Cambridge Natural History Mammalia/Chapter XIII

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CHAPTER XIII

CARNIVORA[257]—FISSIPEDIA

Order VII. CARNIVORA

This order may be thus defined:—Small to large quadrupeds, terrestrial, arboreal, or aquatic, of usually carnivorous habits. The teeth have generally sharp and cutting edges, and the canines are well developed; the incisors are small, and four to six in number. The number of toes is never less than four. There are usually strong and sharp claws. The clavicles are incomplete or absent. In the hand the scaphoid and lunar bones are always united. The brain is well developed, and the hemispheres are well convoluted. The stomach is always simple, while the caecum, if present, is always small. The members of this group have a deciduate and zonary placenta.

The fewness of the characters used in the above definition is chiefly owing to the fact that the Seals and Sea-lions, although they are referable without a doubt to this order, have undergone in their metamorphosis into aquatic animals so many changes that some of the main features in the structure of their terrestrial relatives have been lost. This group will, however, be again characterised. We shall deal at present with the land division of the Carnivora, the Carnivora Fissipedia as they are generally termed. The name is of course given to them to distinguish them from the corresponding division of the Pinnipedia. In the latter group the feet and hands are modified into "fins"; in the other the fingers and toes are cleft, as with terrestrial beasts generally.

Sub-Order 1. FISSIPEDIA.

Cambridge Natural History Mammalia Fig 190.png

Fig. 190.—Under surface of the cranium of a Dog. × ½. apf, Anterior palatine foramen; AS, alisphenoid; as, posterior opening of alisphenoid canal; BO, basioccipital; BS, basisphenoid; cf, condylar foramen; eam, external auditory meatus; ExO, exoccipital; flm, flp, foramen lacerum medium and posterius; fm, foramen magnum; fo, foramen ovale; Fr, frontal; fr, foramen rotundum; gf, glenoid fossa; gp, post-glenoid process; Ma, malar; Mx, maxilla; oc, occipital condyle; op, optic foramen; Per, mastoid portion of periotic; pgf, post-glenoid foramen; Pl, palatine; PMx, premaxilla; pp, paroccipital process; ppf, posterior palatine foramen; PS, presphenoid; Pt, pterygoid; sf, sphenoidal fissure or foramen lacerum anterius; sm, stylomastoid foramen; SO, supraoccipital; Sq, zygomatic process of squamosal; Ty, tympanic bulla; Vo, vomer. (From Flower's Osteology.)

A very marked feature of the terrestrial Carnivora is to be found in the structure of the teeth. The incisors are nearly always six, and are somewhat feebly developed in many cases. The canines are almost invariably very large strong teeth, and are always present. In some of the extinct Cats they reached enormous dimensions. The number of cheek teeth is not always identical; but the last premolar in the upper jaw and the first true molar in the lower jaw, known as the "carnassial" or "sectorial" teeth, mark a difference in structure between the anterior and the posterior crushing teeth; those in front of the carnassial tooth have cutting edges, and are often merely small, conical teeth; those behind have broader crowns and are tuberculate; those of simpler forms often trituberculate; those of others with numerous tubercles. The carnassial tooth is often, but by no means always, very much larger and especially longer than the rest of the molar and premolar series. It is less pronounced in some of the omnivorous Arctoidea. The skull of the Carnivora is longer in the more primitive types, such as the Canidae, and shorter in the more specialised Felidae. The orbit is hardly ever completely shut off by bone, though the postorbital process of the frontal sometimes approaches the corresponding upward process of the zygomatic arch. The palate, which is completely ossified, sometimes reaches back for some distance behind the teeth; it always extends as far as the last molar. The tympanic bulla is often very inflated, and if flatter, as in the Bears, is at any rate large and conspicuous. The lower jaw has a high coronoid process, and the condyle is transversely elongated, this part of the bone being rolled into an almost cylindrical form; it fits very closely into the glenoid cavity, and the articulation is thereby very strict—an obvious advantage in a creature with so great a need for power of jaw.

Cambridge Natural History Mammalia Fig 191.png

Fig. 191.—A, Atlas of Dog. Ventral view, × ½. B, Axis of Dog. Side view, × ⅔. o, Odontoid process; pz, posterior zygapophysis; s, spinous process; sn, foramen for first spinal nerve; t, transverse process; v, vertebrarterial canal. (From Flower's Osteology.)

In the vertebral column the atlas always has large wing-like processes; the spine of the axis vertebra has a long antero-posteriorly elongated form. The transverse processes of the fourth to the sixth cervicals are, as a rule, double. These features, however, though characteristic of the Carnivora are not by any means distinctive. The true sacrum consists of but a single vertebra to which the ilia are attached; but at most two other vertebrae are fused with this. The clavicle is always small and sometimes quite rudimentary, or even absent. The spine of the scapula is well developed, and almost equally divides the surface of that bone. The digits of the Carnivora are mostly five, and are never less than four. The mode of progression may be digitigrade or plantigrade, and the intermediate semidigitigrade mode of walking also occurs. The brain in all Carnivora is large and well convoluted. The arrangement of the convolutions is characteristic. There are three or four gyri disposed round each other, of which the lowest surrounds the Sylvian fissure. The stomach in these creatures is always simple in form, without subdivisions. The caecum is never large, and may be, as in the Bear tribe, completely absent.

Cambridge Natural History Mammalia Fig 192.png

Fig. 192.—Brain of Dog. A, Ventral; B, dorsal; C, lateral aspect. B.ol, Olfactory lobe; Cr.ce, crura cerebri; Fi.p, great longitudinal fissure; HH, HH1, lateral lobes of cerebellum; Hyp, hypophysis; Med, spinal cord; NH, medulla oblongata; Po, pons Varolii; VH, cerebral hemispheres; Wu, middle lobe (vermis) of cerebellum; I-XII, cerebral nerves. (From Wiedersheim's Comparative Anatomy.)

The distribution of the Carnivora is world-wide, excluding only the Australian region, if, as seems probable, the Dingo of that region is an introduced species. The most striking features in their distribution are perhaps the following:—There are no Bears in the Ethiopian region or in Madagascar, and but a single species in the Neotropical. The only Carnivora in Madagascar are the Viverridae, and of the seven genera there found six are peculiar. The Procyonidae are nearly entirely New World in range; out of sixteen genera of Mustelidae only five are New World, and only two of those are peculiar to the American continent. The Hyaenidae are limited to the Old World.

The classification of the Carnivora is a matter which is difficult, and which has therefore been very variously effected. It is unfortunate that the classification of Flower (based upon the researches of H. N. Turner as well as his own, and accepted by Mivart) should fail when applied to fossil forms. For it separates with great clearness the existing genera into three great divisions, the Cynoidea, Aeluroidea, and Arctoidea, definable by visceral as well as by osteological characters. The apparent anomaly, too, of a single supposed Viverrine genus, to wit Bassariscus, existing in America, while all the rest of its kin are Old-World forms, was shown by his characters to be neither an anomaly nor a fact. It will be better, therefore, to divide the Carnivora into the families, Felidae, Machaerodontidae, Viverridae, Hyaenidae, Canidae, Ursidae, Procyonidae, and Mustelidae, indicating at the same time the reasons for and against retaining the three divisions of Sir W. Flower.

Fam. 1. Felidae.[258]—This family includes only the Cats (i.e. Lions, Tigers, "Cats," Hunting Leopard, etc.), and is to be distinguished by the following characters:—In the skull the auditory bulla is much inflated, and there is an internal septum; the paroccipital processes are flattened against the bullae. There is no alisphenoidal canal. The dental formula is I 3, C 1, Pm 3 to 2, M 1. The carnassial tooth of the upper jaw has three lobes to the blade; that of the lower jaw is without an inner cusp. The digits are five on the fore-feet, four on the hind. The caecum is present and small. This family contains but two genera, Felis and Cynaelurus.

Cambridge Natural History Mammalia Fig 193.png

Fig. 193.—Section of auditory bulla of Tiger. am, Auditory meatus; BO, basioccipital; e, Eustachian canal; ic, oc, two chambers of bulla divided by s, septum; *, their aperture of communication; Pt, periotic; Sq, squamosal; t, tympanic ring. (From Flower's Osteology.)

The genus Felis is very wide in its distribution, being common to both the Old and the New Worlds. Its distinctive characters, as opposed to Cynaelurus, are mainly the following:—The claws are retractile, and the retractility is more markedly developed than in the Cheetah. The molar is not so nearly in a line with the other teeth; the upper carnassial, moreover, has an inner tubercle. The legs are relatively shorter.

The complete retractility of the claws is a very distinctive feature of the true Cats. It is brought about in this way: the terminal joint of the toe, which is clad with the claw, folds back into a sheath by the outer side of or above the middle phalanx. It is held in this position by a strong ligament. The flexor muscles straighten the phalanx which bears the claw, so that the natural position for the animal is to be in a state of retracted claws, which of course preserves them from friction; when wanted for aggressive purposes, they are pulled into sight by the action of the muscles already mentioned.

Cambridge Natural History Mammalia Fig 194.png

Fig. 194.—The phalanges of the middle digit of the manus of the Lion (Felis leo). × ½. a, The central portion forming the internal support to the horny claw; b, the bony lamina reflected around the base of the claw; ph1, proximal phalanx; ph2, middle phalanx; ph3, ungual phalanx. (From Flower's Osteology.)

Much has been written as to the shape of the pupil of the Cat's eye. Some careful observations upon the matter have been made by Dr. Lindsay Johnson,[259] who found that out of 180 Domestic Cats 111 had round pupils, and that in 19 the shape was a pointed oval, intermediate conditions being offered by the rest. These 180 comprised males and females of many varieties. When the pupil of the Cat's eye contracts, it forms a vertical slit with two pin holes, one at each end, through which alone light appears to enter. In the Genet and the Civet the contraction of the pupil is as in the Cat. In the Lion, Tiger—in fact apparently in all the large Cats—the pupil retains its circular shape even when contraction is fully effected. Dr. Johnson has, furthermore,[260] made some interesting experiments upon the Seal's eye—a creature which has, of course, to exert its powers of vision in two media, and from one to the other. This is effected by dilatation of the pupil when in the water, and its contraction to a vertical slit with parallel margins and rounded ends when in the air, the contraction being to some extent at least under the influence of the animal's will.

The coloration of these creatures is very varied: spots of black, or bordered with black upon a more or less tawny ground-colour, is the prevailing pattern. Stripes are also met with, as in the Tiger, but these are usually cross stripes,[261] while in the related Viverridae there are many examples of longitudinal stripes. Finally, many Cats, as for instance the Puma and the Eyra, are "self-coloured"—have, that is to say, a uniform tint. Just as the unstriped Horse sometimes shows traces of the former existence of stripes, so the self-coloured Cats are occasionally spotted when young; this is markedly so in the case of the Puma; while the Lion is spotted as a cub, and in the adult—particularly in the lioness—there are distinct indications of these spots. It is evident, therefore, that there are grounds for regarding a spotted condition to be antecedent, at least in some cases, to a uniform colour. There are divers explanations of these hues and of these changes. It is held by many that the coloration has a relation to the habits of the creature: the spotted Cats, it is pointed out, are largely arboreal; this is eminently so with the Jaguar at any rate; and in an arboreal creature the spots, it is said, give the impression of flecks of sunlight broken up by foliage. On the other hand, the self-coloured Cats of a sandy to earthen hue assimilate in tint with a sandy or stony soil. The stripes of the Tiger, it is thought, approximate to the tall parallel stems of grasses and other plants in the dense cover in which it lives. In favour of these views is undoubtedly the fact that in other mammals and other animals belonging to quite different groups the same four plans of coloration are met with. Spots and cross stripes are found in the Marsupials; the young Tapir is spotted while the adult is self-coloured, and so forth. This last fact, however, serves to illustrate another view which has been put forward in explanation of these characteristic markings of the Felidae. Eimer has come to the conclusion that there is and has been a regular series of steps in the evolution of these markings. The primitive condition was, he thinks, a longitudinally striped one; the stripes then broke up into spots, and the spots rearranged themselves as transverse stripes; the self-coloured Puma and Lion are a final stage in this gradual evolution. In support of this is the fact that spots precede self-coloration in the individual growth of these animals. The exact sequence of these markings is, however, contradicted by Dr. Haacke's observations upon a certain Australian fish which is cross striped when young and longitudinally striped when adult, a precise reversal of what ought to occur on Eimer's view.

