north of Africa. The small _U. namadicus_, from the Pleistocene of the Narbada valley, India, may have been allied to _U. malayanus_.

_Melursus._[482]—This differs from the true Bears in the first upper incisor being absent or shed at a very early age, in the very small size of the other teeth, in the very large extensile lips, the deep concavity of the palate, and other minor characters. The one species, _M. labiatus_, the well-known Sloth-Bear of India, feeds chiefly on black ants, termites, beetles, fruit, honey, etc. This species inhabits peninsular India, from near the Himalaya to Cape Comorin and Ceylon, and its remains are found in the cavern-deposits of Madras. The black hair is very long and coarse; there is a light horse-shoe-shaped mark on the chest (as in _Ursus torquatus_), and the extremity of the muzzle is of an ashy gray.

[Illustration: FIG. 256.—_Æluropus melanoleucus._ (From Milne-Edwards.)]

_Æluropus._[483]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₃, _m_ ²⁄₃; total 40. Premolars large, increasing in size from first to last, and two-rooted except the first. First upper molar with quadrate crown, broader than long; second larger than the first. Cranium with zygomatic arches and sagittal crest immensely developed, and ascending ramus of mandible very high, giving greater spaces for attachments of temporal muscle than in any other existing member of the order. Facial portion short. Bony palate not extending behind the last molar tooth. No alisphenoid canal. Feet bear-like, but soles more hairy, and perhaps less completely plantigrade. Fur long and thick. Tail very short. One extremely rare species, _A. melanoleucus_ (Fig. 256), discovered by Père David in 1869, in the most inaccessible mountains of Moupin in Eastern Tibet. Said to feed principally on roots, bamboos, and other vegetables. It is of the size of a small Brown Bear, of a white colour, with ears, spots round the eyes, shoulders and limbs black. In the large size and complex crowns of the upper premolars this genus differs very markedly from the true Bears. The fourth upper premolar (carnassial) makes no approach to the markedly sectorial type presented by the corresponding tooth of _Hyænarctus_, its structure being, on the whole, more like that of _Ælurus_.

[Illustration: FIG. 257.—Palate of _Arctotherium bonariense_, Pleistocene, South America—¼ natural size. (From the _Palæontologia Indica_.)]

_Extinct Genera._—The genus _Arctotherium_ includes some very large Bear-like animals from the Pleistocene of South America and California, in which the dentition departs less widely from a normal carnivorous type than in the true Bears. Thus the upper carnassial (Fig. 257) is relatively larger than in _Ursus_; while the crowns of the upper molars are broader and shorter. The humerus is said to have an entepicondylar foramen. _Hyænarctus_, of the Miocene and Pliocene of Europe and Southern Asia, has the crowns of the upper molars either square or triangular; the upper carnassial having three distinct lobes to the blade, while the lower carnassial is practically indistinguishable from that of the Dog-like _Dinocyon_ (p. 556). The proximal extremity of the ulna differs from that of _Ursus_ in having a long olecranon, and thereby resembles the corresponding bone of the Dogs. Indeed all the characters at present available tend to show a complete passage from the Tertiary Dog-like animals, through _Dinocyon_, _Hyænarctus_, and _Arctotherium_, to the true Bears. Most of the species of _Hyænarctus_ were of very large dimensions, but smaller forms occur in the Miocene. _Cephalogale_, of the Continental Tertiaries, is a genus represented by several species of medium size showing evident signs of affinity with _Hyænarctus_. The upper molars have subtriangular crowns, while the carnassial is short, and has two comparatively low lobes. Here also may be mentioned several other genera, apparently more or less closely allied to the present group, some of which are regarded by Dr. Schlosser as showing marked signs of affinity to the _Procyonidæ_. Among these are _Simocyon_ from the Pliocene of Europe, with _p_ ²⁄₂₋₄, _m_ ²⁄₂; and _Enhydrocyon_ of the North American Miocene, with _p_ ³⁄₃, _m_ ²⁄₂, a secant talon to the lower carnassial, and a very short skull. The Miocene _Ælurodon_ comprises several large North American forms, having a trilobed upper carnassial like that of _Hyænarctus_, and a dental formula similar to that of the latter and _Canis Prohyæna_ is founded upon a much-worn jaw of _Ælurodon_. _Hyænocyon_, of the Miocene of the United States, with _p_ ³⁄₃, _m_ ¹⁄₂, appears to be an allied form, also having a trilobed upper carnassial.

_Family_ PROCYONIDÆ.

True molars ²⁄₂, tuberculated or multicuspid; upper carnassial short and broad. Alisphenoid canal absent, except in _Ælurus_. Feet plantigrade. Tail generally annulated. In some cases an entepicondylar foramen to the humerus. Typically American, but with the outlying Oriental genus _Ælurus_.

_Ælurus._[484]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ³⁄₄, _m_ ²⁄₂; total 38. First lower premolar very minute and deciduous. Molars (Fig. 259) remarkable for their great transverse breadth and the numerous cusps of their crowns. Vertebræ: C 7, D 14, L 6, S 3, C 18. Skull (Fig. 259) high and compressed, very convex, with the facial portion short, the palate convex antero-posteriorly, and the ascending ramus of mandible extremely high. Head round. Face short and broad. Ears large, erect, pointed. Limbs stout, with large sharp semiretractile claws. Tail nearly as long as body, cylindrical, annulated, and clothed with long hairs. Fur long and thick. One existing species, _Æ. fulgens_, the Panda (Fig. 258), an animal rather larger than a Cat, found in the South-East Himalaya, at heights of from 7,000 to 12,000 feet above the sea, among rocks and trees, and chiefly feeding on fruits and other vegetable substances. Its fur is of a remarkably rich reddish-brown colour, darker below.

The genus _Ælurus_ has been made the type of a distinct family, but its relationship to the Raccoons is regarded by Mr. W. T. Blanford[485] as sufficiently close to admit of its being included in the same family. According to this zoölogist the Panda often sleeps coiled up like a Cat, with the bushy tail over its head, but at other times resting on its legs with the head tucked under the chest and between the fore legs, after a manner said to be common with the Raccoons. Although by no means strictly nocturnal, these animals sleep much during the day, and roam out in search of food in the morning and evening. The young are born in a very helpless condition, and remain for a long period concealed in the holes of trees or rocks.

[Illustration: FIG. 258.—The Panda (_Ælurus fulgens_). The dark nasal stripe shown in this figure is generally absent. (From Sclater, _Proc. Zool. Soc._ 1869, p. 408.)]

Fossil remains of a species of _Ælurus_ (_Æ. anglicus_) have been obtained from the English Pliocene Crag deposits which indicate an animal of about one and half times the size of _Æ. fulgens_. The first evidence of this fossil species was afforded by part of the mandible with the last molar in place, and the subsequent discovery of an entire first upper molar renders full confirmation of the generic determination. This distribution of _Ælurus_ is very important, as showing how its area may have once approximated to that of the ancestors of the American representatives of the family. It is probable that the genus existed in India during the Siwalik period.

The whole of the under-mentioned genera are American, and are characterised by the absence of an alisphenoid canal in the skull.

[Illustration: FIG. 259.—Lateral view of skull and right half of palate of _Ælurus fulgens_. (From Blanford, _Mammalia of British India_, p. 190.)]

_Procyon._[486]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₂; total 40. The molar teeth broad and tuberculated (Fig. 259). The upper carnassial with three cusps along the outer margin, and a very broad bicuspid inner tubercle, giving an almost quadrate form to the crown. First molar with a large tuberculated crown, rather broader than long; second considerably smaller, with transversely oblong crown. Lower carnassial with an extremely small and ill-defined blade, placed transversely in front, and a large inner cusp and hind talon. Second molar as long as the first, but narrower behind, with five obtuse cusps. Vertebræ: C 7, D 14, L 6, S 3, C 16-20. Body stout. Head broad behind, but with a pointed muzzle. Limbs plantigrade, but in walking the entire sole is not applied to the ground as it is when the animal is standing. Toes, especially of the fore foot, very free, and capable of being spread wide apart. Claws compressed, curved, pointed, and non-retractile. Tail moderately long, cylindrical, thickly covered with hair, annulated, non-prehensile. Fur long, thick, and soft. The well-known Raccoon[487] (_Procyon lotor_, Fig. 260) of North America is the type of this genus. It is a clumsy thickly-built animal about the size of a Badger, with a coat of long coarse grayish-brown hairs, short ears, and a bushy black and white ringed tail. Its range extends over the whole of the United States, and stretches on the west northwards to Alaska and southwards into Central America, where it attains its maximum size. The following notes on the habits of the Raccoon are taken from Dr. C. H. Merriam’s _Mammals of the Adirondack Region_:—

[Illustration: FIG. 260.—The Raccoon (_Procyon lotor_).]

“Raccoons are omnivorous beasts, and feed upon mice, small birds, birds’ eggs, turtles and their eggs, frogs, fish, crayfish, molluscs, insects, nuts, fruits, maize, and sometimes poultry. Excepting the bats and flying squirrels, they are the most strictly nocturnal of all our mammals, and yet I have several times seen them abroad on cloudy days. They haunt the banks of ponds and streams, and find much of their food in these places, such as crayfish, mussels, and fish, although they are unable to dive and pursue the latter under water, like the otter and mink. They are good swimmers, and do not hesitate to cross rivers that lie in their path.... The Raccoon hibernates during the severest part of the winter, retiring to its nest rather early, and appearing again in February or March, according to the earliness or lateness of the season. It makes its home high up in the hollow of some large tree, preferring a dead limb to the trunk itself. It does little in the way of constructing a nest, and from four to six young are commonly born at a time, generally early in April in this region. The young remain with the mother about a year.”

The South-American _P. cancrivorus_, the Crab-eating Raccoon, is very similar to _P. lotor_, but differs by its much shorter fur, larger size, proportionally more powerful teeth, and other minor characters. It extends over the whole of South America, as far south as the Rio Negro, and is very common in all suitable localities. Its habits are similar to those of the North-American species. Fossil remains of _Procyon_ have been described from the Pleistocene deposits of the United States.

_Bassaris._[488]—A form closely allied to _Procyon_, but of more slender and elegant proportions, with a sharper nose, longer tail, and more digitigrade feet, and with teeth otherwise like, but smaller, and more sharply denticulated. It was formerly, but erroneously, placed among the _Viverridæ_. Two species:—_B. astuta_, from the southern parts of the United States and Mexico, and _B. sumichrasti_, from Central America.

_Bassaricyon._[489]—This name has been given to a distinct modification of the Procyonine type of which at present only two examples are known, one from Costa Rica and the other from Ecuador, which, appearing to be different species, have been named _B. gabbi_ and _B. alleni_. They much resemble the Kinkajou (_Cercoleptes_) in external appearance, but the skull and teeth are more like those of _Procyon_ and _Nasua_.

_Nasua._[490]—Dentition as in _Procyon_, but the upper canines are larger and more strongly compressed, and the molars smaller. The facial portion of the skull is more elongated and narrow. Vertebræ: C 7, D 14, L 6, S 3, C 22-23. Body elongated and rather compressed. Nose prolonged into a somewhat upturned, obliquely truncated, mobile snout. Tail long, non-prehensile, tapering, annulated. These animals, commonly called Coatis or Coati-Mundis, live in small troops of eight to twenty, are chiefly arboreal, and feed on fruits, young birds, eggs, insects, etc. Recent researches have reduced the number of supposed species to two, _N. narica_ of Mexico and Central America, and _N. rufa_ of South America from Surinam to Paraguay. Remains of this genus, mostly referable to the existing species, occur in the cavern-deposits of Brazil.

_Cercoleptes._[491]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ³⁄₃, _m_ ²⁄₂; total 36. Molars with low flat crowns, very obscurely tuberculated. Skull short and rounded, with flat upper surface. Vertebræ: C 7, D 14, L 6, S 3, C 26-29. Clavicles present, but in a very rudimentary condition. Head broad and round. Ears short. Body long and musteline. Limbs short. Tail long, tapering, and prehensile. Fur short and soft. Tongue long and very extensile. But one species of this somewhat aberrant genus is known, _C. caudivolvulus_, the Kinkajou, found in the forests of the warmer parts of South and Central America. It is about the size of a Cat, of a uniform, pale, yellowish-brown colour, nocturnal and arboreal in its habits, feeding on fruit, honey, eggs, and small birds and mammals, and is of a tolerably gentle disposition and easily tamed.

