Part 3

The more typical members of the group, constituting the subfamily _Viverrinae_, are characterized by their sharp, curved and largely retractile claws, the presence of five toes to each foot, and of perineal and one pair of anal glands, and a tympanic bone which retains to a great extent the primitive ring-like form, so that the external auditory meatus has scarcely any inferior lip, its orifice being close to the tympanic ring. The first representatives of the subfamily are the civet-cats, or civets (_Viverra_ and _Viverricula_), and the genets (_Genetta_), in all of which the dentition is _i. 3/3, c. 1/1, p. 4/4, m. 2/2_; total 40. The skull is elongated, with the facial portion small and compressed, and the orbits well-defined but incomplete behind. Vertebrae: C. 7, D. 13, L. 7 (or D. 14, L. 6), S. 3, Ca. 22-30. Body elongated and compressed. Head pointed in front; ears rather small. Extremities short. Feet small and rounded. Toes short, the first 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. The pair of large glands situated on the perineum (in both sexes) secretes an oily substance of a peculiarly penetrating odour. In the true civets, which include the largest members of the group, the teeth are stouter and less compressed than in the other genera; the second upper molar being especially large, and the auditory bulla smaller and more pointed in front; the body is shorter and stouter; the limbs are longer; the tail shorter and tapering. The under side of the tarsus is completely covered with hair, and the claws are longer and less retractile. Fur rather long and loose, and in the middle line of the neck and back especially elongated so as to form a sort of crest or mane. 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 fox, and an inhabitant of intratropical Africa. _V. zibetta_, the Indian civet, of about equal size, approaches in many respects, especially in the characters of the teeth and feet and absence of the crest of elongated hair on the back, to the next section. It inhabits Bengal, China, the Malay Peninsula and adjoining islands. _V. tangalunga_ is a smaller but nearly allied animal from the same part of the world. From these three 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 by scraping the interior of the perineal follicles with a spoon or spatula. The single representative of the genus _Viverricula_ resembles in many respects the genets, but agrees with the civets in having the whole of the under side of the tarsus hairy; the alisphenoid canal is generally absent. _V. malaccensis_, the rasse, inhabiting India, China, Java and Sumatra, is an elegant little animal which affords a favourite perfume to the Javanese. The genets (_Genetta_) are smaller animals, with more elongated and slender bodies, and shorter limbs than the civets. The skull is elongated and narrow; and the auditory bulla large, elongated and rounded at both ends. The teeth are compressed and sharp-pointed, with a lobe on the inner side of the third, upper premolar not present in the previous genera. Pupil contracting to a linear aperture. Tail long, slender, ringed. Fur short and soft, spotted or cloudy. Under side of the metatarsus with a narrow longitudinal bald streak. _Genetta vulgaris_, or _G. genetta_, the common genet, is found in France south of the river Loire, Spain, south-western Asia and North Africa. _G. felina, senegalensis, tigrina, victoriae_ and _pardalis_ are other named species, all African in habitat.

The Malagasy fossane (_Fossa daubentoni_), which has but little markings on the fur of the adult, differs by the absence of a scent-pouch and the presence of a couple of bare spots on the under surface of the metatarsus. The beautiful linsangs (_Linsanga_ or _Prionodon_), ranging from the eastern Himalaya to Java and Borneo, are represented by two or three species, easily recognizable by the broad transverse bands of blackish brown and yellow with which the body and tail are marked. They are specially distinguished by having only one pair of upper molars, thereby resembling the cats, with which, in correlation with their arboreal habits, they agree in their highly retractile claws, and the hairy surface of the under side of the metatarsus. About 15 in. is the length of the type species. In West Africa the linsangs are represented by _Poiana richardsoni_, a small species with a spotted genet-like coat, and also with a narrow naked stripe on the under surface of the metatarsus, as in genets.

Here may be placed the two African spotted palm-civets of the genus _Nandinia_, namely _N. binotata_ from the west and _N. gerrardi_ from the east forest-region. In common with the true palm-civets, they have a dentition numerically identical with that of _Viverra_ and _Genetta_, but the cusps of the hinder premolars and molars are much less sharp and pointed. They are peculiar in that the wall of the inner chamber of the auditory bulla never ossifies, while the paroccipital process is not flattened out and spread over the bulla. In this respect they resemble the Miocene European genus _Amphictis_, as they do in the form of their teeth, so that they may be regarded as nearly related to the ancestral _Viverridae_, and forming in some degree a connecting link between the present and the next subfamily. _Nandinia_ is also peculiar in possessing a kind of rudimentary marsupial pouch. Apparently _Eupleres goudoti_, of Madagascar, which has been generally classed in the _Herpestinae_, is a nearly related animal, characterized by the reduction of its dentition, due to insectivorous habits (fig. 3); the canines being small, the anterior premolars canine-like, and the hinder premolars molariform. It is a uniformly-coloured creature of medium size.

