13, L 6, S 4, C 7. This genus includes several species widely distributed
throughout South America, extending even to the Straits of Magellan. The
Restless Cavy (_C. porcellus_), which is found throughout Uruguay and
Brazil, has been very generally regarded as the ancestral form of the
domesticated Guinea-Pig. It is about 10 inches long, and weighs a little
over a pound; its fur is long and of a nearly uniform grayish-brown
colour. This species is rarely found in dry sandy localities, preferring
marshes covered with aquatic plants, among which it lies concealed,
feeding in the early morning and after sunset in the evening; but when
the soil is dry it forms burrows. It lives in societies of from six to
eighteen individuals, breeding but once a year, with one, or at most
only two, young at a birth. The Guinea-Pig (probably a misnomer of
Guiana-Pig) is larger than _C. porcellus_, and is regarded by Dr. Nehring
as descended from another species, _C. cutleri_. It is white in colour,
with irregular patches of reddish-brown and black. The Bolivian Cavy (_C.
boliviensis_), found throughout the higher regions of Bolivia, usually at
an elevation of 10,000 or 12,000 feet, is exceedingly shy, and lives in
burrows, which in some districts are so numerous as to have completely
undermined the soil. The Rock-Cavy (_C. rupestris_), distinguished by its
short, blunt nails, is found in rocky situations throughout Brazil, and
is much sought after for its flesh. The Southern Cavy (_C. australis_),
common along the coast of Patagonia, forms deep burrows, with several
outlets, in sandy declivities. Remains of existing species of _Cavia_ are
found in the cavern-deposits of Lagoa Santa, Brazil.
_Dolichotis._[416]—Characterised by the great length of the ears and the
short tail. The palate is so much contracted in front that the premolars
of opposite sides touch by their antero-internal edges. Vertebræ: C 7, D
12, L 8, S 3, C 10.
The Patagonian Cavy (_D. patachonica_)—the only living representative of
the genus—is rather larger than a Hare, which it somewhat resembles in
external appearance. It inhabits the dry sterile districts of Patagonia
and La Plata, disappearing wherever the country becomes more humid. This
animal burrows in the earth, although in districts where the Viscacha is
found it is said to avail itself of the works of the latter. Unlike other
cavies, its eyes are protected from the glare of the sun by prominent
eyelashes. The body is covered with a long dense fur of a rusty colour.
Two young are produced at a birth. Three species of _Dolichotis_ have
been described from the Brazilian cave-deposits, one of which is probably
not really separable from the existing form.
_Hydrochœrus._[417]—A large aquatic form with all the feet fully webbed;
the skull (Fig. 213, p. 481) large, with enormous paroccipital processes;
and the molars very complex, the third upper one having some twelve
transverse laminæ. Upper incisors grooved. Vertebræ: C 7, D 14, L 6, S 3,
C 8.
The Capybara (_H. capybara_) is the largest existing Rodent, and the
only living representative of the genus. It is a bulky and stoutly built
animal, and attains a length of about 4 feet. The body is covered with
long and coarse hair, reddish-brown above and brownish-yellow beneath.
Capybaras are found over the whole of the eastern part of South America,
and to the westward range into Bolivia and Peru. They frequent the
borders of rivers and lakes, concealing themselves among reeds and other
water plants. Remains of _Hydrochœrus_ are found in the cavern-deposits
of Brazil, which are probably referable to the existing species; one
extinct species from the Pleistocene of Buenos Ayres is estimated to
have attained a length of 5 feet, while _H. magnus_ of the same deposits
was of still larger dimensions. The genus is also represented in the
Pleistocene of South Carolina and the infra-Pampean beds of Parana.
_Extinct Genera._—A number of South American fossil Rodents have been
referred to extinct genera of _Caviidæ_. Thus _Plexochœrus_, from the
Tertiary of Argentina, differs from _Hydrochœrus_ in having only nine
laminæ in the last upper molar; _Cardiomys_, _Cardiatherium_, etc., from
the infra-Pampeans are also stated to be allied to _Hydrochœrus_, while
_Contracavia_, of the same deposits, is related to _Cavia_, but of larger
size. _Microcavia_, again, from the Pleistocene of Argentina, is regarded
as connecting _Cavia_ with _Dolichotis_. The Tertiary European genera
_Issiodoromys_ and _Nesocerodon_ are apparently referable to the present
family.
_Suborder_ DUPLICIDENTATA.
Two pairs of incisors in the upper jaw (the second very small, and placed
directly behind the large first pair), the enamel of which extends
round to their posterior surfaces. At birth there are three pairs of
these incisors, but the outer one on each side is soon lost. Incisive
foramina large; and usually confluent; bony palate very narrow from
before backwards; no true alisphenoid canal; fibula ankylosed to the
tibia, and articulating with the calcaneum. Testes permanently external.
This suborder includes the Picas, Hares, and Rabbits, all of which are
strictly terrestrial.
_Family_ LAGOMYIDÆ.
Complete clavicles, subequal limbs, no external tail, and short ears.
Skull depressed, frontals contracted and without postorbital processes;
_p_ ¹⁄₁ or ²⁄₂; molars rootless, with transverse enamel-folds. Palæarctic
and Nearctic.
_Lagomys._[418]—Represented by about a dozen species of small
Guinea-Pig-like animals, inhabiting chiefly the mountainous parts of
Northern Asia (from 11,000 to 14,000 feet), one species only being known
from South-East Europe, and another from the Rocky Mountains.
The Picas, or Tailless Hares, live in holes among the rocks of their
native mountains, and are agile and shy little creatures. The genus is
well represented through the upper and middle Tertiaries. It has been
proposed to separate those fossil forms with _p_ ²⁄₁ as _Myolagus_, and
those with _p_ ¹⁄₁ as _Titanomys_, but this seems scarcely advisable.
_Family_ LEPORIDÆ.
Imperfect clavicles, elongated hind limbs, short recurved tail, and
long ears. Skull (Fig. 216) compressed, frontals with large wing-shaped
postorbital processes _p_ ³⁄₂; molars as in the _Lagomyidæ_. Cosmopolitan
(except Australasia). Vertebræ: C 7, D 12, L 7, S 4, C 13-15.
[Illustration: FIG. 216.—Skull of Hare (_Lepus timidus_).]
_Lepus._[419]—The single genus _Lepus_ includes about twenty species,
all of which resemble one another in general external characters. In all
the fore limbs have five and the hind only four digits, and the soles of
the feet are densely clothed with hairs similar to those covering the
legs; the inner surface of the cheeks is also hairy. Although the family
has such a wide distribution, the greater number of the species are
restricted to the Palæarctic and Nearctic regions, only a single species
(_L. brasiliensis_) extending into South America, where it has existed
since the date of the Pleistocene deposits of the Brazilian caves.
The Common Hare (_L. timidus_[420]) may be taken as a typical example of
the genus, and is characterised by the great length of the ears and hind
limbs. It is found in all parts of Europe except the north of Russia,
the Scandinavian peninsula, and Ireland. Its fur is usually of a tawny
gray colour above and white beneath, with the upper surface of the short
tail and the tips of the ears black. The colour of the fur differs,
however, considerably in different latitudes and at different seasons of
the year; showing a tendency to become white during winter in northern
countries, while assuming a reddish-yellow hue in the more genial climate
of southern Europe. The Hare is a nocturnal animal, remaining during the
day on its “form,” as the slight depression is called which it makes in
the open field, usually among grass.
[Illustration: FIG. 217.—The Common Hare (_Lepus timidus_).]
The Mountain Hare (_L. variabilis_) is found throughout the northern part
of the Palæarctic region, ranging from Ireland in the west to Japan in
the east, and also occurring in several of the more southerly mountain
ranges, such as the Pyrenees, the Alps, and the Caucasus. It is smaller
than the common species, with a smaller and more rounded head, and
shorter ears, tail, and hind limbs. In cold climates the colour of the
whole animal changes in the winter to a pure white (as in Fig. 218), with
the exception of the tips of the ears, which remain black. In Ireland no
winter change of colour takes place.
[Illustration: FIG. 218.—The Mountain Hare (_Lepus variabilis_).]
The Rabbit (_L. cuniculus_), speaking of the wild race only, is
distinguished from the Hare externally by its smaller size, shorter ears
and feet, the absence or reduction of the black patch at the tip of
the ears so characteristic of the Hare, and by its grayer colour. The
skull is smaller and lighter, with a slenderer muzzle and a longer and
narrower palate. Besides these characters, however, the Rabbit is sharply
separated from the Hare by the fact that it brings forth its young naked,
blind, and helpless; to compensate for this, it digs a deep burrow in
the earth in which they are born and reared, while the young of the Hare
are born fully clothed with fur, and able to take care of themselves in
the “form” in which they are born. The weight of the Rabbit is from 2½
to 3 lbs., although individuals perfectly wild have been recorded up to
more than 5 lbs. Its general habits are too well known to need a detailed
description here. It breeds from four to eight times a year, bringing
forth each time from three to eight young. Its period of gestation is
about thirty days, and it begins to breed when six months old. It attains
to an age of about seven or eight years.
[Illustration: FIG. 219.—The Rabbit (_Lepus cuniculus_).]
The geographical distribution of the Rabbit presents many most
interesting peculiarities. It is believed to be originally a native of
the western half of the Mediterranean basin only, and still abounds in
Spain, Sardinia, Southern Italy, Sicily, Greece, Tunis, and Algeria; and
many of the Islands adjoining these countries are quite overrun with
it. Thence it has spread, partly by man’s agency, northwards throughout
temperate Western Europe, increasing rapidly wherever it gains a footing;
and this extension is still going on, as is shown by the case of
Scotland, in which sixty years ago Rabbits were little known, while they
are now found in all suitable localities up to the extreme north. It has
also gained admittance into Ireland, and now abounds there as much as in
England. Out of Europe the same extension of range has been going on. In
New Zealand and Australia Rabbits, introduced either for profit or sport,
have increased to such an extent as to form one of the most serious
pests that the farmers have to contend against, as the climate and soil
seem to suit them perfectly, and their natural enemies are too few and
too lowly organised to keep their numbers within reasonable bounds. In
other cases Rabbits introduced into islands have become or remained
more or less distinct from their parent stock; thus the Rabbits both of
the Falkland Islands and of Jamaica still show traces of their descent
from domesticated varieties, and have never reverted to the ordinary
brownish-gray type. And again, as was pointed out by Mr. Darwin,[421]
the Rabbits in the island of Porto Santo, near Madeira, whose ancestors
were introduced from Spain in 1418 or 1419, have formed quite a distinct
diminutive race, barely half the bulk or weight of English Rabbits, and
differing in certain slight details of colour and habits.
_Bibliography of Rodentia._—G. R. Waterhouse, “Observations
of the Rodentia,” _Mag. Nat. Hist._ iii. (1839); Id. _Ann.
Nat. Hist._ viii. and x. (1839-42); Id. “On the Geographical
Distribution of the Rodentia,” _Proc. Zool. Soc._ 1839, pp.
162-174; Id. _Natural History of the Mammalia_, vol. ii.
“Rodentia” (1848); Gervais, _Dic. Univ. d’Hist. Nat._ xi. p.
202 (1848); Brandt, “Untersuchungen über die craniologischen
Entwickelungsstufen und Classification der Nager der Jetzwelt,”
_Mém. de l’Acad. Impér. de St. Pétersbourg_ (1855); Lilljeborg,
_Systematisk Œfversight af de Gnagnde Däggdjuren_, Upsala,
1866; Alston, “On the Classification of the Order _Glires_,”
_Proc. Zool. Soc._ 1876, pp. 61-98; Trouessart, “Catal. de
Rongeurs, Vivants et Fossiles,” _Bullet. Soc. d’Études Scient.
d’Angers_, 1880-1881; Coues and Allen, “Monographs of North
American Rodentia,” _United States Geol. Surv. of Territories_,
vol. xi. (1877); Winge, “Rodentia pa Lagos Santa, Brazil.”
_Mus. Lund._ vol. iii. (1887); various papers by Peters in
_Monatsber. Ak. Berlin_, and by Alston, Anderson, Blanford,
Dobson, Milne-Edwards, Thomas, and others, in _Proc. Zool.
Soc._, _Journ. Asiat. Soc. Beng._, _Ann. Mag. Nat. Hist._, etc.
