[Illustration: FIG. I.--Left upper sectorial or carnassial teeth of
Carnivora. I, _Felis_; II, _Canis_; III, _Ursus_. 1, anterior, 2,
middle, and 3, posterior cusp of blade; 4, inner cusp supported on
distinct root; 5, inner cusp, posterior in position, and without
distinct root, characteristic of the _Ursidae_.]
The typical section of the group, the Carnivora Vera, Fissipedia or
Carnassidentia, includes all the existing terrestrial members of the
order, together with the otters and sea-otters. In this section the
fore-limbs never have the first digit, or the hind-limbs the first and
fifth digits, longer than the others; and the incisors are 3/3 on each
side, with very rare exceptions. The cerebral hemispheres are more or
less elongated; always with three or four convolutions on the outer
surface forming arches above each other, the lowest surrounding the
Sylvian fissure. In the cheek-series there is one specially modified
tooth in each jaw, to which the name of "sectorial" or "carnassial" is
applied. The teeth in front of this are more or less sharp-pointed and
compressed; the teeth behind broad and tuberculated. The characters of
the sectorial teeth deserve special attention, as, though fundamentally
the same throughout the group, they are greatly modified in different
genera. The upper sectorial is the most posterior of the teeth which
have predecessors, and is therefore reckoned as the last premolar (p. 4
of the typical dentition). It consists of a more or less compressed
blade supported on two roots and an inner lobe supported by a distinct
root (see fig. 1). The blade when fully developed has three cusps (i, 2
and 3), but the anterior is always small, and often absent. The middle
cusp is conical, high and pointed; and the posterior cusp has a
compressed, straight, knife-like edge. The inner cusp. (4) varies in
extent, but is generally placed near the anterior end of the blade,
though sometimes median in position. In the _Ursidae_ alone both the
inner cusp and its root are wanting, and there is often a small internal
and posterior cusp (5) without root. In this family also the sectorial
is relatively to the other teeth much smaller than in other Carnivora.
The lower sectorial (fig. 2) is the most anterior of the teeth without
predecessors in the milk-series, and is therefore reckoned the first
molar. It has two roots supporting a crown, consisting when fully
developed of a compressed bilobed blade (1 and 2), a heel (4), and an
inner tubercle (3). The cusps of the blade, of which the hinder (2) is
the larger, are separated by a notch, generally prolonged into a linear
fissure. In the specialized _Felidae_ (I) the blade alone is developed,
both heel and inner tubercle being absent or rudimentary. In _Meles_
(V) and _Ursus_ (VI) the heel is greatly developed, broad and
tuberculated. The blade in these cases is generally placed obliquely,
its flat or convex (outer) side looking forwards, so that the two lobes
or cusps are almost side by side, instead of anterior and posterior. The
inner tubercle (3) is generally a conical pointed cusp, placed to the
inner side of the hinder lobe of the blade. The special characters of
these teeth are more disguised in the sea-otter than in any other
species, but even here they can be traced.
[Illustration: FIG. 2.--Left lower sectorial or carnassial teeth of
Carnivora, I, _Felis_; II, _Canis_; III, _Herpestes_; IV, _Lutra_; V,
_Meles_; VI, _Ursus_. 1, Anterior cusp of blade; 2, posterior cusp of
blade; 3, inner tubercle; 4, heel. It will be seen that the relative
size of the two roots varies according to the development of the portion
of the crown they respectively support.]
The toes are nearly always armed with large, strong, curved and sharp
claws, ensheathing the terminal phalanges and held firmly in place by
broad plates of bone reflected over their attached ends from the bases
of the phalanges. In the _Felidae_ these claws are "retractile"; the
terminal phalange with the claw attached, folding back in the fore-foot
into a sheath by the outer or ulnar side of the middle phalange of the
digit, and retained in this position when at rest by a strong elastic
ligament. In the hind-foot the terminal joint or phalange is retracted
on to the top, and not the side of the middle phalange. By the action of
the deep flexor muscles the terminal phalanges are straightened, the
claws protruded from their sheath, and the soft "velvety" paw becomes
suddenly converted into a formidable weapon of offence. The habitual
retraction of the claws preserves their points from wear.
The land Carnivora are best divided into two subgroups or sections--(A)
the Aeluroidea, or Herpestoidea, and (B) the Arctoidea; the recognition
of a third section, Cynoidea, being rendered untenable by the evidence
of extinct forms.
(A) _Aeluroidea_.--In this section, which comprises the cats
(_Felidae_), civets (_Viverridae_), and hyenas (_Hyaenidae_), the
tympanic bone is more or less ring-like, and forms only a part of the
outer wall of the tympanic cavity; an inflated alisphenoid bulla is
developed; and the external auditory meatus is short. In the nasal
chamber the maxillo-turbinal is small and doubly folded, and does not
cut off the naso-turbinal and adjacent bones from the nasal aperture.
The carotid canal in the skull is short or absent. Cowper's glands are
present, as is a prostate gland and a caecum, as well as a
duodenal-jejunal flexure in the intestine, but an os penis is either
wanting or small.
Cat tribe.
The members of the cat tribe, or _Felidae_, are collectively
characterized by the following features. An alisphenoid is lacking on
the lower aspect of the skull. In existing forms the usual dental
formula is i. 3/3, c. 1/1, p. 3/2, m. 1/1; the upper molar being
rudimentary and placed on the inner side of the carnassial, but the
first premolar may be absent, while, as an abnormality, there may be a
small second lower molar, which is constantly present in some of the
extinct forms. The auditory bulla and the tympanic are divided by an
internal partition. The paroccipital process is separate from, or only
extends to a slight degree upon the auditory bulla. The thoracic
vertebrae number 13; the feet are digitigrade, with five front and
four hind toes, of which the claws are retractile; and the metatarsus
is haired all round. Anal glands are present.
As regards the teeth, when considered in more detail, the incisors are
small, and the canines large, strong, slightly recurved, with
trenchant edges and sharp points, and placed wide apart. The premolars
are compressed and sharp-pointed; the most posterior in the upper jaw
(the sectorial) being a large tooth, consisting of a compressed blade,
divided into three unequal cusps supported by two roots, with a small
inner lobe placed near the front and supported by a distinct root
(fig. 1, I). The upper molar is a small tubercular tooth placed more
or less transversely at the inner side of the hinder end of the last.
In the lower jaw the molar (sectorial) is reduced to the blade, which
is large, trenchant, compressed and divided into two subequal lobes
(fig. 2, I). Occasionally it has a rudimentary heel, but never an
inner tubercle. The skull generally is short and rounded, though
proportionally more elongated in the larger forms; with the facial
portion short and broad, and the zygomatic arches wide and strong. The
auditory bullae are large, rounded and smooth. Vertebrae: C. 7, D. 13,
L. 7, S. 3, Ca. 13-29. Clavicles better developed than in other
Carnivora, but not articulating with either the shoulder-bones or
sternum. Of the five front toes, the third and fourth are nearly equal
and longest, the second slightly, and the fifth considerably shorter.
The first is still shorter, not reaching the metacarpophalangeal
articulation of the second. In the hind-feet the third and fourth toes
are the longest, the second and fifth somewhat shorter and nearly
equal, while the first is represented only by the rudimentary
metatarsal bone. The claws are large, strongly curved, compressed,
very sharp, and exhibit the retractile condition in the highest
degree. The tail varies greatly in length, being in some species a
mere stump, in others nearly as long as the body. The ears are of
moderate size, more or less triangular and pointed; and the eyes
rather large, with the iris mobile, and with a pupillary aperture
which contracts under the influence of light in some species to a
narrow vertical slit, in others to an oval, and in some to a circular
aperture. The tongue is thickly covered with sharp, pointed, recurved
horny papillae; and the caecum is small and simple.
As in structure so in habits, the cat may be considered the most
specialized of all Carnivora, although they exhibit many features
connecting them with extinct types. All the members of the group feed
almost exclusively on warm-blooded animals which they have themselves
killed, but one Indian species, _Felis viverrina_, is said to prey on
fish, and even fresh-water molluscs. Unlike dogs, they never associate
in packs, and rarely hunt their prey on open ground, but from some
place of concealment wait until the unsuspecting victim comes within
reach, or with noiseless and stealthy tread, crouching close to the
ground for concealment, approach near enough to make the fatal spring.
In this manner they frequently attack and kill animals considerably
exceeding their own size. They are mostly nocturnal, and the greater
number, especially the smaller species, more or less arboreal. None
are aquatic, and all take to the water with reluctance, though some
may habitually haunt the banks of rivers or pools, because they more
easily obtain their prey in such situations. The numerous species are
widely diffused over the greater part of the habitable world, though
most abundant in the warm latitudes of both hemispheres. None are,
however, found in the Australian region, or in Madagascar. Although
the Old World and New World cats (except perhaps the northern lynx)
are all specifically distinct, no common structural character has been
pointed out by which the former can be separated from the latter. On
the contrary, most of the groups into which the family may be divided
have representatives in both hemispheres.