The Felidae are almost universally distributed with the exception, of course, of Australia and a good deal of the Australian region; the headquarters of the group are undoubtedly in the tropics of the Old World.

The characteristics of a few species of the Cat tribe will now be given. As there are at any rate forty-five species, this survey will have to be somewhat incomplete.

The Lion, F. leo, differs from all other species by the mane of the male. It is an inhabitant of Africa, India, and certain parts of Western Asia. Within the historic period it ranged into Europe. According to Sir Samuel Baker those of us who have not seen the Lion in his native haunts have never seen a really magnificent specimen of the brute; but other travellers disagree, and state that a captive Lion is often a finer animal—by reason, of course, of good feeding. Unlike the majority of Cats, the Lion cannot climb. His roar (which is so suggestive, towards its end, of that animal who once dressed himself up in his skin) is literally after his prey. The Lion, it is stated, does not roar except upon a full stomach. The Lion is mainly nocturnal in its habits, and is said to be not in the least dangerous if unprovoked in the daytime; but here again opinions differ. The tail of the animal is provided at the extremity with a slight claw, but it can hardly be sufficient for the animal to lash itself into a fury with it. A Lion will live for thirty or forty years, and will breed freely in captivity. The Gardens of the Zoological Society of Dublin have been famed for their success in breeding Lions; but more surprising still, this has been successfully accomplished in travelling menageries. The "desert" colour of the Lion is familiar to all. It is stated that the likeness to the parched soil of certain parts of Africa is greatly heightened by black patches in the mane, for in certain regions of that continent the arid yellow of the general environment is diversified by pieces of black lava. It is apparently a popular delusion to speak of the Maneless Lion of Guzerat. No doubt maneless Lions do come from there, but so do young and maneless Lions from other places; in short, it is simply a question of age, and old Lions from the Asiatic continent are as fully maned as those from Africa.

The Tiger, F. tigris, is an animal of about the same size as the Lion, distinguished, of course, by the stripes. The skeletons are much like those of other Cats; but the skull of the Tiger may be distinguished from that of the Lion by the fact that the nasal bones reach back beyond the frontal processes of the maxillae. The Tiger is an exclusively Asiatic beast, ranging northward into icy Siberia. The northern individuals have a closer fur, and have been quite unnecessarily separated as a distinct variety. Nine feet six inches is the size of the average full-grown Tiger; but the skins will stretch, a fact of which the sportsman will sometimes take advantage. A "man-eater" is a Tiger which has discovered "that it is far easier to kill a native than to hunt for the scarce jungle game." As with the Lion, the accounts of travellers differ enormously, particularly with regard to the strength of the creature. Some have said that a Tiger can easily lift a full-grown bullock and leap with it in the mouth over a considerable obstacle, a statement which is ridiculed by Sir Samuel Baker. Unlike the Lion, the Tiger can climb trees; it will also voluntarily enter the water, and can swim considerable rivers.

Mr. H. N. Ridley[262] observes that Tigers "habitually swim over to Singapore across the Johore Strait, usually by way of the intermediate islands of Pulau Ubin and Pulau Tekong. They make the passage at night, landing in the early morning. As so much of the coast is mangrove swamp, and the animals do not risk going through the mud, they are only able to cross where the shores are sandy, and thus they have regular starting- and landing-places."

The Tiger is mainly nocturnal, but begins its depredations towards five o'clock in the afternoon, before which it remains sleeping in shady thickets. If the weather is rainy and windy it becomes restless and wanders about earlier. Under the provocation of extreme hunger it will hunt during the daytime. Hunger, too, naturally produces extreme boldness. Mr. Ridley relates a story of four Tigers who walked up the steps of a house in search of the master of the house or his dog, and broke into it, the inhabitants retiring in their favour. The Malays have superstitions about Tigers, which are precisely paralleled by the man-and-wolf stories of Europe. "Certain people are supposed to have the power of turning into tigers for a short time, and resuming their human form at pleasure. The transformation commences tail first, and the human tiger is so completely changed that not only has it all the actions and appearance of the tiger, but on resuming its human form it is quite unconscious of what it has been doing in the tiger state." Mr. Ridley disputes the common stories as to man-eaters. If a Tiger has once tasted human flesh it does not always confine itself afterwards to that article of diet, nor is it only aged and comparatively toothless animals which hunt man. That they do take a large toll of coolies is an undoubted fact, and many are the artifices to prevent the rest from knowing the fate of one of their fellow-workmen, or of becoming acquainted with the presence in the neighbourhood of one of the dreaded beasts.

The Leopard or Panther, F. pardus, is, like the Lion, African and Asiatic in range. The animal is spotted with rosettes of black spots surrounding a central field of the tawny colour of the body generally. Some of the spots are solid and black. "The pantere like unto the smaragdyne" seems to be an inapt description of this Cat, unless indeed the eyes be referred to. The ancients ascribed to it a most fragrant odour. As with the Tiger, a northern variety of this Carnivore has a closer and longer fur. There is a tendency towards melanism in this animal, the black Leopard being comparatively common, particularly, it appears, in high lands. Several other variations in colour are known. These have received different specific names; but it seems that there is in reality but one species of Leopard. The Leopard can climb with the agility of any Cat. Sir S. Baker reserves the name Panther for large Leopards, which reach a length of 7 feet 6 inches. But there is no valid distinction between any two such varieties. The Leopard is as ferocious as the Tiger; and Sir Samuel Baker advises that the power of the human eye be not experimented with when meeting unarmed one of these brutes.

Cambridge Natural History Mammalia Fig 195.jpg

Fig. 195.—Snow Leopard. Felis uncia. × 120.

The Snow Leopard or Ounce, F. uncia, is a beautiful creature, confined to the highlands of Central Asia. The ground-colour is white, and the spots are larger than those of the ordinary Leopard. Two examples of this rather rare Carnivore have been recently on view in the Zoological Society's Gardens, London. The Clouded Leopard, F. nebulosa, is an animal of considerable size (6 feet total length).

The Fishing Cat, F. viverrina, of India and China, is about 3 feet 6 inches including the tail. Its black spots upon a grey-brown ground have a tendency to form longitudinal lines. It is in fact, on Eimer's theory, a case of longitudinal stripes breaking up into spots. It differs from the bulk of Cats by preying upon fish, though it is not known how it catches them. It also feeds upon the large snail Ampullaria. In addition to these there are twenty-four species of Cats found in the Old World, mainly in the Oriental region, of small to moderate size.

Cambridge Natural History Mammalia Fig 196.jpg

Fig. 196.—European Lynx. Felis lynx. × 112.

The European Lynx, F. lynx, has rather long legs, a short tail, and tufted, pointed ears. It has only two premolars in the upper jaw instead of the usual three. It seems to be doubtful whether the Asiatic Lynx can be distinguished from the European, but the Spanish form, F. pardina, does appear to be distinct. The Common Lynx, sometimes called F. canadensis, also ranges into America, where some other forms exist, known by the specific names of F. rufa and F. baileyi.

In America there are altogether sixteen species of Cats, if we allow three species of Lynx, none of which, however, does Dr. Mivart allow to be different from the European and Asiatic Lynx (F. lynx).

The largest of American Cats is the Jaguar, F. onca. This is an arboreal creature with a long, heavy body and short limbs. Its pelage is much like that of the Leopard, but the spots are larger and more definitely arranged in groups. There are a number of distinct rows of spots. The length of the body alone is not greater than 4 feet. They prey very largely on the Capybara, and upon turtles, which they surprise upon the sand when about to lay their eggs; the reptiles are turned upon their backs, so as to be incapable of escaping, and the Jaguar then easily devours them. The Jaguar will even pursue the turtle into the water, and will devour its eggs and the newly-hatched young.

Cambridge Natural History Mammalia Fig 197.jpg

Fig. 197.—Jaguar. Felis onca. × 115.

The Ocelot is another spotted American Cat. F. pardalis[263] ranges from Arkansas in North America southwards, its range corresponding with that of the Jaguar. Although small for one of the "larger cats," the Ocelot inspired with considerable respect Captain Dampier, who remarked of it: "The Tigre-cat is about the bigness of a bull-dog, with short truss, body shaped much like a mastiff, but in all things else, its head, the colour of its hair, the manner of its preying, much resembling the tigre, only somewhat less.... But I have wisht them farther off when I have met them in the woods; because their aspect appears so very stately and fierce."

Cambridge Natural History Mammalia Fig 198.jpg

Fig. 198.—Ocelot. Felis pardalis. × 110.

The Puma, F. concolor, the American Lion as it is called in the north, is a rather smaller animal than the last, and of a uniform tawny colour, tending to white on the abdomen and to a dark stripe along the back. The young, as already mentioned, are very distinctly spotted. Like the Tiger, the Puma can endure extremes of heat and cold; it is equally at home in the snow of North America and among the tropical forests and swamps of the south. It is a ferocious creature so far as concerns Deer, Lamas, Raccoons, even Skunks and Rheas, but, according to Mr. W. H. Hudson, will not attack man, and will even defend him against the Jaguar.[264] In captivity the Puma will purr like a Cat.

The Eyra, F. eyra, is another self-coloured American cat, which has a curious likeness to the totally distinct Cryptoprocta of Madagascar.

The Wild Cat of Europe, F. catus, is found over the greater part of Europe, and also in Northern Asia. It was undoubtedly common at one time in this country, though it appears never to have extended its range into Ireland. But the real Wild Cat is now rare in this island, and is confined to certain districts in Scotland. Plenty of alleged wild Cats have been seen and even shot; but these are too frequently merely feral Cats, i.e. domestic tabbies which have reverted to a hunting life. The real Wild Cat differs from the domestic races by the proportionately longer body and limbs, the shorter and thicker tail; the pads of the toes are not quite black. The period of the gestation of the Wild Cat, according to Mr. Cocks, is a week or so longer than that of any domestic Cat.

The Domestic Cat is in fact regarded as the descendant of the Eastern F. caffra, or (perhaps and) the closely-allied F. maniculata. It is highly probable, however, that after introduction into this country as a domestic animal it has interbred with the Wild Cat. Many allied species of Cats will interbreed, even two so far apart as the Lion and the Tiger. There are interesting archaeological and linguistic reasons for regarding the Domestic Cat as an importation. The legend of Dick Whittington's Cat points to it being a rare and valuable animal, which a tamed F. catus would not at that time have been. There was an enactment in Wales of a penalty against him who should kill the king's Cat, again suggestive of its rarity and consequent value. The very name "Puss" is a hint of a foreign origin. Some would derive it from Perse, and upon this is based the notion that the Cat is from Persia. But it seems that Puss is the same as Pasht and Bubastis, showing so far an Egyptian origin for the animal. The ancestral Cats mentioned above are natives of Egypt.[265]

The genus Cynaelurus, which includes but a single species, C. jubatus, the Cheetah or Hunting Leopard, is separated from Felis by a number of characters. In the first place the claws are non-retractile, or at least less retractile than those of the true Cats. It is, moreover, longer legged. The molar is more in a line with the other teeth of the jaw, and the upper carnassial tooth has no inner tubercle. Messrs. Windle and Parsons have lately pointed out many Dog-like features in the muscles. This animal is about as large as a Leopard, but has plain black spots. As its vernacular name implies it is used for sport, and is quite easily tameable. It will purr like the Puma. The Cheetah occurs in India, Persia, Turkestan, and also in Africa; the latter form is sometimes, though quite unnecessarily, separated as C. lanigera. The genus occurs fossil in the Siwalik deposits of India, the species being known as C. brachygnatha.

