Chapter XI: The Order Carnivora (2)

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_.

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.

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.

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.

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.

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_.

_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.

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.

_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_.

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.”

_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.

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.

_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.

_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_.

_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.

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.

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.

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.

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.

_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.