Notwithstanding the considerable diversity in external appearance and
size between different members of this extensive family, the
structural differences are but slight. The principal differences are
to be found in the form of the cranium, especially of the nasal and
adjoining bones, the completeness of the bony orbit posteriorly, the
development of the first upper premolar and of the inner lobe of the
upper sectorial, the length of the tail, the form of the pupil, and
the condition and coloration of the fur, especially the presence or
absence of tufts or pencils of hair on the external ears.
In the typical genus _Felis_, which includes the great majority of the
species, and has a distribution coextensive with that of the family,
the upper sectorial tooth has a distinct inner cusp, the claws are
completely contractile, the tail is long or moderate, and the ears do
not carry distinct tufts of hair. As regards the larger species, the
lion (_F. leo_), tiger (_F. tigris_), leopard (_F. pardus_), ounce or
snow-leopard (_F. uncia_) and clouded leopard (_F. nebulosa_) are
described in separate articles. Of other Old World species it must
suffice to mention that the Tibetan Fontanier's cat (_F. tristis_),
and the Indian marbled cat (_F. marmorata_), an ally of the
above-mentioned clouded leopard, appear to be the Asiatic
representatives of the American ocelots. The Tibetan Pallas's cat (_F.
manul_) has been made the type of a distinct genus, _Trichaelurus_, in
allusion to its long coat. One of the largest of the smaller species
is the African serval, q.v. (_F. serval_), which is yellow with solid
black spots, has long limbs, and a relatively short tail. Numerous
"tiger-cats" and "leopard-cats," such as the spotted _F. bengalensis_
and the uniformly chestnut _F. badia_, inhabit tropical Asia; while
representative species occur in Africa. The jungle-cat (_F. chaus_),
which in its slightly tufted ears and shorter tail foreshadows the
lynxes, is common to both continents. Another African species (_F.
ocreata_) appears to have been the chief progenitor of the European
domestic cat, which has, however, apparently been crossed to some
extent with the ordinary wild cat (_F. catus_). Of the New World
species, _F. concolor_, the puma or couguar, commonly called "panther"
in the United States, is about the size of a leopard, but of a uniform
brown colour, spotted only when young, and is extensively distributed
in both North and South America, ranging between the parallels of 60
deg. N. and 50 deg. S., where it is represented by numerous local
races, varying in size and colour. _F. onca_, the jaguar, is a larger
and more powerful animal than the last, and more resembles the leopard
in its colours; it is also found in both North and South America,
although with a less extensive range, reaching northwards only as far
as Texas, and southwards nearly to Patagonia (see JAGUAR). _F.
pardalis_ and several allied smaller, elegantly-spotted species
inhabiting the intratropical regions of America, are commonly
confounded under the name of ocelot or tiger-cat. _F. yaguarondi_,
rather larger than the domestic cat, with an elongated head and body,
and of a uniform brownish-grey colour, ranges from northern Mexico to
Paraguay; while the allied _F. eyra_ is a small cat, weasel-like in
form, having an elongated head, body and tail, and short limbs, and is
of a uniform light reddish-brown colour. It is a native of South
America and Mexico. _F. pajeros_ is the Pampas cat.
The typical lynxes, as represented by _Lynx borealis_ (_L. lynx_), the
southern _L. pardina_, and the American _L. rufa_, are a northern
group common to both hemispheres, and characterized by their tufted
ears, short tail, and the presence of a rudimentary heel to the lower
carnassial tooth. As a rule, they are more or less spotted in winter,
but tend to become uniformly-coloured in summer. They are connected
with the more typical cats by the long-tailed and uniformly red
caracal, _Lynx (Caracal) caracal_, of India, Persia and Africa, and
the propriety of separating them from _Felis_ may be open to doubt
(see LYNX and CARACAL).
However this may be, there can be no doubt of the right of the
hunting-leopard or chita (cheeta), as, in common with the leopard, it
is called in India, to distinction from all the other cats as a
distinct genus, under the name of _Cynaelurus jubatus_. From all the
other _Felidae_ this animal, which is common to Asia and Africa, is
distinguished by the inner lobe of the upper sectorial tooth, though
supported by a distinct root, having no salient cusp upon it, by the
tubercular molar being more in a line with the other teeth, and by the
claws being smaller, less curved and less completely retractile, owing
to the feebler development of the elastic ligaments. The skull is
short and high, with the frontal region broad and elevated in
consequence of the large development of air-sinuses. The head is small
and round, the body light, the limbs and tail long, and the colour
pale yellowish-brown with small solid black spots (see CHEETA).
Civet tribe.
The family _Viverridae_, which includes the civet-cats, genets and
mongooses, is nearly allied to the _Felidae_, but its members have a
fuller dentition, and exhibit certain other structural differences
from the cats, to the largest of which they make no approach in the
matter of bodily size. As a rule, there is an alisphenoid canal; the
cheek-dentition is p. (3 or 4)/(3 or 4), m. (1 or 2)/(1 or 2). The
bulla is small and the tympanic large, with a low division between
them; and the paroccipital process is leaf-like and spread over the
bulla. The number of dorsal vertebrae, except in the aberrant
_Proteles_, is 13 or 14; the claws may be either completely or
partially retractile or non-retractile; generally each foot has five
toes, but there may be four in front and five behind, the reverse of
this, or only four on each foot; the gait may be either digitigrade or
partially plantigrade; and the metatarsus may be either hairy or naked
inferiorly. Anal, and in some cases also perineal, glands are
developed. The family is limited to the warmer parts of the Old World.
Considerable difference of opinion prevails with regard to the serial
position of the fossa, or foussa (_Cryptoprocta ferox_), of
Madagascar, some writers considering that its affinities are so close
to the _Felidae_ that it ought not to be included in the present
family at all. Others, on the contrary, see no reason to separate it
from the _Viverrinae_ or more typical representatives of the
civet-tribe. As a medium course, it may be regarded as the sole
representative of a special subfamily--_Cryptoproctinae_--of the
_Viverridae_. The subfamily and genus are characterized by possessing
a total of 36 teeth, arranged as i. 3/3, c. 1/1, p. 4/4, m. 1/1. The
teeth generally closely resemble those of the _Felidae_, the first
premolar of both jaws being very minute and early deciduous; the upper
sectorial has a small inner lobe, quite at the anterior part; the
molar is small and placed transversely; and the lower sectorial has a
large trenchant bilobed blade, and a minute heel, but no inner
tubercle. The skull is generally like that of _Felis_, but
proportionally longer and narrower, with the orbit widely open behind.
Vertebrae: C. 7, D. 13, L. 7, S. 3, Ca. 29. Body elongated. Limbs
moderate in size. Feet subplantigrade, with five well-developed toes
on each, carrying sharp, compressed, retractile claws. Ears moderate.
Tail long and cylindrical. The foussa is a sandy-coloured animal with
an exceedingly long tail (see FOUSSA).
The more typical members of the group, constituting the subfamily
_Viverrinae_, are characterized by their sharp, curved and largely
retractile claws, the presence of five toes to each foot, and of
perineal and one pair of anal glands, and a tympanic bone which
retains to a great extent the primitive ring-like form, so that the
external auditory meatus has scarcely any inferior lip, its orifice
being close to the tympanic ring. The first representatives of the
subfamily are the civet-cats, or civets (_Viverra_ and _Viverricula_),
and the genets (_Genetta_), in all of which the dentition is _i. 3/3,
c. 1/1, p. 4/4, m. 2/2_; total 40. The skull is elongated, with the
facial portion small and compressed, and the orbits well-defined but
incomplete behind. Vertebrae: C. 7, D. 13, L. 7 (or D. 14, L. 6), S.
3, Ca. 22-30. Body elongated and compressed. Head pointed in front;
ears rather small. Extremities short. Feet small and rounded. Toes
short, the first on fore and hind feet much shorter than the others.
Palms and soles covered with hair, except the pads of the feet and
toes, and in some species a narrow central line on the under side of
the sole, extending backwards nearly to the heel. Tail moderate or
long. The pair of large glands situated on the perineum (in both
sexes) secretes an oily substance of a peculiarly penetrating odour.
In the true civets, which include the largest members of the group,
the teeth are stouter and less compressed than in the other genera;
the second upper molar being especially large, and the auditory bulla
smaller and more pointed in front; the body is shorter and stouter;
the limbs are longer; the tail shorter and tapering. The under side of
the tarsus is completely covered with hair, and the claws are longer
and less retractile. Fur rather long and loose, and in the middle line
of the neck and back especially elongated so as to form a sort of
crest or mane. Pupil circular when contracted. Perineal glands greatly
developed. These characters apply especially to _V. civetta_, the
African civet, or civet-cat, as it is commonly called, an animal
rather larger than a fox, and an inhabitant of intratropical Africa.
