Part 2

CARNIVORA, the zoological order typified by the larger carnivorous placental land mammals of the present day, such as lions, tigers and wolves, but also including species like bears whose diet is largely vegetable, as well as a number of smaller flesh-eating species, together with the seals and their relatives, and an extinct Tertiary group. Apart from this distinct group (see CREODONTA), the Carnivora are characterized by the following features. They are unguiculate, or clawed mammals, with never less than four toes to each foot, of which the first is never opposable to the rest; the claws, or nails, being more or less pointed although occasionally rudimentary. The teeth comprise a deciduous and a permanent series, all being rooted, and the latter divisible into the usual four series. In front there is a series of small pointed incisors, usually three in number, on each side of both jaws, of which the first is always the smallest and the third the largest, the difference being most marked in the upper jaw; these are followed by strong conical, pointed, recurved canines; the premolars and molars are 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 size and form, but the hemispheres are never destitute of convolutions. The stomach is always simple and pyriform; the caecum is either absent or short and simple; and the colon is not sacculated or much wider than the small intestine. Vesiculae seminales are never developed, but Cowper's glands may be present or absent. The uterus is two-horned, and the teats are abdominal and variable in number; while the placenta is deciduate, and almost always zonary. The clavicle is often absent, and when present never complete. The radius and ulna are distinct; the scaphoid and lunar of the tarsus are united; there is never an os centrale in the adult; and the fibula is distinct.

The large majority of the species subsist chiefly on animal food, though many are omnivorous, and a few chiefly vegetable-eaters. The more typical forms live altogether on recently-killed warm-blooded animals, and their whole organization 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 are capable of being domesticated, and when placed under favourable circumstances exhibit a high degree of intelligence.

I. FISSIPEDIA

[Illustration: FIG. I.--Left upper sectorial or carnassial teeth of Carnivora. I, _Felis_; II, _Canis_; III, _Ursus_. 1, anterior, 2, middle, and 3, posterior cusp of blade; 4, inner cusp supported on distinct root; 5, inner cusp, posterior in position, and without distinct root, characteristic of the _Ursidae_.]

The typical section of the group, the Carnivora Vera, Fissipedia or Carnassidentia, includes all the existing terrestrial members of the order, together with the otters and sea-otters. In this section the fore-limbs never have the first digit, or the hind-limbs the first and fifth digits, longer than the others; and the incisors are 3/3 on each side, with very rare exceptions. The cerebral hemispheres are more or less elongated; always with three or four convolutions on the outer surface forming arches above each other, the lowest surrounding the Sylvian fissure. In the cheek-series there is one specially modified tooth in each jaw, to which the name of "sectorial" or "carnassial" is applied. The teeth in front of this are more or less sharp-pointed and compressed; the teeth behind broad and tuberculated. The characters of the sectorial teeth deserve special attention, as, though fundamentally the same throughout the group, they are greatly modified in different genera. The upper sectorial 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 of a more or less compressed blade supported on two roots and an inner lobe supported by a distinct root (see fig. 1). The blade when fully developed has three cusps (i, 2 and 3), but the anterior is always small, and often absent. The middle cusp is conical, high and pointed; and the posterior cusp has a compressed, straight, knife-like edge. The inner cusp. (4) varies in extent, but is generally placed near the anterior end of the blade, though sometimes median in position. In the _Ursidae_ alone both the inner cusp and its root are wanting, and there is often a small internal and posterior cusp (5) without root. In this family also the sectorial is relatively to the other teeth much smaller than in other Carnivora. The lower sectorial (fig. 2) is the most anterior of the teeth without predecessors in the milk-series, and is therefore reckoned the first molar. It has two roots supporting a crown, consisting when fully developed of a compressed bilobed blade (1 and 2), a heel (4), and an inner tubercle (3). The cusps of the blade, of which the hinder (2) is the larger, are separated by a notch, generally prolonged into a linear fissure. In the specialized _Felidae_ (I) the blade alone is developed, both heel and inner tubercle being absent or rudimentary. In _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 or cusps are almost side by side, instead of anterior and posterior. The inner tubercle (3) is generally a conical pointed cusp, 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 than in any other species, but even here they can be traced.

[Illustration: FIG. 2.--Left lower sectorial or carnassial teeth of Carnivora, I, _Felis_; II, _Canis_; III, _Herpestes_; IV, _Lutra_; V, _Meles_; VI, _Ursus_. 1, Anterior cusp of blade; 2, posterior cusp of blade; 3, inner tubercle; 4, heel. It will be seen that the relative size of the two roots varies according to the development of the portion of the crown they respectively support.]

The toes are nearly always armed with large, strong, curved and sharp claws, ensheathing the terminal phalanges and held firmly in place by broad plates of bone reflected over their attached ends from the bases of the phalanges. In the _Felidae_ these claws are "retractile"; the terminal phalange with the claw attached, folding back in the fore-foot into a sheath by the outer or ulnar side of the middle phalange of the digit, and retained in this position when at rest by a strong elastic ligament. In the hind-foot the terminal joint or phalange is retracted on to the top, and not the side of the middle phalange. By the action of the deep flexor muscles the terminal phalanges are straightened, the claws protruded from their sheath, and the soft "velvety" paw becomes suddenly converted into a formidable weapon of offence. The habitual retraction of the claws preserves their points from wear.