CHAPTER XI
THE ORDER CARNIVORA
Though the existing Carnivora as at present restricted[422] form a very
natural and well-defined order among the Mammalia, it is difficult to
find any important common diagnostic characters by which they can be
absolutely separated; so that, as in the case of so many other natural
groups, it is by the possession of a combination of various characters
that they must be distinguished. Thus they are all unguiculate, and never
have less than four well-developed toes on each foot, with nails more
or less pointed, rarely rudimentary or absent. The pollex and hallux
are never opposable to the other digits. They are regularly diphyodont
and heterodont, and their teeth are always rooted.[423] Their dentition
consists of small pointed incisors, usually three in number, on either
side of each jaw, of which the first is always the smallest and the third
the largest, the difference being most marked in the upper jaw; strong
conical, pointed, recurved canines; cheek-teeth variable, but generally,
especially in the anterior part of the series, more or less compressed,
pointed, and trenchant; if the crowns are flat and tuberculated they
are never complex or divided into lobes by deep inflexions of enamel.
The condyle of the lower jaw is a transversely placed half-cylinder
working in a deep glenoid fossa of corresponding form. The brain varies
much in relative size and form, but the hemispheres are never destitute
of well-marked convolutions (Fig. 23, p. 71). The stomach (Fig. 234)
is always simple and pyriform. The cæcum is either absent or short
and simple (Fig. 235), and the colon is not sacculated, or greatly
wider than the small intestine. Vesiculæ seminales are never present.
Cowper’s glands are present in some, absent in other groups. The uterus
is bicornuate. The mammæ are abdominal, and very variable in number.
The placenta is deciduate, and almost always zonary. The clavicle is
often entirely absent, and when present is never complete. The humerus
often has an entepicondylar foramen. The radius and ulna are distinct.
The scaphoid and lunar bones are united into one, and there is never
a distinct os centrale in the adult. The fibula is always a distinct
slender bone.
Several of these characters are, however, not applicable to all the
members of the extinct group of Carnivores for which the name Creodonta
has been proposed, as will be noticed in the sequel.
The large majority of the species composing this order subsist chiefly
upon some variety of animal food, though many are omnivorous, and some
few chiefly, though not entirely, vegetable eaters. The more typical
forms live altogether on recently killed warm-blooded animals, and their
whole organisation is thoroughly adapted to a predaceous mode of life.
In conformity with this manner of obtaining their subsistence they are
generally bold and savage in disposition, though some species are capable
of being domesticated, and when placed under favourable circumstances
for the development of such qualities exhibit a very high degree of
intelligence and fidelity. The existing representatives of the order
are naturally divided into two suborders, the members of the one being
the more typical, and mainly terrestrial in their mode of life; while
those of the other are aberrant, having the whole of their organisation
specially modified for living habitually in water. These are called
respectively the True, or Fissiped, and the Pinniped Carnivora.
_Suborder_ CARNIVORA VERA.
[Illustration: FIG. 220.—Left upper carnassial teeth of Carnivora. I,
_Felis_; II, _Canis_; III, _Ursus_. 1, Anterior, 2, middle (paracone),
and 3, posterior (metacone) cusp of blade; 4, inner tubercle (protocone)
supported on distinct root; 5, inner cusp posterior in position, and
without distinct root, characteristic of the _Ursidæ_.]
Generally adapted for terrestrial progression and mode of life, though
some may be partially aquatic in their habits. The fore limbs never have
the first digit, or the hind limbs the first and fifth digits, longer
than the others. Incisors ³⁄₃ on each side, with very rare exceptions.
Cerebral hemispheres more or less elongated; always with three or four
gyri on the outer surface forming arches above each other, the lowest
surrounding the Sylvian fissure. The molar series of teeth have not
the uniform characters of those of the Pinnipedia. There is always one
tooth in each jaw which is specially modified, and to which the name of
“sectorial” or “carnassial” tooth has been applied. The teeth in front of
this are more or less sharp pointed and compressed; while those behind
it are broad and tuberculated. The characters of the carnassial teeth
deserve special attention, as, though fundamentally the same throughout
the suborder, they are greatly modified in different genera. The upper
carnassial 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 essentially of a more or less compressed blade
supported on two roots and an inner tubercle supported by a distinct
root (see Fig. 220). The blade when fully developed has three cusps
or lobes (1, 2, and 3), but the anterior is always small, and often
absent. The middle lobe is conical, high, and pointed; the posterior
lobe has a compressed straight knife-like edge. The inner tubercle (4)
varies very much in extent, but is generally placed near the anterior
end of the blade, though sometimes it is median in position. In the
_Ursidæ_ alone both the inner tubercle and root are wanting, and there
is often a small internal and posterior cusp (5) without root. In this
aberrant family also the carnassial is relatively to the other teeth much
smaller than in the rest of the Carnivora. The lower carnassial (see
Fig. 221) is the most anterior of the teeth without predecessors in the
milk-series; it is therefore reckoned the first true molar (_m_ ¹⁄). It
has two roots supporting a crown, consisting when fully developed of a
compressed bilobed blade (1 and 2), a heel, or talon (4), and an inner
cusp (3). The lobes of the blade, of which the hinder (2) is the larger,
are separated by a notch, generally prolonged into a linear fissure. In
the most specialised Carnivora, as the _Felidæ_ (I), the blade alone
is developed, both talon and inner cusp being absent or rudimentary. In
others, as _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 are almost side by side, instead of anterior and posterior. The
inner cusp (3) is generally conical, pointed, and 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 (_Latax_) than in any other
form, but even in it they can be traced.
[Illustration: FIG. 221.—Left lower carnassial teeth of Carnivora. I,
_Felis_; II, _Canis_; III, _Herpestes_; IV, _Lutra_; V, _Meles_; VI,
_Ursus_. 1, Anterior lobe (paraconid) of blade; 2, posterior (protoconid)
lobe of blade; 3, inner cusp (metaconid); 4, talon (hypoconid). It will
be seen that the relative size of the two roots varies according to
the development of the portion of the crown they have respectively to
support.]
The homology of the various parts of the Carnivorous carnassial with the
primitive tritubercular type (p. 30) is indicated in the figures. It may
be observed, however, that the anterior lobe of the three-lobed upper
carnassial is an element added on to the more primitive two-lobed type.
When the talon of the lower carnassial, as in _Canis_, consists of a
large outer and small inner cusp, the latter (not seen in the figure) is
the entoconid.
The toes are nearly always armed with large, strong, curved, and
tolerably sharp claws, ensheathing the ungual phalanges, and held more
firmly in their places by broad laminæ of bone reflected over their
attached ends from the bases of the phalanges. In some forms, most
notably the _Felidæ_, these claws are retractile; that is to say, the
ungual phalanx, with the claw attached, folds back in the fore foot into
a sheath by the outer or ulnar side of the middle phalanx of the digit,
being retained in this position when the animal is at rest by a strong
elastic ligament. In the hind foot the ungual phalanx is retracted on
to the top, and not the side of the middle phalanx. By the action of
the deep flexor muscles, the ungual phalanges are straightened out, the
claws protruded from their sheath, and the soft “velvety” paw becomes
suddenly converted into a most formidable weapon of offence. The habitual
retraction of the claws preserves their points from wear in ordinary
progression.
The skeleton of the Lion represented in Fig. 15 (p. 45) illustrates the
digitigrade mode of progression of the _Felidæ_, as well as the essential
characters of the bony framework of a typical Carnivore.
The Fissipedal Carnivora were divided by Cuvier into two groups,
according to the position of the feet in walking,—the Plantigrada,
or those that place the whole of the soles to the ground, and the
Digitigrada, or those that walk only on the toes; and the difference
between these groups was considered of equal importance to that which
separated the Pinnigrada or Seals from both of them. The distinction is,
however, quite an artificial one, since every intermediate condition
exists between the extreme typical plantigrade gait of the Bears and the
truly digitigrade walk of the Cats and Dogs; in fact, the greater number
of the Carnivora belong to neither one form nor the other, but may be
called “subplantigrade”; often when at rest applying the whole of the
sole to the ground, but keeping the heel raised to a greater or less
extent when walking.
An amended classification of the existing forms is into three distinct
sections, of which the Cats, the Dogs, and the Bears may be respectively
taken as representatives, and which are hence called Æluroidea, Cynoidea,
and Arctoidea. This division is founded mainly on characters exhibited
by the base of the skull, but is corroborated by the structure of other
parts.[424] The presence or absence of a bridge of bone, covering the
external carotid artery in a part of its course by the side of the
alisphenoid bone, and enclosing the “alisphenoid canal” (see Fig. 8, p.
38), a character to which the late Mr. H. N. Turner first drew attention,
might seem unimportant at first sight, but it is curiously constant
in certain groups, which we have other reasons, derived often from a
combination of less easily definable characters, to regard as natural. It
is therefore generally mentioned in the following family definitions.
It must, however, be stated that while the arrangement is a convenient
one as regards the existing Carnivores, it will not hold good when the
fossil forms are included. Thus there is ample evidence to show that
the Dogs and Bears were formerly so intimately connected that in a
palæontological classification the _Canidæ_ cannot be satisfactorily
separated from the _Ursidæ_; while in another direction the _Canidæ_ were
closely allied to the ancestral _Viverridæ_. The most important objection
against this classification is, however, the apparent intimate connection
exhibited by fossil forms between the _Viverridæ_ and the _Mustelidæ_,
which, so far as the present evidence goes, tends to show that the latter
are derived from the former. If this be eventually fully proved, it would
seem to indicate that the Arctoidea are not a natural group; and that the
resemblances between the _Ursidæ_ and _Mustelidæ_ have been independently
acquired, in the course of the descent of the one family from a Canoid,
and of the other from a Viverroid stock.
_Section_ ÆLUROIDEA.
[Illustration: FIG. 222.—Left side of the palatal aspect of the cranium
and mandible of the Suricate (_Suricata tetradactyla_). _c_, Carotid
foramen; _f_, fissure in floor of auditory meatus. From Mivart, _Proc.
Zool. Soc._ 1882, p. 184.]
The Æluroidea or Cat-like Carnivores include the _Felidæ_, _Viverridæ_,
_Proteleidæ_, and _Hyænidæ_. The existing representatives of this section
present the following common features. Auditory bulla (Fig. 222) much
dilated, rounded smooth, thin-walled, and (except in the _Hyænidæ_)
divided into two chambers, by a septum. Bony auditory meatus short.
Paroccipital process applied to, and spread over the hinder part of the
bulla (Fig. 222). Mastoid process never very salient, and often obsolete.
Carotid canal (Fig. 8, p. 38, _car_) small, sometimes very inconspicuous.
Condyloid and glenoid foramina concealed or wanting. Cæcum small, rarely
absent. Os penis generally small and irregular (large in _Cryptoprocta_).
Cowper’s glands present; prostate distinctly lobed. Some details of the
anatomy of the soft parts will be found under the head of _Genetta_.
_Family_ FELIDÆ.
In all the forms, both recent and fossil, which can be included in this
family the canines are strongly developed, there are never more than
one upper and two lower molars, and the three lower incisors are placed
in the same horizontal line. With one exception, the humerus has an
entepicondylar foramen.
The following characters are common to all the existing members. True
molars reduced to one above and below, that of the upper jaw very small
and transversely extended. Only two inferior premolars. Upper carnassial
with three lobes to the blade; lower without talon or inner cusp.
Auditory bulla not externally constricted. No alisphenoid canal. Carotid
canal very minute. Digits 5-4. Dorsal vertebræ 13.
[Illustration: FIG. 223.—Front view of skull of Lion (_Felis leo_).]
_Felis._[425]—The whole structure of the animals of this genus exhibits
the Carnivorous type in its fullest perfection. Dentition: _i_ ³⁄₃, _c_
¹⁄₁, _p_ ³⁄₂, _m_ ¹⁄₁; total 30. A distinctly cusped inner tubercle to
the upper carnassial. Claws completely retractile. The upper anterior
premolar (_p._ 2), always small, and may be absent without any other
modification in the dental or other structures. Such a variation should
not therefore be considered as of generic importance. Incisors very
small. Canines large, strong, slightly recurved, with trenchant edges and
sharp points, and placed wide apart (Fig. 223). Premolars compressed and
sharp pointed. The most posterior in the upper jaw (the carnassial), a
very large tooth, consisting of a subcompressed blade, divided into three
unequal lobes supported by two roots, with a very small inner tubercle
placed near the front end of the tooth and supported by a distinct root
(Fig. 220). The upper true molar a very small tubercular tooth placed
more or less transversely at the inner side of the hinder end of the
last. In the lower jaw the true molar (carnassial) reduced to the blade
alone, which is very large, trenchant, and much compressed, divided
into two subequal lobes. Occasionally it has a rudimentary talon, but
never an inner cusp. The skull is generally short and rounded, though
proportionally more elongated in the larger forms. The facial portion
is especially short and broad, and the zygomatic arches are very
wide and strong. The auditory bullæ are large, rounded, and smooth.