_V. zibetta_, the Indian civet, of about equal size, approaches in
many respects, especially in the characters of the teeth and feet and
absence of the crest of elongated hair on the back, to the next
section. It inhabits Bengal, China, the Malay Peninsula and adjoining
islands. _V. tangalunga_ is a smaller but nearly allied animal from
the same part of the world. From these three species and the next the
civet of commerce, once so much admired as a perfume in England, and
still largely used in the East, is obtained. The animals are kept in
cages, and the odoriferous secretion collected by scraping the
interior of the perineal follicles with a spoon or spatula. The single
representative of the genus _Viverricula_ resembles in many respects
the genets, but agrees with the civets in having the whole of the
under side of the tarsus hairy; the alisphenoid canal is generally
absent. _V. malaccensis_, the rasse, inhabiting India, China, Java and
Sumatra, is an elegant little animal which affords a favourite perfume
to the Javanese. The genets (_Genetta_) are smaller animals, with more
elongated and slender bodies, and shorter limbs than the civets. The
skull is elongated and narrow; and the auditory bulla large, elongated
and rounded at both ends. The teeth are compressed and sharp-pointed,
with a lobe on the inner side of the third, upper premolar not present
in the previous genera. Pupil contracting to a linear aperture. Tail
long, slender, ringed. Fur short and soft, spotted or cloudy. Under
side of the metatarsus with a narrow longitudinal bald streak.
_Genetta vulgaris_, or _G. genetta_, the common genet, is found in
France south of the river Loire, Spain, south-western Asia and North
Africa. _G. felina, senegalensis, tigrina, victoriae_ and _pardalis_
are other named species, all African in habitat.
The Malagasy fossane (_Fossa daubentoni_), which has but little
markings on the fur of the adult, differs by the absence of a
scent-pouch and the presence of a couple of bare spots on the under
surface of the metatarsus. The beautiful linsangs (_Linsanga_ or
_Prionodon_), ranging from the eastern Himalaya to Java and Borneo,
are represented by two or three species, easily recognizable by the
broad transverse bands of blackish brown and yellow with which the
body and tail are marked. They are specially distinguished by having
only one pair of upper molars, thereby resembling the cats, with
which, in correlation with their arboreal habits, they agree in their
highly retractile claws, and the hairy surface of the under side of
the metatarsus. About 15 in. is the length of the type species. In
West Africa the linsangs are represented by _Poiana richardsoni_, a
small species with a spotted genet-like coat, and also with a narrow
naked stripe on the under surface of the metatarsus, as in genets.
Here may be placed the two African spotted palm-civets of the genus
_Nandinia_, namely _N. binotata_ from the west and _N. gerrardi_ from
the east forest-region. In common with the true palm-civets, they have
a dentition numerically identical with that of _Viverra_ and
_Genetta_, but the cusps of the hinder premolars and molars are much
less sharp and pointed. They are peculiar in that the wall of the
inner chamber of the auditory bulla never ossifies, while the
paroccipital process is not flattened out and spread over the bulla.
In this respect they resemble the Miocene European genus _Amphictis_,
as they do in the form of their teeth, so that they may be regarded as
nearly related to the ancestral _Viverridae_, and forming in some
degree a connecting link between the present and the next subfamily.
_Nandinia_ is also peculiar in possessing a kind of rudimentary
marsupial pouch. Apparently _Eupleres goudoti_, of Madagascar, which
has been generally classed in the _Herpestinae_, is a nearly related
animal, characterized by the reduction of its dentition, due to
insectivorous habits (fig. 3); the canines being small, the anterior
premolars canine-like, and the hinder premolars molariform. It is a
uniformly-coloured creature of medium size.
[Illustration: FIG. 3.--Skull of _Eupleres goudoti_.]
The palm-civets, or paradoxures, constituting the Asiatic genus
_Paradoxurus_, have, as already stated, the following dental formula,
viz. i. 3/3, c. 1/1, p. 4/4, m. 2/2, total 40; the cusps of the molars
being low and blunted, and these teeth in the upper jaw much broader
than in the civets. The head is pointed in front, with small rounded
ears; the limbs are of medium length, with the soles of the feet
almost completely naked, and fully retractile claws; while the long
tail is not prehensile and clothed with hair of moderate length. Spots
are the chief type of marking. The vertebrae number C. 7, D. 13, L. 7,
S. 3, Ca. 29-36. Numerous relatively large species ranging from India
to Borneo, Sumatra and Celebes, with one in Tibet, represent the
genus. Nearly allied are _Arctogale leucotis_, with a wide
distribution, and _A. trivirgata_, of Java, both longitudinally
striped species, with small and slightly separated molars, and a
prolonged bony palate (see PALM-CIVET).
The binturong (_Arctictis binturong_) has typically the same dental
formula as the last, but the posterior upper molar and the first lower
premolar are often absent. Molars small and rounded, with a distinct
interval between every two, but formed generally on the same pattern
as _Paradoxurus_. Vertebrae: C. 7, D. 14, L. 5, S. 3, Ca. 34. Body
elongated; head broad behind, with a small pointed face, long and
numerous whiskers, and small ears, rounded, but clothed with a pencil
of long hairs. Eyes small. Limbs short, with the soles of the feet
broad and entirely naked. Tail very long and prehensile. Fur long and
harsh. Caecum extremely small. The binturong inhabits southern Asia
from Nepal through the Malay Peninsula to the islands of Sumatra and
Java. Although structurally agreeing closely with the paradoxures, its
tufted ears, long, coarse and dark hair, and prehensile tail give it a
very different external appearance. It is slow and cautious in its
movements, chiefly if not entirely arboreal, and appears to feed on
vegetables as well as animal substances (see BINTURONG).
_Hemigale_ is another modification of the paradoxure type, represented
by _H. hardwickei_ of Borneo, an elegant-looking animal, smaller and
more slender than the paradoxures, of light grey colour, with
transverse broad dark bands across the back and loins.
_Cynogale_ also contains one Bornean species, _C. bennetti_, a curious
otter-like modification of the viverrine type, having semi-aquatic
habits, both swimming in the water and climbing trees, living upon
fish, crustaceans, small mammals, birds and fruits. The number and
general arrangement of the teeth are as in _Paradoxurus_, but the
premolars are peculiarly elongated, compressed, pointed and recurved,
though the molars are tuberculated. The head is elongated, with the
muzzle broad and depressed, the whiskers are very long and abundant,
and the ears small and rounded. Toes short and slightly webbed at the
base. Tail short, cylindrical, covered with short hair. Fur very dense
and soft, of a dark-brown colour, mixed with black and grey.
In the mongoose group, or _Herpestinae_, the tympanic or anterior
portion of the auditory bulla is produced into an ossified external
auditory meatus of considerable length; while the paroccipital process
never projects below the bulla, on the hinder surface of which, in
adult animals, it is spread out and completely lost. The toes are
straight, with long, unsheathed, non-retractile claws.
In the typical mongooses or ichneumons, _Herpestes_, the dental
formula is i. 3/3, c. 1/1, p. (4 or 3)/(4 or 3), m. 2/2; total 40 or
36; the molars having generally strongly-developed, sharply-pointed
cusps. The skull is elongated and constricted behind the orbits. The
face is short and compressed, with the frontal region broad and
arched. Post-orbital processes of frontal and jugal bones well
developed, generally meeting so as to complete the circle of the orbit
behind. Vertebrae: C. 7, D. 13, L. 7, S. 3, Ca. 21-26. Head pointed in
front. Ears short and rounded. Body long and slender. Extremities
short. Five toes on each foot, the first, especially that on the
hind-foot, very short. Toes free, or but slightly palmated. Soles of
fore-feet and terminal portion of those of hind-pair naked; under
surface of metatarsus clothed with hair. Tail long or moderate,
generally thick at the base, and sometimes covered with more or less
elongated hair. The longer hairs covering the body and tail almost
always ringed. The genus is common to the warmer parts of Asia and
Africa, and while many of the species, like the Egyptian _H.
ichneumon_ and the ordinary Indian mongoose, _H. mungo_, are
pepper-and-salt coloured, the large African _H. albicauda_ has the
terminal two-thirds of the tail clothed with long white hairs (see
ICHNEUMON).
The following distinct African and Malagasy generic representatives of
the subfamily are recognized, viz. _Helogale_, with 3/3 premolars, and
containing the small South African _H. parvula _and a variety of the
same. _Bdeogale crassicauda_ and two allied tropical African species
differ from _Herpestes_ in having only four toes on each foot. The
orbit is nearly complete, and the tail of moderate length and rather
bushy. In _Cynictis_, which has the orbit completely closed, there are
five front and four hind toes; and the skull is shorter and broader
than in _Herpestes_, rather contracted behind the orbits, the face
short, and the anterior chamber of the auditory bulla very large. The
front claws are elongated. Includes only _C. penicillata_ from South
Africa.