_Family_ MUSTELIDÆ.

True molars ¹⁄₂ (or ¹⁄₁ in _Mellivora_[492]). No alisphenoid canal. In the upper molar the inner tubercular portion is always longer in the antero-posterior direction than the secant external portion; the degree of inflation of the auditory bulla is but slight; and the palate is generally much produced behind the last molars, as is the case with the members of the preceding family. The post-glenoid process of the cranium is generally considerably curved over the glenoid fossa, so as to hold very tightly the condyle of the mandible. The humerus may or may not have an entepicondylar foramen. Except in the Otters, the kidneys resemble those of the _Procyonidæ_ in being of simple structure.

This family is a large and widely distributed one, especially in the northern temperate regions of the earth. The different genera, which are very difficult to arrange in any natural order, are rather artificially divided, chiefly according to the characters of their feet and claws, into the Otter-like (Lutrine), Badger-like (Meline), and Weasel-like (Musteline) forms.

Subfamily =Lutrinæ=.—Feet short, rounded (except the hind feet of _Latax_). Toes webbed. Claws small, curved, blunt. Head broad and much depressed. Upper molar large and quadrate, with its inner tubercular portion much expanded antero-posteriorly (Fig. 261). Kidneys conglomerate. Habits aquatic.

_Lutra._[493]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₃, _m_ ¹⁄₂; total 36. Upper carnassial with a trenchant tricuspid blade, and a very large inner lobe, hollowed on the free surface, with a raised sharp edge, and extending along two-thirds or more of the length of the blade. True molar large, with a quadricuspidate crown, broader than long. First upper premolar very small, and in some cases absent (Fig. 261). Skull broad and depressed, contracted immediately behind the orbits. Facial portion very short; brain case large. Vertebræ: C 7, D 14-15, L 6-5, S 3, C 20-26. Body very long. Ears short and rounded. Limbs short. Feet more or less completely webbed; claws usually well developed on all the toes, although they may be rudimentary or absent. Tail long, thick at the base and tapering, rather depressed. Fur short and close. The humerus may or may not have an entepicondylar foramen. In conformity with the shape of the skull, the posterior part of the brain is expanded laterally.

[Illustration: FIG. 261.—Palate of _Lutra cinerea_. (From the _Palæontologia Indica_.)]

The Common British Otter (_L. vulgaris_), as the type of the genus, may be described somewhat fully. It has an elongated, low body, short limbs, short broad feet, with five toes on each, connected together by webs, and all with short, moderately strong, compressed, curved, pointed claws. Head rather small, broad, and flat; muzzle very broad; whiskers thick and strong; eyes small and black; ears short and rounded. Tail a little more than half the length of the body and head together, very broad and strong at the base, and gradually tapering to the end, somewhat flattened horizontally. The fur is of very fine quality, consisting of a short soft under fur of a whitish-gray colour, brown at the tips, interspersed with longer, stiffer, and thicker hairs, very shining, grayish at the base, bright rich brown at the points, especially on the upper parts and outer surface of the legs; the throat, cheeks, under parts and inner surface of the legs brownish-gray throughout. Individual Otters vary much in size; but the total length from the nose to the end of the tail averages about 3½ feet, of which the tail occupies 1 foot 3 or 4 inches. The weight of a full-sized male is from 18 to 24 lbs., that of a female about 4 lbs. less.

As the Otter lives almost exclusively on fish, it is rarely met with far from water, and usually frequents the shores of brooks, rivers, lakes, and, in some localities, the sea itself. It is a most expert swimmer and diver, easily overtaking and seizing fish in the water, but when it has captured its prey it brings it to shore to devour it. When lying upon the bank it holds the fish between its forepaws, commences at the head, and then eats gradually towards the tail, which it is said always to leave. The female produces three to five young ones at a time, in the month of March or April, and brings them up in a nest formed of grass or other herbage, usually placed in a hollow place in the bank of a river, or under the shelter of the roots of some overhanging tree. The Common Otter is found in localities suitable to its habits throughout Great Britain and Ireland, though far less abundantly than formerly, for, being very destructive to fish, and thus coming into keen competition with those who pursue the occupation of fishing either for sport or for gain, it is rarely allowed to live in peace when once its haunts are discovered. Otter-hunting with packs of hounds of a special breed, and trained for the purpose, was formerly a common pastime in the country. When hunted down and brought to bay by the dogs, the Otter is finally despatched by long spears carried for the purpose by the huntsmen.

The Common Otter ranges throughout the greater part of Europe and Asia, the Indian _L. nair_ not being distinct. A closely allied but larger species, _L. canadensis_, is extensively distributed throughout North America, where it is systematically pursued by professional trappers for the value of its fur. The Common Otter is regularly trained by the natives of some parts of Bengal to assist them in fishing, by driving the fish into the nets. In China Otters are taught to catch fish, being let into the water for the purpose attached to a long cord.

Otters are widely distributed over the earth, and, as they are much alike in size and coloration, their specific distinctions are by no means well defined.[494] Besides those mentioned above, the following may be noticed. In the Oriental region there are _L. ellioti_[495] of India, _L. sumatrana_ of the Malay countries, and _L. cinerea_ ranging over the greater part of the region. The latter species (often known as _L. leptonyx_) is of small size, with a short head, and rudimentary claws, which may be absent; it was at one time regarded as generically distinct, under the name of _Aonyx_. The upper true molar (Fig. 261) is characterised by the great development of its inner tubercular portion, and the first upper premolar is absent. In the Ethiopian region there are two species, _L. capensis_ and _L. maculicollis_. Of the Neotropical forms it will suffice to mention the small _L. felina_ and the large _L. brasiliensis_. The latter is by far the largest of the existing forms, and is characterised by the presence of a prominent flange-like ridge along each lateral margin of the tail, on which account it was referred by Dr. Gray to a distinct genus, with the name of _Pteronura sambachi_. It should be observed that all Otters have a very distinct inner cusp to the blade of the lower carnassial, but that the relative size of this cusp varies in the different species.

_Extinct Otters._—Several species of fossil Otters have been described. Thus in the Indian Siwaliks we have _L. palæindica_, which is closely allied to _L. sumatrana_, and a larger form described as _L. bathygnathus_. The Pliocene of Hessen-Darmstadt yields _L. hessica_; while _L. dubia_, of the Middle Miocene of France, is a species characterised by the small size of the inner cusp of the lower carnassial—a character in which it resembles those Tertiary forms described as _Trochictis_, which are believed to connect _Lutra_ with the _Mustelinæ_. Two very large Otters, respectively from the Indian Siwaliks and the Italian Miocene, named _L. sivalensis_ and _L. campanii_, may be regarded either as representing a very distinct _Enhydriodont_ group of _Lutra_ or as referable to a separate genus _Enhydriodon_. They are characterised by certain peculiarities in the structure of the teeth, and the second upper premolar may be absent in the Indian form. Lastly, the genus _Potamotherium_ contains a small Otter (_P. valetoni_) from the Lower Miocene of the Continent, which differs from all other known _Mustelidæ_ in having a minute second upper true molar. This species is evidently a very generalised form approximating to the _Viverridæ_ in its dental formula, and also in the characters of the teeth themselves. The brain, as recently described by Dr. Filhol, differs from that of _Lutra_ and other Mustelines in the great relative width of the anterior extremity of the hemispheres and olfactory lobes, and also in the disposition of the sulci, in both of which respects it more nearly resembles the _Viverridæ_.

_Latax._[496]—Dentition: _i_ ³⁄₂, _c_ ¹⁄₁, _p_ ³⁄₃, _m_ ¹⁄₂; total 32. Differs from all other existing Carnivora in having but two incisors on each side of the lower jaw, the one corresponding to the first (very small in the true Otters) being constantly absent. Though the molar teeth generally resemble those of _Lutra_ in their proportions, they differ very much in the exceeding roundness and massiveness of their crowns and bluntness of their cusps. Feet webbed. Fore feet small, with five subequal toes, furnished with short compressed claws; palms naked. Hind feet very large, depressed, and fin-like. The phalanges flattened as in the Seals. The fifth toe the longest and stoutest, the rest gradually diminishing in size to the first, all with moderate claws. Tail moderate, cylindrical, and obtuse; about one-fourth the length of the head and body.

The Sea-Otter (_L. lutris_, Fig. 262) is the sole representative of this genus. The entire length of the animal from nose to end of tail is about 4 feet, so that the body is considerably larger and more massive than that of the English Otter. The skin is peculiarly loose, and stretches when removed from the animal so as to give the idea of a still larger creature than it really is. The pellage is remarkable for the preponderance of the beautifully soft woolly under fur, the longer stiffer hairs being very scanty. The general colour is a deep liver brown, everywhere silvered or frosted with the hoary tips of the longer hairs. These are, however, removed when the skin is dressed for commercial purposes.

[Illustration: FIG. 262.—The Sea-Otter (_Latax lutris_). From Wolf, _Proc. Zool. Soc._ 1865, pl, vii.]

Sea-Otters are only found upon the rocky shores of certain parts of the North Pacific Ocean, especially the Aleutian Islands and Alaska, extending as far south on the American coast as Oregon; but, owing to the unremitting persecution to which they are subjected for the sake of their skins, which rank among the most valuable known to the furrier, their numbers are greatly diminishing, and, unless some restriction can be placed upon their destruction, such as that which protects the Fur-Seals of the Pribyloff Islands, the species is threatened with extermination, or, at all events, excessive scarcity. When this occurs, the occupation of five thousand of the half-civilised natives of Alaska, who are dependent upon Sea-Otter hunting as a means for obtaining their living, will be gone. The principal hunting grounds at present are the little rocky islets and reefs around the island of Saanach and the Chernobours, where they are captured by spearing, clubbing, or nets, and recently by the more destructive rifle bullet. They do not feed on fish, like the true Otters, but on clams, mussels, sea-urchins, and crabs, for the mastication of which the blunt cusps of their teeth are admirably suited. The female brings forth but a single young one at a time, apparently at any season of the year. They are excessively shy and wary, and all attempts to rear the young ones in captivity have hitherto failed.

Subfamily =Melinæ=.—Feet elongated. Toes straight. Claws non-retractile, slightly curved, subcompressed, blunt; those of the fore foot especially large. Upper molar variable. Kidneys simple. Habits mostly terrestrial and fossorial.

_Mephitis._[497]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ³⁄₃, _m_ ¹⁄₂; total 34. Upper molar larger than the carnassial, subquadrate, rather broader than long. Lower carnassial with talon less than half the length of the whole tooth. Bony palate terminating posteriorly opposite the hinder border of the last molar tooth. Facial portion of skull short and somewhat truncated in front. Vertebræ: C 7, D 16, L 6, S 2, C 21. Head small. Body elongated. Limbs moderate, subplantigrade. Ears short and rounded. Tail long, abundantly clothed with very long fine hair. Anal glands largely developed. The secretion of these glands, which can be discharged at the will of the animal, has an intolerably offensive odour, which circumstance has rendered the Skunks, as they are commonly called, proverbial. They are strictly nocturnal animals, terrestrial and burrowing, feeding chiefly on small mammals, birds, reptiles, insects, worms, roots, and berries. All the known species have a prevalent black colour, varied by white strips or spots on the upper part (Fig. 263). They generally carry the body, much arched, and the tail erect, the long loose hair of which waves like a plume over the back. There are three species, all inhabitants of the American continent, over which they have an extensive range.

[Illustration: FIG. 263.—The Common Skunk (_Mephitis mephitica_).]