Vertebræ: C 7, D 13, L 7, S 3, C 13-29. Clavicles better developed
than in other Carnivora, but not articulating with either the scapulæ
or sternum. Limbs digitigrade. Anterior feet with five toes, the third
and fourth nearly equal and longest, the second slightly and the fifth
considerably shorter; the pollex still shorter, not reaching as far as
the metacarpo-phalangeal articulation of the second. Hind feet with
only four toes. The third and fourth the longest, the second and fifth
somewhat shorter and nearly equal; the hallux represented only by the
rudimentary metatarsal bone. The claws all very large, strongly curved,
compressed, very sharp, and exhibiting the retractile condition in the
highest degree. The tail varies greatly in length, being in some a mere
stump, in others nearly as long as the body. Ears of moderate size, more
or less triangular and pointed. Eyes rather large. Iris very 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. Tongue thickly covered with sharp-pointed,
recurved horny papillæ. Cæcum small and simple.
As in structure so in habits, the Cats may be considered the most
specialised of all the Carnivora. All the known members of the genus
feed, in the natural state, almost exclusively on warm-blooded animals
which they have themselves killed. One Indian species (_F. viverrina_)
preys on fish and even (it is said) on freshwater molluscs. Unlike the
Dogs, they never associate in packs, and rarely hunt their prey in 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 of the genus are very widely diffused over the
greater part of the habitable world, though most abundant in the warm
latitudes of both hemispheres. No species 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 minor
groups into which the genus has been divided have representatives in both
hemispheres.
Notwithstanding the considerable diversity in external appearance and
size between different members of this extensive genus, the structural
differences are but slight, and so variously combined in different
species that the numerous attempts hitherto made to subdivide it are all
unsatisfactory and artificial. 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 tubercle of the upper carnassial,
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. Writing in 1881 Professor
Mivart[426] gave the number of existing species of _Felis_ as 48, but
by Mr. Blanford’s reduction of the number of Indian species[427] the
list may now be diminished to some 41. The following account is chiefly
devoted to some of the more important and better known species.
A. _Old World Species._—The Lion (_F. leo_, Fig. 224) has been well known
to man from the earliest historic times. Its geographical habitat made it
familiar to all the races among whom human civilisation took its origin,
and its strongly marked physical and moral characteristics have rendered
it proverbial, perhaps to an exaggerated degree, and have in all ages
afforded favourite types for poetry, art, and heraldry. The literature
of the ancient Hebrews abounds in allusions to the Lion; and the almost
incredible numbers that are stated to have been provided for exhibition
and destruction in the Roman amphitheatres (as many as six hundred on a
single occasion by Pompey, for example) show how abundant these animals
must have been within accessible distance of the capital of the world.
The geographical range of the Lion was once far more extensive than at
present, even within the historic period covering the whole of Africa,
the south of Asia, including Syria, Arabia, Asia Minor, Persia, and the
greater part of Northern and Central India, and also the south-eastern
portion of Europe, as shown by the well-known story told by Herodotus of
the attacks by Lions on the Camels which carried the baggage of the army
of Xerxes on its march through the country of the Pæonians in Macedonia.
The very circumstantial account of that historian shows that the animal
in his time ranged through the country south of the Balkans, through
Roumania to the west of the River Carasu, and through Thessaly as far
south as the Gulf of Lepanto and the Isthmus of Corinth, having as its
western boundary the River Potamo and the Pindus mountains. The whole
of the evidence relating to the existence of Lions in Europe, and to
their retreat from that continent shortly before the commencement of the
Christian era, has been collected in the article on “_Felis spelæa_” in
Boyd Dawkins and Sandford’s _British Pleistocene Mammalia_ (1868). Fossil
remains attest a still wider range, as it is shown in the same work that
there is absolutely no osteological or dental character by which the
well-known Cave Lion (_F. spelæa_), so abundantly found in cave-deposits
of the Pleistocene age in Western Europe, can be distinguished from the
existing _F. leo_.
[Illustration: FIG. 224.—Lion and Lioness, after a drawing by Wolf in
Elliot’s Monograph of the _Felidæ_.]
At the present day the Lion is found in localities suitable to its
habits, and where not exterminated (as it probably was in Europe) by the
encroachments of man, throughout Africa from Algeria to the Cape Colony,
and in Mesopotamia, Persia, and some parts of the north-west of India.
According to Blanford,[428] Lions are still very numerous in the reedy
swamps bordering the Tigris and Euphrates, and also occur on the west
flanks of the Zagros mountains and the oak-clad ranges near Shiraz, to
which they are attracted by the immense herds of swine which feed on the
acorns. The Lion nowhere exists in the table-land of Persia, nor is it
found in Baluchistan. In India, where it is verging on extinction, it
appears now to be confined to parts of Kattywar and Rajputana, though
within the present century its range extended through the north-west part
of India, from Bahawalpur and Sind to at least the Jumna (about Delhi),
southward as far as Khandesh, and in Central India through the Saugor
and Narbada territories, Bundelkund, and as far east as Palamau. It was
extirpated in Harriana about 1824. One was killed at Rhyli, in the Dumaoh
district, Saugor and Narbada territories, so late as in the cold season
of 1847-48; and one was shot in 1810 near Kot-Deji, Sind.[429]
The great variations in external characters which different Lions
present, especially in the colour and the amount of mane, has given rise
to the idea that there are several species, or at all events distinct
varieties peculiar to different localities. It was at one time supposed,
on the authority of Captain Walter Smee,[430] that the Lion of Gujerat
differed essentially from that of Africa in the absence of a mane, but
subsequent evidence has not supported this view, which was probably
founded upon young specimens having been mistaken for adults. Lions from
that district as well as from Babylonia, which have lived in the gardens
of the London Zoological Society, have had as fully developed manes as
any other of the species. Mr. F. C. Selous[431] has shown that in South
Africa the so-called Black-maned Lion and others with yellow scanty manes
are found, not only in the same locality, but even among individuals of
the same parentage.
The Lion belongs to a well-defined group, containing the largest members
of the genus, and differing from the others in the well-marked character
that the anterior cornu of the hyoid arch is but little ossified, so that
this arch is connected with the cranium by a long ligament, instead of by
a continuous chain of bones, and by the less important one that the pupil
of the eye, when contracted, is a circular hole, instead of a vertical
slit as in the cat. The Lion agrees with the Tiger and the Leopard in
these respects, but differs from them in its uniform style of colouring,
and from all the other _Felidæ_ in the arrangement of its hairy covering;
thus the hair of the top of the head, chin, and neck, as far back as the
shoulder, is not only very much longer, but also differently disposed
from the hair elsewhere, being erect or directed forwards, and so
constituting the characteristic ornament called the mane. There is also a
tuft of elongated hairs at the end of the tail, one upon each elbow, and
in most lions a copious fringe along the middle line of the under surface
of the body, wanting, however, in some examples.[432] It must, however,
be observed that these characters are peculiar to the adults of the male
sex only, and that young lions show indications of the darker stripes and
mottlings so characteristic of the greater number of the members of the
genus.
The usual colour of the adult is yellowish-brown, but it may vary from a
deep red or chestnut brown to an almost silver gray. The mane, as well as
the long hair of the other parts of the body, sometimes scarcely differs
from the general colour, but it is usually darker and not unfrequently
nearly black. The mane begins to grow when the animal is about three
years old, and is fully developed at five or six.
In size the Lion is only equalled or exceeded by the Tiger among the
existing _Felidæ_; though both species present great variations, the
largest specimens of the latter appear to surpass the largest Lions. A
full-sized South African Lion, according to Selous, measures slightly
less than 10 feet from nose to tip of tail, following the curves of
the body. Harris gives 10 feet 6 inches, of which the tail occupies 3
feet. The Lioness is about a foot less. The tongue, like that of the
other species of the genus, is long and flat, and remarkable for the
development of the papillæ of the anterior part of the dorsal surface,
which (except near the edge) are modified so as to resemble long,
compressed, recurved, horny spines or claws; these, near the middle line,
attaining the length of one-fifth of an inch. They give the part of the
tongue on which they occur the appearance and feel of a coarse rasp, and
serve the purpose of such an instrument in cleaning the flesh from the
bones of the animals on which the Lions feed.
The habits of the Lion in a state of nature are fairly well known from
the united observations of numerous travellers and sportsmen who have
explored those districts of the African continent in which it is still
common. It lives chiefly in sandy plains and rocky places interspersed
with dense thorn-thickets, or frequents the low bushes and tall rank
grass and reeds that grow along the sides of streams and near the
springs where it lies in wait for the larger herbivorous animals on
which it feeds. Although it is occasionally seen abroad during the
day, especially in wild and desolate regions, where it is subject to
but little molestation, the night is, as in the case of so many other
predaceous animals, the period of its greatest activity. It is then that
its characteristic roar is chiefly heard, as thus graphically described
by Gordon Cumming:—
“One of the most striking things connected with the Lion is his voice,
which is extremely grand and peculiarly striking. It consists at
times of a low, deep moaning, repeated five or six times, ending in
faintly audible sighs; at other times he startles the forest with loud,
deep-toned, solemn roars, repeated in quick succession, each increasing
in loudness to the third or fourth, when his voice dies away in five
or six low muffled sounds very much resembling distant thunder. At
times, and not unfrequently, a troop may be heard roaring in concert,
one assuming the lead, and two, three, or four more regularly taking
up their parts, like persons singing a catch. Like our Scottish stags
at the rutting season, they roar loudest in cold frosty nights; but on
no occasions are their voices to be heard in such perfection, or so
intensely powerful, as when two or three troops of strange Lions approach
a fountain to drink at the same time. When this occurs, every member
of each troop sounds a bold roar of defiance at the opposite parties;
and when one roars, all roar together, and each seems to vie with his
comrades in the intensity and power of his voice. The power and grandeur
of these nocturnal concerts are inconceivably striking and pleasing to
the hunter’s ear.”
“The usual pace of a Lion,” C. J. Andersson[433] says, “is a walk, and,
though apparently rather slow, yet, from the great length of his body, he
is able to get over a good deal of ground in a short time. Occasionally
he trots, when his speed is not inconsiderable. His gallop—or rather
succession of bounds—is, for a short distance, very fast—nearly or quite
equal to that of a horse. Indeed, unless the steed has somewhat the start
when the beast charges, it will be puzzled to escape. Many instances
are on record of horsemen who have incautiously approached too near to
the Lion, prior to firing, who have been pulled down by him before they
could get out of harm’s way. Happily, however, the beast soon tires of
the exertion of galloping, and unless his first rush succeeds he, for
the most part, soon halts and beats a retreat.” “The Lion, as with other
members of the feline family,” the same writer tells us, “seldom attacks
his prey openly, unless compelled by extreme hunger. For the most part he
steals upon it in the manner of a cat, or ambushes himself near to the
water or a pathway frequented by game. At such times he lies crouched
upon his belly in a thicket until the animal approaches sufficiently
near, when, with one prodigious bound, he pounces upon it. In most cases
he is successful, but should his intended victim escape, as at times
happens, from his having miscalculated the distance, he may make a second
or even a third bound, which, however, usually prove fruitless, or he
returns disconcerted to his hiding-place, there to wait for another
opportunity.” His food consists of all the larger herbivorous animals of
the country in which he resides—buffaloes, antelopes, zebras, giraffes,
or even young elephants or rhinoceroses, though the adults of these
latter he dare not attack. In cultivated districts the cattle, sheep,
and even human inhabitants are never safe from his nocturnal ravages.
He appears, however, as a general rule, only to kill when hungry or
attacked, and not for the mere pleasure of killing, as with some other
carnivorous animals. Moreover, he by no means limits himself to animals
of his own killing, but, according to Selous, often prefers eating game
that has been killed by man, even when not very fresh, to taking the
trouble to catch an animal himself. All books of African travel and sport
abound with stories, many of which are apparently well authenticated, of
the lion’s prodigious strength, as, exemplified by his being able to drag
off a whole ox in his mouth to a long distance, even leaping fences and
dykes with it.