Fam. 2. Machaerodontidae.—This is a family of totally-extinct Cats which range from the Eocene down to the Pleistocene. Their general structure is like that of the Felidae; but they differ in a number of skeletal features. Thus there is an alisphenoid canal, and, as in Bears, there is a postglenoid foramen. There is also a distinct carotid foramen, which does not occur in the true Cats. The teeth are often distinguished by the huge size of the superior canines, which are "weapons for penetrating wounds, without rival among carnivorous animals." These must have been displayed at the sides of the chin when the mouth was closed, and it has even been suggested that the animal possessing these exaggerated canines could hardly have properly closed its mouth. The lower canines were often on the contrary much reduced, and in fact incisor-like. In tracing the series of these Cats we find a gradual reduction of the teeth from a more nearly complete number down to the specialised dentition of the existing Cats. The genus Proaelurus, Miocene in range, had four premolars in each jaw, and two molars in the lower and one in the upper. This is the greatest number of teeth found in any member of the group.

The resemblance of this genus to Cryptoprocta has been insisted upon. Archaelurus has suffered a reduction, since one premolar in the lower jaw has disappeared, its formula being thus I 3/3 C 1/1 Pm 4/3 M 1/2. The next stage is shown by Dinictis with three premolars in both jaws. There are a good many species of this genus which are all American and Miocene. This genus has five toes upon the hind-feet, and was probably plantigrade. It had retractile claws.

In the genus Nimravus the dental formula is still further reduced. Another premolar of the lower jaw has gone, the formula being thus I 3/3 C 1/1 Pm 3/2 M 1/2. Nimravus gomphodus was a Carnivore about the size of a Panther. It has no third trochanter upon the femur, which process is present in the corresponding bone of Dinictis. Pogonodon was an equally large animal in which the premolars were three in each jaw, but the molars have become reduced to one in the lower, as they have in this and other genera in the upper jaw. Finally, Hoplophoneus has acquired the dentition of existing Cats.

The Machaerodons, however, show examples with a yet more reduced dentition than that of the most reduced existing Cat, viz. the Lynx, which has only two premolars in each jaw and one molar. In Eusmilus the molar in both jaws is single, and there is but one premolar in the lower jaw.

The genus Machaerodus itself, which appears to include Smilodon, is referred by Cope to the true Cats, and not to the Nimravidae, as he terms the family which we have called here the Machaerodontidae. These creatures are known as "Sabre-toothed Tigers," and were of very wide distribution, occurring in South America as well as in Europe and North America. "As nothing," remarks Professor Cope, "but the characters of the canine teeth distinguished these from typical felines, it is to these that we must look for the cause of their failure to continue. Professor Flower's suggestion appears to be a good one, viz. that the length of these teeth became an inconvenience and a hindrance to their possessors. I think there can be no doubt that the huge canines in the Smilodons must have prevented the biting off of flesh from large pieces, so as to greatly interfere with feeding, and to keep the animals in poor condition. The size of the canines is such as to prevent their use as cutting instruments excepting with the mouth closed; for the latter could not have been opened sufficiently to allow any object to enter it from the front. Even when it opens so far as to allow the mandible to pass behind the apices of the canines, there would appear to be some risk of the latter being caught on the point of one or the other canine, and forced to remain open, causing early starvation. Such may have been the fate of the fine individual of the S. neogaeus, Lund, whose skull was found in Brazil by Lund, and which is familiar to us through the figures of de Blainville."

Machaerodus is placed among the Felidae on account of the fact that the condyloid and carotid foramina unite with the foramen lacerum posterius. But as in at least one species, M. palmidens, there is an alisphenoid canal, which, however, has disappeared in the more recent American forms, it seems permissible to retain the genus in the family Machaerodontidae though its existence reduces the differential character of that family to a minimum. The genus goes back to the Eocene.

Fam. 3. Viverridae.—The Civets, Genets, and their kind differ from the Cats in a number of points. They form, however, by no means so uniform an assemblage as do the Cats; so that the difficulty is, as Dr. Mivart has remarked, not to divide them into sub-families, but to avoid making too many. But before proceeding to subdivide the family we shall describe the characters of the family and contrast them with those of the Felidae.

All the Viverridae are comparatively small creatures. The head and body are more elongate than in the Cats. The fingers and toes are generally five; but there are some (e.g. Cynictis) where the formula of the toes is as in the Cats, i.e. four on the hind-foot. In the Suricate the fingers are also reduced to four. The claws are perhaps never completely retractile,[266] and often are not at all so. The dental formulae of the genera differ considerably; but in the majority there are more teeth than in the Felidae. The well-known sharp-pointed, conical papillae of the Cat's tongue are not present. The majority have a scent gland beneath the tail, from which the perfume civet is derived. There are a number of osteological characters which differentiate the two families; thus the alisphenoid canal is sometimes present. The bulla is divided, as in the Cats, but is externally constricted.

It seems clear from some at any rate of the characters, i.e. the more complete dentition, the five-fingered hands and feet, the non-retractile claws, etc., that the Civets are on a lower level of specialisation than are the Cats.

Sub-Fam. 1. Euplerinae.—The genus Eupleres is in many ways the most aberrant type of Viverrid, and is placed in a sub-family, Euplerinae. Its salient feature is the very peculiar dentition: peculiar in the small size of the canines, the canine-like character of the anterior premolars, and the resemblance of the premolars to molars. In some of the characters of the teeth, Eupleres is Insectivore-like, and was formerly grouped with that family. There are four premolars and two molars in each jaw on each side. It has five toes upon both fore- and hind-limbs; the skull is very slender. It has no alisphenoid canal. The only species, E. goudotii, is of an olive-grey colour, with dark bands across the shoulders in the young. The nose and upper lip are grooved. There are no scent glands. It appears to burrow in the ground, and possibly contents itself with a diet of worms. Eupleres is a native of Madagascar, where all the most peculiar Viverridae live.

Sub-Fam. 2. Galidictiinae.—Mivart has placed in this sub-family the three Mascarene genera, Galidia, Hemigalidia, and Galidictis. In them the orbit is not enclosed by bone; there is no alisphenoid canal, and there are five toes and fingers.

Galidia consists of but one species, G. elegans, of a chestnut brown colour, with a tail ringed with black. The claws are not retractile. The scent gland is absent. There are five digits upon both hand and foot. There are three premolars and two molars on each side of each jaw. The caecum is (for an Aeluroid) long, and pointed at the apex; it is quite twice the length of that of Genetta.

Closely allied to Galidia is the genus Hemigalidia, of which there are two species. It is distinguished from the last genus by the non-annulated tail. It also differs in the dental formula, which is for the molars Pm 4/3 M 2/1. This animal is termed by Buffon the Vansire. He correctly enumerates its grinders, and distinguishes it from the Ferret!

Galidictis is a third genus from Madagascar containing two species, one of which has been unfortunately named G. vittata, leading perhaps to some confusion with the totally distinct Galictis vittata. As in the last two genera the digits are five. The dental formula is that of Galidia. It is distinguished from the other two genera of its sub-family by the longitudinal brown striping of the upper part of the greyish body.

Sub-Fam. 3. Cryptoproctinae.Cryptoprocta[267] represents a special sub-family, Cryptoproctinae, and includes only a single species, the Fossa (C. ferox) of Madagascar. It is the largest Carnivore of Madagascar, being about twice the size of a Cat, but with an elongated body; the colour is a tawny brown with no striping. The animal is active and lithe in its motions, and is said to be of almost unexampled ferocity in disposition. Its exact systematic position has been much discussed. By Zittel it is placed in a sub-family (including the extinct Proaelurus and Pseudaelurus) of the Felidae. Mivart and Lydekker, on the other hand, regard it as a genus of the Viverridae. The dental formula of the molars, Pm 3/3 M 1/1, is more like that of the Felidae than of the Viverridae, and the teeth are more Feline in structure. The claws of the feet are retractile. As to internal structure the Fossa agrees largely with the Viverridae, but then this family has no very marked points of difference from the Felidae; but where the anatomy does diverge from that of the Felidae it approaches the Viverridae, especially in the muscular system.

Cambridge Natural History Mammalia Fig 199.jpg

Fig. 199.—Fossa. Cryptoprocta ferox. × 16.

The remaining and by far the larger number of genera of Civets are grouped by Professor Mivart in two sub-families: the Viverrinae, including the genera Viverra, Viverricula, Fossa, Genetta, Prionodon, Poiana, Paradoxurus, Arctogale, Hemigale, Arctictis, Nandinia, and Cynogale; and the Herpestinae, including the genera Herpestes, Helogale, Cynictis, and probably Bdeogale and Rhynchogale. In the Viverrinae the digits are always five, the claws are more or less retractile, the prescrotal scent glands are usually present, and the anus does not open into a sac. On the other hand, the Herpestinae are characterised by the non-retractility of the claws, the absence of the glands in question, and the fact that the anus does open into a terminal sac.

Sub-Fam. 4. Viverrinae.Viverra includes the true Civets. The genus, save for one African species, is Oriental in range. The molar formula is the complete one for the Viverridae, viz. Pm 4/4 M 2/2. The secretion of the prescrotal gland of V. civetta yields the civet of commerce.

The "Rasse," genus Viverricula, has been separated generically from the true Civets. It is, remarkably enough, common to both Madagascar[268] and many parts of the Oriental region. It is, moreover, capable of climbing trees, which its relatives are not. It has no mane like Viverra and is of slighter build.

Cambridge Natural History Mammalia Fig 200.jpg

Fig. 200.—Civet Cat. Viverra civetta. × 16.

Prionodon or Linsang differs from the last two genera in the loss of an upper molar. It thus approaches the Cats, with which it also agrees in the furry feet. It is a purely Oriental genus. It also resembles the Cats in that the claws are apparently quite retractile, a feature not common among the group. There are three species of the genus. P. pardicolor has large black spots and a ringed tail. Its body is some 15 inches in length. Dr. Mivart has commented upon the particularly small caecum, which, like that of Arctictis, seems to be on the verge of disappearance.

Genetta, including the Genets, is almost purely African. It has the full tooth formula of Viverra; but is to be distinguished by the absence of a scent pouch, and by a naked strip of skin running up the metatarsus. These animals are all brownish yellowish to greyish with darker spots. The Common Genet, G. vulgaris, is South European, and just gets into Asia; it is also North African. The Genet, an animal "with an appetite for petty carnage," is one of those smaller Carnivora which are possibly to be regarded as meant by the word γαλῆ, and appear to have "functioned" as Cats among the Greeks. So recently as the times of Belon we are told (by him) that Genets were common and tame at Constantinople.

Poiana, containing a single African species, a spotted and entirely Genet-like animal, has been separated as a distinct genus. Dr. Mivart, however, holds it to be a Prionodon which has acquired a Genet-like tarsus.

Arctictis, containing but one species, A. binturong, the Binturong, is in some ways an exceptional form. It is a black arboreal creature of not very wide range in the Oriental region, with a fully prehensile tail. This feature and its plantigrade foot with naked sole have led to its being regarded as more allied to the Arctoidea. It is, however, undoubtedly an ally of Paradoxurus. The caecum is small, or may be quite absent. The dentition is I 3/3 C 1/1 Pm 4/3 M 2/2. The structure of the animal has been investigated by Garrod.[269]

The genus Fossa is a Viverrine confined to Madagascar. There is but one species, F. daubentoni, the "Fossane." It is distinguished from Viverra by the presence of two bare spots on the under surface of the metatarsus in the hind-limb, and by the absence of a scent pouch. The animal is not much spotted and striped, but the striping in the young is much more marked.

Of the genus Paradoxurus there are some ten or a dozen species, belonging entirely to the Oriental region. The teeth are as in Viverra, but occasionally the molars are reduced to one. The pupils are vertical. The tail though long is not prehensile, "but the animal appears to have the power of coiling it to some extent, and in caged specimens the coiled condition not unfrequently becomes confirmed and permanent" (Blanford). This fact accounts for the name Paradoxurus; for a prehensile tail is hardly to be expected in an animal of the zoological position of the Palm Civets, and yet its occasional twisting led originally to the view that it was so. The genus has scent glands. The dentition is I 3/3 C 1/1 Pm 4/4 M 2/2. P. niger, the Indian Palm Civet, is, like other species, not often to be seen in a wild condition. It is arboreal, and, like other members of the genus, feeds upon a mixed diet, consisting of all kinds of small Vertebrata and insects, varied by fruit. Another species, P. grayi, is so distinctly vegetarian in its habits that it makes considerable havoc in pine-apple beds in the Andaman Islands.