[Illustration: FIG. 3.--Skull of _Eupleres goudoti_.]

The palm-civets, or paradoxures, constituting the Asiatic genus _Paradoxurus_, have, as already stated, the following dental formula, viz. i. 3/3, c. 1/1, p. 4/4, m. 2/2, total 40; the cusps of the molars being low and blunted, and these teeth in the upper jaw much broader than in the civets. The head is pointed in front, with small rounded ears; the limbs are of medium length, with the soles of the feet almost completely naked, and fully retractile claws; while the long tail is not prehensile and clothed with hair of moderate length. Spots are the chief type of marking. The vertebrae number C. 7, D. 13, L. 7, S. 3, Ca. 29-36. Numerous relatively large species ranging from India to Borneo, Sumatra and Celebes, with one in Tibet, represent the genus. Nearly allied are _Arctogale leucotis_, with a wide distribution, and _A. trivirgata_, of Java, both longitudinally striped species, with small and slightly separated molars, and a prolonged bony palate (see PALM-CIVET).

The binturong (_Arctictis binturong_) has typically the same dental formula as the last, but the posterior upper molar and the first lower premolar are often absent. Molars small and rounded, with a distinct interval between every two, but formed generally on the same pattern as _Paradoxurus_. Vertebrae: C. 7, D. 14, L. 5, S. 3, Ca. 34. Body elongated; head broad behind, with a small pointed face, long and numerous whiskers, and small ears, rounded, but clothed with a pencil of long hairs. Eyes small. Limbs short, with the soles of the feet broad and entirely naked. Tail very long and prehensile. Fur long and harsh. Caecum extremely small. The binturong inhabits southern Asia from Nepal 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 is slow and cautious in its movements, chiefly if not entirely arboreal, and appears to feed on vegetables as well as animal substances (see BINTURONG).

_Hemigale_ is another modification of the paradoxure type, represented by _H. hardwickei_ of Borneo, an elegant-looking animal, smaller and more slender than the paradoxures, of light grey colour, with transverse broad dark bands across the back and loins.

_Cynogale_ also contains one Bornean species, _C. bennetti_, a curious otter-like modification of the viverrine type, having semi-aquatic habits, both swimming in the water and climbing trees, living upon fish, crustaceans, small mammals, birds and fruits. The number and general arrangement of the teeth are as in _Paradoxurus_, but the premolars are peculiarly elongated, compressed, pointed and recurved, though the molars are tuberculated. The head is elongated, with the muzzle broad and depressed, the whiskers are very long and abundant, and the 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 grey.

In the mongoose group, or _Herpestinae_, the tympanic or anterior portion of the auditory bulla is produced into an ossified external auditory meatus of considerable length; while the paroccipital process never projects below the bulla, on the hinder surface of which, in adult animals, it is spread out and completely lost. The toes are straight, with long, unsheathed, non-retractile claws.

In the typical mongooses or ichneumons, _Herpestes_, the dental formula is i. 3/3, c. 1/1, p. (4 or 3)/(4 or 3), m. 2/2; total 40 or 36; the molars having generally strongly-developed, sharply-pointed cusps. The skull is elongated and constricted behind the orbits. The face is short and compressed, with the frontal region broad and arched. Post-orbital processes of frontal and jugal bones well developed, generally meeting so as to complete the circle of the orbit behind. Vertebrae: C. 7, D. 13, L. 7, S. 3, Ca. 21-26. Head pointed in front. Ears short and rounded. Body 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. Soles of fore-feet and terminal portion of those of hind-pair naked; under surface of metatarsus clothed with hair. 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 ringed. The genus is common to the warmer parts of Asia and Africa, and while many of the species, like the Egyptian _H. ichneumon_ and the ordinary Indian mongoose, _H. mungo_, are pepper-and-salt coloured, the large African _H. albicauda_ has the terminal two-thirds of the tail clothed with long white hairs (see ICHNEUMON).