The Common Skunk (_M. mephitica_, Fig. 263) is an animal of about the size of a small Cat, ranging from Hudson’s Bay to Guatemala. The following account of its habits is given by Dr. C. H. Merriam in his _Mammals of the Adirondack Region_:—

“The skunk preys upon mice, salamanders, frogs, and the eggs of birds that nest on or within reach from the ground. At times he eats carrion, and if he chances to stumble upon a hen’s nest the eggs are liable to suffer; and once in a while he acquires the evil habit of robbing the hen-roost, but as a rule skunks are not addicted to this vice. Of all our native mammals perhaps no one is so universally abused and has so many unpleasant things said about it as the innocent subject of the present biography; and yet no other species is so valuable to the farmer. Pre-eminently an insect-eater, he destroys more beetles, grasshoppers, and the like than all our other mammals together, and in addition to these he devours vast numbers of mice. He does not evince that dread of man that is so manifest in the great majority of our mammals, and when met during any of his circumambulations rarely thinks of running away. He is slow in movement and deliberate in action, and does not often hurry himself in whatever he does. His ordinary gait is a measured walk, but when pressed for time he breaks into a low shuffling gallop. It is hard to intimidate a skunk, but when once really frightened he manages to get over the ground at a very fair pace. Skunks remain active throughout the greater part of the year in this region, and hibernate only during the severest portion of the winter. They differ from most of our hibernating mammals in that the inactive period is apparently dependent solely on the temperature, while the mere amount of snow has no influence whatever upon their movements. Skunks, particularly when young, make very pretty pets, being attractive in appearance, gentle in disposition, interesting in manners, and cleanly in habits—rare qualities indeed! They are playful, sometimes mischievous, and manifest considerable affection for those who have the care of them. Their flesh is white, tender, and sweet, and is delicious eating. Skunks have large families, from six to ten young being commonly raised each season; and as a rule they all live in the same hole until the following spring.”

The two ducts leading from the anal glands open at the tips of two small conical papillæ placed in such a position that the animal can protrude them externally, and can thus guide the direction of the jet of nauseous fluid, which can be propelled by the powerful muscles surrounding the glands to a distance of from 8 to 12 feet.

The Long-tailed Skunk (_M. macrura_), from Central and Southern Mexico, has two lateral stripes, and a longer and more bushy tail than the common species. _M. putorius_, of the Southern United States and thence southwards to Yucatan and Guatemala, is of a much smaller size, with four interrupted white lateral stripes, and a skull differing considerably in form from that of the type species. It is regarded by some writers as representing a distinct genus, _Spilogale_; and has been recently divided by Dr. C. H. Merriam into several nominal species.

_Conepatus._[498]—The Skunk of tropical America (_C. mapacito_), ranging from Texas to Chili and Patagonia, differs considerably from the true Skunks, although in colour it is almost precisely similar to the common species, with which it also agrees in the variation of the relative development of the black and white. Its build is heavier than that of _Mephitis_; the snout and head are more Pig-like; and the nostrils open downwards and forwards instead of laterally on the sides of the muzzle. The skull also has many special characters, and the teeth are different in shape and, as, a rule, in number also, the first minute premolar of _Mephitis_ being almost invariably absent, so that the dental formula is _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ²⁄₃, _m_ ¹⁄₂; total 32.

Remains of _Conepatus_, which have been referred to three species, are found in the cavern-deposits of Brazil.

_Arctonyx._[499]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ¹⁄₂; total 38. Incisor line curved, the outer teeth being placed posteriorly to the others. Lower incisors proclivous. First premolars often rudimentary or absent. Upper molar much larger than the carnassial, longer in the antero-posterior direction than broad; lower carnassial with a very large, low, tuberculated talon. Cranium elongated and depressed; face long, narrow, and concave above. Bony palate extending as far backwards as the level of the glenoid fossa; palatal bones dilated; suborbital foramina very large. Vertebræ: C 7, D 16, L 4, S 4, C 20. Snout long, naked, mobile, and truncated, with large terminal nostrils, much like that of a Pig. Eyes small. Ears very small and rounded. Body compressed rather than depressed. Limbs of moderate length and digitigrade in walking. Tail moderate, tapering. A full soft under fur, with longer, bristly hairs interspersed. The best-known species is _A. collaris_, the Sand-Badger, or _Bhálu-soor_[500] (_i.e._ Bear-pig) of the natives, found in the mountains of the north-east of India and Assam. It is rather larger than the English Badger, higher on its legs, and very Pig-like in general aspect, of a light gray colour, with flesh-coloured snout and feet; and is nocturnal and omnivorous in habits. The imperfectly known _A. taxoides_ from Assam and Arakan, and perhaps China, is a much smaller species. A third form probably exists in Eastern Tibet. Professor Mivart remarks that the brain-case of _Arctonyx_ is narrower than in any other Arctoid; while the palate is relatively longer than in any other Carnivore except _Procyon_; and the metatarsus is relatively shorter than in any other member of the order.

_Mydaus._[501]—Dentition as in the last genus, but the cusps of the teeth more acutely pointed. Cranium elongated, face narrow and produced. Suborbital foramen small, and the palate, as in all the succeeding genera of this group, produced backwards about midway between the last molar tooth and the glenoid fossa. Vertebræ: C 7, D 14-15, L 6-5, S 3, C 12. Head pointed in front; snout produced, mobile, obliquely truncated, the nostrils being inferior. Limbs rather short and stout. Tail extremely short, but clothed with rather long bushy hair. Anal glands largely developed, and emitting an odour like that of the American Skunks. One species, _M. meliceps_, the Teledu, a small burrowing Badger, found in the mountains of Java at an elevation of 7000 or more feet above sea-level.

_Meles._[502]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ¹⁄₂; total 38. The first premolar in both jaws extremely minute and often deciduous. Upper molar very much larger than the carnassial, subquadrate, as broad as long. Lower carnassial with a broad, low, tuberculated talon, more than half the length of the whole tooth. The post-glenoid processes of the skull are so strongly developed, and the glenoid fossa is so deep, that the condyle of the lower jaw is firmly held in its place even after all the surrounding soft parts are removed. Vertebræ: C 7, D 15, L 5, S 3, C 18. Muzzle pointed. Ears very short. Body stout, broad. Limbs short, strong, subplantigrade. Tail short. The best-known species is the common Badger (_M. taxus_) of Europe and Northern Asia, still found in many parts of England, where it lives in woods, is nocturnal, burrowing, and very omnivorous, feeding on mice, reptiles, insects, fruit, acorns, and roots. Other nearly allied species, _M. leucurus_ and _M. chinensis_, are found in continental Asia, _M. canescens_ in Persia, and _M. anakuma_ in Japan.

The appearance of the common Badger is too well known to need description, but, it may be mentioned that a full-grown individual stands about a foot in height at the shoulder, and measures from 2½ to 3 feet in length. The young are born in a naked and blind condition, usually in litters of three or four. It appears that the usual period of gestation is about eleven and a half months, but instances are recorded where the period has been protracted to upwards of fifteen months.

Fossil remains of the common Badger are found in the Pleistocene deposits of Europe, while extinct species have been described from the Lower Pliocene beds of Maragha, in Persia.

_Taxidea._[503]—Dental formula as in _Meles_, except that the rudimentary anterior premolar appears to be always wanting in the upper jaw. The upper carnassial much larger in proportion to the other teeth. Upper molar about the same size as the carnassial, triangular, with the apex turned backwards. Talon of lower carnassial less than half the length of the tooth. Skull very wide in the occipital region; the lambdoidal crest very greatly developed, and the sagittal but slightly, contrary to what obtains in _Meles_. Vertebræ: C 7, D 15, L 5, S 3, C 16. Body very stoutly built and depressed. Tail short. The animals of this genus are peculiar to North America, where they represent the Badgers of the Old World, resembling them much in appearance and habits. _T. americana_ is the common American Badger of the United States; _T. berlandieri_, the Mexican Badger, is perhaps only a local variety.

_Mellivora._[504]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ³⁄₃, _m_ ¹⁄₁; total 32. Upper carnassial large, with its inner tubercle quite at the anterior end of the blade, as in the following genera; molar much smaller and transversely extended, having a very small outer and a larger rounded inner lobe. Talon of lower carnassial very small, scarcely one-fourth of the whole length of the tooth, and with but one cusp; lower tubercular molar absent. Vertebræ: C 7, D 14, L 4, S 4, C 15. Body stout, depressed. Limbs short, strong. Head depressed, nose rather pointed. External ears rudimentary. Tail short. The animals of this genus are commonly called Ratels. _M. indica_ from India, and, _M. ratel_ (Fig. 264) from South and West Africa, have nearly the same general appearance and size, being rather larger than a common Badger. Their coloration is peculiar, all the upper surface of the body, head, and tail being ashy gray, while the lower parts, separated by a distinct longitudinal boundary line, are black. The two species may be distinguished by the circumstance that the African one has a distinct white line round the body at the junction of the gray of the upper side with the black of the lower, while in the Indian form this line is absent; the teeth also of the former are, on the whole, larger, rounder, and heavier than those of the latter. In spite of these differences the two are, however, so nearly allied that they might almost be considered as local races of a single widely spread species.

[Illustration: FIG. 264.—The African Ratel (_Mellivora ratel_).]

The following account of the Indian species is extracted from Dr. Jerdon’s _Mammals of India_: “The Indian badger is found throughout the whole of India, from the extreme south to the foot of the Himalayas, chiefly in hilly districts, where it has greater facilities for constructing the holes and dens in which it lives; but also in the north of India in alluvial plains, where the banks of large rivers afford equally suitable localities wherein to make its lair. It is stated to live usually in pairs, and to eat rats, birds, frogs, white ants, and various insects, and in the north of India it is accused of digging out dead bodies, and is popularly known as the grave-digger. It doubtless also, like its Cape congener, occasionally partakes of honey. It is often very destructive to poultry, and I have known of several having been trapped and killed whilst committing such depredations in Central India and in the northern Circars. In confinement the Indian badger is quiet and will partake of vegetable food, fruits, rice, etc.”

A fossil species of _Millivora_, apparently closely allied to the existing forms, occurs in the Pliocene Siwaliks of India. The same deposits have also yielded remains of an extinct genus described as _Mellivorodon_.

[Illustration: FIG. 265.—_Helictis personata._ (From Blanford, _Mammalia of British India_, p. 175.)]

_Helictis._[505]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ¹⁄₂; total 38. Upper carnassial with a large bicuspid inner tubercle; upper molar smaller, wider transversely than in the antero-posterior direction. Lower carnassial with talon about one-third the length of the tooth. Skull elongated, rather narrow and depressed. Facial portion especially narrow. Infraorbital foramen very large. Head rather small and produced in front, with an elongated, obliquely truncated, naked snout. Ears small. Body elongated. Limbs short. Tail short or moderate, bushy. Several species are described (_H. orientalis_, _personata_ [Fig. 265], _moschata_, _subaurantiaca_), all from Eastern Asia; they are all small animals compared with the other members of the subfamily, climbing trees with agility and living much on fruit and berries as well as on small mammals and birds. The two first named species occur in British India, _H. orientalis_ also ranging into Java; the Chinese _H. subaurantiaca_ is brilliantly coloured in the region of the throat.[506]

[Illustration: FIG. 266.—Left lateral and superior aspect of the brain of _Helictis subaurantiaca_. (From Garrod, _Proc. Zool. Soc._ 1879, p. 307.)]

The brain of _Helictis_, represented in the accompanying figure, shows the general type of cerebral structure characteristic of the _Mustelidæ_. The brain of this genus differs, however, from that of every other Carnivore in that the hippocampal gyrus rises to the surface on either side of the great longitudinal fissure, in consequence of which there is no crucial fissure, and the so-called “Ursine lozenge,” so characteristic of the Arctoidea, is incomplete behind. The superior gyrus, as in _Ictonyx_ and _Mustela_, ceases at the superior posterior angle of the hemisphere.

_Ictonyx._[507]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ³⁄₃, _m_ ¹⁄₂; total 34. In general characters the teeth much resemble those of the Polecats (_Mustela_), being more delicately cut and sharply cusped than in most of the foregoing. Upper molar smaller than the carnassial, narrow from before backwards. Lower carnassial with a small narrow talon and distinct inner cusp. General form of body Musteline. Limbs short. Fore feet large and broad, with five stout, nearly straight, blunt, and non-retractile claws, of which the first and fifth are considerably shorter than the others. Tail moderate, with longer hairs towards the end, giving it a bushy appearance. Hairs generally long and loose. The best known species of this genus, _I. zorilla_, the Cape Polecat, was placed by Cuvier in the genus _Mustela_, and by Lichtenstein in _Mephitis_; and in many characters it forms a transition between these genera. It is about the size of an English Polecat, but conspicuous by its coloration, having broad, longitudinal bands of dark brown, alternating with white. Its odour is said to be as offensive as that of the American Skunks. From the Cape of Good Hope it ranges as far north as Senegal. Another species, _I. frenata_, from Sennaar and Egypt, has been described.