Arctogale is another Oriental genus with very small teeth, those of the molar series being hardly in contact. The soles of the feet are more naked than in the last genus, and the scent glands, if present, appear to be small and ill developed. It has also a long tail, and is arboreal in way of life. There is "nothing particular recorded" as to its habits. The species are A. leucotis and A. stigmatica.

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Fig. 201.—Hardwicke's Civet Cat. Hemigale hardwicki. × 15. (From Nature.)

Closely allied to both the last genera is Hemigale, also an Oriental genus. It is to be distinguished from Paradoxurus by having the soles of the feet much less naked, though they are more so than in Viverra or Prionodon. The coloration of the species, H. hardwicki (a Malayan animal), is very peculiar. The body is banded with five or six broad transverse stripes, and the basal portion of the tail is also ringed, an uncommon feature in the group. A second species of this genus is H. hosei, from Borneo. It is blackish in colour, but is not a melanic variety of the last.

Nandinia appears never to possess a caecum.[270] It is also peculiar among Carnivora in the non-ossification of the hinder part of the bulla. It is an African genus, containing two species which are spotted. The tail is ringed.

Cynogale is at any rate a partially aquatic, short-tailed, web-footed, reddish brown-coloured Civet, which lives upon fish and Crustacea, and inhabits the Malay Peninsula, Sumatra, and Borneo. It has long "moustaches," and is said to have a head bearing a singular resemblance to the head of the Insectivorous "Otter" Potamogale. The metatarsus is bald, and the pollex and hallux are very well developed.

Sub-Fam. 5. Herpestinae.—There are over twenty species of Herpestes (Mongooses) divided between the Ethiopian and Oriental regions, one species, H. ichneumon, being also found in Europe. The fur has a "pepper and salt" appearance; the feet are plantigrade. There are five fingers and toes. The pollex and hallux are small; the tail is long. The tarsus and metatarsus are usually naked. The Egyptian species "has been injudiciously denominated the Cat of Pharaoh." It is perhaps better known as Pharaoh's Mouse. The beast is so far Cat-like that it will destroy Rats and Mice; and it has been exported to sugar plantations for that very purpose. More famous are its combats with venomous serpents. According to Aristotle and Pliny the Ichneumon first coats its body with a coating of mud, in which it wallows, and then with this armour can defy the serpent. Topsell tells the tale better. The Ichneumon burrows in the sand, and "when the aspe espyeth her threatening rage, presently turning about her taile, provoketh the ichneumon to combate, and with an open mouth and lofty head doth enter the list, to her owne perdition. For the ichneumon being nothing afraid of this great bravado, receiveth the encounter, and taking the head of the aspe in his mouth biteth that off to prevent the casting out of her poison." In the West Indies the animal has been described as fearlessly attacking the deadly Fer de Lance and receiving its bites with impunity; it is also added that it will eat the leaves of a particular plant as an antidote! The real explanation of the result of these encounters is of course the agility of the Ichneumon[271]fort cauteleuse beste, as Belon says.

Another species, H. albicauda, is distinguished, as the name denotes, by its white tail. A species of this genus, H. urva, sometimes raised to generic rank as Urva, is partly aquatic in habit; it feeds upon crabs and frogs, but is quite willing to take to poultry and their eggs.

Helogale is a genus whose validity appears doubtful (to Dr. Mivart). It is African, and contains two species.

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Fig. 202.—White-tailed Ichneumon. Herpestes albicauda. × 15.

Cynictis is an African genus, with five digits on the fore-limbs and four on the hind. As in Herpestes, the orbit is completely encircled by bone. There is but a single species, C. penicillata, which is of a reddish colour and has a bushy tail.

Bdeogale, also African, has the toes still further reduced; there are only four on both limbs. The tarsus is hairy and the tail bushy. They are "very rare animals, and nothing is known of their habits." It is known, however, that they will kill poisonous snakes, for Dr. Peters took a Rhinoceros Viper out of the stomach of one.

Rhynchogale[272] differs from all other genera of Viverridae, except Crossarchus and Suricata, in having no groove upon the muzzle. There are five digits. There is the full Viverrine dentition, with five premolars in the upper jaw; but this may be an abnormality.[273]

Crossarchus differs from the last in only having three premolars on each side of each jaw. It is also African, and there are several species.

Suricata is the last genus of Viverridae; it is also African, and contains a single species, Suricata tetradactyla, the "Meerkat" of the Cape. The Suricate has but four toes on each foot; the tarsus and the metatarsus are naked below. The body is banded posteriorly. There are fifteen dorsal vertebrae, and the orbit is closed by bone. The Suricate lives in caves and rock crevices, and will dig burrows. It is distinctly a diurnal animal, and sits upon its hind-legs after the fashion of a Marmot. As Buffon noticed in a tame specimen (thought by him to be a native of Surinam), the animal barks like a dog. The Suricate is largely vegetarian, living upon roots.

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Fig. 203.—Suricate. Suricata tetradactyla. × ¼.

Fam. 4. Hyaenidae.—Unlike though the Hyaenas appear to be to the last family—mainly perhaps on account of size—they are, nevertheless, very nearly akin to them, more so than to the Cat tribe. It will be remembered that the striping and spotting of the Hyaenas is very Genet- and Suricate-like.

There are admittedly two genera among the Hyaenidae, Hyaena itself with three species,[274] and the Aard Wolf, Proteles, with but one. But Dr. Mivart considers that the Spotted Hyaena should form a genus apart, Crocuta—a proceeding which was initiated by the late Dr. Gray of the British Museum. The Hyaenidae are to be distinguished by the following characters:—There are generally four toes, always so in the hind-foot. The claws are non-retractile. The nose and upper lip are grooved. The molar formula is Pm 4/3 M 1/1. The soles of the feet are covered with hairs upon the tarsus and metatarsus. No scent glands. Tail short. Dorsal vertebrae more numerous than in other Aeluroids, i.e. fifteen. The bulla is divided by a rudimentary septum only.

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Fig. 204.—Spotted Hyaena. Crocuta maculata. × 112.

The genera Hyaena and Crocuta, the Striped and Spotted Hyaena respectively, are African and Asiatic in range, Crocuta being limited to South Africa. There is neither hallux nor pollex.

Cambridge Natural History Mammalia Fig 205.jpg

Fig. 205.—Striped Hyaena. Hyaena striata. × 112.

The Hyaenas, stigmatised by Sir Samuel Baker as "low-caste creatures," are mainly carrion feeders. Much Arab superstition is associated with them. Certain peculiarities in the structure of the organs of reproduction have led to the belief that a Hyaena changes its sex every year. Its almost human-sounding howls are supposed to be a deliberate trap for the unwary traveller. There is also a legend that in the eye of the Hyaena is a stone which if placed under the tongue of a man endows him with the gift of prophecy.

Proteles presents many resemblances to the Hyaenas, but also certain differences; by many it is placed in a separate family. There is but one species, P. cristata, the Aard Wolf of South Africa. In outward aspect it is very Hyaena-like, the coat being striped, and the ears, though longer, resembling those of a Hyaena. There is also a mane. There are, however, five toes on the fore-feet. The teeth are feebler, particularly the molars, which are also reduced in number. The skull, as in Hyaena, has no alisphenoid canal, but the bulla tympani is divided by a septum. The animal seems to feed largely upon insects, particularly Termites, and also upon carrion.[275]

Of extinct Hyaenoids Ictitherium seems to be transitional between them and the Viverridae. Its dentition, 3/3, 1/1, 4/3, 2/1, is that of a Viverrid, and the feet are five-toed. The upper carnassial tooth, however, is like that of Hyaena in having a strong inner cusp. Other extinct genera of Hyaenas are Lycyaena and Hyaenictis. The genus Hyaena itself goes back as far as to the Miocene, and occurred in Europe until the Pleistocene. The Cave Hyaena of this country seems to be indistinguishable from Crocuta maculata, though it has received the name of H. spelaea.

Fam. 5. Canidae.[276]—This family cannot be divided into more than five genera, and is universally distributed with the exception of New Zealand. The auditory bulla is smooth and rounded, and has internally a very incomplete septum, extending through about one-fourth or one-third of the cavity. The meatus has a fairly prominent under lip. The paroccipital process is long and prominent. The mastoid is distinct, though but slightly developed. The glenoid foramen is large; the condyloid foramen is conspicuous, and the carotid canal is deep within the foramen lacerum posterius. The last three characters are Bear-like; the form of the bulla is Aeluroid. The teeth vary somewhat in number, and the following table will serve to indicate the gradual reduction observable in the number of molars:—

Otocyon I 3/3 C 1/1 Pm 4/4 M (3 or 4)/4
Canis generally I 3/3 C 1/1 Pm 4/4 M (3 or 2)/(4 or 3)
Cyon I 3/3 C 1/1 Pm 4/4 M 2/2
Icticyon I 3/3 C 1/1 Pm 4/4 M (2 or 1)/2

All the Dogs have a caecum[277] of simple cylindrical form. In C. cancrivorus, C. jubatus, and Nyctereutes procyonides this organ is straight or only very faintly curved; in other Dogs it is coiled into an LetterS.svg-like form, sometimes with an additional twist. The Dogs have, as a rule, five toes, one being dropped in Lycaon. The tail is fairly long and distinctly bushy. There is in a number of species a gland at the root of the tail, the presence of which can frequently be detected by the wet appearance due to the oozing secretion. The great majority of existing Canidae belong to the genus Canis. But certainly three, and more doubtfully four, other genera can be distinguished.

The genus Icticyon contains but one recent species, the Bush Dog (I. venaticus, Lund) of British Guiana. The animal has a somewhat Paradoxure-like, at any rate a distinctly un-dog-like, aspect, being longish in the body (some 2 feet long), shortish in the legs, and big-headed. It is blackish in colour, verging towards golden brown on the head and back. Sir W. Flower, to whom we owe our chief knowledge of its structure, characterises it as like a young Fox, and with the playful manners of a puppy. The animal appears to hunt in packs and by scent, and has a reputation for ferocity. Icticyon differs from Canis and agrees with the Indian Cuon in having but forty teeth, the last molar having disappeared from the upper and lower jaws. The caecum, unlike that of the majority of Canidae, is only slightly curved. The brain, oddly enough, shows a Cat-like peculiarity. It has been pointed out that in their long bodies and short legs the genera Cuon and Icticyon resemble the primitive dogs.[278]

A genus Nyctereutes is usually separated from Canis for the inclusion of N. procyonides only. The separation is based upon the strikingly unusual coloration of this Dog. It is a small animal, with numerous long white hairs dorsally. The face, chest, and much of the belly are black. Its aspect distinctly recalls that of a Raccoon,[279] especially in the black patches below the eyes, whence of course the scientific name and the pseudo-vernacular "Raccoon-like Dog." It inhabits China and Japan. As to structure, there is hardly anything that justifies its exclusion from the genus Canis. Garrod, however, mentions the unusually large size of the Spigelian lobe of the liver.

Cambridge Natural History Mammalia Fig 206.jpg

Fig. 206.—Raccoon-like Dog. Nyctereutes procyonides. × 16.

Wortman and Malkens[280] have instituted a genus Nothocyon for Dr. Mivart's species C. urostictus[281] and C. parvidens, which are both South American forms.

The genus Otocyon contains but one species, O. megalotis, an African species, ranging pretty widely in that continent (from the Cape to Somaliland, in sandy districts), and sometimes confused with the Fennec on account of its long ears. Its principal structural difference from other Dogs is that there is an additional molar in each jaw, the molar formula being thus M 3/4 or even 4/4. Moreover the carnassial teeth are not so pronounced, and Professor Huxley laid especial stress upon the likeness of some of the cheek teeth to those of the more primitive Arctoids. The angle of the lower jaw is inflected, a character, however, which seems to be more general than is usually allowed among animals not referable to the Marsupials. It is possible that Otocyon is a persistent Creodont-like form which has developed in a direction curiously, and in a most detailed fashion, parallel to the Dogs. If, however, we may assume the addition of the molar, then this anomalous but not necessarily untenable conclusion is obviated.