All the foregoing herpestines have the nose short, with its under
surface flat, bald, and with a median longitudinal groove. The
remaining forms have the nose more or less produced, with its under
side convex, and a space between the nostrils and the upper lip
covered with closely pressed hairs, and without any median groove. The
South African _Rhynchogale muelleri_, a reddish animal with five toes
to each foot and 4/4 (abnormally 5/5) premolars, alone represents the
first genus. The cusimanses (_Crossarchus_), which differ by having
only 3/3 premolars, and thus a total of 36 teeth, include, on the
other hand, several species. The muzzle is elongated, the claws on the
fore-feet are long and curved, the first front toe is very short; the
under surface of the metatarsus naked; and the tail shorter than the
body, tapering. Fur harsh. Includes _C. obscurus_, the cusimanse, a
small burrowing animal from West Africa, of uniform dark-brown colour,
_C. fasciatus, C. zebra, C. gambianus_ and others. Lastly, we have
_Suricata_, a more distinct genus than any of the above. The dental
formula is as in the last, but the teeth of the molar series are
remarkably short in the antero-posterior direction, corresponding with
the shortness of the skull generally. Orbits complete behind.
Vertebrae: C. 7, D. 15, L. 6, S. 3, Ca. 20. Though the head is short
and broad, the nose is pointed and rather produced and movable, while
the ears are very short. Body shorter and limbs longer than in
_Herpestes_. Toes 4-4. Claws on fore-feet very long and narrow,
arched, pointed and subequal. Hind-feet with shorter claws, soles
hairy. Tail rather shorter than the body. One species only is known,
the meerkat or suricate, _S. tetradactyla_, a small grey-brown animal,
with dark transverse stripes on the hinder part of the back, from
South Africa.
The names _Galidictis, Galidia_ and _Hemigalidia_ indicate three
generic modifications of the _Herpestinae_, all inhabitants of
Madagascar. The best-known, _Galidia elegans, is _a lively
squirrel-like little animal with soft fur and a long bushy tail, which
climbs and jumps with agility. It is of a chestnut-brown colour, the
tail being ringed with darker brown. _Galidictis vittata_ and _G.
striata_ chiefly differ from the ichneumons in their coloration, being
grey with parallel longitudinal stripes of dark brown.
Considerable diversity of opinion prevails with regard to the serial
position of the aard-wolf, or maned jackal (_Proteles cristatus_), of
southern and eastern Africa, some authorities making it the
representative of a family by itself, others referring it to the
_Hyaenidae_, while others again regard it as a modified member of the
_Viverridae_. After all, the distinction either way cannot be very
great, since the two families just named are intimately connected by
marks of the extinct _Ictitherium_, With the _Viverridae_ it agrees in
having the auditory bulla divided, while in the number of dorsal
vertebrae it is hyena-like. The cheek-teeth are small, far apart, and
almost rudimentary in character (see fig. 4), and the canines long and
rather slender. The dental formula is i. 3/3, c. 1/1, p. m. 4/(3 or 4);
total 30 or 32. Vertebrae: C. 7, D. 15, L. 5, S. 2, Ca. 24. The
fore-feet with five toes; the first, though short, with a distinct
claw. The hind-feet with four subequal toes; all, like those of the
fore-foot, furnished with strong, blunt, non-retractile claws (see
AARD-WOLF).
Hyena tribe.
The hyenas or hyaenas (_Hyaenidae_) differ from the preceding family
(_Viverridae_) in the absence of a distinct vertical partition between
the two halves of the auditory bulla; and are further characterized by
the absence of an alisphenoid canal, the reduction of the molars to
1/1, and the presence of 15 dorsal vertebrae. The dental formula in
the existing forms (to which alone all these remarks apply) is i. 3/3,
c. 1/1. p. 4/3 m. 1/1; total 34; the teeth, especially the canines and
premolars, being very large, strong and conical. Upper sectorial with
a large, distinctly trilobed blade and a moderately developed inner
lobe placed at the anterior extremity of the blade. Molar very small,
and placed transversely close to the hinder edge of the last, as in
the _Felidae_. Lower sectorial consisting of little more than the
bilobed blade. Zygomatic arches of skull very wide and strong; and
sagittal crest high, giving attachment to very powerful biting
muscles. Orbits incomplete behind. Vertebrae: C. 7, D. 15, L. 5, S. 4,
Ca. 19. Limbs rather long, especially the anterior pair, digitigrade,
four subequal toes on each, with stout non-retractile claws, the first
toes being represented by rudimentary metacarpal and metatarsal bones.
Tail rather short. A large post-anal median glandular pouch, into
which the largely developed anal scent glands pour their secretion.
[Illustration: FIG. 4.--Skull and Dentition of Aard-Wolf (_Proteles
cristatus_.)]
The three well-characterized species of _Hyaena_ are divisible into
two sections, to which some zoologists assign generic rank. In the
typical species the upper molar is moderately developed and
three-rooted; and an inner tubercle and heel more or less developed on
the lower molar. Ears large and pointed. Hair long, forming a mane on
the back and shoulders. Represented firstly by _H. striata_, the
striped hyena of northern and eastern Africa and southern Asia; and
_H. brunnea_ of South Africa, in some respects intermediate between
this and the next section. In the second section, forming the subgenus
_Crocuta_, the upper molar is extremely small, two- or one-rooted,
often deciduous; the lower molar without trace of inner tubercle, and
with an extremely small heel. Ears moderate, rounded. Hair not
elongated to form a mane. The spotted hyena, _Hyaena (Crocuta)
crocuta_, of which, like the striped species, there are several local
races, represents this group, and ranges all over Africa south of the
Sahara. In dental characters the first section inclines more to the
_Viverridae_, the second to the _Felidae_; or the second may be
considered as the more specialized form, as it certainly is in its
visceral anatomy, especially in that of the reproductive organs of the
female. (See HYENA.)
(B) _Arctoidea_.--So far as the auditory region of the skull is
concerned, the existing representatives of the dog tribe or _Canidae_
are to a great extent intermediate between the cat and civet group
(_Aeluroidea_) on the one hand, and the typical representatives of the
bear and weasel group on the other. They were consequently at one time
classed in an intermediate group--the Cynoidea; but fossil forms show
such a complete transition from dogs to bears as to demonstrate the
artificial character of such a division. Consequently, the dogs are
included in the bear-group. In this wider sense the Arctoidea will be
characterized by the tympanic bone being disk-shaped and forming the
whole of the outer wall of the tympanic cavity; the large size of the
external auditory meatus or tube; and the large and branching
maxillo-turbinal bone, which cuts off the naso-turbinal and two adjacent
bones from the anterior nasal chamber. The tympanic bulla has no
internal partition. There is a large carotid canal. Cowper's glands are
lacking; and there is a large penial bone.
Dog tribe.
From all the other members of the group the _Canidae_ are broadly
distinguished (in the case of existing forms) by the large and
well-developed tympanic bulla, with which the paroccipital process is
in contact. An alisphenoid canal is present. The feet are digitigrade,
usually with five (in one instance four) front and always four
hind-toes. The molars--generally 2/3--have tall cusps, and the
sectorials are large and powerful (figs. 1 and 2). The intestine has
both a duodeno-jejunal flexure and a caecum. A prostate gland is
present; but there are no glands in the vasa deferentia; the penial
bone is grooved; and anal glands are generally developed. The
distribution of the family is cosmopolitan. The normal dentition is i.
3/3, c. 1/1, p. 4/4, m. 2/3; total 42; thus differing from the typical
series only by the loss of the last pair of upper molars (present in
certain extinct forms). In the characters of the teeth the group is
the most primitive of all Carnivora. Typically the upper secterial
(fig. 1, II) consists of a stout blade, of which the anterior cusp is
almost obsolete, the middle cusp large, conical and pointed backwards,
and the posterior cusp in the form of a compressed ridge; the inner
lobe is very small, and placed at the fore part of the tooth. The
first molar is more than half the antero-posterior length of the
sectorial, and considerably wider than long; its crown consists of two
prominent conical cusps, of which the anterior is the larger, and a
low, broad inward prolongation, supporting two more or less distinct
cusps and a raised inner border. The second molar resembles the first
in general form, but is considerably smaller. The lower sectorial
(fig. 2, II) is a large tooth, with a strong compressed bilobed blade,
the hinder lobe being considerably the larger and more pointed, a
small but distinct inner tubercle placed at the hinder margin of the
posterior lobe of the blade, and a broad, low, tuberculated heel,
occupying about one-third of the whole length of the tooth. The second
molar is less than half the length of the first, with a pair of cusps
placed side by side anteriorly, and a less distinct posterior pair.