Subfamily =Mustelinæ=.—Toes short, partially webbed; claws short, compressed, acute, curved, often semiretractile. Upper molar of moderate size, wide transversely. Kidneys simple. Terrestrial and arboreal in habits.

_Galictis._[508]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ³⁄₃, _m_ ¹⁄₂; total 34. Molars small but stout. Upper carnassial with the inner tubercle near the middle of the inner border of the tooth. Lower carnassial with talon small, and inner cusp small or absent. Body long. Limbs short; claws non-retractile. Palms and soles naked. Head broad and depressed. Tail of moderate length. The best-known species are _G. vittata_, the Grison (genus _Grisonia_, Gray), and _G. barbara_, the Tayra (genus _Galera_, Gray), both South American; _G. allamandi_ is an intermediate form.

Remains of _Galictis_ occur in the Pleistocene cavern-deposits of Brazil, and also in the Pleistocene of North America.

_Mustela._[509]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ³⁻⁴⁄₃₋₄, _m_ ¹⁄₂; total 34 or 38. Upper carnassial with inner tubercle close to the anterior edge of the tooth. Molar nearly as large as carnassial. Lower carnassial with small or no inner cusp. Vertebræ: C 7, D 14, L 6, S 3, C 18-23. Body long and slender. Limbs short, digitigrade. Feet rounded; toes short, with compressed, acute, semiretractile claws. Tail moderate or long, more or less bushy.

The genus _Mustela_, as restricted by Cuvier (_Règne Animal_, 1817), contains a very natural assemblage of animals commonly called Martens, Sables, Polecats, Stoats, Ermines, and Weasels, all closely allied in structure and habits. A structural division, however, occurs between the two first-named and all the others, especially shown in the presence of an additional small premolar tooth on each side of the jaw; and, availing himself of this and some other minor characters, Cuvier divided the genus into two subgenera, for the first of which he retained the name of _Mustela_, and to the second assigned that of _Putorius_. Three years later Nilsson (_Skand. Fauna_, 1820) definitely constituted the two groups into genera, applying to the first the name of _Martes_, by which the animals composing it had been generally designated by the Latin writing zoologists of the preceding century, and keeping _Mustela_ for the more typical Weasels and their immediate allies. Later zoologists have been divided between the nomenclature of Cuvier, which has the priority, and that of Nilsson, which on other grounds is preferable. Those who adopt the latter affirm that Cuvier’s names, being only used by him in a subgeneric sense, and not binominally, need not be applied generically, but this is contrary to the practice usually followed in such cases; and therefore, if the original genus be divided, the name _Mustela_ should be retained for the Martens, and _Putorius_ for the Polecats and Weasels. Here, however, the genus will be employed in its wider sense, and divided into two groups.

The typical group of the Martens[510] presents the following distinctive features. Body long, slender, and very flexible, though less so than in the true Weasels. Head somewhat triangular; muzzle pointed, the nose extending a little beyond the lips; eyes large and prominent; ears conspicuous, broad, somewhat triangular, rounded at the ends, furred outside and in. Limbs short; feet rounded; toes short, five on each foot, all with short, compressed, curved, sharp-pointed claws; soles densely furred between the naked pads. Tail moderately long, more or less bushy. Outer fur long, strong, and glossy; a very abundant soft under fur. Skull elongated and depressed. Facial portion moderate and rather compressed. Zygomata arched and wide, but slender. Postorbital processes small. Auditory bullæ large, but not very globose. Mandible with a strong triangular vertical coronoid process and a well-developed angular process. Premolars ⁴⁄₄. Upper incisors in a straight transverse line, rather long and compressed; first and second subequal, third considerably larger. Lower incisors very small, especially the first, and crowded together, the second placed rather behind the others. Canines long and sharp-pointed. Upper premolars: first very small, with simple crown and one root; second and third nearly equal in size and two-rooted, with simple compressed sharp-pointed crowns, with very slightly developed accessory cusps; fourth (the carnassial) with blade consisting chiefly of the central and posterior lobes, the anterior being rudimentary, inner tubercle small and confined to the anterior part of the tooth. True molar tubercular, about twice as wide transversely as in the antero-posterior direction, having an outer, more elevated, but smaller portion, bearing three blunt tubercles; to the inner side of this the crown is contracted, and its surface deeply hollowed; it then expands again into a broad low lobe, with the central part elevated, and a raised, even, semicircular, slightly crenated internal border. Lower premolars: first very small, simple, and one-rooted; second, third, and fourth increasing slightly in size, with high compressed pointed crowns and posterior accessory cusps, best marked in the third. First molar (carnassial) with well-marked bilobed blade, talon scarcely more than one-third of the length of the tooth, and a very small inner cusp. Second molar small, single-rooted, with a low, flattened, subcircular or oval tubercular crown.

In geographical distribution the Martens are limited to the northern hemisphere, ranging throughout the greater part of the temperate regions of both Old and New Worlds, as far north as conditions of existence suited to their habits are met with, and southwards in America to 35° N. lat., while in Asia one species is met with as far in this direction as the island of Java.

The various species appear to be very similar in their habits. They live in woods and rocky places, and are thoroughly arboreal, spending most of their time in trees, although descending to the ground in quest of prey. They climb with great facility, and are agile and graceful in their movements. Some species are said occasionally to resort to berries and other fruit for food, but as a rule they are strictly carnivorous, feeding chiefly on birds and their eggs, small mammals, as squirrels, hares, rabbits, and moles, but chiefly mice of various kinds, of which they destroy great numbers, and occasionally snakes, lizards, and frogs. In proportion to their size they are among the most bloodthirsty of animals, though less so than the true Weasels. The female usually makes her nest of moss, dried leaves, and grass in the hollow of a tree, but sometimes in a hole among rocks or ruined buildings, and produces several young at a birth, usually from four to six. Though wild and untameable to a great degree if captured when fully grown, when taken young they are very docile, and have frequently been made pets of, not having the strong unpleasant odour of the smaller _Mustelidæ_. The common European Marten appears to have been partially domesticated by the Greeks and Romans, and to have been used to keep houses clear from rats and mice before cats were introduced.[511] In the same way, according to Hodgson, the Yellow-bellied Weasel (_M. cathia_) “is exceedingly prized by the Nipalese for its service in ridding houses of rats. It is easily tamed; and such is the dread of it common to all Murine animals that not one will approach a house where it is domiciled.” It is, however, to the great value attached to the pelts of these animals that their importance to man is chiefly due. Though all yield fur of serviceable quality, the commercial value varies immensely, not only according to the particular species from which it is obtained, but according to individual variation, depending upon age, sex, season, and other trifling circumstances. The skins from northern regions are more full and of a finer colour and gloss than those from more temperate climates, as are those of animals killed in winter compared with the same individuals in the summer season. The caprices of fashion have, moreover, set wholly factitious values upon slight shades of colour, recognised and named by experienced furriers, but not indicating any specific or other distinctions of which zoologists have any cognisance. Enormous numbers of animals are annually caught, chiefly in traps, to supply the demand of the fur trade, Siberia and North America being the principal localities from which they are obtained.

With the exception of the Pekan (_M. pennanti_) all the Martens are so much alike in size, general colouring, and cranial and dental characters that the discrimination of the species, and assignment of the proper geographical distribution to each, has been a subject which has sorely perplexed the ingenuity and patience of zoologists. The following description by Dr. Elliott Coues of the external characters of the American Pine Marten (_M. americana_) will apply almost equally well to most of the others: “It is almost impossible to describe the colour of the Pine Marten, except in general terms, without going into the details of the endless diversities occasioned by age, sex, season, or other incidents. The animal is ‘brown,’ of various shades from orange or tawny to quite blackish; the tail and feet are ordinarily the darkest, the head lightest, often quite whitish; the ears are usually rimmed with whitish; on the throat there is usually a large tawny-yellowish or orange-brown patch, from the chin to the fore legs, sometimes entire, sometimes broken into a number of smaller, irregular blotches, sometimes wanting, sometimes prolonged on the whole under surface, when the animal is bicolor like a Stoat in summer. The general ‘brown’ has a grayish cast, as far as the under fur is concerned, and is overlaid with rich lustrous blackish-brown in places where the long bristly hairs prevail. The claws are whitish; the naked nose pad and whiskers are black. The tail occasionally shows interspersed white hairs, or a white tip.”

The species generally recognised as distinct are the following, the first five belonging to the Old and the last two to the New World:—

_M. foina_, the Beech Marten, Stone Marten, or White-breasted Marten.—Distinguished from the following by the greater breadth of the skull, and some minute but constant dental characters, by the dull grayish-brown colour of the fur of the upper parts, and the pure white of the throat and breast. It inhabits the greater part of the continent of Europe, but is more southern than the next in its distribution, not being found in Sweden or Norway, nor, according to the investigations of Mr. Alston, in the British Isles, although included in their fauna by all earlier writers; to the eastward it ranges into Afghanistan and the Himalaya.

[Illustration: FIG. 267.—The Pine Marten (_Mustela martes_).]

_M. martes_, the Pine Marten (Fig. 267).—Outer fur rich dark brown; under fur reddish-gray, with clear yellow tips; breast spot usually yellow, varying from bright orange to pale cream-colour or yellowish-white. Length of head and body 16 to 18 inches; of tail (including the hair) 9 to 12 inches. This species is extensively distributed throughout northern Europe and Asia, and was formerly common in most parts of Great Britain and Ireland. Though commonly called “Pine Marten,” it does not appear to have any special preference for coniferous trees, except that, inasmuch as they constitute the greater proportion of the forests of the countries which it inhabits, it is more often met with in them than in any other. With regard to its recent occurrence in the British Isles, Mr. Alston writes in the _Proc. Zool. Soc._ 1879:—

“Although greatly reduced in numbers by persecution, it still maintains its ground in the wilder districts of Scotland, the north of England, Wales, and Ireland; and occasionally specimens are killed in counties where the species was thought to have been long extinct. In Scotland it is still found, though comparatively rarely, in the Lews and in most of the Highland mainland counties, being perhaps most abundant in Sutherland and Ross-shire, especially in the deer forests. In the Lowlands a Marten is now a very great rarity; but a fine example was killed in Ayrshire in the winter of 1875-76. In the north of England Mr. W. A. Durnford says the species is still plentiful in the wilder parts of Cumberland, Westmoreland, and Lancashire, and in Lincolnshire several have been recorded, the latest killed in 1865, by Mr. Cordeaux. In Norfolk one was shot last year; and I have myself examined a fine example which was shot in Hertfordshire, within 20 miles of London, in December 1872. In Dorsetshire the last is said to have been killed in 1804; but a specimen occurred in Hampshire about forty years ago, and another in Surrey in 1847. In Ireland the following counties were enumerated by Thompson as habitats of this species: Donegal, Londonderry, Antrim, Down, Armagh, Fermanagh, Longford, Galway, Tipperary, Cork, and Kerry. The _Cat-crann_ is probably now a rarer animal in Ireland than it was when Thompson wrote; but it still exists in various districts, especially in County Kerry, whence the society has received several living examples; and Professor A. Leith Adams states that it has been seen of late years even in county Dublin.”

_M. zibellina_, the Sable (German, _Zobel_ and _Zebel_; Swedish, _sabel_; Russian, _sobel_, a word probably of Turanian origin).—Closely resembling the last, if indeed differing from it except in the quality of the fur, which is the most highly valued of that of all the group. Found chiefly in Eastern Siberia.

_M. flavigula_, the Indian Marten.—Inhabits the southern slopes of the Himalaya, the Nilgiri Hills, the interior of Ceylon, the Malay Peninsula, and Java. The coloration of this species is very striking, the upper parts being blackish-brown, and the throat and breast yellow or orange, in the bright coloured variety. It differs from the other species in having the soles of the feet more or less naked.

_M. melampus._—Japan.