The Lion appears to be monogamous, a single male and female continuing
attached to each other irrespectively of the pairing season. At all
events the Lion remains with the Lioness while the cubs are young and
helpless, and assists in providing her and them with food, and in
educating them in the art of providing for themselves. The number of cubs
at a birth is from two to four, usually three. They are said to remain
with their parents till they are about three years old. The following
account by an eye-witness gives a good idea of Lion family life[434]:—
“I once had the pleasure of, unobserved myself, watching a lion family
feeding. I was encamped on the Black Umfolosi in Zululand, and towards
evening, walking out, about half a mile from camp, I saw a herd of zebra
galloping across me, and when they were nearly 200 yards off, I saw a
yellow body flash towards the leader, and saw him fall beneath the lion’s
weight. There was a tall tree about 60 yards from the place, and anxious
to see what went on, I stalked up to it, while the lion was still too
much occupied to look about him, and climbed up. He had by this time
quite killed the beautifully striped animal, but instead of proceeding
to eat it, he got up and roared vigorously, until there was an answer,
and in a few minutes a lioness, accompanied by four whelps, came trotting
up from the same direction as the zebra, which no doubt she had been to
drive towards her husband. They formed a fine picture as they all stood
round the carcase, the whelps tearing it and biting it, but unable to get
through the tough skin. Then the lion lay down, and the lioness driving
her offspring before her did the same four or five yards off, upon
which he got up, and, commencing to eat, had soon finished a hind leg,
retiring a few yards on one side as soon as he had done so. The lioness
came up next and tore the carcase to shreds, bolting huge mouthfuls, but
not objecting to the whelps eating us much as they could find. There was
a good deal of snarling and quarrelling among these young lions, and
occasionally a stand-up fight for a minute, but their mother did not take
any notice of them, except to give them a smart blow with her paw if
they got in her way.... There was now little left of the zebra but a few
bones, which hundreds of vultures were circling round waiting to pick,
while almost an equal number hopped awkwardly about on the ground within
50 or 60 yards of it, and the whole lion family walked quietly away, the
lioness leading, and the lion, often turning his head to see that they
were not followed, bringing up the rear.”
Though not strictly gregarious, Lions appear to be sociable towards their
own species, and often are found in small troops, sometimes consisting of
a pair of old Lions, with their nearly full-grown cubs, but occasionally
of adults of the same sex; and there seems to be good evidence that
several Lions will associate together for the purpose of hunting upon
a preconcerted plan. As might be supposed, their natural ferocity and
powerful armature are sometimes turned upon one another; combats, often
mortal, occur among male Lions under the influence of jealousy; and
Andersson relates an instance of a quarrel between a hungry Lion and
Lioness over the carcase of an Antelope which they had just killed, and
which did not seem sufficient for the appetite of both, ending in the
Lion not only killing, but even devouring his mate. Old Lions, whose
teeth have become injured with constant wear, often become “man-eaters,”
finding their easiest means of obtaining a subsistence in lurking in
the neighbourhood of villages, and dashing into the tents at night and
carrying off one of the sleeping inmates. Lions differ from most of the
smaller _Felidæ_ in never climbing trees; indeed, as mentioned before,
they are rarely found in forests.
With regard to the character of the Lion, those who have had
opportunities of observing it in its native haunts differ greatly. The
exaggerated accounts of early writers as to its courage, nobility, and
magnanimity have led to a reaction, which causes some modern authors
to speak of it in language quite the reverse, and to accuse it of
positive cowardice and all kinds of meanness. Livingstone goes so far
as to say, “Nothing that I ever learned of the lion could lead me to
attribute to it either the ferocious or noble character ascribed to
it elsewhere,” and he adds that its roar is not distinguishable from
that of the ostrich. Of course these different estimates depend to
a great extent upon the particular standard of the writer, and also
upon the circumstance that Lions, like other animals, undoubtedly show
considerable individual differences in character, and behave differently
under varying circumstances. They are certainly not so reckless as to be
entirely devoid of the instinct of self-preservation, and if one, perhaps
satiated with a good meal the night before, unexpectedly disturbed in the
daytime, will occasionally retreat when confronted, even by an unarmed
man, that is scarcely a reason for assigning cowardice as one of the
characteristics of the species. The latest authority, Selous, while never
denying the daring courage of the Lion when hungry or provoked, and
vindicating the awe-inspiring character of the roar of several Lions in
unison, when heard at close quarters, as the grandest sound in nature,
says with regard to its outward aspect:—
“It has always appeared to me that the word ‘majestic’ is singularly
inapplicable to the lion in its wild state, as when seen by daylight
he always has a stealthy furtive look that entirely does away with the
idea of majesty. To look majestic a lion should hold his head high. This
he seldom does. When walking he holds it low, lower than the line of
his back, and it is only when he first becomes aware of the presence of
man that he sometimes raises his head and takes a look at the intruder,
usually lowering it immediately, and trotting away with a growl. When
at bay, standing with open mouth and glaring eyes, holding his head
low between his shoulders, and keeping up a continuous low growling,
twitching his tail the while from side to side, no animal can look more
unpleasant than a lion; but there is then nothing majestic or noble in
his appearance.”
Notwithstanding this evidently truthful description of the animal when
seen under what may be called unfavourable circumstances, no one with an
eye for beauty can contemplate the form of a fine specimen of a Lion,
at all events in a state of repose, even though in the confinement of a
menagerie, without being impressed with the feeling that it is a grand
and noble-looking animal.
The Tiger (_F. tigris_) is so closely related to the Lion that it is
chiefly by external characters that the two species are distinguished.
There are, however, slight distinctions in the proportionate size of the
lower teeth, the general form of the cranium, and the relative length of
the nasal bones and ascending processes of the maxillaries by which the
skull of the Lion and Tiger can be easily discriminated by the practised
observer.
Although examples of both species present considerable variations in
size, and reliance cannot always be placed upon alleged dimensions,
especially when taken from skins stripped from the body, it seems well
ascertained that the length of the largest-sized Bengal Tiger may exceed
that of any Lion. According to Mr. W. T. Blanford,[435] adult males
measure from 5½ to 6½ feet from the nose to the root of the tail; the
tail itself measuring some 3 feet in length. Measured along the curves
of the head and back to the tip of the tail, males usually give a length
of from 9 to 10 feet, but some specimens reach to 12 feet. The female
is somewhat smaller, and has a lighter and narrower head. The Tiger
has no mane, but in old males the hair of the cheeks is rather long
and spreading. The ground colour of the upper and outer parts of the
head, body, limbs, and tail is a bright rufous fawn, and these parts
are beautifully marked with transverse stripes of a dark, almost black
colour. The markings vary much in different individuals, and even on the
two sides of the same individual. The under parts of the body, the inside
of the limbs, the cheeks, and a large spot over each eye are nearly
white. The Tigers which inhabit hotter regions, as Bengal and the south
Asiatic islands, have shorter and smoother hair, and are more richly
coloured and distinctly striped than those of Northern China and Siberia,
in which the fur is longer, softer, and lighter coloured.
[Illustration: FIG. 225.—The Tiger (_Felis tigris_).]
The Tiger is exclusively Asiatic, but has a very wide range in that
continent, having been found in almost all suitable localities south of
a line drawn from the river Euphrates, passing along the southern shores
of the Caspian and Sea of Aral by Lake Baikal to the Sea of Okhotsk.
Its most northern range is the territory of the Amur, its most southern
the islands of Sumatra, Java, and Bali. Westward it reaches to Turkish
Georgia and eastward to the island of Saghalin. It is absent, however,
from the great elevated plateau of Central Asia, nor does it inhabit
Ceylon, Borneo, or the other islands of the Indo-Malayan Archipelago,
except those above mentioned. Its absence from Ceylon leads Mr. Blanford
to conclude that the Tiger has only recently migrated into Southern India.
The principal food of the Tiger in India is cattle, deer, wild hog, and
pea-fowl, and occasionally human beings. The regular “man-eater” is
generally an old Tiger whose vigour is passed, and whose teeth are worn
and defective; it takes up its abode in the neighbourhood of a village,
the population of which it finds an easier prey than the larger or
wilder animals named above. Though chiefly affecting grassy plains or
swamps, it is also found in forests, and seems to be fond of haunting the
neighbourhood of old ruins. As a rule, Tigers do not climb trees; but
when pressed by fear, as during an inundation, they have been known to
do so. They take to the water readily and are good swimmers. The Tigers
of the Sundarbans (Ganges delta) continually swim from one island to the
other to change their hunting-grounds for deer. The following extract on
the habits of the Tiger is taken from Sir J. Fayrer’s _Royal Tiger of
Bengal_ (1875):—
“The tigress gives birth to from two to five, even six cubs; but three is
a frequent number. She is a most affectionate and attached mother, and
generally guards and trains her young with the most watchful solicitude.
They remain with her until nearly full grown, or about the second year,
when they are able to kill for themselves and begin life on their own
account. Whilst they remain with her she is peculiarly vicious and
aggressive, defending them with the greatest courage and energy, and
when robbed of them is terrible in her rage; but she has been known to
desert them when pressed, and even to eat them when starved. As soon
as they begin to require other food than her milk, she kills for them,
teaching them to do so for themselves by practising on small animals,
such as deer and young calves or pigs. At these times she is wanton and
extravagant in her cruelty, killing apparently for the gratification
of her ferocious and bloodthirsty nature, and perhaps to excite and
instruct the young ones, and it is not until they are thoroughly capable
of killing their own food that she separates from them. The young tigers
are far more destructive than the old. They will kill three or four
cows at a time, whilst the older and more experienced rarely kill more
than one, and this at intervals of from three or four days to a week.
For this purpose the tiger will leave its retreat in the dense jungle,
proceed to the neighbourhood of a village or gowrie, where cattle feed,
and during the night will steal on and strike down a bullock, drag it
into a secluded place, and then remain near the ‘marrie,’ or ‘kill,’ for
several days, until it has eaten it, when it will proceed in search of a
further supply, and, having found good hunting ground in the vicinity of
a village or gowrie, continue its ravages, destroying one or two cows or
buffaloes a week. It is very fond of the ordinary domestic cattle, which
in the plains of India are generally weak, half-starved, under-sized
creatures. One of these is easily struck down and carried or dragged off.
The smaller buffaloes are also easily disposed of; but the buffalo bulls,
and especially the wild ones, are formidable antagonists, and have often
been known to beat the Tiger off, and even to wound him seriously.”
In many districts of India the number of Tigers has been very
considerably diminished of late years. In some other countries they
appear, however, to be on the increase; thus according to one of the
administration reports of Java laid before the Dutch Chambers, portions
of that island are being depopulated through Tigers. In 1882 the
population of a village in the south-west of the Bantam province was
removed and transferred to an island off the coast in consequence of the
trouble caused to the people by Tigers. These animals have now become
an intolerable pest in parts of the same province. The total population
is about 600,000, and, in 1887, sixty-one were killed by Tigers, and in
consequence of the dread existing among the people, it has been proposed
to deport the inhabitants of the villages most threatened to other parts
of the country where Tigers are not so common, and where they can pursue
their agricultural occupations with a greater degree of security. At
present they fear to go anywhere near the borders of the forest. The
people seem disinclined, or they lack the means and courage, to attack
and destroy their enemy, although considerable rewards are offered by
Government for the destruction of beasts of prey. In 1888 the reward for
killing a Royal Tiger was raised to two hundred florins. It appears also
that the immunity of the Tiger is in part due to superstition, for it is
considered wrong to kill one unless he attacks first or otherwise does
injury.
The Leopard (_F. pardus_, Fig. 226), although belonging to the same
restricted group as the two preceding species, is distinguished from both
by its inferior size, and its coloration. The animal now commonly known
as the Leopard was called Pard (πάρδος and πάρδαλις) or Panther (πάνθηρ)
by the ancients. Leopard (_leo-pardus_) is a later term, originally
applied, it is believed, to the Cheeta or Hunting Leopard, upon the
supposition that it was a creature intermediate between the Lion and the
true Pard. If so it has been completely transferred to the more common
species, and though in this sense a perfectly unnecessary and unmeaning
term, has gradually superseded those by which this was originally known.
Pard, so commonly used by Elizabethan authors, is now nearly obsolete
in the English language, and Panther has either become synonymous with
Leopard, or is used vaguely for any similar large feline animal, even the
Puma of America.
[Illustration: FIG. 226.—The Leopard (_Felis pardus_).]
Owing to their extensive geographical range, and the great variations,
both in size, form, and coloration to which Leopards are subject,
zoologists have scarcely decided whether all the forms popularly referred
to this animal should be regarded as specifically alike, or whether they
should constitute several distinct species, but the prevailing opinion
is in favour of the former view. The attempts to separate a larger and
more robust variety, under the name of Panther, from a smaller and more
graceful form, to which the term Leopard might properly be restricted,
have failed, owing to the existence of intermediate conditions which
cannot be assigned definitely to either one or the other form.[436] The
most marked anatomical difference yet noted in different varieties of
leopard is in the length of the tail as compared with that of the body,
even the number of the caudal vertebræ showing variation, though within
what limits, and whether correlated with other characters, has not yet
been clearly ascertained. The fur of those specimens which inhabit the
most northern confines of its range of distribution, as North China, is
longer and softer, and the markings are consequently less distinct than
on those from more congenial climates, and the well-marked variation thus
produced has given rise to the idea of specific distinction.
