Chapter XI: The Order Carnivora (3)

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

_Section_ ARCTOIDEA.

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

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

_Family_ URSIDÆ.

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

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

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

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

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

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

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

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

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

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

_Family_ PROCYONIDÆ.

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

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

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

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

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

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

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

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

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

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

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

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

_Family_ MUSTELIDÆ.

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

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

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

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

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