_M. americana_, the North-American Sable or Marten.—A species so closely allied to the European Pine Marten and Asiatic Sable that it is very difficult to assign constant distinguishing characters between them. The importance of the fur of this animal as an article of commerce may be judged of from the fact that 15,000 skins were sold in one year by the Hudson’s Bay Company as long ago as 1743, and the more recent annual imports into Great Britain have exceeded 100,000. It is ordinarily caught in wooden traps of very simple construction, being little enclosures of stakes or brush in which the bait is placed upon a trigger, with a short upright stick supporting a log of wood, which falls upon its victim on the slightest disturbance. A line of such traps, several to a mile, often extends many miles. The bait is any kind of meat, a mouse, squirrel, piece of fish, or bird’s head. It is principally trapped during the colder months, from October to April, when the fur is in good condition, as it is nearly valueless during the shedding in summer. Dr. Coues tells us that, notwithstanding the persistent and uninterrupted destruction to which the American Sable is subjected, it does not appear to diminish materially in numbers in unsettled parts of the country. It holds its own partly in consequence of its shyness, which keeps it away from the abodes of men, and partly because it is so prolific, bringing forth six to eight young at a litter. Its home is sometimes a den under ground or beneath rocks, but oftener the hollow of a tree, and it is said frequently to take forcible possession of a squirrel’s nest, driving off or devouring the rightful proprietor.

_M. pennanti_, the Pekan or Pennant’s Marten, also called Fisher Marten, though there appears to be nothing in its habits to justify the appellation.—This is the largest species of the group, the head and body measuring from 24 to 30 inches, and the tail 14 to 18 inches. It is also more robust in form than the others, its general aspect being more that of a Fox than a Weasel; in fact, its usual name among the American hunters is “Black Fox.” Its general colour is blackish, lighter by mixture of brown or gray on the head and upper fore part of the body, with no light patch on the throat, and unlike the other Martens generally darker below than above. It was generally distributed in wooded districts throughout the greater part of North America, as far north as Great Slave Lake, 63° N. lat., and Alaska, and extending south to the parallel of 35°; but at the present time it is almost exterminated in the settled parts of the United States east of the Mississippi.

Fossil remains of a Marten from the Pliocene Siwaliks of India indicate a species which cannot be distinguished from those now inhabiting the same region; while remains of _M. martes_ occur in European cavern-deposits, and in the fens of Cambridgeshire.

With the _Putoriine_ group (genus _Putorius_) we come to those smaller forms distinguished by having only three premolars in each jaw, by the absence of an inner cusp to the blade of the lower carnassial, as well as by certain external characters. This group contains a few species known as Minks, differing from the rest by slight structural modifications, and especially by their semiaquatic habits. They are distinguished from the Polecats, Stoats, and Weasels, which constitute the remainder of the group, by the facial part of the skull being narrower and more approaching in form that of the Martens, by the premolar teeth (especially the anterior one in the upper jaw) being larger, by the toes being partially webbed, and by the absence of hair in the intervals between the naked pads of the soles of the feet. The two best-known species, so much alike in size, form, colour, and habits that although they are widely separated geographically some zoologists question their specific distinction, are _M. lutreola_, the _Nörz_ or _Sumpfotter_ (Marsh-Otter) of Eastern Europe, and _M. vison_, the Mink of North America. The former inhabits Finland, Poland, and the greater part of Russia, though not found east of the Ural Mountains. Formerly it extended westward into Central Germany, but is now very rare, if not extinct, in that country. The latter is found in places which suit its habits throughout the whole of North America. Another form, _M. sibirica_, from Eastern Asia, of which much less is known, appears to connect the true Minks with the Polecats.

For the following description, chiefly taken from the American form (though almost equally applicable to that of Europe), we are mainly indebted to Dr. Coues’s _Fur-bearing Animals of North America_. In size it much resembles the English Polecat,—the length of the head and body being usually from 15 to 18 inches, that of the tail to the end of the hair about 9 inches. The female is considerably smaller than the male. The tail is bushy, but tapering at the end. The ears are small, low, rounded, and scarcely project beyond the adjacent fur. The pellage consists of a dense, soft, matted under fur, mixed with long, stiff, lustrous hairs on all parts of the body and tail. The gloss is greatest on the upper parts; on the tail the bristly hairs predominate. Northern specimens have the finest and most glistening pellage; in those from southern regions there is less difference between the under and over fur, and the whole pellage is coarser and harsher. In colour different specimens present a considerable range of variation, but the animal is ordinarily of a rich dark brown, scarcely or not paler below than on the general upper parts; but the back is usually the darkest, and the tail is nearly black. The under jaw, from the chin about as far back as the angle of the mouth, is generally white. In the European Mink the upper lip is also white, but as this occasionally occurs in American specimens it fails as an absolutely distinguishing character. Besides the white on the chin, there are often other irregular white patches on the under parts of the body. In very rare instances the tail is tipped with white. The fur, like that of most of the animals of the group to which it belongs, is an important article of commerce.

The principal characteristic of the Mink in comparison with its congeners is its amphibious mode of life. It is to the water what the other Weasels are to the land, or Martens to the trees, being as essentially aquatic in its habits as the Otter, Beaver, or Musk-Rat, and spending perhaps more of its time in the water than it does on land. It swims with most of the body submerged, and dives with perfect ease, remaining long without coming to the surface to breathe. It makes its nest in burrows in the banks of streams, breeding once a year about the month of April, and producing five or six young at a birth. Its food consists of frogs, fish, freshwater molluscs and crustaceans, as well as mice, rats, musk-rats, rabbits, and small birds. In common with the other animals of the genus, it has a very peculiar and disagreeable effluvium, which, according to Coues, is more powerful, penetrating, and lasting than that of any animal of the country except the Skunk. It also possesses the courage, ferocity, and tenacity of life of its allies. When taken young, however, it can be readily tamed, and lately Minks have been extensively bred in captivity in America, both for the sake of their fur and for the purpose of using them in like manner as Ferrets in England, to clear buildings of rats.

[Illustration: FIG. 268.—The Common Polecat (_Mustela putorius_).]

The Polecats include four species confined to the northern hemisphere, the best known of which is the Common Polecat (_M. putorius_, Fig. 268). The Ferret is a domesticated variety of this species, generally of a yellowish-white colour; whereas the Wild Polecat is dark brown above and black beneath, the face being variegated with dark brown and white markings.

The skull is rough, strongly ridged, and of a far more powerful type than that of the Stoats, Weasels, or Martens; being in the female much smaller and lighter than in the male. The fur, which is long, coarse, and of comparatively small value, changes its colour very little, if at all, at the different seasons of the year.

The distribution and habits of this species have been described by Blasius, the following being an abstract of his account. The Polecat ranges over the greater part of Europe, reaching northwards into Southern Sweden, and in Russia to the region of the White Sea. It does not occur in the extreme South, but is common everywhere throughout Central Europe. In the Alps it ranges far above the tree-line during the summer, but retreats in winter to lower ground. In fine weather it lives either in the open air, in holes, fox-earths, rabbit-warrens, under rocks, or in wood-stacks, while in winter it seeks the protection of deserted buildings. During the day it sleeps in its hiding-place, sallying forth at night to plunder dovecots and hen-houses. It climbs but little, and shows far less activity than the Marten. It feeds ordinarily on small mammals, such as rabbits, hamsters, rats, and mice, on such birds as it can catch, especially poultry and pigeons, and also on snakes, lizards, frogs, fish, and eggs. Its prey is devoured only in its lair, but, even though it can carry away but a single victim, it commonly kills everything that comes in its way, often destroying all the inhabitants of a hen-house in order to gratify its passion for slaughter. The pairing time is towards the end of the winter, and the young, from three to eight in number, are born in April or May, after a period of gestation of about two months. The young, if taken early, may be easily trained, like Ferrets, for rabbit catching. The Polecat is very tenacious of life, and will bear many severe wounds before succumbing; it is also said to receive with impunity the bite of the adder. Its fetid smell has become proverbial.

Four other species of Polecats are known, viz.—The Siberian Polecat (_M. eversmanni_) of Western and Northern Asia is nearly allied to the European species, but the head and back are almost white, and the skull is stouter and more constricted behind the orbits. The Tibetan _M. larvata_ is distinguished from the last by the presence of a process connecting the pterygoid with the auditory bulla, and by a difference in the shape of the upper molar. The American Polecat (_M. nigripes_), inhabiting the central plateau of the United States, and extending southwards into Texas, is another closely allied species, although some zoologists have made it the type of the genus _Cynomyonax_. Finally, the Mottled Polecat (_M. sarmatica_) is a species sparsely distributed in Eastern Europe and parts of Western Asia, but common in Southern Afghanistan. Its skull, although smaller, resembles that of the common species; but the coloration is very different, all the upper parts being mottled with large irregular reddish spots on a white ground, and the under side, limbs, and tail deep shining black. The tail is long.

The Common Polecat occurs in a fossil condition in the cave-deposits of Europe.

[Illustration: FIG. 269.—The Common Weasel (_Mustela vulgaris_).]

The remaining members of the genus comprise the true Weasels and Stoats, which are of almost cosmopolitan distribution. In the Common Weasel (_M. vulgaris_, Fig. 269) the upper parts, outside of limbs and tail, are a uniform reddish-brown, the under parts pure white. In very cold regions, both in Europe and America, it turns completely white in winter, but less regularly and at a lower temperature than the Stoat, from which it is easily distinguished by its smaller size, and by its wanting the black end of the tail. The length of the head and body of the male is usually about 8 inches, that of the tail 2½ inches; the female is smaller.

This species is pretty generally distributed throughout Europe, Northern and Central Asia, British North America, and the northern portions of the United States. It possesses in a full degree all the active, courageous, and bloodthirsty disposition of the rest of the genus, but its diminutive size prevents it attacking and destroying any but the smaller mammals and birds. Mice, rats, voles, moles, and frogs constitute its principal food. It is generally found on or near the surface of the ground, but it can not only pursue its prey through very small holes and crevices of rocks and under dense tangled herbage, but follow it up the stems and branches of trees, or even into the water, swimming with perfect ease. It constructs a nest of dried leaves and herbage, placed in a hole in the ground or a bank or hollow tree, in which it brings up its litter of four to six (usually five) young ones. The mother will defend her young with the utmost desperation against any assailant, having been often known to sacrifice her own life rather than desert them.

The Stoat or Ermine (_M. erminea_) has nearly the same distribution as the Weasel, but in Asia it is said to extend into parts of the Kashmir Himalaya. Its size, as already mentioned, considerably exceeds that of the Weasel; and its most distinctive feature is the black tip at the end of the tail, which remains when the rest of the pellage turns white. The white winter skins from the northern regions of its habitat, where the fur is thick and close, form the well-known and valuable ermine of commerce. Remains of the Stoat are found in the Pleistocene cavern-deposits of Europe. The other species of Weasels are very numerous and widely distributed.

_Extinct Mustelines._—A number of European Miocene Carnivores may be referred to the genus _Mustela_ in its wider sense, and serve to confirm the propriety of this use of the term. Thus _M. sectoria_ is a species of somewhat larger size than the Stoat, with _p_ ⁴⁄₄, while in _M. angustifrons_ the number of premolars is ³⁄₄, and in _M. mustelina_ ⁴⁄₃; the latter species agreeing very closely in size with the Stoat. The extinct _Plesictis_, in which there are _p_ ⁴⁄₄ and the lower carnassial has a large inner cusp, is distinguished from _Mustela_ by the circumstance that the temporal ridges of the skull never unite to form a sagittal crest. Moreover, the inner tubercular portion of the upper molar (as in some of the Miocene species of _Mustela_) is shorter in an antero-posterior direction than the secant outer moiety; and the auditory bulla is more inflated than in _Mustela_, although it has no septum. Both these features indicate a decided approximation to the Viverroid genus _Stenoplesiotis_ (p. 539); and since there are no well-marked characters of family value by which these two genera can be distinguished the available evidence points to a transition from the Viverroid to the Musteloid type. _Mustela larteti_, of the Middle Miocene of France, should perhaps be referred to _Ictonyx_.

_Pœcilogale._[512]—This genus has been made for the reception of the South African _Mustela albinucha_, in which the coloration is similar to that of _Ictonyx_, but the number of cheek-teeth is usually reduced to _p_ ²⁄₂, _m_ ¹⁄₁, although there may be a second lower molar. The auditory bulla is quite flat.

_Lyncodon._[513]—This name has been proposed for a small Musteline from Patagonia, with _p_ ²⁄₂, _m_ ¹⁄₁, which Mr. O. Thomas suggests may be nothing more than an aberrant southern form of _Mustela_ (_Putorius_) _brasiliensis_. The auditory bulla is more inflated than in the typical Weasels. This animal is somewhat larger than the Stoat.

[Illustration: FIG. 270.—The Wolverene (_Gulo luscus_).]