The following distinct African and Malagasy generic representatives of the subfamily are recognized, viz. _Helogale_, with 3/3 premolars, and containing the small South African _H. parvula _and a variety of the same. _Bdeogale crassicauda_ and two allied tropical African species differ from _Herpestes_ in having only four toes on each foot. The orbit is nearly complete, and the tail of moderate length and rather bushy. In _Cynictis_, which has the orbit completely closed, there are five front and four hind toes; and the skull is shorter and broader than in _Herpestes_, rather contracted behind the orbits, the face short, and the anterior chamber of the auditory bulla very large. The front claws are elongated. Includes only _C. penicillata_ from South Africa.

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 closely pressed hairs, and without any median groove. The South African _Rhynchogale muelleri_, a reddish animal with five toes to each foot and 4/4 (abnormally 5/5) premolars, alone represents the first genus. The cusimanses (_Crossarchus_), which differ by having only 3/3 premolars, and thus a total of 36 teeth, include, on the other hand, several species. The muzzle is elongated, the claws on the fore-feet are long and curved, the first front toe is very short; the under surface of the metatarsus naked; and the tail shorter than the body, tapering. Fur harsh. Includes _C. obscurus_, the cusimanse, a small burrowing animal from West Africa, of uniform dark-brown colour, _C. fasciatus, C. zebra, C. gambianus_ and others. Lastly, we have _Suricata_, a more distinct genus than any of the above. The dental formula is as in the last, but the teeth of the molar series are remarkably short in the antero-posterior direction, corresponding with the shortness of the skull generally. Orbits complete behind. Vertebrae: C. 7, D. 15, L. 6, S. 3, Ca. 20. Though the head is short and broad, the nose is pointed and rather produced and movable, while the ears are very short. Body shorter and limbs longer than in _Herpestes_. Toes 4-4. Claws on fore-feet very long and narrow, arched, pointed and subequal. Hind-feet with shorter claws, soles hairy. Tail rather shorter than the body. One species only is known, the meerkat or suricate, _S. tetradactyla_, a small grey-brown animal, with dark transverse stripes on the hinder part of the back, from South Africa.

The names _Galidictis, Galidia_ and _Hemigalidia_ indicate three generic modifications of the _Herpestinae_, all inhabitants of Madagascar. The best-known, _Galidia 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 ringed with darker brown. _Galidictis vittata_ and _G. striata_ chiefly differ from the ichneumons in their coloration, being grey with parallel longitudinal stripes of dark brown.

Considerable diversity of opinion prevails with regard to the serial position of the aard-wolf, or maned jackal (_Proteles cristatus_), of southern and eastern Africa, some authorities making it the representative of a family by itself, others referring it to the _Hyaenidae_, while others again regard it as a modified member of the _Viverridae_. After all, the distinction either way cannot be very great, since the two families just named are intimately connected by marks of the extinct _Ictitherium_, With the _Viverridae_ it agrees in having the auditory bulla divided, while in the number of dorsal vertebrae it is hyena-like. The cheek-teeth are small, far apart, and almost rudimentary in character (see fig. 4), and the canines long and rather slender. The dental formula is i. 3/3, c. 1/1, p. m. 4/(3 or 4); total 30 or 32. Vertebrae: C. 7, D. 15, L. 5, S. 2, Ca. 24. The fore-feet with five toes; the first, though short, with a distinct claw. The hind-feet with four subequal toes; all, like those of the fore-foot, furnished with strong, blunt, non-retractile claws (see AARD-WOLF).

Hyena tribe.

The hyenas or hyaenas (_Hyaenidae_) differ from the preceding family (_Viverridae_) in the absence of a distinct vertical partition between the two halves of the auditory bulla; and are further characterized by the absence of an alisphenoid canal, the reduction of the molars to 1/1, and the presence of 15 dorsal vertebrae. The dental formula in the existing forms (to which alone all these remarks apply) is i. 3/3, c. 1/1. p. 4/3 m. 1/1; total 34; the teeth, especially the canines and premolars, being very large, strong and conical. Upper sectorial with a large, distinctly trilobed blade and a moderately developed inner lobe 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 _Felidae_. Lower sectorial consisting of little more than the bilobed blade. Zygomatic arches of skull very wide and strong; and sagittal crest high, giving attachment to very powerful biting muscles. Orbits incomplete behind. Vertebrae: C. 7, D. 15, L. 5, S. 4, Ca. 19. Limbs rather long, especially the anterior pair, digitigrade, four subequal toes on each, with stout non-retractile claws, the first toes being 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. 4.--Skull and Dentition of Aard-Wolf (_Proteles cristatus_.)]