The genus Cuon, or Cyon, has been instituted for the two or three species of Eastern Dogs (C. primaevus, C. dukkunensis, etc.) which agree with each other in the constant loss of a molar in the lower jaw, or, it should be said, almost constant loss, for the missing tooth is occasionally represented. The latter of the two species mentioned, the Dhole, is, like its congeners, an animal which hunts in packs; it is said to hunt even the ferocious Tiger, and to be thus one of the few animals which can face the largest and fiercest of the Carnivora.

The genus Lycaon is a very distinct type, being differentiated from other Dogs by the possession of only four toes on both fore- and hind-limbs, and by the dental formula, which is Pm 4/4 M 2/3. The one species is L. pictus, the Cape Hunting Dog. It is singularly like a Hyaena[282] in general appearance; the ochraceous grey ground-colour with black markings and the long ears produce this likeness. The animal has got its vernacular name from the habit of hunting in packs. Its range is over a good part of Africa. The occurrence of this species (or at least genus, for the name L. anglicus has been used) in caves in Glamorganshire seems to show that it is a comparatively recent immigrant into Africa. As to its visceral structures, Lycaon[283] does not differ widely from other Dogs. It has, however, no lytta beneath the tongue. The intestines are thus divided: large, 9 feet 1 inch; small, 1 foot 3 inches. This contrasts with the proportions observable in some other Dogs. While other Dogs have but a cartilaginous rudiment of the clavicle, Lycaon has a considerably larger representative of this bone.

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Fig. 207.—Fennec Fox. Canis zerda. × 15.

Cambridge Natural History Mammalia Fig 208.jpg

Fig. 208.—Prairie Wolf or Coyote. Canis latrans. × ⅛.

The bulk of the Dogs, Wolves, Foxes, and Jackals are thus left over for inclusion in the genus Canis. But the numerous members of this genus can, according to Professor Huxley, be sorted into two series by certain cranial characters. The two series he termed the "Alopecoid" or Fox-like, and the "Thooid" or Wolf-like. It was suggested that the generic name Vulpes be used for the former, and Canis for the second. The characters which will be dealt with immediately are also to be noted among the Dogs belonging to genera that have already been separated off. Thus Lycaon is distinctly Thooid. The characters in question are these:—In the Fox series, the frontal air-sinus of the Thooids is absent; the cranial cavity is pear-shaped, without an abrupt angle coinciding with the supra-orbital sulcus, such as exists in the other group; the coronoid process of the mandible is rather higher and more turned back in the Foxes, while the depth of the mandible at the level of the first molar is greater.

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Fig. 209.—Japanese Wolf. Canis hodophylax. × ⅛ (From Nature.)

To the Fox series belong among others the species C. lagopus (Arctic Fox), C. zerda (the Fennec), C. chama (the Silver-backed Fox of Africa), C. virginianus (the Virginian Fox), C. velox (the Kit Fox), and of course the Common Fox of this country. On the other hand, the Dogs proper (such as C. dingo), the Wolves (C. lupus, C. pallipes, C. niger), the Japanese Wolf (C. hodophylax), the Red Wolf of America (C. jubatus), the Jackals (C. aureus, C. anthus, etc.), the Prairie Wolf (C. latrans), and a number of American forms, such as C. azarae, its close ally C. cancrivorus ( = C. rudis), C. antarcticus, C. magellanicus, etc., are decidedly Wolves rather than Foxes.

The Arctic Fox, Canis lagopus, is known by its bluish summer and pure white winter dress as "Blue Fox" and "White Fox" respectively. It is an inhabitant of the Arctic north; but in former days, as its remains show, it descended to such southern latitudes as Germany and this country. The most southern point which it now inhabits is Iceland. This small Fox is well known as being one of the few animals which change their dress to a complete white in winter. This change is, however, not absolutely universal; and M. Trouessart has even stated that the supposed change does not exist, but that the colours are a question of age and sex. This Fox feeds on birds and cast-up carcases of Whales and Seals; it is also said to devour shell-fish, and actually to store up food when abundant for seasons of scarcity. A Fox has been observed to "carry off eggs in his mouth from an eider duck's nest, one at a time, until the whole were removed"; and in winter to "scratch a hole down through very deep snow to a cache of eggs beneath." These anecdotes are told by Sir Leopold M‘Clintock; but others have also asserted the storing habits of this Fox, which really has only a short time of the year in which it can catch suitable living food.

Canis vulpes, the Fox, is not only a native of England, but extends as far to the east as Egypt, the so-called C. aegyptiacus being at most a mere variety. Varieties indeed occur in these islands; the English Fox being redder, the Scotch greyer. Not only is the Fox a truly indigenous English beast, but its remains go back a very long way into past time. Its bones occur in the Red Crag, a deposit of Pliocene times. Its prevalence now is no doubt due to its preservation as a beast of chase. It lives in burrows, either excavating them itself or taking possession of those of some other animal; the Badger suffers in this way, and is said to be vanquished not by the teeth of the burglarious Fox, but by its far fouler habits! It is curious that the expression "foxing" is not so suitable to this animal as to many others. The habit of "shamming death" is a widely-spread one in the animal world, but at least not common with our Fox. The sagacity of the Fox appears to be a little more proverbial than actual; literature teems with its accomplishments. The worthy Archbishop of Upsala, Olaus Magnus, figured Foxes dipping their tails in the streams, and then pulling out inquisitive crayfishes which had seized upon them. "It is a crafty, lively, and libidinous creature," observed a writer of the last century.

Of Jackals there are many species, both African and Oriental. Mr. de Winton allows the following list of African species[284]:—C. anthus, C. variegatus, C. mesomelas, C. lateralis. C. mesomelas is distinguished by the broad black patch in the middle of the back. These animals do not appear to go in packs as so many Canidae do; they live upon carrion, but also rob hen-roosts, and commit other depredations upon the live stock of farmers. The "Quaha," C. lateralis, is distinguished from the last by its sharp bark, and by the obvious side stripe which has given to it its name. It is curious that it should live in apparent amity with C. mesomelas, since the habits of the two are identical and would lead, one might suppose, to a severe struggle for existence, in which one of the two would disappear. Of Indian Jackals C. aureus is the most familiar type.

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Fig. 210.—Wolf. Canis lupus. × ⅛.

The European Wolf, Canis lupus, was once, but is no longer, an inhabitant of the British Islands. Their former prevalence is indicated by many names of towns and villages, such as Ulceby and Usselby in Lincolnshire, the town of Wolverton, and Woolmer Forest. In Saxon times Wolves were very abundant; and even so recently as the reign of Elizabeth they were to be seen on Dartmoor and in the Forest of Dean. In the New Forest they were hunted in the twelfth century. It would seem that the last English Wolf was slain some time during the reign of Henry VII. In Scotland, however, they persisted very much longer. So recently as 1743 was the last killed. But before this period they had begun to get exceedingly scarce, for the price of a skin in 1620 is quoted at £6:13:4. In Ireland Wolves lingered yet longer; about 1770 is believed to be the date of their final extinction in that island. The Wolf nowadays is distributed over the greater part of Europe, Northern Asia, and North America, the American form not being considered to be distinct from its European ally. Much legend has collected round this fierce Carnivore. Aristotle, usually accurate in the main, still "states more of wolves than experience warranted." Pliny, unable to sift truth from falsehood, was in this matter "an eager listener to all old women's tales." Aelian added to his marvels and asserted that the Wolf cannot bend its head back; if it should happen to tread on the flower of the squill it at once becomes torpid. So the wily fox, fearing his more powerful enemy, takes care to strew his path with squills! The conversion of men into Wolves was a well-known superstition, dating from Grecian and Roman times; it formed the basis of much of the witchcraft persecutions of the Middle Ages and onwards, and has left its mark in folklore, e.g. the Wolf in "Red Riding Hood."

The Indian Wolves, C. pallipes, C. chanco, and C. laniger, are hardly, if at all, different from C. lupus. Professor Huxley has remarked upon the likeness of C. pallipes to a Jackal, thus bridging over the very inconsiderable gap that may be held to divide Jackals and Wolves.

The Dingo, Canis dingo, is an interesting and somewhat mysterious species of Dog or Wolf. As is well known, it is an Australian species; but it does not seem to be certain whether it was tamed and brought over to Australia by the native races, or is a true and indigenous Australian species.

The colour of this species varies, but is usually of a reddish brown; it is, however, often grey and indeed almost black. Whether indigenous or introduced, the Dingo is a plague to Australian settlers, devouring Sheep, which it generally destroys by tearing out the paunch. It does not as a rule hunt in packs. The Dingo is stated to feign death with so much persistence that an individual has been known to be partly flayed before moving. Dingo remains have been found in river-gravels in Australia where no human remains have been detected. This argues for its indigeneity; but, on the other hand, it has been pointed out that man himself in the Australian continent goes back a very long way into time, and may thus still have imported this companion with him. Anyhow it is quite a wild creature now. Dr. Nehring, an expert investigator into the subject of domestic animals, has stated that the skeleton of the Dingo does not suggest a feral animal at all but a purely wild race.

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Fig. 211.—Dingo. Canis dingo. × ⅛.

The Domestic Dog is usually spoken of as Canis familiaris; but to remains in bone caverns the name of C. ferus or C. mikii has been given. There seems to be no doubt that the Dog was the "friend of man" in very early times. Its remains have been met with in Danish kitchen-middens, in the lake-dwellings of the Swiss lakes, and during the Bronze Age in Europe generally. But "there are few more vexed questions in the archaeology of natural history than the origin of the dog." Its remains already referred to may in many cases have argued its use as food. But in a Neolithic barrow a Dog was found buried with a woman, the skeletons of both being in situ; this animal was about the size of a Shepherd Dog. The actual Dog of to-day is divisible into more than 180 different breeds; but in a work upon "Natural History" it would seem out of place to enumerate and characterise these artificial products. Authors vary in their opinion as to what stock gave rise to the domestic races of the past and of to-day. The Jackal, the Bunasu (C. primaevus), the Indian Wolf (C. pallipes), have been proposed as likely ancestors. It is more probable that there is much admixture, and that various wild types have been selected by man in various countries.

Extinct Canidae.—Many of the existing species of Canidae are also to be found in Pleistocene deposits of the countries which they now inhabit. A few show a wider range in the immediate past than in the present. Thus Lycaon (L. anglicus) has been met with in caves in Glamorganshire, while Icticyon of South America appears to be congeneric with Speothos of the Brazilian caves. The African Otocyon seems to occur in deposits in India. There are also numerous extinct species belonging to the genus Canis, which extend as far back as the Pliocene.

The earlier types of Dogs have been placed in different genera. Cynodictis is an Eocene form from European strata. The skull is decidedly Civet-like, with a short snout. The fore- and hind-feet were five-toed, with well-developed pollex and hallux. The dentition was that of modern Dogs, the molars being two in the upper and three in the lower jaw. The general aspect of the creature and the form of the skeleton was much like that of the Viverrine genus Paradoxurus, of which, as well as of the Dogs, Cynodictis might have been an ancestor.

Simocyon of the Upper Miocene serves as the type of a separate sub-family of Dogs, Simocyoninae. The skull is short, broad, and high; the shortening of the skull affecting the jaws has reduced the teeth greatly; the first three premolars are very small, fall out soon, and are thus often deficient. There are only two molars in each jaw. This type is of course nowhere near the ancestral Dog. It is a much-specialised branch of an early type. Cephalogale is less specialised; there are the usual four premolars. Enhydrocyon is an intermediate form; it has lost one premolar in each jaw.

Amphicyon, forming the type of another sub-family, Amphicyoninae, though usually placed among the Dogs, presents us with many Bear-like features in its organisation. The feet, for instance, were plantigrade and five-toed. The ulna and the radius are specially compared with the same bones in the Bear tribe. The skull on the other hand is as distinctly Dog-like in form. The molars are large, broad, and crushing, and Bear-like. The largest known species, A. giganteus, is of about the size of the Brown Bear. Amphicyon is a Miocene genus. Eocene and allied to it is Pseudamphicyon. This genus has, like Amphicyon, the complete dentition of forty-four teeth. In the Amphicyoninae generally the feet are five-toed, the humerus has an entepicondylar foramen and the femur a third trochanter. The upper molars are large.