_Gulo._[514]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ¹⁄₂; total 38. Crowns of the teeth very stout. Upper molar very much smaller than the carnassial. Lower carnassial large, with very small talon and no inner cusp. Third upper incisor unusually large, almost like a canine. The dentition, though really but a modification of that of the Weasels, presents a great general resemblance to that of the Hyæna. Palate prolonged somewhat behind the last molar. Humerus with an entepicondylar foramen. Vertebræ: C 7, D 15, L 5, S 3, C 15. Body and limbs stoutly made. Feet large and powerful, subplantigrade, with large, compressed, much curved, and sharp-pointed claws. Soles of the feet (except the pads of the toes) covered with thick bristly hairs. Ears very small, nearly concealed by the fur. Eyes small. Tail short, thick, and bushy. Fur full, long, and rather coarse. The one species, the Wolverene or Glutton (_G. luscus_, Fig. 270), an inhabitant of the forest regions of Northern Europe, Asia, and America, much resembles a small Bear in appearance. It is a very powerful animal for its size, climbs trees, and lives on grouse, squirrels, hares, foxes, beavers, reindeer, and is said to attack even horses and cows. The Wolverene has a curious habit of stealing and secreting articles of which it can make no possible use, as is exemplified in the following instance related by Dr. Coues: “A hunter and his family, having left their lodge unguarded during their absence, on their return found it completely gutted—the walls were there, but nothing else. Blankets, guns, kettles, axes, cans, knives, and all the other paraphernalia of a trapper’s tent had vanished, and the tracks left by the beast showed who had been the thief. The family set to work, and, by carefully following up all his paths, recovered, with some trifling exceptions, the whole of the lost property.” The pairing season occurs in March, and the female, secure in her burrow, produces her young, four or five at a birth, in June or July. In defence of these she is exceedingly bold, and the Indians, according to Coues, “have been heard to say that they would sooner encounter a she-bear with her cubs than a carcajou (the Indian name of the glutton) under the same circumstances.”

Fossil remains of the Wolverene are found in cavern and other Pleistocene deposits in various parts of Europe.

_Suborder_ PINNIPEDIA.

The Eared-Seals, Walruses, and Seals differ from the rest of the Carnivora mainly in the structure of their limbs, which are modified for aquatic progression,—the two proximal segments being very short and partially enveloped in the general integument of the body; while the third segment, especially in the hinder extremities, is elongated, expanded, and webbed. There are always five well-developed digits on each limb. In the hind limb the two marginal digits (first and fifth) are stouter and generally longer than the others. The teeth also differ from those of the more typical Carnivora. The incisors are always fewer than ³⁄₃. The cheek series consists generally of four premolars and one molar of very uniform characters, with never more than two roots, and with conical, more or less compressed, pointed crowns, which may have accessory cusps, placed before or behind the principal one, but are never broad and tuberculated; and there is no differentiated carnassial tooth. The milk-teeth are very small and simple, and are shed or absorbed at a very early age, usually either before or within a few days after birth. The brain is relatively large; the cerebral hemispheres being broad in proportion to their length, with numerous and complex convolutions. There is a very short cæcum. The kidneys are divided into numerous distinct lobules. There are no Cowper’s glands. The mammæ are either two or four, and abdominal in position. No clavicles. Tail always very short. Eyes very large and exposed, with flat cornea.

The animals of this group are all aquatic in their mode of life, spending the greater part of their time in the water, swimming and diving with great facility, feeding mainly on fish, crustaceans, and other marine animals, and progressing on land with difficulty. They always come on shore, however, for the purpose of bringing forth their young. They are generally marine, but they occasionally ascend large rivers, and some inhabit inland seas and lakes, as the Caspian and Baikal. Though not numerous in species, they are widely distributed over the world, but occur most abundantly on the coasts of lands situated in cold and temperate zones. The suborder is divisible into three well-marked families: the _Otariidæ_, Fur-Seals or Sea-Bears, which form a transition from the Fissiped Carnivora to the Seals; the _Trichechidæ_, containing the Walrus; and the _Phocidæ_ or typical Seals.

The resemblances between the skull and other parts of the body of the Fur Seals and the Ursoid true Carnivora is suggestive of some genetic relationship between the two groups, and Professor Mivart[515] expresses the opinion that the one group is the direct descendant of the other. The same writer goes on to suggest that if the Eared-Seals have been derived from Bear-like Carnivores this need not necessarily hold good with the true Seals, which may have had another, and possibly Lutrine, origin. The presence of an alisphenoid canal in _Ursus_ and the _Otariidæ_, and its absence in _Lutra_ and the _Phocidæ_, together with other osteological features, are cited in support of this view; but although these resemblances and differences are certainly remarkable, yet much more evidence is required before the hypothesis can be accepted as even a probable one. It must, moreover, be borne in mind that the true Bears are a very modern group; and there is a possibility that the Pinnipeds may prove to have been independently derived from the extinct Carnivora noticed below under the name of Creodonta.

_Family_ OTARIIDÆ.

When on land the hind feet are turned forwards under the body, and aid in supporting and moving the trunk as in ordinary mammals. A small external ear. Testes suspended in a distinct external scrotum. Skull with postorbital processes, and an alisphenoid canal. Angle of mandible inflected. Palms and soles of feet naked.

_Otaria._[516]—Dentition: _i_ ³⁄₂, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ¹⁻²⁄₁; total 34 or 36. First and second upper incisors small, with the summits of the crowns divided by a deep transverse groove into an anterior and a posterior cusp of nearly equal height; the third large and canine-like. Canines large, conical, pointed, recurved. Molars and premolars usually ⁵⁄₅, of which the second, third, and fourth are preceded by milk-teeth shed a few days after birth; sometimes (as in Fig. 271) a sixth upper molar (occasionally developed on one side and not the other); all with similar characters, generally uniradicular; crown moderate, compressed, pointed, with a single principal cusp, and sometimes a cingulum, and more or less developed anterior and posterior accessory cusps. Vertebræ: C 7, D 15, L 5, S 4, C 9-14. Head rounded. Eyes large. Pinna of ear small, narrow, and pointed. Neck long. Skin of all the feet extended far beyond the nails and ends of the digits, with a deeply-lobed margin. The nails small and often quite rudimentary, especially those of the first and fifth toes of both feet, the best developed and most constant being the three middle claws of the hind foot, which are elongated, compressed, and curved.

[Illustration: FIG. 271.—Skull of _Otaria forsteri_. (From Gray, _Proc. Zool. Soc._ 1872, p. 660.)]

The Eared-Seals, commonly called Sea-Bears or Sea-Lions, are widely distributed, especially in the temperate regions of both hemispheres, though absent from the coasts of the North Atlantic. As might be inferred from their power of walking on all fours, they spend more of their time on shore, and range inland to greater distances, than the true Seals, especially at the breeding time, though they are obliged always to return to the water to seek their food. They are gregarious and polygamous, and the males are usually much larger than the females, a circumstance which has given rise to some of the confusion existing in the specific determination of the various members of the genus. Some of the species possess, in addition to the stiff, close, hairy covering common to all the group, an exceedingly fine, dense, woolly under fur. The skins of these, when dressed and deprived of the longer harsh outer hairs, constitute the “seal-skin” of commerce, so much valued for wearing apparel, which is not the product of any of the true Seals. The best-known species are _O. stelleri_, the Northern Sea Lion, the largest of the genus, from the North Pacific, about 10 feet in length; _O. jubata_, the Patagonian or Southern Sea Lion (Fig. 272), from the Falkland Islands and Patagonia; _O. californiana_, from California, frequently exhibited alive in menageries in Europe; _O. ursina_, the common Sea-Bear or Fur-Seal of the North Pacific, the skins of which are imported in immense numbers from the Prybiloff Islands; _O. pusilla_, from the Cape of Good Hope; _O. forsteri_ and others, from the coasts of Australia and various islands scattered over the southern hemisphere. These have been grouped by some zoologists into many genera, founded upon very trivial modifications of teeth and skull. In a recent memoir Mr. Beddard[517] concludes that if the genus be split up at all, it should be divided into _Otaria_, containing only _O. jubata_ (with its numerous synonyms), and _Arctocephalus_, comprising all the other species. The latter group is distinguished by the more narrow and pointed nose, the longer ears, the palate not excavated nor truncated posteriorly, the presence of a hook-like process to the pterygoids, and by the posterior border of the nasals extending behind the zygoma.

[Illustration: FIG. 272.—The Patagonian Sea-Lion (_Otaria jubata_). From Sclater, _Proc. Zool. Soc._ 1866, p. 80.]

The following account of _O. ursina_ in the Prybiloff Islands is taken, with slight verbal alteration, from Nordenskiöld’s _Voyage of the Vega_: “The Sea-Bears are found year after year during summer at certain parts of the coast, known as ‘rookeries,’ where, collected in hundreds of thousands, they pass several months without the least food. The males or ‘bulls’ come first to the place, most of them in the month of May or in the beginning of June. The most violent conflicts, often with a deadly issue for one of the parties, now arise regarding the space of about a hundred square feet which each bull-seal considers necessary for his home. The strongest and most successful in fight retain the best places near the shore; the weaker have to crawl farther up on land, where the chances of getting a sufficient number of spouses are not particularly great. The fighting goes on with many feigned attacks and parades. At first the contest concerns only the proprietorship of the soil. The attacked, therefore, never follows his opponent beyond the area he has once taken up, but haughtily lays itself down, when the enemy has retired, in order to collect strength for a new combat. The animal in such a case grunts with satisfaction, throws himself on his back, scratches himself with his fore feet, attends to his toilet, or cools himself by slowly fanning with one of his hind feet; but he is always on the alert and ready for a new fight, until he is tired out and meets his match and is driven farther up from the beach. In the middle of June the females come up from the sea. At the water’s edge they are received in a very gallant way by some strong bulls that have succeeded in securing for themselves places next the shore, and now are bent by fair means or foul on annexing the females for their harem. But scarcely is the female that has come up out of the water established with male No. 1 than he rushes towards a new female on the surface of the water. Male No. 2 now stretches out his neck and without ceremony lays hold of the female of No. 1, to be afterwards exposed to a similar trick by No. 3. In such cases the females are quite passive, never fall out with each other, and bear with patience the severe wounds they often get when they are pulled about by the combatants, now in one direction, now in another. All the females are finally distributed in this way after furious combats among the males, those of the latter who are nearest the beach getting from 12 to 15 consorts to their share. Soon after landing the females bring forth their young, which are treated with great indifference, and are protected by their adopted father only within the limits of the harem. Next comes the pairing season, and when it has passed there is an end to the arrangement and distribution into families at first so strictly maintained. The males, rendered lean by three months’ absolute fasting, by degrees leave the rookery, which is left in possession of the Walruses and the young Sea Bears, including a number of young males that have not ventured to the place before. In the middle of September, when the young have learned to swim, the place is quite abandoned, with the exception of single animals that have for some reason remained behind.”

_Family_ TRICHECHIDÆ.

In many characters the single genus representing this family is intermediate between the _Otariidæ_ and _Phocidæ_, but it has a completely aberrant dentition. It has no external ears, as in the _Phocidæ_; but when on land the hind feet are turned forwards and used in progression, though less completely than in the _Otariidæ_. The upper canines are developed into immense tusks, which descend a long distance below the lower jaw. All the other teeth (Fig. 273), including the lower canines, are much alike, small, simple, and one-rooted, the molars with flat crowns. The skull is without postorbital process, but has an alisphenoid canal.

[Illustration: FIG. 273.—Diagram of the dentition of the Walrus (_Trichechus rosmarus_). The denticles placed apart from the others are milk-teeth, and disappear soon after birth. The small teeth in connection with the jaws frequently persist throughout life.]

_Trichechus._[518]—Dentition of young: _i_ ²⁄₂, _c_ ¹⁄₁, _p_ and _m_ ⁵⁄₄. Many of these teeth are, however, lost early or remain through life in a rudimentary state concealed by the gums. The teeth which are usually developed functionally are _i_ ¹⁄₀, _c_ ¹⁄₁, _p_ ³⁄₃, _m_ ⁰⁄₀; total 18. Vertebræ: C 7, D 14, L 6, S 4, C 12. Head round. Eyes rather small. Muzzle short and broad, with on each side a group of long, very stiff, bristly whiskers. The remainder of the hair-covering very short and adpressed. Tail very rudimentary. Fore feet with subequal toes, carrying five minute flattened nails. Hind feet with subequal toes, the fifth slightly the largest, having cutaneous lobes projecting beyond the ends as in _Otaria_; first and fifth with minute flattened nails; second, third, and fourth with large, elongated, subcompressed pointed nails.