The size of different individuals, as before said, varies greatly, the
head and body usually measuring from 3½ to 4½ feet in length, and the
tail from 2½ to 3 feet, but specimens have been met with which fall short
of or exceed these limits. The ground colour of the fur varies from a
pale fawn to a rufous buff, graduating into a pure white on the under
parts and inside of the limbs. This is spotted over with dark brown or
black; the spots on the back and sides being arranged in rosettes or
broken rings, which vary greatly in size and distinctness in different
individuals, but are without the central spot seen in those of the
Jaguar. The spots on the under parts and limbs are simple and blacker
than those on the other parts of the body. The bases of the ears behind
are black, the tips buff. The upper side of the tail is buff, spotted
with broken rings like the back, its under surface white with simple
spots. The hair of the cubs is longer than that of the adults, its
ground colour less bright, and its spots less distinct. Perfectly black
Leopards, which, however, in certain lights show the characteristic
markings on the fur, are not uncommon. These appear to be examples of
melanism, occurring as individual variations, sometimes in one cub out
of a litter of which the rest are normally coloured, and therefore not
indicating a distinct race, much less a species. These are met with
chiefly in Southern Asia. We are not aware of any recorded case from
Africa, though there seems no reason why they should not occur.
In habits the Leopard resembles the other large Cat-like animals,
yielding to none in the ferocity and bloodthirstiness of its disposition.
It is exceedingly quick and active in its movements, but seizes its
prey by waiting in ambush or stealthily approaching to within springing
distance, when it suddenly rushes upon it and tears it to the ground with
its powerful claws and teeth. It preys upon almost any animal it can
overcome, such as antelopes, deer, sheep, goats, monkeys, peafowls, and
is said to have a special liking for dogs. It not unfrequently attacks
human beings in India, chiefly children and old women, but instances have
been known of a Leopard becoming a regular “man-eater.” When favourable
opportunities occur, it often kills many more victims than it can devour
at once, apparently to gratify its propensity for killing, or only for
the sake of their fresh blood. It generally inhabits woody districts,
and can climb high trees with facility if necessary for its safety when
hunted, but usually lives on or near the ground, among rocks, bushes, and
roots and low branches of large trees.
The present geographical range of the Leopard is very extensive, as it is
met with in various suitable localities, where not too much interfered
with by human cultivation, throughout the greater part of Africa from
Algeria to the Cape Colony, and through the whole of the South of Asia
from Palestine to China, including all India south of the Himalaya,
and the islands of Ceylon, Java, Sumatra, and Borneo. Fossil bones and
teeth, indistinguishable from those of existing Leopards, have been
found in cave-deposits of Pleistocene age in Spain, France, Germany, and
England. The evidence of the former existence of the Leopard in England
is described at length by Boyd Dawkins and Sanford in their _British
Pleistocene Mammalia_.[437]
The Ounce, or Snow Leopard (_F. uncia_), inhabits the highlands of
Central Asia, from the lofty mountains of Tibet to the southern parts
of Siberia, at altitudes of from 9000 to 18,000 feet above the sea.
It is about the size of the common Leopard, but lighter in colour,
with longer fur, less distinct spots, and a long thick tail. Its skull
differs in shape from that of all the other _Felidæ_; the facial portion
being very broad, the nasal bones especially being wide and depressed,
and the zygomatic arches very strong and deep. The Clouded Tiger (_F.
nebulosa_[438]) is a beautifully marked species, with elongated head
and body, long tail, and rather short limbs. The canine teeth are
proportionally longer than in any existing member of the genus. It is
thoroughly arboreal, and is found in the forests of South-East Asia and
the islands of Sumatra, Java, Borneo, and Formosa. _F. serval_, the
Serval, from South Africa, is yellow with black spots, and has a short
tail and large ears. Numerous smaller species called Tiger Cats and Wild
Cats, of which the Oriental _F. marmorata_ (Fig. 227) is a good example,
are found throughout the warmer parts of Asia and Africa. The Wild Cat of
Europe, _F. catus_, still inhabits the mountainous and wooded parts of
Great Britain.
[Illustration: FIG. 227.—The Marbled Cat (_Felis marmorata_). From
Blanford, _Mammalia of British India_, p. 74, after Elliot.]
The Caffre Cat (_F. caffra_[439]), of Africa and Southern Asia, was the
species held in veneration by the ancient Egyptians, and immense numbers
of its mummified remains have recently been found in Egypt, whence they
have been imported in large quantities to this country for manure. This
species is generally regarded as the main ancestral stock from which the
European Domestic Cat has been derived; one of the arguments in support
of this opinion being that the whole of the sole of the hind foot of
_F. caffra_ is black, and that the same feature obtains in the darker
varieties of the Domestic Cat; while in _F. catus_ there are only spots
of black upon this portion of the limb. Remains of the Caffre Cat occur
in the Pleistocene cave-deposits at Gibraltar. The Indian _F. rubiginosa_
is the smallest species of Cat.
The Caracal or Persian Lynx (_F. caracal_) is an animal about the size
of a fox, of slender build, with a moderately long tail, reaching down
to the heels. It is of a uniform vinous or bright fulvous brown colour
above, and is paler, sometimes almost white, beneath. It is quite or
almost entirely unspotted. The tail has a black tip, and the ears are
black externally, long, upright, pointed, and surmounted by a pencil
of fine black hairs. It inhabits Central and North-West India, Persia,
Arabia, Syria, and the greater part of Africa.
The true Lynxes comprise various species or varieties found in the
northern and temperate regions of both the Old and New World, all larger
than the true Wild Cats, with long limbs, short stumpy tail, ears tufted
at the tip, and pupil of the eye linear when contracted. Their fur is
generally long and soft, varying, however, according to season and
locality, and always longish upon the cheeks. Their colour is always
light brown or gray, and generally more or less spotted with a darker
shade. The naked pads of the feet are more or less covered by the hair
that grows between them. The skull and skeleton do not differ markedly
from those of the other cats, but the small anterior upper premolar tooth
found in many other species is usually wanting; and the lower carnassial
has a rudimental talon. Their habits are exactly those of the other Wild
Cats, and they are exceeded by none in the untameable savageness of their
disposition. They capture their prey in the same manner, either lying in
wait, or noiselessly stealing within reach, and then making a sudden rush
or spring upon it. Their food consists of any mammals or birds which they
can overpower. In inhabited countries they commit extensive ravages upon
sheep, lambs, and poultry. Lynxes generally frequent rocky places and
forests, being active climbers, and passing much of their time among the
branches of the trees. Their skins are of considerable commercial value.
Zoologists are by no means agreed at present as to the specific
distinctions, if any really exist, between the various modifications
of this group. As many as eight species are sometimes recognised, four
belonging to the Old and four to the New World. The former are _F.
lynx_, of Scandinavia, Russia, Northern Asia, and till lately the forest
regions of Central Europe; though not an inhabitant of Britain during
the historic period, its remains have been found in cave-deposits of
Pleistocene age; _F. cervaria_, Siberia; _F. pardina_, Turkey, Greece,
Sicily, Sardinia, and Spain; and _F. isabellina_, Tibet. The American
varieties are _F. canadensis_, the most northern species, and _F.
rufa_, the American Wild Cat or Bay Lynx, extensively distributed from
the Atlantic to the Pacific throughout nearly the whole latitude of
the United States, but replaced in Texas and southern California by
_F. maculata_, and in northern Oregon and Washington territory by _F.
fasciata_.
[Illustration: FIG. 228.—European Lynx (_Felis lynx_). From a drawing by
Wolf in Elliot’s _Monograph of the Felidæ_.]
In both cases, as might be supposed, specimens obtained from the more
southern climates are shorter in their fur, more brightly coloured,
and more distinctly spotted than those from colder regions. When only
a few individuals of each most markedly different form are examined
the distinctions are sufficiently evident. The occurrence, however,
of transitional or intermediate forms makes it extremely difficult to
draw the line between the different varieties or species, or to assign
definite characters by which they can be separated. Wherefore it is best
at present to accept the so-called species as only provisional, and wait
until more abundant materials, with fuller knowledge of the localities
from which they are derived, and of the variations due to age, sex,
season, and climate, have been more carefully studied. We shall then
probably come to the conclusion that all or nearly all the existing forms
of northern Lynxes, whether American or Eurasian, belong to what may
fairly be called a species, which is becoming by degrees differentiated
into several more or less strongly marked local varieties. Mr. W. T.
Blanford has indeed shown that the Tibetan Lynx (_F. isabellina_) is
inseparable from _F. lynx_; the specimens from Gilgit being intermediate
in colour between the typical forms of the two races. On the other hand,
from the evidence of cranial characters, Professor Mivart is disposed to
regard _F. pardina_ as a valid species.
[Illustration: FIG. 229.—The Puma (_Felis concolor_).]
B. _New World Species._—The Puma or Couguar (_F. concolor_, Fig. 229),
commonly called “Panther” in the United States, is about the size of a
Leopard, but of an uniform brown colour. It usually measures from nose
to root of tail about 40 inches, the tail being rather more than half
that length. The head is rather small compared with that of other Cats
and has no mane. The ears are large and rounded. The tail is cylindrical,
with some bushy elongation of the hairs near the end, but not forming a
distinct tuft as in the Lion. The general colour of all the upper parts
and sides of the adult is a tawny yellowish-brown, sometimes having a
gray or silvery shade, but in some individuals dark or inclining to
red. The lower parts of the body, inner surface of the limbs, the
throat, chin, and upper lip are dirty white; the outside of the ears,
particularly at their base, and a patch on each side of the muzzle black;
the end of the tail dusky. The young are, when born, spotted with dusky
brown and the tail ringed; these markings gradually fading, and quite
disappearing before the animal becomes full-grown.
The Puma has an exceedingly wide range of geographical distribution,
extending over a hundred degrees of latitude, from Canada in the north
to Patagonia in the south, and was formerly pretty generally diffused
in suitable localities from the Atlantic to the Pacific Ocean, but the
advances of civilisation have in recent years considerably curtailed
the extent of the districts which it inhabits. In Central America it is
still common in the dense forests which clothe the mountain ranges as
high as 8000 or 9000 feet above the sea-level, where the hideous sound
of its howling is said to be almost continuously heard at night during
the breeding season. Though an expert climber, it is by no means confined
to wooded districts, being frequently found in scrub and reeds along the
banks of rivers, and even in the open pampas and prairies. Its habits
much resemble those of the rest of the group to which it belongs; and,
like the Leopard, when it happens to come within reach of an abundant
and easy prey, as the sheep or calves of an outlying farming station, it
kills far more than it can eat, either for the sake of the blood only
or to gratify its propensity for destruction. It rarely attacks man,
and, when pursued, escapes if possible by ascending lofty trees. Several
instances have occurred of Pumas becoming tame in captivity. Edmund Kean,
the celebrated actor, had one which followed him about like a dog. When
caressed they express their pleasure by purring like a domestic cat.
_F. onca_, the Jaguar, is a larger and more powerful animal than the
last, and more resembles the Leopard in its colours. It also is found in
both North and South America, but with less extensive range, reaching
northwards only as far as Texas, and southwards nearly to Patagonia.
It climbs as well as the Puma, and preys to a great extent upon
monkeys. Several allied smaller elegantly spotted forms inhabiting the
intratropical regions of America are commonly included under the name
of Ocelot or Tiger Cat, though zoologists are still undecided whether
under this designation several distinct species have not been confused,
or whether all the Ocelots are to be referred to a single species (_F.
pardalis_) showing great individual or racial variation. Their fur has
always a tawny yellow or reddish-gray ground colour, and is marked with
black spots, aggregated in streaks and blotches, or in elongated rings
enclosing an area which is rather darker than the general ground colour.
They range through the wooded parts of tropical America, from Arkansas
in the north as far south as Paraguay, and in their habits resemble
the other smaller members of the Cat tribe, being ready climbers and
exceedingly bloodthirsty.
_F. yaguarundi_, rather larger than the Domestic Cat, with an elongated
head and body, and of a uniform brownish-gray colour, ranges from
Matamoras to Paraguay. _F. eyra_ is a small Cat, very Musteline in form,
having an elongated head, body, and tail, and short limbs, and is also
of a uniform light reddish-brown colour. It is a native of South America
and Mexico. _F. pajeros_ is the Pampas Cat. The American Lynxes have been
already noticed with those of the Old World.
[Illustration: FIG. 230.—The Ocelot (_Felis pardalis_).]
C. _Fossil Species._—It has been already incidentally mentioned that
several of the existing species of _Felis_, such as the Lion, Leopard
and Caffre Cat, are met with in a fossil condition in the European
Pleistocene deposits, and it may be added that the Pardine Lynx has left
its remains in the cavern-deposits of Gibraltar. The caves of Brazil have
yielded remains of the Jaguar and Ocelot; while the Puma is found in the
Pleistocene of the United States. Existing species now inhabiting India
are met with in cavern-deposits in Madras. In the Pliocene Siwaliks of
Northern India the huge extinct _F. cristata_ shows characters connecting
it both with the Tiger and the Jaguar; and the same deposit also
contains the remains of a small species of the size of _F. bengalensis_.