The closely allied and American genus Daphaenus has also plantigrade feet, and has in its structure many reminiscences of the Creodonts. So, too, has the Eocene Uintacyon.

Cynodesmus is closely allied to Cynodictis. It has ancient features combined with quite modern ones. The skull is described as being Creodont-like, but the dentition is that of the microdont modern Dogs. In accordance with its age the cerebral convolutions of this Dog are much simpler than in existing Dogs, and the hemispheres do not cover the cerebellum so much.

The Bear-like Carnivora or Arctoidea.—That division of the Carnivora which is typically represented by the Bears embraces three recent families, which are united by a number of characters. These Carnivora are always plantigrade or nearly so. They have nearly always five toes. The claws are not retractile, or at most semi-retractile as in the Panda. In the skull the tympanic bulla is often depressed, and is not so globular and obvious as in the Cats. Its cavity is not divided by a septum. The paroccipital processes are not applied to it. The carnassial tooth is less emphasised in this group than in the Cats.

These characters, however, have to be used with caution, as they are hardly universally applicable. A fairly typical Arctoid bulla is seen in such a form as Cercoleptes. The bulla itself is a little more swollen than in Ursus, but it is flattened off in the same way towards the bony meatus. The paroccipital processes, slightly developed, are at a distance of ¼-inch from the posterior margin of the bulla. In the Raccoon the bullae are much more swollen, and the paroccipital processes are closer to them. In the Marbled Polecat, Putorius sarmaticus, the bullae are fairly swollen, and there is but little flattening towards the meatus: the paroccipital processes, though slight, are in contact with the bullae basally, though their free tips are turned away from them. Finally, in Ictonyx the bullae are much swollen; there is but little flattening towards the meatus, and the paroccipital processes, themselves much swollen, are pressed closely against the bullae. The Mustelidae, therefore, in this as in other characters, approach the Aeluroids.

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Fig. 212.—Section of the left auditory bulla and surrounding bones of a Bear (Ursus ferox). am, External auditory meatus; BO, basioccipital; Car, carotid canal; e, Eustachian canal; g, glenoid canal; Sq, squamosal; T, tympanic; t, tympanic ring. (From Flower, Proc. Zool. Soc. 1869.)

There is no caecum, a feature which marks off the Arctoidea from all Carnivora except the Viverrids Nandinia and Arctictis (occasionally). The brain is characterised by the possession of what Dr. Mivart has described as the "ursine lozenge," a tract about the middle of the hemispheres, defined posteriorly by the crucial sulcus, and formed by the emergence on to the surface of the brain of the hippocampal gyrus.

The Arctoidea are very widely distributed. But there are some curious exceptions. Thus there are no representatives of the group (as might be expected) in the Australian region; they are completely absent from Madagascar; while the true Bears (family Ursidae) are totally absent from Ethiopian Africa, and are only represented by a single species, Ursus ornatus, in the Neotropical region.

It is noteworthy that the Arctoidea never show spots or cross stripes (save rings on the tail), which are so common a feature of the coloration of the Cat-like forms.

In bracketing together the three families which are described in the following pages, emphasis is laid upon a number of undoubtedly common features. Palaeontology seems, however, to suggest that the Mustelidae come nearer to the Viverridae. That the Bears and Dogs are connected by extinct annectent genera does not interfere with their present distinctness.

The systematic arrangement of these Carnivora is not easy. It may be useful, however, to give a method of arrangement for the convenient placing of the genera.

The most primitive group is perhaps that of the true Bears, family Ursidae; for in them the molars are two above and three below, and have thus not become diminished in number as in some of the other members of the order. Moreover, the Bears have lobate kidneys, which character, often occurring in the young of animals which when adult have smooth kidneys, may be looked upon as a primitive character. The feet furthermore are completely plantigrade. This family will contain only three genera, Ursus, Melursus, and Aeluropus.

Next comes the family Procyonidae, in several members of which one molar is lost below, though in others the more archaic formula is retained. The kidneys are simple. This family contains the American genera Procyon, Nasua, Bassariscus, Bassaricyon, Cercoleptes, and the Old-World form Aelurus.

The third family, Mustelidae, has the molar formula reduced to 1/2 or 1/1. The kidneys are simple except in the Otters. To this family are assigned the following genera:—Arctonyx, Conepatus, Meles, Mephitis, Taxidea, Mydaus, Mellivora, Helictis, Ictonyx, Mustela, Galictis, Grisonia, Putorius, Gulo, and the aquatic Lutra, Enhydris, and Aonyx.

Fam. 6. Procyonidae.—This family is mainly American in range, the genus Aelurus alone being a native of the Old World. But Zittel would include with the genera of this family the Viverrine and Oriental genus Arctictis, a proceeding which is perhaps hardly admissible, though the occasional absence of a caecum in that animal is so far in favour of such an alliance. The largely vegetable nature of its food and its arboreal habits cause a certain amount of likeness to some of the members of the present group of Carnivores. The Procyonidae have two molars in either half of each jaw. The carnassial teeth are not typically developed, and the molars are broad and tuberculate. The tail is long, often prehensile, and often ringed in the disposition of its colour pattern. The alisphenoid canal is absent save in the aberrant Aelurus. Both condyloid and postglenoid foramina are present. The members of this family are plantigrade.

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Fig. 213.—Raccoon. Procyon lotor. × 15.

The genus Procyon includes at least two species of Raccoon, the northern form, P. lotor, and the South American, P. cancrivorus. To these may possibly be added a third, P. nigripes. This genus is characterised by the length and the mobility of the fingers, and indeed it uses its hands greatly. It has no median groove upon the muzzle, which is found in many other Arctoids; the ears are moderately large; the tail is not long, being about one-third of the entire length of the animal, including the tail. The soles of the feet are naked. Its limbs are very long (for an Arctoid), and this gives to the animal a bunched-up appearance when walking. There are four premolars and two molars on each side of each jaw. There are fourteen pairs of ribs, of which ten pairs reach the sternum. The latter is composed of nine pieces.

The first-named species has received its name from the fact—of which there is abundant proof—that it dips its food into water. As a matter of fact, the animal frequents the margins of streams, and hunts in the shallow water beneath stones for crayfish, and it also captures fish. Not only is this animal partially aquatic, but it can climb well—"they make their homes in trees, but carry on their business elsewhere." The animal can be readily tamed, but is a tiresome pet on account of its insatiable curiosity and its skill in the use of its hands, which enables it to unlatch doors and generally to pry about everywhere. The Raccoons are mostly nocturnal creatures.

The genus Bassaricyon[285] includes two species, both American, B. alleni being from Ecuador, and B. gabbii from Costa Rica. They have so much the aspect of a Kinkajou that a specimen, which arrived at the Zoological Gardens, was presented and entered as one of those animals. Nevertheless there are many differences between the two genera. The tail of Bassaricyon is not prehensile, and the animal, as will be seen from Fig. 214, has a sharper snout; the brain is more like that of Bassariscus. The likeness to Cercoleptes can hardly perhaps be regarded as an example of "mimicry" since the forms are so nearly related, and the advantage of such an imitation remains to be proved. The muzzle of Bassaricyon is grooved; the ears are fairly large; the soles of the feet are naked; there is but a single pair of teats. There are two molars and four premolars to each half jaw.

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Fig. 214.—Bassaricyon. Bassaricyon alleni. × 15.

The dorsal vertebrae are thirteen in number; nine of the ribs reach the sternum. The slenderness and convexity of the lower margin of the lower jaw, as well as the feeble angular process, distinguish this genus from its undoubtedly near ally Cercoleptes. The dental formula also is different.

Bassariscus has a ringed tail like a Raccoon, and is also American in range; it furthermore agrees with the Raccoon in being nocturnal and mainly arboreal in habit. There are apparently three species, of which B. astutus is the best known, having been on several occasions exhibited at the Zoological Society's Gardens, the last examples so lately as 1900. The animal was for a long time believed to be allied to the Oriental Paradoxures, and its occurrence in America was therefore puzzling. The real affinities of the creature were, however, definitely set at rest by Sir W. Flower, and later accounts of its anatomy have confirmed this opinion.[286] The vertebrae are more numerous than in Procyon, and the teeth are slightly different; otherwise it presents many likenesses to its nearest ally. The ears are long; the nose is grooved; and the palms and soles are naked.

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Fig. 215.—Cunning Bassarisc. Bassariscus astutus. × 15. (From Nature.)

The Kinkajou, Cercoleptes, is likewise an American Arctoid. It ranges from Central Mexico down to the Rio Negro in Brazil. It was at one time confounded, and, considering its external appearance, not unnaturally, with the Lemurs. Sir R. Owen dispelled this view by a careful dissection of the creature. Nevertheless, there are certain anatomical features in which it differs from Carnivora and resembles Lemuroids.[287] It has been pointed out that the form of the lower jaw "much resembles that of the Lemuroid Microrhynchus." There is, however, no doubt that it is rightly placed in the present group. The tail is very prehensile, and the animal is therefore, as might be supposed from this circumstance, purely arboreal. It has some twenty-eight vertebrae. This genus has a median groove upon the nose. The claws are long and sharply pointed, and the palms and soles of the feet are naked. The premolars are three, the molars two. There are fourteen dorsal vertebrae, of which nine are united to the nine-jointed sternum by ribs. There is but one species, C. caudivolvulus, of a uniform yellowish-brown colour.

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Fig. 216.—Kinkajou. Cercoleptes caudivolvulus. × 16.

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Fig. 217.—Coati. Nasua rufa. × 16.

Nasua, the Coati, ranges from Texas to Paraguay, and has two species. In Guatemala it reaches a height of 9000 feet on the mountains. The nose is produced into a short and very mobile proboscis, hence its name. The native Mexican name for the creature is "Quanhpecotl."

The Coati is largely arboreal, and hunts iguanas in large bands, some of them being on the trees and some on the ground beneath. It also grubs up worms and larvae, for which purpose its long snout is suited. The molars of the genus resemble those of Procyon.

There is not a median groove upon the nose. The palms and soles are naked. Six teats occur. There are thirteen dorsal vertebrae. Nasua nasica[288] and N. rufa are the best known and perhaps the only species. The colour of the fur varies a good deal, and has led to the use of other names for supposed species.

Aelurus, the Panda, is a largish animal found in the south-eastern Himalayas up to a height of 12,000 feet. It has a glossy fur of a reddish colour, and a "white somewhat cat-like face." The molar formula which distinguishes it from the New-World Arctoids belonging to the Procyonidae, as well as from its possible ally Aeluropus, is Pm 3/4 M 2/2. The anatomy of the animal has been described by Sir W. Flower.[289] Dr. Mivart has pointed out that the muzzle though short is upturned in a way distinctly recalling that of Nasua. The animal inhabits forests, and feeds almost entirely upon vegetable food. It eats eggs, however, and insects. Though living to a great extent upon the ground, it is also arboreal, and has sharp semi-retractile claws. It is said to be dull of sight, hearing, and smell, and yet with these disadvantages is also unprovided with cunning or ferocity. Its habits have been compared with those of a Kinkajou.

Fossil Procyonidae.—In addition to several of the existing genera, the remains are known of various extinct forms of Procyonidae. Leptarctus, with one species, L. primaevus, is of Pliocene age, but is known only by one ramus of the lower jaw. It appears to "offer a number of transitional characters between the more typical Procyonidae and the aberrant Cercoleptes."[290]

Fam. 7. Mustelidae.—Contrary to what has been stated with regard to the habits of the Procyonidae, the Mustelidae are for the most part "bloodthirsty robbers," and are spread over the whole surface of the world, with the exception of Australia and Madagascar. The molar teeth are generally reduced to one in the upper jaw, and sometimes to one in the lower jaw, which thus gives "a sort of prima facie resemblance to the feline dentition." There is no alisphenoid canal; postglenoid and condyloid foramina are found.

Sub-Fam. 1. Melinae.—Of this sub-family there are representatives both in the Old and New Worlds.

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Fig. 218.—Badger. Meles taxus. × 16.