_Trichechus_ is the almost universally accepted generic name by which the Walrus or Morse[519] is known to zoologists, but some confusion has been introduced into literature by the revival of the nearly obsolete terms _Rosmarus_ by some authors and _Odobænus_ by others. _T. rosmarus_ is the name of the species met with in the Arctic seas; that of the North Pacific, if distinct, is _T. obesus_. The preceding and following descriptions will apply equally to both. A full-grown male Walrus measures from 10 to 11 feet from the nose to the end of the very short tail, and is a heavy, bulky animal, especially thick about the shoulders. The soles of both fore and hind feet are bare, rough, and warty. The surface of the skin generally is covered with short, adpressed hair of a light, yellowish-brown colour, which, on the under parts of the body and base of the flippers, passes into dark reddish-brown or chestnut. In old animals the hair becomes more scanty, sometimes almost entirely disappearing, and the skin shows ample evidence of the rough life and pugnacious habits of the animal in the innumerable scars with which it is usually covered. It is everywhere more or less wrinkled, but especially over the shoulders, where it is thrown into deep and heavy folds.

[Illustration: FIG. 274.—The Walrus (_Trichechus rosmarus_).]

The tusks are formidable weapons of defence, but their principal use seems to be scraping and digging among the sand and shingle for the molluscs and crustaceans on which the Walrus feeds. They are said also to aid in climbing up the slippery rocks and ledges of ice on which so much of the animal’s life is passed. Although this function of the tusks is affirmed by numerous authors, some of whom appear to have had opportunities of actual observation, it is explicitly denied by Malmgren.

Walruses are more or less gregarious in their habits, being met with generally in companies or herds of various sizes. They are only found near the coast or on large masses of floating ice, and rarely far out in the open sea; and, though often moving from one part of their feeding ground to another, they have no regular seasonal migrations. Their young are born between the months of April and June, usually but one at a time, never more than two. Their strong affection for their young, and their sympathy for each other in times of danger, have been particularly noticed by all who have had the opportunity of observing them in their native haunts. When one of their number is wounded, the whole herd usually join in a concerted and intelligent defence. Although harmless and inoffensive when not molested, they exhibit considerable fierceness when attacked, using their great tusks with tremendous effect either on human enemies who come into too close quarters or on Polar Bears, the only other adversaries they can meet with in their own natural territory. Their voice is a loud roaring, and can be heard at a great distance; it is described by Dr. Kane as “something between the mooing of a cow and the deepest baying of a mastiff, very round and full, with its bark or detached notes repeated rather quickly seven or nine times in succession.”

The principal food of the Walrus consists of bivalved molluscs, especially _Mya truncata_ and _Saxicava rugosa_, two species very abundant in the Arctic regions, which it digs up from the mud and sand in which they lie buried at the bottom of the sea by means of its tusks. It crushes and removes the shells by the aid of its grinding teeth and tongue, swallowing only the soft part of the animal. It also feeds on other molluscs, sand-worms, star-fishes, and shrimps. Portions of various kinds of algæ or sea-weeds have been found in its stomach, but whether swallowed intentionally or not is still doubtful.

The commercial products of the Walrus are its oil, hide (used to manufacture harness and sole-leather and twisted into tiller ropes), and tusks. The ivory of the latter is, however, inferior in quality to that of the Elephant. Its flesh forms an important article of food to the Eskimo and Tchuktchis. Of the coast tribes of the last-named people the Walrus forms the chief means of support. “The flesh supplies them with food, the ivory tusks are made into implements used in the chase and for other domestic purposes, as well as a affording a valuable article of barter, and the skin furnishes the material for covering their summer habitations, harness for their dog-teams, and lines for their fishing gear” (Scammon).

Geographically the Walrus is confined to the northern circumpolar regions of the globe, extending apparently as far north as explorers have penetrated, but its southern range has been much restricted of late in consequence of the persecutions of man. On the Atlantic coast of America it was met with in the sixteenth century as low as the southern coast of Nova Scotia, and in the last century it was common in the Gulf of St. Lawrence and on the shores of Labrador. It still inhabits the coast round Hudson’s Bay, Davis Straits, and Greenland, where, however, its numbers are daily decreasing. It is not found on the Arctic coast of America between the 97th and 158th meridians. In Europe occasional stragglers have reached the British Isles, and it was formerly abundant on the coasts of Finmark. It is rare in Iceland, but Spitzbergen, Nova Zembla, and the western part of the north coast of Siberia are still constant places of resort, in all of which a regular war of extermination is carried on. The North Pacific, including both sides of Behring’s Strait, northern Kamschatka, Alaska, and the Pribyloff Islands, are also the haunts of numerous Walruses, which are isolated from those of the North Atlantic by the long stretches of coast, both of Siberia and North America, where they do not occur. The Pacific Walrus appears to be as large as, if not larger than, that of the Atlantic; its tusks are longer and more slender, and curved inwards; the whiskers are smaller, and the muzzle (of the skull) relatively deeper and broader. These and certain other minor differences have induced some naturalists to consider it specifically distinct under the name of _Trichechus obesus_. Its habits appear to be quite similar to those of the Atlantic form. Though formerly found in immense herds, it is rapidly becoming scarce, as the methods of destruction used by the American whalers, who have systematically entered upon its pursuit, are far more certain and deadly than those of the native Tchuktchis, to whom, as mentioned before, the Walrus long afforded the principal means of subsistence.

Fossil remains of Walruses and closely allied animals have been found in the United States, and in England, Belgium, and France, in deposits of Pliocene age.

_Family_ PHOCIDÆ.

The true Seals are the most completely adapted for aquatic life of all the Pinnipeds. When on land the hind limbs are extended behind them and take no part in progression, which is effected by a series of jumping movements produced by the muscles of the trunk, in some species aided by the fore limbs only. The palms and soles of the feet are hairy. There is no pinna to the ear, and no scrotum, the testes being abdominal. The upper incisors have simple, pointed crowns, and vary in number in the different groups. All the forms have well-developed canines and ⁵⁄₅ teeth of the cheek-series. In those species of which the milk-dentition is known, there are three milk molars (Fig. 275), which precede the second, third, and fourth permanent molars; the dentition is therefore _p_ ⁴⁄₄, _m_ ¹⁄₁, the first premolar having as usual no milk-predecessor. The skull has no postorbital process and no alisphenoid canal; and the angle of the mandible is not inflected. The fur is stiff and adpressed, without woolly under fur.

Subfamily =Phocinæ=.—Incisors ³⁄₂. All the feet with five well-developed claws. The toes on the hind feet subequal, the first and fifth not greatly exceeding the others in length, and with the interdigital membrane not extending beyond the toes.

_Halichœrus._[520]—Dentition: _i_ ³⁄₂, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ¹⁄₁; total 34. Crowns of molars large, simple, conical, recurved, slightly compressed, with sharp anterior and posterior edges, but without accessory cusps, except sometimes in the two hinder ones of the lower jaw. With the exception of the last one or two in the upper jaw and the last in the lower jaw they are all uniradicular. Vertebræ: C 7, D 15, L 5, S 4, C 14.

One species, _H. grypus_, the Gray Seal of the coasts of Scandinavia and the British Isles (see page 604.)

[Illustration: FIG. 275.—Upper permanent and deciduous dentition of the Greenland Seal (_Phoca grœnlandica_). The first and second deciduous incisors are already absorbed.]

_Phoca._[521]—Dental formula as the last. Teeth smaller and more pointed. Molars (Figs. 275 and 276) with two roots (except the first in each jaw); and their crowns with accessory cusps. Vertebræ: C 7, D 15, L 5, S 4, C 12-15. Head round and short. Fore feet short, with five very strong, subcompressed, slightly curved, rather sharp claws, subequal in length. On the hind feet the claws much narrower and less curved. The species of this genus are widely distributed throughout the northern hemisphere, and include _P. barbata_, the Bearded Seal; _P. grœnlandica_, the Greenland Seal; _P. vitulina_, the Common Seal (Fig. 277); and _P. hispida_, the Ringed Seal of the North Atlantic; _P. caspica_, from the Caspian and Aral Seas; and _P. sibirica_, from Lake Baikal.

[Illustration: FIG. 276.—Skull of Common Seal, showing form of teeth.]

Although the members of this subfamily swim and dive with the greatest ease, often remaining as much as a quarter of an hour or more below the surface, and are dependent for their sustenance entirely on living prey captured in the water, yet they frequently resort to sandy beaches, rocks, or ice-floes, either to sleep or to bask in the sun, and especially for the purpose of bringing forth their young. The latter appears to be the universal habit, and, strange as it may seem, the young seals—of some species at least—take to the water at first very reluctantly, and have actually to be taught to swim by their parents. The number of young produced is usually one annually, though occasionally two. They are at first covered with a coat of very thick, soft, nearly white fur, and until it falls off they do not usually enter the water. This occurs in the Greenland and Gray Seal when from two to three weeks old, but in the Common Seal apparently much earlier. One of this species born in the London Zoological Gardens had shed its infantile woolly coat and was swimming and diving about in its pond within three hours after its birth. The movements of the true Seals upon the ground or ice are very different from those of the Eared-Seals. Thus the hinder limbs (by which mainly they propel themselves through the water) are on land always perfectly passive, stretched backwards, with the soles of the feet applied to each other, and often raised to avoid contact with the ground. Sometimes the fore limbs are equally passive, being placed close to the sides of the body, and motion is then effected by a shuffling or wriggling action produced by the muscles of the trunk. When, however, there is any necessity for a more rapid mode of progression the animals use the fore paws, either alternately or simultaneously, pressing the palmar surface on the ground and lifting and dragging the body forwards in a succession of short jumps. In this way they manage to move so fast that a man has to step out beyond a walk to keep up with them; but such rapid action costs considerable effort, and they very soon become heated and exhausted. These various modes of progression appear to be common to all species so far as has been observed.

Most kinds of Seals are gregarious and congregate, especially at the breeding season, in immense herds. Such is the habit of the Greenland Seal (_Phoca grœnlandica_), which resorts in the spring to the ice-floes of the North Sea, around Jan Mayen Island, where about 200,000 are killed annually by the crews of the Scotch, Dutch, and Norwegian sealing vessels. Others, like the Common Seal of the British islands (_P. vitulina_), though having a wide geographical range, are never met with in such large numbers or far away from land. This species is stationary all the year round, but some have a regular season of migration, moving south in winter and north in summer. They are usually harmless, timid, inoffensive animals, though, being polygamous, the old males often fight desperately with each other, their skins being frequently found covered with wounds and scars. They are greatly attached to their young, and remarkably docile and easily trained when in captivity; indeed, although there would seem little in the structure or habits of the Seal to fit it by nature to be a companion of man, yet there is perhaps no wild animal which attaches itself so readily to the person who takes care of and feeds it. Seals appear to have much curiosity, and it is a very old and apparently well-attested observation that they are strongly attracted by musical sounds. Their sense of smell is very acute, and their voice varies from a harsh bark or grunt to a plaintive bleat. Seals feed chiefly on fish, of which they consume enormous quantities; some, however, subsist largely on crustaceans, especially species of _Gammarus_, which swarm in the northern seas, also on molluscs, echinoderms, and even occasionally sea-birds, which they seize when swimming or floating on the water.

[Illustration: FIG. 277.—The Common Seal (_Phoca vitulina_).]

Although the true Seals do not possess the beautiful under fur (“seal-skin” of the furriers) which makes the skin of the Sea-Bears so precious, yet their hides are still sufficiently valuable as articles of commerce, together with the oil yielded by their fat, to subject them to a devastating persecution, by which their numbers are being continually diminished.