In Europe numerous species occur in the Upper and Lower Pliocene, some
of which were as large as a Leopard. _F. atrox_ and _F. augusta_, of the
Pliocene of the United States, were of the dimensions of the Lion.
_Cynælurus._[440]—The Cheeta or Hunting Leopard (_C. jubatus_) is
distinguished from the other _Felidæ_ by the inner tubercle of the upper
carnassial, though supported by a distinct root, having no salient cusp
upon it; by the tubercular molar being more in a line with the other
teeth; and by the claws being smaller, less curved, and less completely
retractile, owing to the feebler development of the elastic ligaments.
The skull is short and high, with the frontal region broad and elevated
in consequence of the large development of the frontal air-sinuses. The
head is small and round, the body light, the limbs and tail long. Its
colour is pale yellowish-brown with small black spots. The Cheeta is
less savage and more easily tamed than most of the Cats. In Asia it has
been trained for the chase of the Antelope. It has rather an extensive
geographical range from the Cape of Good Hope, throughout Africa and the
south-western parts of Asia, as far as Southern India.
_Extinct Genera._—A number of forms are gradually becoming known,
especially through the researches of American palæontologists, which,
though evidently animals of the same general type, and therefore to be
placed in or near the family _Felidæ_, depart so much in various details
of structure that they must be referred to different genera. As one of
the points in which _Felis_ manifests its specialisation is the reduction
of the number of the molar series of teeth, with concomitant shortening
of the jaws, it might be supposed that in the earlier and perhaps
ancestral forms these teeth would be more numerous and approach more
nearly to the primitive or typical number of the heterodont mammals, viz.
seven on each side. This is actually the case. Similarly we find that
many of these forms exhibit a less specialised structure of the teeth
themselves, as is shown by the absence of the anterior lobe of the upper
carnassial, and the retention of the hind talon in the corresponding
lower tooth. Again, some of them have an alisphenoid canal in the skull;
while the femur may have a third trochanter, and the claws be very
imperfectly retractile.
An extremely generalised form is the small _Proælurus_, from the Upper
Eocene and Lower Miocene, with _p_ ⁴⁄₄, _m_ ¹⁄₂, an alisphenoid canal,
and a third trochanter to the femur. _Dinictis_, of the North American
Miocene, is a larger allied form, with _p_ ³⁄₃, _m_ ¹⁄₂; the upper
carnassial having no anterior lobe, and the ungual phalanges being devoid
of bony sheaths. The characters of the base of the skull, and the form
and relations of the astragalus, differ very considerably from _Felis_.
_Pseudælurus_, from the French Miocene, is another very generalised
Feline, in which there may be either three or four premolars, and the
lower carnassial may retain its inner cusp. _Ælurictis_, of the French
Phosphorites, with _p_ ³⁄₃₋₄, _m_ ¹⁄₁₋₂, together with several American
Miocene genera, such as _Nimravus_ (_p_ ³⁄₂, _m_ ¹⁄₂), _Archælurus_ (_p_
³⁄₃₋₄, _m_ ¹⁄₂), _Pogonodon_ (_p_ ³⁄₃, _m_ ¹⁄₁), and _Hoplophoneus_ (_p_
²⁻³⁄₂, _m_ ¹⁄₁), approach more closely to the modern Cats, although many
or all of them retain the alisphenoid canal, and have not yet developed
the anterior lobe to the upper carnassial, or lost the talon to the lower
one. _Hoplophoneus_ has a descending flange to the mandible; and its
scapholunar bone has a line indicating its dual origin; while the femur
still retains the third trochanter, of which all traces are lost in the
modern Cats.
On the other hand, some of the extinct _Felidæ_ show a most remarkable
tendency towards a specialisation not occurring in any of the surviving
members of the family, viz. an enormous development of the upper canines,
with which is usually associated an expansion downwards and flattening
of the anterior part of the ramus of the lower jaw, on the outer side
of which the canine lies, when the mouth is closed. In _Machærodus
næogeus_, the Sabre-toothed Tiger, from the caves of Brazil and also
from Pleistocene deposits near Buenos Ayres, an animal about the size
of a Tiger, these teeth are 7 inches in length, greatly compressed, and
finely serrated on the trenchant anterior edges. Similar serrations are
seen on a much fainter scale in the unworn teeth of modern Tigers. Many
modifications of this commonly-called “machærodont” type have been met
with both in the Old and New World. In _M. cultridens_, of the Upper
Pliocene of Italy and France, the upper canine is long and narrow, with
smooth cutting edges; the smaller form described as _M. meganthereon_
being apparently the female of this species. _M. crenatidens_, of
the same deposits, is distinguished by the shorter and broader upper
canine, in which both edges are strongly serrated; the same feature
occurring in the closely allied or identical _M. latidens_ of the English
cavern-deposits. The Italian Pliocene form described as _M. nestianus_
has serrations only on the hinder edge of the upper canine, and the third
lower premolar is separated by a long interval from the fourth. _M.
necator_, of the Pleistocene of South America, is remarkable as being
the only member of the family in which the humerus has no entepicondylar
foramen. A very remarkable form, _Eusmilus_, from the Upper Eocene
Phosphorites of Central France, differs from all other known Felines in
having only two pairs of incisors in the lower jaw, and a small canine
separated by a very long diastema from the cheek-teeth, which consist
only of one premolar and one sectorial true molar. The lower jaw is
enormously expanded towards the symphysis to protect the large upper
canines. This animal then, although of Eocene age, appears to form the
culminating development of the sabre-toothed or machærodont dentition,
the most specially carnivorous type of structure known.
Other species of _Machærodus_ are found in the Pliocene deposits of
Europe and Asia. The accompanying woodcut exhibits the last two upper
teeth of the Indian _M. sivalensis_, from which it will be seen that the
inner tubercle of the carnassial is much reduced in size, while the molar
is very minute.
_Family_ VIVERRIDÆ.
[Illustration: FIG. 231.—Oral surface of the left upper carnassial and
molar of _Machærodus sivalensis_.]
Premolars ³⁄₃ or ⁴⁄₄. Molars ¹⁄₁ or ²⁄₂. Upper carnassial usually without
an anterior lobe, and the lower one with a well-developed talon; second
lower incisor (as in all the following families) raised above the level
of the first and third. Auditory bulla externally constricted, and
divided by a septum. An alisphenoid canal (with very rare exceptions).
Carotid canal distinct as a groove on the side of the bulla. Humerus
usually with an entepicondylar foramen. Digits usually 5-5, but sometimes
the pollex or hallux or both may be wanting. Dorsal vertebræ 13 or 14.
Limited in distribution to the Old World.
The subfamily =Cryptoproctinæ= contains the single genus
_Cryptoprocta_.[441] Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ¹⁄₁; total
36. The teeth generally closely resemble those of the _Felidæ_. The first
premolar of both jaws is very minute and early deciduous. The upper
carnassial has a very small inner tubercle, quite at the anterior part
of the tooth. The true molar is very small and placed transversely. The
lower carnassial has a large trenchant bilobed blade, and a very minute
talon, but no inner cusp. Skull generally like that of _Felis_, but
proportionately longer and narrower. Orbit widely open behind. Vertebræ:
C 7, D 13, L 7, S 3, C 29. Body elongated. Limbs moderate in size. Feet
subplantigrade; five well-developed toes on each, with sharp, compressed,
retractile claws. Ears moderate. Tail long and cylindrical.
The only known species, _C. ferox_, the “Foussa” of the Malagasy, is
peculiar to Madagascar, being the largest carnivorous animal in the
island. It is about twice the size of the common Cat (5 feet from nose
to end of tail), with short close fur of nearly uniform pale brown.
Little is as yet known of its habits, except that it is nocturnal,
frequently attacks and carries off goats, and especially kids, and shows
great ferocity when wounded, on which account it is much dreaded by the
natives.
The remaining numerous specific and generic modifications found in the
existing animals belonging to this family seem to arrange themselves
mainly into two tolerably distinct groups, distinguishable by the
characters of the auditory bulla and neighbouring parts of the base of
the skull, and by the structure of the feet. The one form has the genus
_Viverra_ or Civet Cats for its most typical representative, and the
other _Herpestes_ or the Ichneumons.
Subfamily =Viverrinæ=.—Auditory bulla oval, or rather conical, broad
and truncated and not everted behind, narrow in front and more or less
compressed at the sides. The outer or anterior chamber very small and
flat. The meatus with scarcely any inferior lip, its orifice being
close to the tympanic ring. Paroccipital process triangular, its apex
projecting slightly beyond the bulla. Claws strongly curved and more or
less retractile. Perineal scent-glands generally present.
This subfamily includes both Ethiopian and Oriental forms, but the former
are the more numerous.
The typical section, which includes five genera, has the following
characters. Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₂ (¹⁄₂ in
_Prionodon_); total 40. Skull elongated; facial portion small and
compressed. Orbits well-defined but incomplete behind. Teeth always
sectorial, never very small. Vertebræ: C 7, D 13, L 7 (or D 14, L 6), S
3, C 22-30. Body elongated and compressed. Head pointed in front; ears
rather small. Extremities short. Feet small and rounded. Toes short, five
on each foot. First toe both on fore and hind feet much shorter than the
others. Palms and soles covered with hair, except the pads of the feet
and toes, and in some species a narrow central line on the under side
of the sole, extending backwards nearly to the heel. Tail moderate or
long; usually marked with dark and light rings. A pair of large glandular
follicles situated on the perineum (in both sexes), and secreting in most
species an oily substance of a peculiarly penetrating odour.
The numerous species of this section form a large series, the two
extremes of which differ considerably, but the several genera into which
they may be divided blend so into one another that it is difficult to
differentiate them sharply.
All the animals of this section are, for their size, extremely active,
fierce, and rapacious. They feed chiefly on small mammals and birds.
_Viverra._[442]—This includes the largest species. The teeth (Fig. 232)
are stouter and less compressed than in the other genera; the second
upper molar being especially larger. The auditory bulla smaller and more
pointed in front. Body shorter and stouter; limbs longer; tail shorter,
tapering. Under side of tarsus completely covered with hair. Claws
longer and less retractile. Fur rather long and loose, and in the middle
line of the neck and back usually elongated so as to form a sort of
crest or mane; neck with a black gorget. Pupil circular when contracted.
Perineal glands greatly developed. These characters apply especially
to _V. civetta_, the African Civet, or “Civet-Cat” as it is commonly
called, an animal rather larger than a common Fox, and an inhabitant of
intratropical Africa. _V. zibetha_, the Indian Civet, of about equal
size, inhabits Bengal, China, the Malay Peninsula, and adjoining islands.
_V. tangalunga_, from Java, Sumatra, Borneo, and the Philippines, and _V.
megaspila_, from Burma, are smaller but nearly allied animals; the latter
being more distinctly spotted than either of the others. From these
species and the next the civet of commerce, once so much admired as a
perfume in England, and still largely used in the East, is obtained. The
animals are kept in cages, and the odoriferous secretion collected from
the interior of the perineal follicles with a spoon or spatula.
[Illustration: FIG. 232.—The left upper dentition of the Indian Civet
(_Viverra zibetha_). From the _Palæontologia Indica_.]
The Rasse or Lesser Indian Civet (_V. malaccensis_) may be regarded as
the representative of a distinct group of _Viverra_, although often
referred to a separate genus (_Viverricula_). The size of this animal
is smaller than in the typical group, the build is slighter, the muzzle
finer, the claws sharper and more curved, and there is no erectile mane
along the back. Generally there is an alisphenoid canal in the skull; and
the anterior chamber of the auditory bulla is much more inflated than the
hinder one, so that the apparent length of the whole bulla is increased.
This species is found over the greater part of India, and extends to the
Malay Peninsula and Southern China.
Large species of _Viverra_ occur in the Pleistocene and Pliocene of
India, and also in the Pliocene of France, which approximate in some
characters of the dentition to the extinct genus _Ictitherium_, mentioned
at the end of the family. Species of this genus have also been described
from the Miocene and Upper Eocene of Europe. The Lower Miocene _V.
antiqua_ has an alisphenoid canal, and all the other cranial characters
of the typical forms.
_Fossa._[443]—The Fossa of Madagascar comes so close to the Rasse that
its right to generic distinction seems doubtful. There is, however, no
scent-pouch. The limbs are slender; and there are two small bare spots
on the sole of the hind foot, above the plantar pads. There is no dark
line along the back; the throat gorget of _Viverra_ is absent; and in the
tail the spots only tend to form rings, which are not complete. The skull
has an alisphenoid canal, and a large bulla as in the typical group of
_Viverra_.