Meles, the Badger, is exclusively Palaearctic in range.[291] Dr. Mivart says that Meles has a relatively longer dorsal region than any other Carnivore, and that it is most nearly approached by its allies Ictonyx and Conepatus. The molar formula is, as in Arctonyx, Mydaus, and Helictis, Pm 4/4 M 1/2. The molars differ from those of any other Carnivore in the much greater size of the first molars than of the last premolars. The nose is not grooved; the soles of the feet are naked. The claws of the fore-feet are much longer than those of the hind-feet.

The genus Arctonyx is a "pig-like badger" from Hindostan, Assam, and North China. The epithet "pig-like" is derived from the long and mobile snout, which is truncated and has terminal nostrils. It is remarkable for having a part of the palate formed by the pterygoids, as in Whales and certain Edentata (e.g. Myrmecophaga). There are sixteen dorsal vertebrae. A. collaris lives in fissures of rocks, or in holes dug by itself. It is a purely nocturnal beast.

The singular genus Mydaus, containing the species M. meliceps, the Teledu or Javanese Skunk, is an inhabitant of Java and Sumatra. It frequents the mountains of these islands, into the soil of which it burrows in search of worms and larvae. There is but one species, which is "like a miniature badger, of rather eccentric colours." It is blackish brown, with a yellowish-white top to its head, and a stripe of the same colour down the back. It may be distinguished by its elongated snout, obliquely truncated, and with inferiorly-placed nostrils. As to osteological characters, it has a more oblique symphysis of the mandible than in any other Carnivore. The secretion of the anal glands is said to rival that of the Skunk in offensiveness and in the distance to which it can be propelled.

Sub-Fam. 2. Mustelinae.—Representatives occur in both the Old and New Worlds; but the genera and even the species are in one or two cases common to both.

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Fig. 219.—Tayra. Galictis barbara. × 17.

Galictis barbara,[292] the Tayra, is a brown, elongated, and Weasel-like animal from Mexico and South America. As is the case with the Weasel, it is sometimes gregarious, a herd of twenty having been observed. The soles of the feet are naked, and the molar formula is Pm 3/3 M 1/2. In these characters the Grison (G. vittata) agrees with G. barbara; but it has been referred to a different genus, Grisonia.

The Grison, "this savage and diabolical-looking weasel," as Mr. Aplin terms it,[293] is known also as the "Hurón." It almost rivals the Skunk in the power of the odour which it can emit when enraged. A trapped specimen was placed in a cage 50 yards or so from the house, and even at this distance it was disagreeably easy to tell when any one visited the animal—at least when the wind set in the right direction. It is greyish yellow above and blackish beneath, presenting, as has been remarked, a curious similarity to the Ratel. The nose of this animal is destitute of a median groove, which is present in the Tayra; the soles of the feet, however, are naked as in that animal, and it is nearly plantigrade in walk. It differs also from Galictis in having sixteen[294] instead of fourteen dorsal vertebrae. Eleven of the ribs reach the sternum. Considering the differences that exist between some other genera of Arctoids, it may be fairly allowed that a genus Grisonia is tenable.

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Fig. 220.—Grison. Grisonia vittata. × 17.

G. allamandi is darker coloured than the Grison, with a white band from the forehead to the neck. Mr. T. Bell described a tame individual as eating eggs, frogs, and even a young alligator.

A third genus of this group has recently been founded by Mr. Oldfield Thomas[295] for a small African animal, which is Grison-like in its coloration. The name given to the genus, Galeriscus, is intended to suggest its likeness to the Grison (Galera or Grisonia). The chief distinctive feature of this genus, whose skeleton is not yet known, is the presence of only four digits on each limb; the pollex and the hallux being entirely absent. The ears of this Grison are short.

The genus Mustela includes the Martens and Sables, which are distinguished from the following genus by the molar formula, which is Pm 4/4 M 1/2. The same character separates them from Galictis, and also the generally hairy under surface of the feet. In more southern latitudes, however, the palms are sometimes naked. The nose is grooved, and the ears are short and broad. The genus is widely distributed, being common to the Old and New Worlds. In the Old World it extends from Europe to Java, Sumatra, and Borneo. The largest species of the genus is the American Pekan, an animal which may be 46 inches in length, including the tail. There are two species of Sable, one European (M. zibellina), the other American.

The only British species of the genus is the Pine Marten, M. martes. It is dark brown, with a brownish-yellow throat, and reaches a length of some 17 inches, with an eight-inch tail. It is getting rare, but is still fairly common in the Lake country. The animal is largely arboreal in habit, whence the vernacular name. It is also called Marten Cat. The allied M. foina, the Beech Marten, has been stated to be, but apparently is not, an inhabitant of these islands. The colour of the animal is a rich brown. It has small eyes and ears and a short tail. The palms of the hands and the soles of the feet are hairy; the muzzle is naked, and has a groove as in Cercoleptes, etc.

The Glutton, Gulo, is a well-marked genus, containing but one species, which is circumpolar in range. The dentition is Pm 4/4 M 1/2. The ferocity but not the voracity of this animal appears to have been exaggerated. It mainly feeds on carcases, and is not really a successful hunter. As to the carcases, Olaus Magnus tells in straightforward language the way in which the animal dilates in size during a meal, and presently, after following the practice of the ancient Romans, returns to the banquet: "Creditur a natura creatum ad ruborem hominum qui vorando bibendoque vomunt redeuntque ad mensam"!

This is one of the few land animals which ranges completely round the pole. There is no difference to be noted between the Old-World and the New-World specimens. It is now an entirely northern form, but in Pleistocene times it reached as far south as this country. The fossil species seems to be Gulo luscus, and to be quite indistinguishable from the living forms.

Putorius, the genus which embraces the Weasel tribe, contains many species known popularly as Weasels, Ermines, Stoats, Ferrets, Polecats, Minks, and Vison. Not only is the genus common to both Old and New Worlds, but in a few cases the species (e.g. P. erminea) range from Asia to America. The molar formula is Pm 3/3 M 1/2. The form of the body is an exaggerated one, the length of the trunk to the limbs being very great. The feet are more or less hairy beneath, and the animals are digitigrade. The nose is grooved. The dorsal vertebrae vary from thirteen to sixteen.

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Fig. 221.—Polecat. Mustela putorius. × 16.

There are four British representatives of this genus:—

The Polecat, P. foetidus, is a dark brown-coloured animal. Its total length is about 2 feet, of which the tail occupies some 7 inches. It is a species banned by the gamekeeper, and hence is approaching extinction in this country. It is excessively bloodthirsty, as are apparently all the members of this genus, and kills out of mere wantonness. The Ferret is simply a domesticated variety of the Polecat.

The Stoat or Ermine, P. erminea, is reddish brown above, white beneath. In winter, in certain localities, it becomes white with the exception of the black tip of the tail. This colour-change bears some relation to the degree of latitude. It is universal in the north of Scotland, rare in the south of England. As is the case with some other animals that generally change their colour in the winter, there are individuals which seem to have lost the power of change, and others which change in an apparently capricious manner, not influenced by season or cold. Like so many other animals, the Stoat appears at times to migrate, which it does in large parties. Such parties are said to be dangerous, and will attack a man who crosses their path.

The Weasel, P. vulgaris, has much the same colour as the Stoat, but is a smaller animal; it differs also by undergoing no seasonal change. It is equally agile and ferocious, and ought to be encouraged, as it vents its ferocity largely upon Voles and Moles, which it can pursue underground. Like other species of Putorius, it seems to kill its prey by biting through the brain-case.

The fourth British species is the recently-described Irish Stoat, P. hibernicus. It is somewhat intermediate between the last two.

Poecilogale is a genus recently instituted by Mr. Thomas for a small South African Weasel, P. albinucha, coloured like the Zorilla, i.e. with whitish stripes upon black, but differing in its reduced molar formula, which is Pm 2/2 M 1/1 or 1/2.

Lyncodon[296] is thought to be more doubtful; it is South American (Patagonian), with the same molar formula as the most reduced forms of the last genus, i.e. Pm 2/2 M 1/1. The ears are short and almost invisible; the claws of the anterior limbs are long, those of the hind limbs short. It is not quite certain that it is not "an aberrant southern form of Putorius brasiliensis." That its distinction is justifiable appears to be shown by the discovery in the same region of a fossil species, L. luganensis. Matschie places it near Galictis.

The Ratel, Mellivora, is common to India and West and South Africa. It is a black animal with a grey back and grey on the top of the head, the contrast of colour suggesting a dorsal carapace. It runs with a swift trot. The animal lives much on the ground, but can climb trees. It is exclusively nocturnal in its habits. It has the reputation in India of feeding upon dead bodies, a view which has probably no foundation in fact save that it can burrow. The molar formula is Pm 3/3 M 1/1. There are fourteen dorsal vertebrae. The African and Indian species are hardly to be distinguished from each other. The ears are very minute. The tail is short. The muzzle is rather pointed, and the soles and palms are naked.

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Fig. 222.—Ratel. Mellivora capensis. × ⅛.

The structure of Helictis has been described by the late Professor Garrod,[297] as well as by Sir W. Flower in his general account of the Carnivorous skeleton. The animal, which is a native of East Asia, is sometimes gaily coloured. H. subaurantiaca, the species dissected and figured by Garrod, is a varied black and orange. The genus is arboreal, and the tail may be moderately long and bushy. The ears are small; the nose is grooved; the palms are naked, but the soles of the feet are hairy. There are fourteen dorsal vertebrae. The molar formula is Pm 4/4 M 1/2.

The Zorilla, Ictonyx, is the last of the Old-World genera of Melinae. It is African, ranging from the tropical parts of the continent to the Cape. "In colour and markings," remarks Dr. Mivart, "as well as in the odour of the secretion of its anal glands, the one or two species which form this genus resemble the skunks; so much so that did they inhabit the same region, and were they devoid of an offensive secretion, they would certainly be said to mimic the skunks." The molar formula of the genus is Pm 3/3 M 1/2. There are fifteen dorsal vertebrae. The nose is grooved and the soles partly hairy.

The American Badger, Taxidea, is a burrower of omnivorous tastes, and correlated with the former habit are the immense claws of the fore-paws. It is North American, but gets into Mexico. The molar formula is as in the American genera Mephitis and Conepatus, and as in the Old-World Ictonyx, and it thus differs from that of Meles. Besides the great size of the claws upon the hand, which are larger relatively than those of any other Carnivore, the genus Taxidea is to be distinguished from all Arctoids (indeed, from all Carnivora) except Mydaus, by the fact that the pelvic limb is of the same length as the pectoral. The muzzle is furry except at the very extremity; this is grooved. The animal is carnivorous, subsisting upon the following very varied kinds of food—"Spermophiles, Arvicolas, birds' eggs, and snails, also honey-comb, wax, and bees."

The Skunk, Mephitis, is an American animal with several species, which range from North to Central America. The black-and-white colour distinguishes the genus, which is furthermore marked by the fact that the third digit of the hand is relatively longer than in any other Carnivore except Taxidea. The soles are partly hairy. It is a terrestrial fossorial animal with well-known powers of protecting itself from aggression. But nevertheless the Skunk has its enemies, and is not quite so unmolested as is sometimes popularly supposed. The Puma, Harpy Eagle, and the Great Horned Owl will at least occasionally attack and devour it. The molar formula is Pm 3/3 M 1/2. There are sixteen dorsal vertebrae.

Conepatus is a more southern form of Skunk, extending down into South America. Its dentition is like that of Mephitis save for the loss of an upper premolar. This genus, which has been further subdivided, differs from Mephitis in the fact that the soles of the feet are wholly naked, whereas in Mephitis those of the hind-limbs are partially hairy. It has no groove on the nose. Its tail is shorter than that of Mephitis. This Skunk has the same habits as the last. In certain parts of South America the animals are so abundant and their odour so powerful that in the evening there is generally a recognisable smell about. This is said to be good for the headache!