The land Carnivora are best divided into two subgroups or sections--(A) the Aeluroidea, or Herpestoidea, and (B) the Arctoidea; the recognition of a third section, Cynoidea, being rendered untenable by the evidence of extinct forms.

(A) _Aeluroidea_.--In this section, which comprises the cats (_Felidae_), civets (_Viverridae_), and hyenas (_Hyaenidae_), the tympanic bone is more or less ring-like, and forms only a part of the outer wall of the tympanic cavity; an inflated alisphenoid bulla is developed; and the external auditory meatus is short. In the nasal chamber the maxillo-turbinal is small and doubly folded, and does not cut off the naso-turbinal and adjacent bones from the nasal aperture. The carotid canal in the skull is short or absent. Cowper's glands are present, as is a prostate gland and a caecum, as well as a duodenal-jejunal flexure in the intestine, but an os penis is either wanting or small.

Cat tribe.

The members of the cat tribe, or _Felidae_, are collectively characterized by the following features. An alisphenoid is lacking on the lower aspect of the skull. In existing forms the usual dental formula is i. 3/3, c. 1/1, p. 3/2, m. 1/1; the upper molar being rudimentary and placed on the inner side of the carnassial, but the first premolar may be absent, while, as an abnormality, there may be a small second lower molar, which is constantly present in some of the extinct forms. The auditory bulla and the tympanic are divided by an internal partition. The paroccipital process is separate from, or only extends to a slight degree upon the auditory bulla. The thoracic vertebrae number 13; the feet are digitigrade, with five front and four hind toes, of which the claws are retractile; and the metatarsus is haired all round. Anal glands are present.

As regards the teeth, when considered in more detail, the incisors are small, and the canines large, strong, slightly recurved, with trenchant edges and sharp points, and placed wide apart. The premolars are compressed and sharp-pointed; the most posterior in the upper jaw (the sectorial) being a large tooth, consisting of a compressed blade, divided into three unequal cusps supported by two roots, with a small inner lobe placed near the front and supported by a distinct root (fig. 1, I). The upper molar is a small tubercular tooth placed more or less transversely at the inner side of the hinder end of the last. In the lower jaw the molar (sectorial) is reduced to the blade, which is large, trenchant, compressed and divided into two subequal lobes (fig. 2, I). Occasionally it has a rudimentary heel, but never an inner tubercle. The skull generally is short and rounded, though proportionally more elongated in the larger forms; with the facial portion short and broad, and the zygomatic arches wide and strong. The auditory bullae are large, rounded and smooth. Vertebrae: C. 7, D. 13, L. 7, S. 3, Ca. 13-29. Clavicles better developed than in other Carnivora, but not articulating with either the shoulder-bones or sternum. Of the five front toes, the third and fourth are nearly equal and longest, the second slightly, and the fifth considerably shorter. The first is still shorter, not reaching the metacarpophalangeal articulation of the second. In the hind-feet the third and fourth toes are the longest, the second and fifth somewhat shorter and nearly equal, while the first is represented only by the rudimentary metatarsal bone. The claws are large, strongly curved, compressed, very sharp, and exhibit the retractile condition in the highest degree. The tail varies greatly in length, being in some species a mere stump, in others nearly as long as the body. The ears are of moderate size, more or less triangular and pointed; and the eyes rather large, with the iris 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. The tongue is thickly covered with sharp, pointed, recurved horny papillae; and the caecum is small and simple.

As in structure so in habits, the cat may be considered the most specialized of all Carnivora, although they exhibit many features connecting them with extinct types. All the members of the group feed almost exclusively on warm-blooded animals which they have themselves killed, but one Indian species, _Felis viverrina_, is said to prey on fish, and even fresh-water molluscs. Unlike dogs, they never associate in packs, and rarely hunt their prey on 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 are widely diffused over the greater part of the habitable world, though most abundant in the warm latitudes of both hemispheres. None 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 groups into which the family may be divided have representatives in both hemispheres.

Notwithstanding the considerable diversity in external appearance and size between different members of this extensive family, the structural differences are but slight. 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 lobe of the upper sectorial, 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.