The third is an extremely small and simple tooth with a subcircular
tuberculated crown and single root.
Views differ in regard to the best classification of the _Canidae_,
some writers adopting a number of generic groups, while others
consider that very few meet the needs of the case. In retaining the
old genus _Canis_ in the wide sense, that is to say, inclusive of the
foxes, Professor Max Weber is followed. The best cranial character by
which the different members of the family may be distinguished is that
in dogs, wolves and jackals the post-orbital process of the frontal
bone is regularly smooth and convex above, with its extremity bent
downwards, whereas in foxes the process is hollowed above, with its
outer margin (particularly of the anterior border) somewhat raised.
This modification coincides in the main with the division of the group
into two parallel series, the Thooids or Lupine forms and Alopecoids
or Vulpine forms, characterized by the presence of frontal air-sinuses
in the former, which not only affects the external form but to a still
greater degree the shape of the anterior part of the cranial cavity,
and the absence of such sinuses in the latter. The pupil of the eye
when contracted is round in most members of the first group, and
vertically elliptical in the others, but more observations are
required before this character can be absolutely relied upon. The form
and length of the tail is often used for the purposes of
classification, but its characters do not coincide with those of the
cranium, as many of the South American _Canidae_ have the long bushy
tails of foxes and the skulls of wolves.
[Illustration: FIG. 5.--The African Hunting-Dog (_Lycaon pictus_).]
The most aberrant representative of the thooid series is the African
hunting-dog (_Lycaon pictus_, fig. 5), which differs from the other
members of this series by the teeth being rather more massive and
rounded, the skull shorter and broader, and the presence of but four
toes on each limb, as in _Hyena_. The hunting-dog, from south and east
Africa, is very distinct externally from all other _Canidae_; being
nearly as large as a mastiff, with large, broadly ovate erect ears and
a singular colouring, often consisting of unsymmetrical large spots of
white, yellow and black. It presents some curious superficial
resemblances to _Hyena crocuta_, perhaps a case of mimetic analogy,
and hunts its prey in large packs. Several local races, one of which
comes from Somaliland, differing in size and colour, are recognized
(see HUNTING-DOG). Nearly related to the hunting-dog are the dholes or
wild dogs of Asia, as represented by the Central Asian _Cyan
primaevus_ and the Indo-Malay _C. javanicus_. They have, however, five
front-toes, but lack the last lower molar; while they agree with
_Lycaon_ and _Speothos_ in that the heel of the lower sectorial tooth
has only a single compressed cutting cusp, in place of a large outer
and a smaller inner cusp as in _Canis_. Dholes are whole-coloured
animals, with short heads; and hunt in packs. The bush-dog
(_Speothos_, or _Icticyon venaticus_) of Guiana is a small,
short-legged, short-tailed and short-haired species characterized by
the molars being only (2 or 1)/2; the carnassial having no inner cusp.
The long-haired raccoon-dog (_Nyctereutes procyonoides_) of Japan and
China agrees essentially in everything but general appearance (which
is strangely raccoon-like) with _Canis_. The typical group of the
latter includes some of the largest members of the family, such as the
true wolves of the northern parts of both Old and New Worlds (_C.
lupus, &c._), and the various breeds of the domestic dog (_C.
familiaris_), the origin of which is still involved in obscurity.
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 half-wild 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 is
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 (see
DOG). The dingo or Australian dog is met with wild, and also as the
domestic companion of the aboriginal race of the country, by whom it
appears to have been originally introduced. It is nearly related to a
half-wild dog inhabiting Java, and also to the pariah dogs of India
and other eastern countries. 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 18th century. The slender-jawed _C. simensis_ of Abyssinia and the
South American _C. jubatus_ and _C. antarcticus_ are also generally
placed in this group. On the other hand, the North American coyote
(_C. latrans_), with its numerous subspecies, and the Old World
jackals, such as the Indo-European _C. aureus_ the Indian _C.
pallipes_, and the African _C. lupaster, C. anthus, C. adustus, C.
variegatus_ and _C. mesomelas_ (the black-backed jackal), although
closely related to the wolves, have been placed in a separate group
under the name of _Lupulus_. Again, _Thous_ (or _Lycalopex_), is a
group proposed for certain South American _Canidae_, locally known as
foxes, and distinguished from all the foregoing by their fox-like
aspect and longer tails, although with skulls of the thooid type.
Among these are the bright-coloured colpeo, _C. magellanicus_, the
darker _C. thous, C. azarae, C. griseus, C. cancrivorus_ and _C.
brasiliensis_. Some of these, such as _C. azarae_ and _C. griseus_,
show a further approximation to the fox in that the pupil of the eye
forms a vertical slit. More distinct from all the preceding are the
members of the alopecoid or vulpine section, which are unknown in
South America. The characteristic feature of the skull has been
already mentioned. In addition to this, reference may be made to the
elliptical (in place of circular) pupil of the eye, and the general
presence of ten (rarely eight) teats instead of a smaller number. The
typical groups constituting the subgenus (or genus) _Vulpes_, is
represented by numerous species and races spread over the Old World
and North America. Foremost among these is the European fox (_C.
vulpes_--otherwise _Vulpes alopex_, or _V. vulpes_), represented in
the Himalaya by the variety _C. v. montanus_ and in North Africa by
_C. v. niloticus_, while the North American _C. pennsylvanicus_ or
_fulvus_, can scarcely be regarded as more than a local race. On the
other hand, the Asiatic _C. bengalensis_ and _C. corsac_, and the
North American _C. velox_ (kit-fox) are smaller and perfectly distinct
species. From all these the North American _C. cinereo-argentatus_
(grey fox) and _C. littoralis_ are distinguished by having a fringe of
stiff hairs in the tail, whence they are separated as _Urocyon_.
Again, the Arctic fox (_C. lagopus_), of which there is a blue and a
white phase, has the tail very full and bushy and the soles of the
feet thickly haired, and has hence been distinguished as _Leucocyon_.
Lastly, we have the elegant little African foxes known as fennecs
(_Fennecus_), such as _C. zerda_ and _C. famelicus_ of the north, and
the southern _C. chama_, all pale-coloured animals, with enormously
long ears, and correspondingly inflated auditory bullae to the skull
(see WOLF, JACKAL, FOX).
Whatever differences of opinion may obtain among naturalists as to the
propriety of separating generically the foxes from the wolves and
dogs, there can be none as to the claim of the long-eared fox
(_Otocyon megalotis_) of south and east Africa to represent a genus by
itself. In this animal the dental formula is i. 3/3, c. 1/1, p. 4/4, m.
(3 or 4)/4; total 46 or 48. The molar teeth being in excess of almost
all other placental mammals with a differentiated series of teeth.
They have the same general characters as in _Canis_, with very pointed
cusps. The lower sectorial shows little of the typical character,
having five cusps on the crown-surface; these can, however, be
identified as the inner tubercle, the two greatly reduced and
obliquely placed lobes of the blade, and two cusps on the heel. The
skull generally resembles that of the smaller foxes, particularly the
fennecs. The auditory bullae are very large. The hinder edge of the
lower jaw has a peculiar form, owing to the great development of an
expanded, compressed and somewhat inverted subangular process.
Vertebrae: C. 7, D. 13, L. 7, S. 3, Ca. 22. Ears very large. Limbs
rather long, with the normal number of toes. The two parietal ridges
on the skull remain widely separated, so that no sagittal crest is
formed. The animal is somewhat smaller than an ordinary fox. In the
year 1880 Professor Huxley suggested that in the long-eared fox we
have an animal nearly representing the stock from which have been
evolved all the other representatives of the dog and fox tribe. One of
the main grounds for arriving at this conclusion was the fact that
this animal has very generally four true molars in each jaw, and
always that number in the lower jaw; whereas three is the maximum
number of these teeth to be met with in nearly all placental mammals,
other than whales, manatis, armadillos and certain others. The
additional molars in _Otocyon_ were regarded as survivals from a
primitive type when a larger number was the rule. Palaeontology has,
however, made great strides since 1880, and the idea that the earlier
mammals had more teeth than their descendants has not only received no
confirmation, but has been practically disproved. Consequently Miss
Albertina Carlsson had a comparatively easy task (in a paper published
in the _Zoologisches Jahrbuch_ for 1905) in demonstrating that the
long-eared fox is a specialized, and to some extent degraded, form
rather than a primitive type. This, however, is not all, for the lady
points out that, as was suggested years previously by the present
writer, the creature is really the descendant of the fossil _Canis
curvipalatus_ of northern India. This is a circumstance of
considerable interest from a distributional point of view, as
affording one more instance of the intimate relationship between the
Tertiary mammalian fauna of India and the existing mammals of Africa.