[Illustration: FIG. 233.—The Common Genet (_Genetta vulgaris_).]
_Genetta._[444]—The Genettes are smaller animals, with more elongated
and slender bodies, and shorter limbs than the Civets. Skull elongated
and narrow. Auditory bulla large, elongated, rounded at both ends. Teeth
compressed and sharp pointed. The inner side of the third upper premolar
has a tubercle not present in the previous genus, and the talon of the
lower carnassial is larger. Pupil contracting to a linear aperture. Tail
long, slender. Fur short and soft, spotted or cloudy. Under side of the
tarso-metatarsus with a narrow longitudinal bald streak. No pouch for
storing the secretion of the scent-gland. _G. vulgaris_, the common
Genet (Fig. 233), is found in France south of the river Loire, Spain,
South-Western Asia, and Africa from Barbary to the Cape. _G. felina_,
_senegalensis_, _tigrina_, and _pardalis_ are other named species, all
African in habitat.
[Illustration: FIG. 234.—Stomach of Genet cut open. _œ_, Œsophagus; _pv_,
pyloric valve; _x_, sudden bend where the internal folds are interrupted.
(From Mivart, _Proc. Zool. Soc._ 1882, p. 505.)]
[Illustration: FIG. 235.—Cæcum of Genet. (After Mivart, _loc. cit._ p.
508.)]
A few details (taken from Professor Mivart’s memoirs on the Æluroidea) of
the anatomy of the soft parts of the Genet may be given as illustration
of these parts in the Carnivora generally, and of this family and genus
in particular. The salivary glands are shown in Fig. 19 (p. 56), and
these conform to the general type prevalent in the Æluroidea. Thus
there is a distinct zygomatic gland; the parotid with its (Steno’s)
duct is well developed; and there is a small submaxillary gland. The
stomach (Fig. 234), while conforming to the simple type characteristic
of the Carnivora, is much larger than in the Cat; it is characterised
by the presence of some strongly marked internal folds near the pyloric
extremity, which stop suddenly at a point where the stomach makes an
abrupt constriction and flexure. Beyond this point there are three other
longitudinal folds; and the pyloric valve is small. The allied genera
present modifications from this form of stomach. The cæcum (Fig. 235) is
short, thick, and pointed. The liver (Fig. 236) much resembles that of
the Cat, but differs in that the left lateral lobe is undivided, although
having a small groove on its posterior or abdominal aspect, while the
cystic fissure is less deep, and situated more to the right. The caudate
lobe is relatively longer, has a deep concavity, and runs uninterruptedly
into the Spigelian; the latter being relatively somewhat larger than in
the Cat, with a deep groove dividing the proximal third from the distal
two-thirds. In _Viverra_ the right lateral and right central lobes are
nearly equal in size. The variations in the form of the liver of the
allied genera are detailed in Professor Mivart’s memoir. The brain of the
Genet is shown in Fig. 23 (p. 71); the small depression _d_ placed on
the superior lateral gyrus appears to be the sole representative of the
distinct crucial sulcus which distinguishes the brains of the _Felidæ_
from those of all other members of the Æluroidea.
[Illustration: FIG. 236.—Abdominal aspect of the liver of the Genet. _c_,
Caudal lobe; _gb_, gall-bladder; _ha_, hepatic artery; _hd_, hepatic
duct; _LC_, left central lobe; _LL_, left lateral lobe; _pv_, portal
vein; _RC_, right central lobe; _RL_, right lateral lobe; _Sp_, Spigelian
lobe; _vc_, vena cava. (From Mivart, _Proc. Zool. Soc._ 1882, p. 510.)]
[Illustration: FIG. 237.—Cæcum of _Prionodon_. (From Mivart, _Proc. Zool.
Soc._ 1882, p. 508.)]
_Prionodon._[445]—This and the following genus comprise the beautiful
Linsangs (Fig. 238), which are distinguished from the preceding genera
by the loss of the second upper molar, which is, however, very small in
some of the Genets. In the present genus the ground colour is whitish
or yellowish with brown or black markings, which may either form broad
continuous patches across the hinder part of the body, or may be broken
up into spots. The tail is very long, the limbs comparatively short, and
the fur very short and close. The pollex and hallux are well developed;
the claws are almost completely retractile; and the tarsus and metatarsus
are completely haired. The pupil is round. The cæcum (Fig. 237) is
remarkably small. This genus is exclusively Oriental, and comprises _P.
gracilis_ from Borneo, Java, and (?) Sumatra, _P. pardicolor_ from Nipal,
and _P. maculosus_ from Tenasserim; the head and body of the latter
measuring from 18 to 20 inches in length. Speaking of _P. pardicolor_,
Mr. Hodgson observes that it is “equally at home on trees and on the
ground; it dwells and breeds in the hollows of decayed trees. It is not
gregarious at all, and preys chiefly upon small birds, which it is wont
to pounce upon from the cover of the grass. The times of breeding are
said to be February and August, and the litter to consist of two young,
there being two litters each year.”
_Poiana._[446]—This African genus, represented solely by one species,
_P. poënsis_ (Fig. 238), from Fernando Po, is very closely allied to
the preceding, but the spots are smaller, and show no tendency to run
into transverse bands or stripes, except in the region of the head and
shoulder; while the sole of the foot has a narrow bald band running up
towards the tarsus, as in _Genetta_. The length of the head and body is
38 inches, and that of the tail about 40 inches. It is probable that this
animal should really be regarded as a slightly aberrant species of the
genus _Prionodon_.
[Illustration: FIG. 238.—The African Linsang (_Poiana poënsis_). From
Mivart, _Proc. Zool. Soc._ 1882, p. 160.]
The five following genera differ in several important respects from all
the preceding, and collectively constitute the _Paradoxurine_ section
of Professor Mivart. With the exception of one African form, they are
mainly Oriental. In this section the auditory bulla is frequently in two
portions, the posterior moiety in one case being unossified, and it is
always much narrowed in front (Fig. 239). The palate (as in the figure)
may be much produced behind the molars; and the teeth are often but
slightly sectorial, and may be very small. The long tail is in most cases
not ringed.
_Paradoxurus._[447]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₂; total
40. The blunt and rounded form of the cusps of the hinder premolar and
the molar teeth distinguishes this genus from most of the members of the
family. Vertebræ: C 7, D 13, L 7, S 3, C 29-36. Head pointed in front.
Ears small, rounded. Body long. Limbs moderate. Palms and soles almost
entirely naked, and joining the foot-pads without the intervention of
any hairy space. Claws completely retractile. Pupil vertical. Tail long,
non-prehensile; in the Indian species without rings. The Paradoxures
or Palm-Civets are less strictly carnivorous than the other members of
the family. They are mostly about the size of the common Cat, or rather
larger, and are partly arboreal in their habits. The species are rather
numerous, and present considerable variations in the details of the
form and size of their molar teeth; in only a few does the bony palate
extend behind the molars. They are restricted geographically to Southern
Asia and the Indo-Malayan archipelago. The best known species[448] are
_P. niger_, _P. hermaphroditus_, _P. jerdoni_, _P. aureus_, _P. grayi_
from India and Burma, _P. philippinensis_ of the Philippines, _P.
larvatus_ of Southern China and Formosa, _P. leucomystax_ of the Malay
Peninsula, Sumatra, and Borneo, and _P. musschenbroeki_ of Celebes. The
name _Paradoxurus_ was applied from the mistaken notion that the tail
was prehensile. Mr. Blanford[449] gives the following account of the
habits of _P. niger_: “The common Palm-Civet, Tree-Cat, or Toddy-Cat,
is a familiar animal in most parts of India, though, being thoroughly
nocturnal in its habits, it is but rarely seen in the daytime. It is
arboreal, passing the day generally in trees, either coiled up in the
branches, or in a hole in the trunk, and in places where cocoa-nut
palms are common it frequently selects one of them for a residence.
Mango groves are also a favourite resort. It not unfrequently takes up
its abode in the thatched roofs of houses; Jerdon found a large colony
established in the rafters of his own house in Tellicheri. It even
occurs in large towns; I have known of one being caught in the middle of
Calcutta.”
[Illustration: FIG. 239.—Palatal aspect of the left side of the
cranium and mandible of _Arctogale leucotis_. _a_, Anterior opening of
alisphenoid canal; _o_, foramen ovale; _c_, carotid canal ¹⁄₁. (From
Mivart, _Proc. Zool Soc._ 1882, p. 165.)]
_Arctogale._[450]—This genus—represented only by _A. trivirgata_ of Java,
and _A. leucotis_ of Burma, Tenasserim, Sumatra, Java, etc.—is chiefly
distinguished from _Paradoxurus_ by the extremely small size of the
cheek-teeth (Fig. 239), which are often not in contact with one another;
the upper carnassial being almost triangular in shape. Palate frequently
convex longitudinally between the carnassials, and greatly produced
behind the last molar, with a very narrow bony aperture of the posterior
nares. The soles of the feet are still more naked than in _Paradoxurus_;
and the pollex and hallux are more divergent. In _A. leucotis_ the length
of the head and body is 26·5 inches, and the tail 27 inches. In many
specimens the three dorsal stripes are much less distinctly marked than
in others, and tend to break up into spots; while the general coloration
is considerably lighter.
_Hemigale_,[451] another modification of the Paradoxure type, contains
one species, _H. hardwickei_, from Borneo and Malacca, an elegant-looking
animal, smaller and more slender than the Paradoxures, of light gray
colour, with transverse broad dark bands across the back and loins;
the proximal portion of the tail being ringed. The tarsus is hairy. The
general cranial characters are those of _Paradoxurus_, but the auditory
bulla is ankylosed into a single piece.
_Arctictis._[452]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₂; total
40. The posterior upper molar and the first lower premolar very often
absent. Cheek-teeth generally small and rounded, with a distinct
interval between them, but formed generally on the same pattern as
_Paradoxurus_. Vertebræ: C 7, D 14, L 5, S 3, C 34. Body elongated. Head
broad behind, with a small pointed face. Whiskers long and numerous. Ears
small, rounded, but clothed with a pencil of long hairs. Eyes small.
Limbs short. Soles and palms broad, entirely naked. Tail very long and
prehensile; thickly covered with long hair. Fur long and harsh. Cæcum
extremely small. But one species is known, _A. binturong_, the Binturong,
an inhabitant of Southern Asia from Nipal through the Malay Peninsula to
the islands of Sumatra and Java. Although structurally agreeing closely
with the Paradoxures, its tufted ears, long, coarse, and dark hair, and
prehensile tail give it a very different external appearance. It may
be regarded as a very aberrant Paradoxure, connected, so far as dental
characters are concerned, with _Paradoxurus_ by means of _Arctogale_. The
bony palate also extends considerably behind the last molar, as in the
latter. The Binturong is slow and cautious in its movements, chiefly if
not entirely arboreal, and appears to feed on vegetable as well as animal
substances.
_Nandinia_[453] contains one species, _N. binotata_, a somewhat aberrant
Paradoxure, from West Africa. It is rather smaller than the true
Paradoxures, with smaller and more pointed molar teeth, and no cæcum. The
wall of the hinder chamber of the auditory bulla remains through life
unossified.
The dentition appears to be of a more decidedly carnivorous type than in
the other members of the section.
_Cynogale._[454]—This remarkable genus is regarded by Professor Mivart as
representing a third section of the _Viverrinæ_; it contains one species,
_C. bennetti_ (described by S. Müller under the name of _Potamophilus
barbatus_), from Borneo, Sumatra, and the Malay Peninsula. This is a
curious Otter-like modification of the Viverrine type, having semiaquatic
habits, both swimming in the water and climbing trees, living upon fish,
crustacea, small mammals, birds, and fruit. The number and general
arrangement of its teeth are as in _Paradoxurus_, but the premolars are
peculiarly elongated, compressed, pointed and recurved, somewhat as in
the Seals, though the molars are tuberculated. The head is elongated, the
muzzle broad and depressed. Whiskers very long and abundant. Ears small
and rounded. Toes short and slightly webbed at the base. Tail short,
cylindrical, covered with short hair. Fur very dense and soft, of a dark
brown colour, mixed with black and gray. Humerus without entepicondylar
foramen.
Subfamily =Herpestinæ=.—Auditory bulla very prominent, and somewhat
pear-shaped, the posterior chamber being large, rounded, and generally
with its greatest prominence to the outer side. The anterior chamber
considerably dilated, and produced into a short inferior wall to the
auditory meatus, in which is a depression or vacuity just below the
centre of the opening of the meatus. Sometimes this vacuity is continued
into the meatus, forming a narrow fissure. The paroccipital process
does not project beyond the bulla, but is spread out and lost (in adult
animals) on its posterior surface. Toes straight; claws lengthened,
exserted, non-retractile. No perineal glands. The dentition is always of
a markedly sectorial type; and the orbit may be surrounded by bone. Very
generally the anus opens into a sac-like depression. The majority of the
genera are Ethiopian; the type genus alone extending into the Oriental
and Palæarctic regions.