Sub-Fam. 3. Lutrinae.—Of this sub-family there are at least two genera. Enhydris (Latax),[298] the Sea-Otter, is confined to the shores of the North Pacific. It is more purely aquatic than are other Otters. Specimens have been seen swimming fifteen miles from land. The gait of the creature when on land is suggestive of a marine animal; the webbed hind-feet are doubled back upon the knuckles during progression upon land, and locomotion is effected by a series of short springs from these feet; the Otter does not walk "in ordinary acceptance of the term." The tail is flattened, being twice as broad as it is thick, and ends in a bluntish point. Enhydris feeds mainly upon crabs and sea-urchins, but also upon fish. Its dental formula is peculiar by reason chiefly of the reduction of the lower incisors. The formula runs as follows: I 3/2 C 1/1 Pm 3/3 M 1/2.

The molar teeth of this creature, in accordance with its diet, have lost the sharp points of the Mustelidae in general; the crowns are flattened, and the tubercles very blunt. In this it contrasts with Lutra, and presents some resemblance to the Crab-eating Raccoon, Procyon cancrivorus; but the teeth are still further blunted. Enhydris feeds largely upon sea-urchins and shell-fish, and needs blunt teeth for the crushing of the hard shells of its prey. It is interesting to notice that the habits of this animal have been altered by the interference of man. The creature has been hotly pursued for a long time on account of its valuable fur. Instead of feeding and breeding upon the shore in places readily accessible to its pursuers, the Sea-Otter has now taken to the open sea in a greater degree. It utilises masses of floating seaweed for those purposes, and hunts for its food in the deeper water at a greater distance from the shore. In conjunction with the increasing rarity of the Sea-Otter the price of its skin has enormously increased: whereas in 1888 the average price per skin was £21:10s., the value of a fine skin now is at least £100, and as much as £200 and even £250 has been given. The animal is captured by netting and by clubbing and spearing.[299] From the Miocene Siwalik beds remains of an allied form, Enhydridon, have been obtained, whose teeth are somewhat intermediate in their crowns between Lutra and Enhydris.

Lutra, including the Otters, is widely distributed. Both manus and pes are webbed. The ears are small and hairy. The nose is not grooved, and the naked part is very circumscribed; the claws upon the hind-feet are flattened and somewhat nail-like. There are about ten species, but of course, as is so universally the case, a great many more names have been given. The molar formula is like that of Enhydris save that there is an extra premolar in the upper jaw. There are fourteen pairs of ribs, of which eleven pairs reach the ten-jointed sternum. The caudals are twenty-three. The Cape Otter, the "clawless" Otter, has been separated as a genus Aonyx. So too has the South American Pteronura brasiliensis. But in neither case is the separation allowed by Mr. Thomas in a recent revision of the genus.[300] The latter species has the reputation of being very fierce, and is known in Uruguay by the name of "Lobo de pecho blanco." The British species, L. vulgaris, reaches a length of 2 feet or so, with a tail of 16 inches; it ranges over the whole of Europe and a large portion of Asia. This Otter often burrows in the banks of the streams which it frequents; and in the burrow in March or April the female brings forth her young, three to five in number. It will also frequent the sea-coast.

Cambridge Natural History Mammalia Fig 223.jpg

Fig. 223.—Otter. Lutra vulgaris. × 16.

Fossil Mustelidae.—Besides a number of the existing genera there are fossil members of this family which cannot be referred to existing genera. These latter extend back into time as far as the Eocene. Stenoplesictis, one of these Eocene forms referable to the sub-family Mustelinae, is to be distinguished from living Mustelines by its comparatively long legs. In this genus as in several others there are two upper molars.

Fam. 8. Ursidae.—This family is nearly universal in distribution, and consists of but three genera, Ursus, Melursus, and Aeluropus.

Ursus has the palms and soles naked except in the Polar Bear, which needs a furry sole to walk with ease upon ice surfaces. The ears are fairly large, and the nose may or may not be traversed by a median groove.[301] The molar formula[302] is Pm 4/4 M 2/3. The brain is naturally (because of the size of the animals of this genus) richly convoluted. The lobate kidneys have already been mentioned in defining this family (see p. 426).

Cambridge Natural History Mammalia Fig 224.jpg

Fig. 224.—Himalayan Bear. Ursus tibetanus. × 115.

A very large number of species of Bears have been described. But it is the opinion of Mr. Lydekker[303] and of others that many of these are really to be referred to the European Brown Bear; in this event the Grizzly of North America, the Isabelline Bear, the Syrian Bear, a Bear from Algeria, the Kamschatkan and Japanese Bears, besides the extinct Ursus fossilis of Pleistocene caves, are to be regarded as slight modifications of Ursus arctos. On the other hand, the great Cave Bear, U. spelaeus, and the Thibetan Blue Bear (U. pruinosus) are distinct species, not to be confounded with U. arctos. Neither, of course, are the Peruvian U. ornatus and the Sun Bear, U. malayanus.

Cambridge Natural History Mammalia Fig 225.jpg

Fig. 225.—Malayan Bear. Ursus malayanus. × 112.

The Polar Bear has even been placed in a separate genus, Thalassarctos, a proceeding which is quite unnecessary. The white colour of this Bear tends to become browner with age. It is one of the few mammals which extend right round the pole; the Polar Bear is of course a purely Arctic animal. The chief food of the Polar Bear is Seal. Out of thirty Bears examined, Mr. Koettlitz found that only fifteen had animal remains in their stomachs, and these remains were invariably Seal. The animal apparently hunts by scent rather than by sight or hearing, both of which senses seem to be somewhat dull. The males and females wander separately, except of course during the breeding season. The Bears dig holes in which they may remain for some time, but there is no hibernation. In Pleistocene times, the Polar Bear extended as far south as Hamburg. The female has four mammae, pectoral in position.

Melursus includes only M. labiatus, the Sloth Bear of India. This animal has an upturned snout, which is described as closely resembling that of Mydaus, the Teledu. The snout has no groove. All Bears are largely vegetarian and insect feeders; but this Bear is especially so. It delights in the nests of Termites, and its energy in destroying these hills for the sake of their inhabitants is so great that the name of "sloth" appeared to Sir Samuel Baker to be an entire misnomer.

Aeluropus, a rare Carnivore with but one species, A. melanoleucus, is not inferior in size to the Brown Bear, and is distinguished by its largely white coloration. It was discovered in the mountains of East Thibet by Père David, and described by Milne-Edwards[304] as a distinct and new genus, the discoverer himself having named it as a species of Ursus. It is a vegetable-feeding creature and bulky in form, with a rudimentary tail and a short broad head; in fact, more like a Bear than a Procyonid (with which group it is placed by some). The width of the head, however, is greater than in any other Carnivore; it is most closely approached in this by Aelurus and by Hyaena. The molar formula is Pm 4/3 M 2/3. The soles are hairy. There is no alisphenoid canal. The molars are especially large and multicuspid.

Cambridge Natural History Mammalia Fig 226.jpg

Fig. 226.—Aeluropus melanoleucus. × 112.

Fossil Ursidae.—The genus Ursus itself goes back to Pliocene times. The well-known Cave Bear, Ursus spelaeus of Pleistocene times, was one of the commonest of Carnivorous creatures during the very early times of the present era. It was as huge as a Polar Bear or a Grizzly. The skull is remarkable for the fact that the first three premolars, which are small in all Bears, dropped out early in life. An immense number of names have been given to what are in all probability the same species as this Cave Bear of remote antiquity.

Hyaenarctos is the oldest genus of true Ursidae. It goes back into Middle Miocene times, and ranged over Europe and North Africa.

Arctotherium is an American genus of Pleistocene times. The likeness of some of the extinct Canidae to Bears has been already commented upon.


257 ^  For a general account of the osteology, see Flower, Proc. Zool. Soc. 1869, p. 4; and for muscular anatomy, Windle and Parsons, Proc. Zool. Soc. 1897, p. 370, and 1898, p. 152.
258 ^  See St. G. Mivart "On the Aeluroidea," Proc. Zool. Soc. 1882, p. 135: and The Cat, London, J. Murray, 1881.
259 ^  "On the Pupils of the Felidae," Proc. Zool. Soc. 1894, p. 481.
260 ^  "Observations ... on the Seal's Eye," Proc. Zool. Soc. 1893, p. 719.
261 ^  It is noteworthy that in the Tiger some of the stripes have pale centres and are thus like spots pulled out, while there are also small black spots.
262 ^  Natural Science, vi. 1895, p. 89.
263 ^  For an account of this and of other mammals which occur in Central America, see Alston in Messrs. Godman and Salvin's Biologia Centrali-Americana, 1879-1882.
264 ^  But Mr. Belt says that the "Tigre" never attacks man unless it be provoked.
265 ^  See E. Hamilton, The Wild Cat of Europe, London, Porter, 1896; and M. G. Watkins, Gleanings from the Natural History of the Ancients, London, Elliot Stock, 1896.
266 ^  The retractility is most marked in the Linsangs.
267 ^  Beddard in Proc. Zool. Soc. 1895, p. 430.
268 ^  Where it has probably been introduced.
269 ^  Proc. Zool. Soc. 1873, p. 196.
270 ^  Flower, Proc. Zool. Soc. 1872, p. 683.
271 ^  See also vol. viii. p. 591.
272 ^  The original name was Rhinogale.
273 ^  That it is an abnormality has been recently stated.
274 ^  For the anatomy of Hyaenas see Morrison Watson in Proc. Zool. Soc. 1877, p. 369; 1878, p. 416; and 1879, p. 79.
275 ^  Flower, Proc. Zool. Soc. 1869, p. 457.
276 ^  For a general account of the Canidae see Mivart, A Monograph of the Canidae, London, 1890.
277 ^  Flower, Proc. Zool. Soc. 1879, p. 766.
278 ^  Proc. Zool. Soc. 1880, p. 70.
279 ^  The relationship between the Canidae and the Procyonidae must not be lost sight of in considering this point of external likeness.
280 ^  Bull. Amer. Mus. Nat. Hist. xii. 1900, p. 109.
281 ^  Proc. Zool. Soc. 1890, p. 98.
282 ^  Temminck, its original describer, placed it in the genus Hyaena.
283 ^  See Garrod, Proc. Zool. Soc. 1878, p. 373.
284 ^  Proc. Zool. Soc. 1899, p. 533.
285 ^  See Beddard, Proc. Zool. Soc. 1900, p. 661, for anatomy.
286 ^  Beddard, Proc. Zool. Soc. 1898, p. 129.
287 ^  It is a curious fact that a native name for the creature is "Pottos" (cf. of course Potto); and indeed the generic name Potos seems to have the priority over Cercoleptes.
288 ^  "Narica" is generally written, after Linnaeus. But this was, according to Mr. Alston, probably an error for nasica.
289 ^  Proc. Zool. Soc. 1870, p. 752.
290 ^  See Wortman, Bull. Amer. Mus. Nat. Hist. vi. 1894, p. 229.
291 ^  As a small point of likeness between this Mustelid and the Procyonidae may be mentioned the colours of the face. M. anakuma is particularly Raccoon-like.
292 ^  See Trans. Zool. Soc. ii. 1841, p. 201.
293 ^  Proc. Zool. Soc. 1894, p. 306.
294 ^  I found fifteen.
295 ^  Ann. Nat. Hist. (6) xiii. 1893, p. 522.
296 ^  See Matschie, SB. Ges. Naturf. Berlin, 1895, p. 171.
297 ^  Proc. Zool. Soc. 1879, p. 305.
298 ^  Lydekker, "Note on the Structure and Habits of the Sea-Otter (Latax lutris)," Proc. Zool. Soc. 1895, p. 421; and ibid. 1896, p. 235.
299 ^  See an article by Mr. Lydekker in Knowledge, April 1898, from which many of the above facts have been taken.
300 ^  "Preliminary Notes on the Characters and Synonymy of the different Species of Otter," Proc. Zool. Soc. 1889, p. 190.
301 ^  Even apparently in the same species.
302 ^  The number of premolars is reduced in the Polar Bear.
303 ^  "The Blue Bear of Thibet," etc., Proc. Zool. Soc. 1897, p. 412.
304 ^  Nouv. Arch. Mus. vii. 1872, Bull. p. 92; and Recherches pour servir à l'histoire naturelle des Mammifères, 1868-1874, p. 321. This genus has quite recently (Lankester, Trans. Linn. Soc. viii. 1901, p. 163) been definitely referred to the Procyonidae.