In regard to the members of the dog-tribe as a whole, it may be stated
that 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 upper surface of the tail of the fox has a gland
covered with coarse straight hair. This gland, which emits an aromatic
odour, is found in all _Canidae_, with possibly the exception of
_Lycaon pictus_. Although the bases of the hair covering the gland are
usually almost white, the tips are always black; this colour being
generally extended to the surrounding hairs, and often forming dark
bars on the buttocks. The dark spot on the back of the tail is
particularly conspicuous, notably in such widely separated species as
the wolves, Azara's dog and the fennec.
Bear tribe.
Although its existing representatives are very different, the
bear-family or _Ursidae_, as will be more fully mentioned in the
sequel, was in past times intimately connected with the _Canidae_. In
common with the next two families, the modern _Ursidae_ are
characterized by the very small tympanic bulla, and the broad
paroccipital process, which is, however, independent of the bulla. The
feet are more or less completely plantigrade and five-toed. The
intestine has neither duodeno jejunal flexure nor a caecum; the
prostate gland is rudimentary; but glands occur in the vasa
deferentia; and the penial bone is cylindrical. As distinctive
characteristics of the _Ursidae_, may be mentioned the presence of an
alisphenoid canal on the base of the skull; the general absence of a
perforation on the inner side of the lower end of the humerus; the
presence of two pairs of upper and three of lower molars, which are
mostly elongated and low-cusped; and the non-cutting character and
fore-and-aft shortening of the upper sectorial, which has no inner
root and one inner cusp (fig. I, III.). Anal glands are apparently
wanting. The short tail, bulky build, completely plantigrade feet and
clumsy gait are features eminently characteristic of the bears.
The great majority of existing bears may be included in the typical
genus _Ursus_, of which, in this wide sense, the leading
characteristics will be as follows. The dentition is i. 3/3, c. 1/1,
p. 4/4, m. 2/3 = 42; but the three anterior premolars, above and
below, are 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 cheek-series. The fourth (upper sectorial) differs
essentially from the corresponding tooth of other Carnivora in that
the inner lobe is not supported by a distinct root; its sectorial
characters being very slightly marked. The crowns of both true molars
are longer than broad, with flattened, tuberculated, grinding
surfaces; the second having a large backward prolongation or heel. The
lower sectorial 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; while the third is
smaller. The milk-teeth are comparatively small, and shed at an early
age. The skull is more or less elongated, with the orbits small and
incomplete behind, and the palate prolonged considerably behind the
last molar. Vertebrae: C. 7, D. 14, L. 6, S. 5, Ca. 8-10. Body heavy.
Feet broad, completely plantigrade; the five toes on each well
developed, and armed with long compressed and moderately curved,
non-retractile claws, the soles being generally naked. Tail very
short. Ears moderate, erect, rounded, hairy. Fur generally long, soft
and shaggy.
Bears are 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, although absent from Africa
south of the Sahara and Australasia. As a rule, they are omnivorous,
or vegetable feeders, even the polar bear, which subsists for most of
the year on flesh and fish, eating grass in summer. On the other hand,
many of the brown bears live largely on salmon in summer. Among the
various species the white polar bear of the Arctic regions, _Ursus
(Thalassarctus) maritimus_, differs from the rest by its small and low
head, small, narrow and simple molars, and the presence of a certain
amount of hair on the soles of the feet. The typical group of the
genus is represented by the brown bear (_U. arctus_) of Europe and
Asia, of which there are many local races, such as the Syrian _U. a.
syriacus_, the Himalayan _U. a. isabellinus_, the North Asiatic _U. a.
collaris_, and the nearly allied Kamchadale race, which is of great
size. In Alaska the group is represented by huge bears, which can
scarcely claim specific distinctness from _U. arctus_; and if these
are ranked only as races, it is practically impossible to regard the
Rocky Mountain grizzly bear (_U. horribilis_) as of higher rank,
although it naturally differs more from the Asiatic animal. On the
other hand, the small and light-coloured _U. pruinosus_ of Tibet may
be allowed specific rank. More distinct is the North American black
bear _U. americanus_, and its white relative _U. kermodei_ of British
Columbia; and perhaps we should affiliate to this group the Himalayan
and Japanese black bears (_U. torquatus_ and _U. japonicus_). Very
distinct is the small Malay sun-bear _U. (Helarctus) malayanus_,
characterized by its short, smooth fur, extensile tongue, short and
wide head, and broad molars. Finally, the spectacled bear of the
Andes, _U. (Tremarctus) ornatus_, which is also a broad-skulled black
species, differs from all the rest in having a perforation, or
foramen, on the inner side of the lower end of the humerus. A second
genus, _Melursus_, represented by the Indian sloth-bear (_M.
ursinus_), differs from the preceding in having only two pairs of
upper incisors, the small size of the cheek-teeth, and the extensile
lips. Ants, white-ants, fruits and honey form the chief food of this
shaggy black species,---a diet which accounts for its feeble dentition
(see BEAR).
[Illustration: FIG. 6.--The Parti-coloured Bear, or Giant Panda
(_Aeluropus melanoleucus_).]
The parti-coloured bear or giant panda (_Aeluropus melanoleucus_, fig.
6) of eastern Tibet and north-west China forms in some degree a
connecting link between the bears and the true panda, although placed
by Professor E.R. Lankester in the same family as the latter. In the
number of the teeth, and to some extent in the character of the
molars, as well as in the abbreviated tail, _Aeluropus_ resembles the
bears, but in the structure of the sectorial tooth, the presence of an
extra radial carpal bone, and the osteology generally, it is more like
the panda. In the absence of an alisphenoid canal to the skull it
differs both from the latter and the bears, and thereby resembles the
raccoons; while in having a perforation at the lower end of the
humerus, it agrees with the spectacled bear, the panda and raccoons.
The dentition is i. 3/3, c. 1/1, p. 4/3, m. 2/3; total 40; premolars
increasing in size from first to last, and two-rooted except the
first; the first upper molar with quadrate crown, broader than long;
and the second larger than the first. Skull with the zygomatic arches
and sagittal crest immensely developed, ascending branch of lower jaw
very high, giving great space for attachment of temporal muscle, and
facial portion short. Bony palate not extending behind the last molar.
No alisphenoid canal. Feet bear-like, but soles more hairy, and
perhaps less completely plantigrade. Fur long and thick; and tail
extremely short. Humerus with a perforation on the inner side of the
lower end; a very large extra radial carpal bone. The colour of this
strange animal is black and white (fig. 6).
With the panda (_Aelurus fulgens_) we reach an undoubted
representative of the _Procyonidae_, or raccoon tribe, differing,
however, from all the rest except the doubtful _Aeluropus_, in its
Asiatic habitat. If the latter be included, the family may be defined
as follows. Molars 3/2, except in _Aeluropus_, with blunt or sharp
cusps; no alisphenoid canal, except in _Aelurus_; humerus generally
with a foramen; feet plantigrade; tail, except in _Aeluropus_, long
and generally ringed.
In the panda the dentition is i. 3/3, c. 1/1, p. 3/4, m. 2/2; total
38; the first lower molar being minute and deciduous, and the upper
molars broad with numerous and complicated cusps. Vertebrae: C. 7,
D. 14, L. 6, S. 3, Ca. 18. Skull high and compressed, with an
alisphenoid canal, a short facial portion, and the ascending branch of
the lower jaw, as in _Aeluropus_, very tall. Face cat-like, with
moderate, erect, pointed ears. Claws blunt. Tail cylindrical and
ringed. Fur long and thick. Extra radial carpal bone moderate. The
panda is a bright golden red animal, with black under-parts, ranging
from the eastern Himalaya to north-western China, where it is
represented by a distinct race. Fossil species occur in the later
Tertiary deposits of Europe (see PANDA).
The raccoons (_Procyon_) are the first and typical representatives of
the American section of the family, in which an alisphenoid canal is
always wanting. In this genus the dentition is i. 3/3, c. 1/1, p. 4/4,
m. 2/2; total 40; the upper molars being broad and tuberculated; the
upper sectorial (like that of _Aeluropus_ and _Aelurus_) having three
outer cusps and a broad bicuspid inner lobe, giving an almost quadrate
form to the crown. First upper molar with a large tuberculated crown,
rather broader than long; second considerably smaller, with
transversely oblong crown. Lower sectorial (first molar) with an
extremely small and ill-defined blade, placed transversely in front,
and a large inner tubercle and heel; second molar as long as the
first, but narrower behind, with five obtuse cusps. Vertebrae: C. 7,
D. 14, L. 6, S. 3, Ca. 16-20. Body stout. Head broad behind, but with
a pointed muzzle. In walking the entire sole 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 and pointed. Tail moderately long, cylindrical,
thickly covered with hair, ringed, non-prehensile. Fur long, thick and
soft. The common raccoon (_Procyon lotor_) of North America is the
type of this genus; it is replaced in South America by _P.
cancrivorus_ (see RACCOON). The cacomistles (_Bassariscus_) are nearly
allied to _Procyon_, but of more slender and elegant proportions, with
sharper nose, longer tail, and more digitigrade feet, and teeth
smaller and more sharply cusped. The typical _B. astuta_ is from the
southern parts of the United States and Mexico, while _B. (Wagneria)
annulata_ is Mexican and Central American.