The three well-characterized species of _Hyaena_ are divisible into two sections, to which some zoologists assign generic rank. In the typical species the upper molar is moderately developed and three-rooted; and an inner tubercle and heel more or less developed on the lower molar. Ears large and pointed. Hair long, forming a mane on the back and shoulders. Represented firstly by _H. striata_, the striped hyena of northern and eastern Africa and southern Asia; and _H. brunnea_ of South Africa, in some respects intermediate between this and the next section. In the second section, forming the subgenus _Crocuta_, the upper molar is extremely small, two- or one-rooted, often deciduous; the lower molar without trace of inner tubercle, and with an extremely small heel. Ears moderate, rounded. Hair not elongated to form a mane. The spotted hyena, _Hyaena (Crocuta) crocuta_, of which, like the striped species, there are several local races, represents this group, and ranges all over Africa south of the Sahara. In dental characters the first section inclines more to the _Viverridae_, the second to the _Felidae_; or the second may be considered as the more specialized form, as it certainly is in its visceral anatomy, especially in that of the reproductive organs of the female. (See HYENA.)

(B) _Arctoidea_.--So far as the auditory region of the skull is concerned, the existing representatives of the dog tribe or _Canidae_ are to a great extent intermediate between the cat and civet group (_Aeluroidea_) on the one hand, and the typical representatives of the bear and weasel group on the other. They were consequently at one time classed in an intermediate group--the Cynoidea; but fossil forms show such a complete transition from dogs to bears as to demonstrate the artificial character of such a division. Consequently, the dogs are included in the bear-group. In this wider sense the Arctoidea will be characterized by the tympanic bone being disk-shaped and forming the whole of the outer wall of the tympanic cavity; the large size of the external auditory meatus or tube; and the large and branching maxillo-turbinal bone, which cuts off the naso-turbinal and two adjacent bones from the anterior nasal chamber. The tympanic bulla has no internal partition. There is a large carotid canal. Cowper's glands are lacking; and there is a large penial bone.

Dog tribe.

From all the other members of the group the _Canidae_ are broadly distinguished (in the case of existing forms) by the large and well-developed tympanic bulla, with which the paroccipital process is in contact. An alisphenoid canal is present. The feet are digitigrade, usually with five (in one instance four) front and always four hind-toes. The molars--generally 2/3--have tall cusps, and the sectorials are large and powerful (figs. 1 and 2). The intestine has both a duodeno-jejunal flexure and a caecum. A prostate gland is present; but there are no glands in the vasa deferentia; the penial bone is grooved; and anal glands are generally developed. The distribution of the family is cosmopolitan. The normal dentition is i. 3/3, c. 1/1, p. 4/4, m. 2/3; total 42; thus differing from the typical series only by the loss of the last pair of upper molars (present in certain extinct forms). In the characters of the teeth the group is the most primitive of all Carnivora. Typically the upper secterial (fig. 1, II) consists of a stout blade, of which the anterior cusp is almost obsolete, the middle cusp large, conical and pointed backwards, and the posterior cusp in the form of a compressed ridge; the inner lobe is very small, and placed at the fore part of the tooth. The first molar is more than half the antero-posterior length of the sectorial, and considerably wider than 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 sectorial (fig. 2, II) is a large tooth, with a strong compressed bilobed blade, the hinder lobe being considerably the larger and more pointed, a small but distinct inner tubercle placed at the hinder margin of the posterior lobe of the blade, and a broad, low, tuberculated 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.

Views differ in regard to the best classification of the _Canidae_, some writers adopting a number of generic groups, while others consider that very few meet the needs of the case. In retaining the old genus _Canis_ in the wide sense, that is to say, inclusive of the foxes, Professor Max Weber is followed. The best cranial character by which the different members of the family may be distinguished is that in dogs, wolves and jackals the post-orbital process of the frontal bone is regularly smooth and convex above, with its extremity bent downwards, whereas in foxes the process is hollowed above, with its outer margin (particularly of the anterior border) somewhat raised. This modification coincides in the main with the division of the group into two parallel series, the Thooids or Lupine forms and Alopecoids or Vulpine forms, characterized by the presence of frontal air-sinuses in the former, which not only affects the external form 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, as many of the South American _Canidae_ have the long bushy tails of foxes and the skulls of wolves.

[Illustration: FIG. 5.--The African Hunting-Dog (_Lycaon pictus_).]