In the typical genus _Felis_, which includes the great majority of the species, and has a distribution coextensive with that of the family, the upper sectorial tooth has a distinct inner cusp, the claws are completely contractile, the tail is long or moderate, and the ears do not carry distinct tufts of hair. As regards the larger species, the lion (_F. leo_), tiger (_F. tigris_), leopard (_F. pardus_), ounce or snow-leopard (_F. uncia_) and clouded leopard (_F. nebulosa_) are described in separate articles. Of other Old World species it must suffice to mention that the Tibetan Fontanier's cat (_F. tristis_), and the Indian marbled cat (_F. marmorata_), an ally of the above-mentioned clouded leopard, appear to be the Asiatic representatives of the American ocelots. The Tibetan Pallas's cat (_F. manul_) has been made the type of a distinct genus, _Trichaelurus_, in allusion to its long coat. One of the largest of the smaller species is the African serval, q.v. (_F. serval_), which is yellow with solid black spots, has long limbs, and a relatively short tail. Numerous "tiger-cats" and "leopard-cats," such as the spotted _F. bengalensis_ and the uniformly chestnut _F. badia_, inhabit tropical Asia; while representative species occur in Africa. The jungle-cat (_F. chaus_), which in its slightly tufted ears and shorter tail foreshadows the lynxes, is common to both continents. Another African species (_F. ocreata_) appears to have been the chief progenitor of the European domestic cat, which has, however, apparently been crossed to some extent with the ordinary wild cat (_F. catus_). Of the New World species, _F. concolor_, the puma or couguar, commonly called "panther" in the United States, is about the size of a leopard, but of a uniform brown colour, spotted only when young, and is extensively distributed in both North and South America, ranging between the parallels of 60 deg. N. and 50 deg. S., where it is represented by numerous local races, varying in size and colour. _F. onca_, the jaguar, is a larger and more powerful animal than the last, and more resembles the leopard in its colours; it is also found in both North and South America, although with a less extensive range, reaching northwards only as far as Texas, and southwards nearly to Patagonia (see JAGUAR). _F. pardalis_ and several allied smaller, elegantly-spotted species inhabiting the intratropical regions of America, are commonly confounded under the name of ocelot or tiger-cat. _F. yaguarondi_, rather larger than the domestic cat, with an elongated head and body, and of a uniform brownish-grey colour, ranges from northern Mexico to Paraguay; while the allied _F. eyra_ is a small cat, weasel-like in form, having an elongated head, body and tail, and short limbs, and is of a uniform light reddish-brown colour. It is a native of South America and Mexico. _F. pajeros_ is the Pampas cat.

The typical lynxes, as represented by _Lynx borealis_ (_L. lynx_), the southern _L. pardina_, and the American _L. rufa_, are a northern group common to both hemispheres, and characterized by their tufted ears, short tail, and the presence of a rudimentary heel to the lower carnassial tooth. As a rule, they are more or less spotted in winter, but tend to become uniformly-coloured in summer. They are connected with the more typical cats by the long-tailed and uniformly red caracal, _Lynx (Caracal) caracal_, of India, Persia and Africa, and the propriety of separating them from _Felis_ may be open to doubt (see LYNX and CARACAL).

However this may be, there can be no doubt of the right of the hunting-leopard or chita (cheeta), as, in common with the leopard, it is called in India, to distinction from all the other cats as a distinct genus, under the name of _Cynaelurus jubatus_. From all the other _Felidae_ this animal, which is common to Asia and Africa, is distinguished by the inner lobe of the upper sectorial tooth, 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 air-sinuses. The head is small and round, the body light, the limbs and tail long, and the colour pale yellowish-brown with small solid black spots (see CHEETA).

Civet tribe.

The family _Viverridae_, which includes the civet-cats, genets and mongooses, is nearly allied to the _Felidae_, but its members have a fuller dentition, and exhibit certain other structural differences from the cats, to the largest of which they make no approach in the matter of bodily size. As a rule, there is an alisphenoid canal; the cheek-dentition is p. (3 or 4)/(3 or 4), m. (1 or 2)/(1 or 2). The bulla is small and the tympanic large, with a low division between them; and the paroccipital process is leaf-like and spread over the bulla. The number of dorsal vertebrae, except in the aberrant _Proteles_, is 13 or 14; the claws may be either completely or

## partially retractile or non-retractile; generally each foot has five

toes, but there may be four in front and five behind, the reverse of this, or only four on each foot; the gait may be either digitigrade or

## partially plantigrade; and the metatarsus may be either hairy or naked

inferiorly. Anal, and in some cases also perineal, glands are developed. The family is limited to the warmer parts of the Old World.

Considerable difference of opinion prevails with regard to the serial position of the fossa, or foussa (_Cryptoprocta ferox_), of Madagascar, some writers considering that its affinities are so close to the _Felidae_ that it ought not to be included in the present family at all. Others, on the contrary, see no reason to separate it from the _Viverrinae_ or more typical representatives of the civet-tribe. As a medium course, it may be regarded as the sole representative of a special subfamily--_Cryptoproctinae_--of the _Viverridae_. The subfamily and genus are characterized by possessing a total of 36 teeth, arranged as i. 3/3, c. 1/1, p. 4/4, m. 1/1. The teeth generally closely resemble those of the _Felidae_, the first premolar of both jaws being very minute and early deciduous; the upper sectorial has a small inner lobe, quite at the anterior part; the molar is small and placed transversely; and the lower sectorial has a large trenchant bilobed blade, and a minute heel, but no inner tubercle. The skull is generally like that of _Felis_, but proportionally longer and narrower, with the orbit widely open behind. Vertebrae: C. 7, D. 13, L. 7, S. 3, Ca. 29. Body elongated. Limbs moderate in size. Feet subplantigrade, with five well-developed toes on each, carrying sharp, compressed, retractile claws. Ears moderate. Tail long and cylindrical. The foussa is a sandy-coloured animal with an exceedingly long tail (see FOUSSA).