The name _Bassaricyon_ has been given to a distinct modification of
the procyonine type of which at present two species are known, one
from Costa Rica and the other from Ecuador respectively, named _B.
gabbi_ and _B. alleni_. They much resemble the kinkajou in external
appearance, but the skull and teeth are more like those of _Procyon_
and _Nasua_. In the coatis, _Nasua_, the dentition is as in _Procyon_,
but the upper canines are larger and more strongly compressed, and the
molars smaller; while the facial portion of the skull is more
elongated and narrow. Vertebrae: C. 7, D. 14, L. 6, S. 3, Ca. 22-23.
Body elongated and rather compressed. Nose prolonged into a somewhat
upturned, obliquely-truncated, mobile snout. Tail long,
non-prehensile, tapering and ringed. Coatis, or coati-mundis, live in
small troops of eight to twenty, are chiefly arboreal, and feed on
fruits, young birds, eggs, insects, &c. The two best-known species are
_N. narica_ of Mexico and Central America, and _N. rufa_ of South
America from Surinam to Paraguay (see COATI).
In the kinkajou (q.v.), an animal long known as _Cercoleptes
caudivolvulus_, but whose designation it has been proposed to change
to the unclassical _Potos flavus_, the dentition is i. 3/3, c. 1/1, p.
3/3, m. 2/2 = 36. Molars with low flat crowns, very obscurely
tuberculated. Skull short and rounded, with flat upper surface.
Vertebrae: C. 7, D. 14, L. 6, S. 3, Ca. 26-28. 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.
Weasel tribe.
The last existing family of the land Carnivora is that typified by the
martens and weasels, and hence known as the _Mustelidae_. The group is
characterized by the absence of an alisphenoid canal in the skull, the
reduction of the molars to 1/2 or even 1/1, the medium size of the
sectorial tooth in each jaw, the absence or presence of a perforation
in the humerus, and the presence of anal glands. The family is
cosmopolitan in distribution, with the exception of Australasia and
Madagascar.
The first section of the family, forming the subfamily _Mustelinae_,
is typically characterized by the short and partially webbed toes,
furnished with short, compressed, sharp, curved and often partially
retractile claws. The upper molar is always of moderate size and
elongated in the transverse direction. In the martens and sables
(_Mustela_) the dentition is i. 3/3, c. 1/1, p. 4/3, m. 1/2; total 38;
the upper sectorial having its inner lobe close to the anterior edge
of the tooth; and the upper molar being nearly as large as the
sectorial. Lower sectorial with small inner tubercle. Vertebrae: C. 7,
D. 14, L. 6, S. 3, Ca. 18-23. Body long and slender. Limbs short,
partially digitigrade, with the feet rounded and the toes short, with
compressed, acute, semi-retractile claws. Tail moderate or long, more
or less bushy. One species, _M. martes_, the pine-marten, is British;
the remainder inhabit the northern regions of Europe, Asia and
America. Many of the species, as the sable (_M. zibellina_), yield fur
of great value (see MARTEN).
The dentition of _Putorius_ differs from that of _Mustela_ chiefly in
the absence of the anterior premolars of both jaws. The teeth are more
sharply cusped, and the lower sectorial wants the inner tubercle.
External characters generally similar to those of the martens, but the
body longer and more slender, and the limbs even shorter. All the
species are small animals, of active, bloodthirsty and courageous
disposition, living chiefly on birds and small mammals, and rather
terrestrial than arboreal, dwelling among rocks, stones and
out-buildings. Some of the species, as the stoat or ermine (_P.
ermineus_), inhabiting cold climates, undergo a seasonal change of
colour, being brown in summer and white in winter, though the change
does not affect the whole of the fur, the end of the tail remaining
black in all seasons. This is a large genus, having a very extensive
geographical range throughout the Old and New Worlds, and includes the
animals commonly known as weasels, polecats, ferrets and minks (q.v.).
In the glutton (_Gulo luscus_) the dentition is i. 3/3, c. 1/1, p.
4/4, m. 1/2; total 38; the crowns of the teeth being stout, and the
upper molar much smaller than the sectorial. Lower sectorial large,
with small heel and no inner tubercle. The dentition, though really
but a modification of that of the weasels, presents a general
resemblance to that of hyena. Vertebrae: C. 7, D. 15, L. 5, S. 3, Ca.
15. Body and limbs stoutly made; feet large and powerful,
subplantigrade, with large, compressed, much-curved and sharp-pointed
claws. Soles of the feet (except the pads of the toes) covered with
thick bristly hairs. Ears very small, nearly concealed by the fur.
Eyes small. Tail short, thick and bushy. Fur full, long and rather
coarse. The one species, the wolverine or glutton, is an inhabitant of
the forest regions of northern Europe, Asia and America, and much
resembles a small bear in appearance. It is a very powerful animal for
its size, climbs trees and lives on squirrels, hares, beavers,
reindeer, and is said to attack even horses and cows.
The South American grison and tayra represent the genus _Galictis_, in
which the dentition is i. 3/3, c. 1/1, p. 3/3, m. 1/2; total 34; the
molars being small but stout, and the upper sectorial with the inner
lobe near the middle of the inner border. Lower sectorial with heel
small, and inner tubercle small or absent. Body long; limbs short,
with non-retractile claws and naked soles. Head broad and depressed.
Tail of moderate length. The species include the grison (_G.
vittata_), _G. allamandi_, and the tayra (_G. barbara_); the last,
which extends northward into Central America, being sub-generically
separated as _Galera_. Nearly allied to these is the smaller and more
weasel-like _Lyncodon patagonicus_. All the foregoing South American
carnivores display a marked tendency to being darker on the lower than
on the upper surface. The same feature obtains in the African and
Indian ratels, or honey-badgers, constituting the genus _Mellivora_,
distinguished from all the other members of the family by having only
a single pair of lower molars, the dentition being i. 3/3, c. 1/1, p.
3/3, m. 1/1; total 32; the upper sectorial is large, with its inner
cusp at the anterior end of the blade, the molar much smaller and
transversely extended, having a small outer and a larger rounded inner
lobe. Heel of lower sectorial very small, scarcely one-fourth of the
whole length of the tooth, with but one cusp. Vertebrae: C. 7, D. 14,
L. 4, S. 4, Ca. 15. Body stout, depressed; limbs short, strong; head
depressed; nose rather pointed; ears rudimentary. Tail short. _M.
indica_, from India, and _M. ratel_, from south and west Africa, have
nearly the same general appearance and size, being rather larger than
a common badger, and may be only races of the same species. Their
coloration is peculiar, all the upper surface of the body, head and
tail being ash-grey, while the lower parts, separated by a distinct
longitudinal boundary line, are black. They live chiefly on the
ground, into which they burrow, but can also climb trees. They feed on
small mammals, birds, reptiles and insects, and are partial to honey.
In the Indo-Malay ferret-badger, _Helictis_, the dentition is i. 3/3,
c. 1/1, p. 4/4, m. 1/2; total 38. Upper sectorial with a large
bicusped inner lobe, molar smaller, wider transversely than in the
antero-posterior direction. Lower sectorial with heel about one-third
the length of the tooth. Skull elongated, rather narrow and depressed;
facial portion especially narrow; infraorbital foramen very large.
Head rather small and produced in front, with an elongated, obliquely
truncated, naked snout and small ears. Body elongated, limbs short.
Tail short or moderate, bushy. Several species are described, such as
_H. orientalis, moschata, nipalensis_, and _subaurantiaca_, from
eastern Asia, all small animals, climbing trees with agility and
living on fruits and berries as well as on small mammals and birds.
The African striped zorilles, or _Muis-honds_ (_Ictonyx_), have a
dental formula of i. 3/3, c. 1/1, p. 3/3, m. 1/2; total 34; the teeth
much resembling those of the polecats, and the upper molar being
smaller than the sectorial, and narrow from before backwards. Lower
sectorial with a smalt narrow heel and distinct inner tubercle.