Two species of seals only are met with regularly on the British coasts, the Common Seal and the Gray Seal. The former (Fig. 277) is a constant resident in all suitable localities round the Scottish, Irish, and English coasts, from which it has not been driven away by the molestations of man. Although, naturally, the most secluded and out-of-the-way spots are selected as their habitual dwelling-places, there are few localities where they may not be occasionally met with. Within the writers’ knowledge one was seen not many years ago lying on the shingly beach at so populous a place as Brighton, and another was caught in the river Welland, near Stamford, 30 miles from the sea. They frequent bays, inlets, and estuaries, and are often seen on sandbanks or mudflats left dry at low tide, and, unlike some of their congeners, are not found on the ice-floes of the open sea, nor, though gregarious, are very large numbers ever seen in one spot. The young are produced at the end of May or beginning of June. They feed chiefly on fish, and the destruction they occasion among salmon is well known to Scottish fishermen. The Common Seal is widely distributed, being found not only on the European and American coasts bordering the Atlantic Ocean, but also in the North Pacific. It is from 4 to 5 feet in length, and variable in colour, though usually yellowish-gray, with irregular spots of dark brown or black above and yellowish-white beneath. The Gray Seal (_Halichœrus grypus_) is of considerably larger size, the males attaining when fully adult a length of 8 feet from nose to end of hind feet. It is of a yellowish-gray colour, lighter beneath, and with dark gray spots or blotches, but, like most other Seals, is liable to great variations of colour according to age. This species appears to be restricted to the North Atlantic, having been rarely seen on the American coasts, but not farther south than Nova Scotia; it is chiefly met with on the coasts of Ireland, England, Scotland, Norway, and Sweden, including the Baltic and Gulf of Bothnia, and Iceland, though it does not appear to range farther north. It is apparently not migratory, and its favourite breeding places are rocky islands; the young being born in the end of September or beginning of October.

Subfamily =Monachinæ=.—Incisors ²⁄₂. Cheek-teeth two-rooted, except the first. On the hind feet the first and fifth toes greatly exceeding the others in length, with nails rudimentary or absent.

_Monachus._[522]—Dentition: _i_ ²⁄₂, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ¹⁄₁; total 32. Crowns of molars strong, conical, compressed, hollowed on the inner side, with a strongly marked lobed cingulum, especially on the inner side, and slightly developed accessory cusps before and behind. The first and last upper and the first lower molar considerably smaller than the others. Vertebræ: C 7, D 15, L 5, S 2, C 11. All the nails of both fore and hind feet very small and rudimentary. One species, _M. albiventer_, the Monk-Seal of the Mediterranean and adjacent parts of the Atlantic.

The other genera[523] of this section have the same dental formula, but are distinguished by the characters of the cheek-teeth and the feet. They are all inhabitants of the shores of the southern hemisphere.

_Ogmorhinus._[524]—All the teeth of the cheek-series with three distinct pointed cusps, deeply separated from each other; of these the middle or principal cusp is largest and slightly recurved; the other two (anterior and posterior) are nearly equal in size, and have their apices directed towards the middle one. Skull much elongated. One species, _O. leptonyx_, the Sea-Leopard, widely distributed in the Antarctic and southern temperate seas.

_Lobodon._[525]—Cheek-teeth with much-compressed elongated crowns and a principal recurved cusp, rounded and somewhat bulbous at the apex, and one anterior, and one, two, or three posterior, very distinct accessory cusps. One species, _L. carcinophaga_.

_Pœcilophoca._[526]—Cheek-teeth small, with simple, subcompressed, conical crowns, having a broad cingulum, but no distinct accessory cusps. One species, _P. weddelli_.

_Ommatophoca._[527]—All the teeth very small; those of the cheek-series with pointed recurved crowns, and small posterior and still less developed anterior accessory cusps. Orbits very large. Nails quite rudimentary on front, and absent on hind feet. The skull bears a considerable resemblance to that of the members of the next subfamily, towards which it may form a transition. There is one species, _O. rossi_, of which very little is known.

Subfamily =Cystophorinsæ=.—Incisors ²⁄₁. Teeth of cheek-series generally one-rooted. Nose of males with an appendage capable of being inflated. First and fifth toes of hind feet greatly exceeding the others in length, with prolonged cutaneous lobes, and rudimentary or no nails.

_Cystophora._[528]—Dentition: _i_ ²⁄₁, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ¹⁄₁; total 30. The last molar has generally two distinct roots. Beneath the skin over the face of the adult male, and connected with the nostrils, is a sac which, when inflated, forms a kind of hood covering the upper part of the head. Nails present, though small, on the hind feet. One species, _C. cristata_, the Hooded or Bladder-Nose Seal of the Polar Seas.

_Macrorhinus._[529]—Dentition as the last, but cheek-teeth of simpler character, and all one-rooted. All the teeth, except the canines, very small relatively to the size of the animal. Hind feet without nails. Vertebræ: C 7, D 15, L 5, S 3, C 11. Nose of adult male produced into a short tubular proboscis, ordinarily flaccid, but capable of dilatation and elongation under excitement. One species, _M. leoninus_, the Elephant Seal, or Sea-Elephant of the whalers, the largest of the whole family, attaining the length of nearly 20 feet. Formerly abundant in the Antarctic Seas, and also found on the coast of California.

_Extinct Seals._—Remains of animals of this group have been found in late Miocene and Pliocene strata in Europe and America, the most abundant and best-preserved being those of the Pliocene Antwerp Crag, the subject of an illustrated monograph by Van Beneden. Nothing has, however, yet been discovered which throws any light upon the origin of the group, since all the extinct forms at present known come within the definition of the existing families; and, though annectant forms between these occur, there are as yet no transitions to a more generalised type of mammal. Indeed, all those of which the characters are best known belong to the completely developed Phocine or Trichechine, and not to the Otariine, type. The typical genus _Phoca_ occurs in the Antwerp Crag, while remains of Seals provisionally referred to this genus are found in the Pliocene of the Crimea and the Miocene of Malta and Virginia. Of the other Antwerp forms _Callophoca_ is said to be allied to _Phoca grœnlandica_, _Platyphoca_ to _Phoca barbata_, _Phocanella_ to _Phoca foetida_, _Gryphoca_ to _Halichœrus_, _Palæophoca_ and _Monatherium_ to _Monachus_, and _Mesotaria_ to _Cystophora_; while _Prophoca_ does not appear to come very close to any existing form. It should be observed that it is extremely doubtful whether all these fossil Seals are really entitled to generic distinction.

_Bibliography of Pinnipedia._—J. A. Allen, _History of North American Pinnipeds_, 1880; St. George Mivart, “Notes on the Pinnipedia,” _Proc. Zool. Soc._ 1885, p. 484; P. J. Van Beneden, _Ossements fossiles d’Anvers_, in the _Mém. Acad. Roy. d. Belgique_.

_Suborder_ CREODONTA.

The discovery of fossil remains in Eocene and early Miocene formations both in Europe and North America shows that numerous species of terrestrial carnivorous animals existed upon the earth during those periods which cannot be referred to either of the sections into which the order has now become broken up. By some zoologists these have been supposed to be Marsupials, or at least to show transitional characters between the Metatherian and Eutherian subclasses. By others they are looked upon as belonging altogether to the latter group, and as the common ancestors of existing Carnivores and Insectivores, or perhaps rather as descendants or relatives of such common ancestors, retaining more of the generalised characters than any of the existing species. They shade off almost insensibly into numerous other forms less distinctly carnivorous, to the whole of which, including the modern Insectivora, Cope (to whom we are indebted for much of our knowledge of the American extinct species) gives the name of BUNOTHERIA, those more specially related to the existing Carnivora forming the suborder Creodonta. These are instances, however, in which the application of the principles of classification adopted in the case of existing species, of which the entire structure is known, and which have become divided into isolated groups by the extinction of intermediate forms, is almost impossible. If the generally accepted view of evolution is true, and the extreme modifications pass insensibly into each other by minute gradations (a view the palæontological proof of which becomes strengthened by every fresh discovery), there must be many of these extinct forms which cannot be assigned to definitely characterised groups. There are, however, some which stand out prominently from the others as formed on distinct types, having no exact representatives at present living on the earth.

[Illustration: FIG. 278.—Anterior portion of the skull of _Hyænodon leptorhynchus_. (After Filhol.)]

The more typical Creodonts appear, however, to be so closely related to the true Carnivora through the extinct _Miacidæ_ (p. 539), that it is on the whole advisable to regard them as representing a distinct suborder of Carnivora. In the strong development of the canines (Fig. 278) they are distinguished from the modern Insectivora; and they also differ from the latter and resemble the true Carnivores in the form of the incisors, the second one in the lower jaw (when three are present) being thrust up above the level of the other two in the manner obtaining in most of the modern Carnivora. Some of the most generalised forms included in the present group approximate so closely to the Condylarthrous Ungulates as to indicate that both groups have probably had a common origin.

The Creodonta as a whole are characterised by the small size of the brain, the absence of a single differentiated carnassial tooth, and the triangular form or secant character of their upper molars. In the carpus the scaphoid and lunar were usually distinct; the femur has a third trochanter; the upper or tibial surface of the astragalus usually wants the groove found in modern Carnivores: and the feet were plantigrade. The curious resemblance of the molars of many of these forms to those of the Marsupials may indicate a genetic relationship between the two groups; but, on the other hand, the presence of a full set of milk-teeth and the absence of palatal vacuities, or of an inflection of the angle of the mandible, sharply distinguishes them from that order. Space permits of a notice only of the more interesting forms.

_Hyænodontidæ._—This family is taken to include some of the more specialised types, such as the European and American _Hyænodon_ and _Oxhyæna_ and the European _Pterodon_. In _Hyænodon_ (Fig. 278) the dental formula is _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₃; the fourth premolar above and the first true molar below being formed upon the “carnassial” plan, but the teeth behind these, instead of being tuberculated as in all existing Carnivora, repeat the characters of the carnassial, and also increase in size, especially in the lower jaw, from before backwards. The last lower molar differs from the two preceding teeth, and is very like the carnassial of _Felis_. The scaphoid and lunar of the carpus were fused together. Some species, as _H. leptorhynchus_, were as large as a Wolf, while others did not exceed a Fox in size. _Pterodon_ is readily distinguished by having _m_ ³⁄₃, by the larger size of the inner tubercles of the upper molars, and the similarity in the form of the three lower molars. In some species there were only two upper incisors, and the first lower premolar may be wanting. _Oxhyæna_ is a specialised form with _i_ ²⁻³⁄₀, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₂, and a very long mandibular symphysis.

_Proviverridæ._—The European and American genus _Proviverra_ (_Cynohyænodon_ or _Stypolophus_) may be regarded as representing a second family. The dental formula in this genus is the typical _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ³⁄₃, the upper molars have a large inner tubercle, while the lower molars are differentiated into a blade and talon, the blade having a large inner cusp. The upper teeth closely resemble the molars of _Dasyurus_, while the lower molars are like the lower carnassial of _Cynodictis_ and _Viverra_; and thus indicate how the Creodonts may have passed into the true Carnivores through the extinct _Miacidæ_.

[Illustration: FIG. 279.—The three right upper molars of _Arctocyon dueli_ (_a_), and the second of _A. gervaisi_ (_b_); from the lowest Eocene of Rheims. _pr_, protocone; _pa_, paracone; _me_, metacone; _hy_, hypocone; _ml_, metaconule; _pl_, paraconule. (From Osborn.)]

_Arctocyonidæ and Mesonychidæ._—The first of these families is represented by _Arctocyon primævus_, one of the oldest known Tertiary mammals, from the lowest Eocene beds of La Fère, department of Aisne, France, and also by other species from corresponding beds at Rheims. The dental formula is _i_ ³⁄₃, _c_ ¹⁄₁, _p_?⁄₃₋₄, _m_ ³⁄₃. The upper molars (Fig. 279) are tritubercular, with an incipient postero-internal column (hypocone); the lower are quadritubercular; and the premolars simple. The typical species was of large size, but the two of which the teeth are figured were considerably smaller. In the American _Mesonyx_ the dental formula was the typical one, the jaws were comparatively short, the mandibular symphysis was elongated, the cheek-teeth were of simple structure, and resembled the premolars of many of the true Carnivora, and the astragalus had a grooved tibial surface and distinct distal facets for the cuboid and navicular, resembling in the latter respect the corresponding bone of a Perissodactyle Ungulate. The terminal phalanges had deeply fissured extremities, and are said to be more like those of Rodents than true Carnivores. _Mesonyx ossifragus_ was larger than a Grizzly Bear. _Amblyctonus_, of the same deposits, differs by the smooth tibial face of the astragalus and the development of an anterior cusp to the lower molars.