General form of body musteline. Limbs short, fore-feet large and
broad, with five stout, nearly straight, blunt and non-retractile
claws, of which the first and fifth are considerably shorter than the
others. Tail moderate, with longer hairs towards the end, giving it a
bushy appearance. Hair generally long and loose. The best-known
species of this genus, the Cape polecat, _Ictonyx capensis_ (or
_Zorilla zorilla_), is about the size of a polecat, but conspicuous by
its broad, longitudinal bands of dark-brown, alternating with white.
Its odour is said to be as offensive as that of the American skunks.
From the Cape of Good Hope it ranges as far north as Senegal. Another
species, _I. lybicus_, from Sennaar, has been described. The small
striped polecat of southern Africa, _Poecilogale albinucha_,
represents a genus by itself, and is a shorter-haired animal.
The skunks of America are very similar to the two genera last
mentioned in their colouring, and with the latter serve to form a
connecting link with the more typical _Mustelinae_, and the badger
group, or _Melinae_, in which the feet are elongated, with straight
toes and non-retractile, slightly curved, subcompressed, blunt claws,
especially large on the fore-foot. In all cases the upper molar is
larger than the sectorial, and in the more typical genera is much
longer than broad.
In the North American skunks of the genus _Mephitis_ the dentition is
i. 3/3, c. 1/1, p. 3/3, m. 1/2; total 34. Upper molar larger than the
sectorial, subquadrate, rather broader than long; lower sectorial with
heel less than half the length of the whole tooth. Bony palate
terminating posteriorly opposite the hinder border of the last molar.
Facial portion of skull short and somewhat truncated in front.
Vertebrae: C. 7, D. 16, L. 6, S. 2, Ca. 21. Head small. Body
elongated. Limbs moderate, subplantigrade. Ears short and rounded.
Tail long, abundantly clothed with long fine hair. Anal glands largely
developed; their secretion, which can be discharged at the will of the
animal, has an intolerably offensive odour and has rendered skunks
proverbial. The South American species, which have only two upper
premolars, and differ in some other characters, are generically
separated under the name of _Conepatus_; while the small North
American arboreal skunks are distinguished as _Spilogale_ (see SKUNK).
Badger tribe.
Passing on to the more typical members of the badger group, we have
first the genus _Arctonyx_, with the dentition i. 3/3, c. 1/1, p. 4/4,
m. 1/2; total 38. The incisor line is curved, the outer teeth being
placed posteriorly to the others: lower incisors inclined forwards.
First premolars often rudimentary or absent; upper molar much larger
than the sectorial, longer in the antero-posterior direction than
broad; lower sectorial with a very large, low, tuberculated heel.
Skull elongated and depressed; face long, narrow and concave above;
bony palate extending as far backwards as the level of the glenoid
fossa; and palatal bones dilated. Suborbital foramina very large.
Vertebrae: C. 7, D. 16, L. 4, S. 4, Ca. 20. Snout long, naked, mobile
and truncated, with large terminal nostrils, much like those of a pig.
Eyes small; ears very small and rounded. Body compressed, rather than
depressed. Limbs of moderate length, and partially digitigrade in
walking. Tail moderate, tapering. A full soft under-fur, with longer
bristly hairs interspersed. The longest-known species is _A.
collaris_, the _bhalu-soor_ (bear-pig) or _bali-soor_ (sand-pig) of
the natives of the mountains of north-eastern India, Burma and Borneo.
It is rather larger than the badger, higher on its legs, and very
pig-like in general aspect, of a light grey colour, with
flesh-coloured snout and feet; nocturnal and omnivorous. Other species
or local varieties have been described from north China and Burma.
In the genus _Mydaus_ the dentition is as the last, but the cusps of
the teeth are more acutely pointed. Skull elongated, face narrow and
produced. Suborbital foramen small, and the palate, as in all the
succeeding genera of this group, produced backwards about midway
between the last molar and the glenoid fossa. Vertebrae: C. 7, D.
14-15, L. 6-5, S. 3, Ca. 12. Head pointed in front; snout produced,
mobile, obliquely truncated, the nostrils being inferior. Limbs rather
short and stout. Tail extremely short, but clothed with rather long
bushy hair. Anal glands largely developed, and emitting an odour like
that of the skunks. One species, _M. meliceps_, the teledu, a small
burrowing animal from the mountains of Java, at an elevation of 7000
or more ft. above the sea-level; and a second (_M. marchei_) from the
Philippines.
In the true badger of the genus _Meles_ the dentition is i. 3/3, c.
1/1, p. 4/4, m. 1/2; total 38. The first premolar in both jaws is
extremely minute and often deciduous; while the upper molar is much
larger than the sectorial, subquadrate, and as broad as long. Lower
sectorial with a broad, low, tuberculated heel, more than half the
length of the whole tooth. The postglenoid process of the skull so
strongly developed, and the glenoid fossa so deep, that the condyle of
the lower jaw is firmly held in place after the soft parts are
removed. Vertebrae: C. 7, D. 15, L. 5, S, 3, Ca. 18. Muzzle pointed.
Ears very short. Body stout, broad. Limbs short, strong,
subplantigrade. Tail short. Typified by the common badger (_M. taxus_
or _M. meles_) of Europe and northern Asia, still found in many parts
of England, where it lives in woods, is nocturnal, burrowing and very
omnivorous, feeding on mice, reptiles, insects, fruit, acorns and
roots. Other nearly allied species, _M. leucurus_ and _M. chinensis_,
are found in continental Asia, and _M. anakuma_ in Japan.
In the nearly-allied genus _Taxidea_ the dental formula is as in
_Meles_, except that the rudimentary anterior premolars appear to be
always wanting in the upper jaw. The upper sectorial is much larger in
proportion to the other teeth; and the upper molar about the same size
as the sectorial, triangular, with the apex turned backwards. Heel of
lower sectorial less than half the length of the tooth. Skull very
wide in the occipital region; the lambdoidal crest greatly developed,
and the sagittal but slightly, contrary to what obtains in _Meles._
Vertebrae: C. 7. D. 15. L. 5, S. 3, Ca. (?). Body stoutly built and
depressed. Tail short. The animals of this genus are peculiar to North
America, where they represent the badgers of the Old World, resembling
them much in appearance and habits. _T. americana_ is the common
American badger of the United States, _T. berlandieri_, the Mexican
badger, being a local variety.
Otter tribe.
The third and last subfamily is that of the otters, or _Lutrinae_, in
which the feet (with the exception of the hind pair in the sea-otter)
are short and rounded, with the toes webbed, and the claws small,
curved and blunt. The head is broad and much depressed. The upper
posterior cheek-teeth are large and quadrate. The kidneys are
conglomerate. Habits aquatic.
In the true otter of the genus _Lutra_ the dentition is i. 3/3, c.
1/1, p. 4/3, m. 1/2; total 36. Upper sectorial with a trenchant
tricusped blade, and a very large inner lobe, hollowed on the free
surface, with a raised sharp edge, extending along two-thirds or more
of the length of the blade. Upper molar large, with a quadricuspidate
crown, broader than long. Skull broad and depressed, contracted
immediately behind the orbits; with the facial portion very short and
the brain-case large. Vertebrae: C. 7, D. 14-15, L. 6-5, S. 3, Ca.
20-26. Body very long. Ears short and rounded. Limbs short. Feet
completely webbed, with well-developed claws on all the toes. Tail
long, thick at the base and tapering, rather depressed. Fur short and
close.
Otters are more or less aquatic, living on the margins of rivers,
lakes, and in some cases the sea; are expert divers and swimmers, and
feed chiefly on fish. They have an extensive geographical range, and
so much resemble each other in outward appearance, especially in the
nearly uniform brown colouring, that in some cases the species are by
no means well-defined. The Brazilian otter (_L. brasiliensis_) is a
very large species from Brazil, Demerara and Surinam, with a prominent
ridge along each lateral margin of the tail. In two small species the
feet are only slightly webbed; claws exceedingly small or altogether
wanting on some of the toes; the first upper premolar very small,
sometimes wanting; and the molars very broad and massive. The species
in question are _L. inunguis_ of South Africa, and _L. leptonyx_ or
_cinerea_ of India, Java and Sumatra, and have been separated as a
distinct genus, Aonyx.
The sea-otter, _Latax_ (or _Enhydra_) _lutra_, with a dentition of i.
3/2, c. 1/1, p. 3/3, m. 1/2, total 32, differs from other Carnivora in
having but two incisors on each side of the lower jaw, the one
corresponding to the first (very small in the true otters) being
absent. Though the molar teeth generally resemble those of _Lutra_ in
their proportions, they differ in the exceeding roundness and
massiveness of their crowns and bluntness of their cusps. Feet webbed;
fore-feet short, with five subequal toes, with short compressed claws;
hind-feet very large, depressed and fin-like, their phalanges
flattened as in seals. The fifth toe the longest and stoutest, the
rest gradually diminishing in size to the first, all with moderate
claws. Tail moderate, cylindrical (see OTTER).