_Herpestes._[455]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, sometimes ³⁄₃,
_m_ ²⁄₂; total 40 or 36. Teeth of molar series generally with strongly
developed, sharply-pointed cusps. Skull elongated, constricted behind
the orbits. Face short and compressed. Frontal region broad and arched.
Postorbital processes of frontal and jugal bones well developed,
generally meeting so as to complete the circle of the orbit behind.
Vertebræ: C 7, D 13, L 7, S 3, C 21-26. Head pointed in front. Ears short
and rounded. Body very long and slender. Extremities short. Five toes on
each foot, the first, especially that on the hind foot, very short. Toes
free, or but slightly palmated. Palms generally naked. Distal portion of
soles naked, under surface of tarsus and metatarsus usually clothed with
hair, but considerable specific variation in this respect. Tail long or
moderate, generally thick at the base, and sometimes covered with more or
less elongated hair. The longer hairs covering the body and tail almost
always annulated. This genus contains a very large number of animals
commonly called Ichneumons, or in India Mungooses, varying in size from
that of a large Cat down to a Weasel. They are widely distributed over
the African continent and the southern parts of Asia, especially India
and the Indo-Malayan archipelago, one species occurring also in Spain.
They are mostly terrestrial in their habits, feeding on small mammals
and birds, reptiles, especially snakes, eggs of birds and reptiles, and
also insects. Some species are partially domesticated, being used to
keep houses clear of rats, mice, and snakes. _H. ichneumon_ was a sacred
animal to the ancient Egyptians. They vary considerably in appearance,
some, as _H. galera_ and _H. urva_ (Fig. 240), are larger and heavier,
with stouter body, longer limbs, and stronger teeth. The common Indian
Mungoose (_H. mungo_) is considerably smaller than the Egyptian form;
its fur is of a pale gray colour, the hairs being largely white ringed,
while the cheeks and throat are more or less reddish. Like the Egyptian
species, it is frequently domesticated, and put to a similar use. It is
especially serviceable in India as a serpent-killer, destroying not only
the eggs and young of these creatures, but attacking without hesitation
and killing the most venomous adult snakes. The fact that it invariably
survives those encounters has led to the belief that it either enjoys
immunity from the effects of snake-poison, or that after being bitten
it has recourse, as the natives maintain, to the root of a plant as an
antidote. Neither of these suppositions has stood the test of scientific
examination, for it has been found that when actually bitten it falls
a victim to the poison as rapidly as other mammals, while there is no
trustworthy evidence of its seeking a vegetable antidote. The truth
seems to be that the Mungoose, by its exceeding agility and quickness
of eye, avoids the fangs of the snake while fixing its own teeth in
the back of the reptile’s neck. One large species, believed to be from
Africa, recently described as _H. grandis_, is remarkable for the extreme
complexity of the cusps on the molars, and also for the absence of an
entepicondylar foramen to the humerus; the latter feature also occurring
in the allied _H. albicaudatus_. The Oriental _H. urva_ (Fig. 246) is
stated to be somewhat aquatic in habits, and to feed on frogs and crabs.
[Illustration: FIG. 240.—The Crab-eating Mungoose (_Herpestes urva_).
From Blanford, _Mammalia of British India_, p. 130.]
Remains of the small _H. nipalensis_ occur in the cavern-deposits of
Madras. Viverroids from the Miocene and Upper Eocene of Europe, which
agree with _Herpestes_ in the presence of an inner tubercle to the
third upper premolar and of a hinder cusp to the fourth lower premolar,
have been referred to the existing genus. The species which have been
separated generically under the three following names are very closely
allied to _Herpestes_.
_Helogale_,[456] premolars ³⁄₃, without diastema between first and
second; soles of feet completely naked. Contains two small South-African
species, _H. parvula_ and _H. undulata_.
_Bdeogale_[457] contains also two small Ichneumon-like animals, _B.
crassicauda_ and _puisa_, differing from _Herpestes_ proper in having
only four toes on each foot, both pollex and hallux being absent. The
orbit is nearly complete, the tail of moderate length and rather bushy.
_Cynictis._[458]—Pollex present, but hallux absent. Skull shorter and
broader than in _Herpestes_, rather contracted behind the orbits, which
are large and complete behind. Face short. Anterior chamber of the
auditory bulla very large. Front claws elongated. _C. penicillata_, from
South Africa. The cæcum (Fig. 241) of this genus is longer than in any
other member of the family.
All the foregoing Herpestines have the nose short, with its under surface
flat, bald, and with a median longitudinal groove. The remaining forms
have the nose more or less produced, with its under side convex, and a
space between the nostrils and the upper lip covered with close adpressed
hairs, and without any median groove.
[Illustration: FIG. 241.—Cæcum of _Cynictis penicillata_. (From Mivart,
_Proc. Zool. Soc._ 1882, p. 508.)]
_Rhinogale._[459]—Toes 5-5. Claws of fore feet short, compressed, acute.
Under surface of tarsus hairy. Palate flat. Founded on a single specimen
from East Africa, _R. melleri_.
_Crossarchus._[460]—Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ³⁄₃, _m_ ²⁄₂; total
36. Snout elongated. Toes 5-5. Claws on fore feet long and curved.
Hallux very short. Under surface of tarsus naked. Tail shorter than the
body, tapering. Palate flat. Fur harsh. Species: _C. obscurus_, the
Kusimanse, a small burrowing animal from West Africa, of uniform dark
brown colour; _C. fasciatus_; _C. zebra_; and _C. gambianus_.
_Suricata._[461]—A more distinct genus than any of the above. The dental
formula as in the last, but the teeth of the cheek-series remarkably
short in the antero-posterior direction, corresponding with the shortness
of the skull generally (Fig. 222). Orbits complete behind. Vertebræ: C
7, D 15, L 6, S 3, C 20. Though the head is short and broad, the nose is
pointed and rather produced and movable. Ears very short. Body shorter
and limbs longer than in _Herpestes_. Toes 4-4, the pollex and hallux
being absent. Claws on fore feet very long and narrow, arched, pointed,
and subequal. Hind feet with much shorter claws, soles hairy. Tail rather
shorter than the body. One species only is known, the Suricate, _S.
tetradactyla_, a small gray-brown animal, with dark transverse stripes on
the hinder part of the back, from South Africa. The cæcum is short.
[Illustration: FIG. 242.—Cæcum of _Galidea elegans_. (From Mivart, _Proc.
Zool. Soc._ 1882, p. 508.)]
_Galidictis_,[462] _Galidea_,[463] and _Hemigalidea_[464] are names
of three slight generic modifications of the Viverrine type, allied
to the _Herpestinæ_, but placed by Mivart in a distinct subfamily,
_Galidictiinæ_. They are all characterised by the absence of the
alisphenoid canal in the skull, as well as of the entepicondylar foramen
to the humerus; and are inhabitants of Madagascar. The best known,
_Galidea elegans_, is a lively Squirrel-like little animal with soft fur
and a long bushy tail, which climbs and jumps with agility. It is of a
chestnut-brown colour, the tail being annulated with darker brown. The
cæcum (Fig. 242) is remarkable for its comparative length and pointed
termination. _Hemigalidea_ is distinguished by the absence of rings on
the tail. _Galidictis vittata_ and _striata_ chiefly differ from the
Ichneumons in their coloration, being gray with parallel longitudinal
stripes of dark brown.
_Eupleres_[465] is another form, also from Madagascar, which has been
placed in a subfamily apart. It differs remarkably from all the other
_Viverridæ_ in the weak development of the jaws and the small size of the
teeth (Fig. 243), in consequence of which it was, when first discovered,
placed in the order Insectivora. Dentition: _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄,
_m_ ²⁄₂; total 40. Vertebræ: C 7, D 13, L 7, S 3, C 20. No alisphenoid
canal; an entepicondylar foramen to the humerus. But one species is
known, _E. goudoti_.
[Illustration: FIG. 243.—Skull of _Eupleres goudoti_. ⅘ natural size.
Mus. Roy. Coll. Surgeons.]
_Extinct Genera._—The Tertiaries of the Old World have yielded
several genera allied to the existing Viverroids, some of which show
decided signs of affinity with other families. Of these the Lower
Miocene _Amphictis_ appears to be nearly related to _Viverra_, but is
distinguished by the form of the second lower molar, which is longer and
has two distinct roots. _Palæoprionodon_, of the French Phosphorites, has
a dentition very like that of _Prionodon_, the molars being reduced to
¹⁄₂; the skull has an alisphenoid canal and the general basal characters
of the _Viverridæ_, but resembles the _Mustelidæ_ in the presence of
a glenoid foramen and in the position of the condylar foramen. In
_Stenoplesictis_, of the same deposits, the dental formula is _i_ ³⁄₃,
_c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₂; and although the skull has a complete septum
in the bulla, yet some of the cranial and dental features approximate
so decidedly towards those of the extinct _Mustelidæ_, as to lead
some authorities to refer the genus to that family. The most probable
explanation of this resemblance is that the Musteloids have originated
from generalised Viverroids allied to _Stenoplesictis_. The Lower
Pliocene _Ictitherium_ differs from all other Viverroids in the presence
of three distinct lobes to the upper carnassial, and thereby connects
the other members of the family so closely with the _Hyænidæ_ that it is
practically impossible to draw up a definition which will distinguish the
two families.
The North American Eocene genera _Miacis_ and _Didymictis_ are generally
regarded as representing a separate family—_Miacidæ_—with affinities both
to the _Viverridæ_ and _Canidæ_.
_Family_ PROTELEIDÆ.
Skull with no alisphenoid canal; and the auditory bulla divided into two
distinct chambers. Dorsal vertebræ 15. Molars ¹⁄₁. Premolar and molar
teeth very small and simple in character.
_Proteles._[466]—This genus contains but a single species, _P.
cristatus_, the Aard-Wolf or Earth-Wolf of the Dutch colonists of the
Cape, an animal nearly allied to the Hyænas, but remarkably modified
in its dentition, the molar teeth being very small, placed far apart,
and almost rudimentary in character (Fig. 244). The canines are long
and rather slender. The dental formula is _i_ ³⁄₃, _c_ ¹⁄₁, _p_ and _m_
⁴⁄₃₋₄; total 30 or 32. Vertebræ: C 7, D 15, L 5, S 2, C 24. The fore feet
with five toes; the pollex though short, with a distinct claw. The hind
feet with four subequal toes. Claws all strong, blunt, subcompressed, and
non-retractile. The general external appearance is very like that of a
small Striped Hyæna, but the muzzle is more pointed and the ears larger.
It has a copious mane of long hair, capable of being erected when the
animal is excited, along the middle line of the neck and back. It is a
native of South Africa, and is a burrowing nocturnal animal, feeding on
decomposing animal substances, larvæ, and termites. Observations upon
specimens in captivity indicate that it has neither inclination nor power
to attack or feed upon living vertebrated animals.
[Illustration: FIG. 244.—Skull and Dentition of the Aard-Wolf (_Proteles
cristatus_). ½ natural size.]
Some writers regard _Proteles_ as representing a subfamily of the
_Hyænidæ_.[467]
_Family_ HYÆNIDÆ.
Skull with no alisphenoid canal; and the auditory bulla not divided by a
septum into two chambers. Dorsal vertebræ 15. Molars usually ¹⁄₁, but in
some fossil forms ¹⁄₂, or ²⁄₂, the second lower molar being very small;
upper carnassial with three distinct lobes; lower carnassial with a
large blade and small talon. No entepicondylar foramen to the humerus.
This family is confined to the Old World, where it is now represented
by a single genus, which, although evidently nearly related to the
_Viverridæ_, is sufficiently distinct to be regarded as not referable to
that family. The extinct _Ictitherium_, however, as already mentioned,
connects the more generalised members of the _Hyænidæ_ very closely with
the _Viverridæ_.
_Hyæna._[468]—Dentition in existing forms usually _i_ ³⁄₃, _c_ ¹⁄₁,
_p_ ⁴⁄₃, _m_ ¹⁄₁; total 34. Teeth, especially canines and premolars,
very large, strong, and conical. Upper carnassial (Fig. 245) with a
very large, distinctly trilobed blade and a moderately developed inner
tubercle placed at the anterior extremity of the blade. Molar very
small, and placed transversely close to the hinder edge of the last, as
in the _Felidæ_. Lower carnassial consisting of little more than the
bilobed blade. Zygomatic arches of cranium very wide and strong. Sagittal
crest high, giving attachment to very powerful biting muscles. Orbits
incomplete behind. Vertebræ: C 7, D 15, L 5, S 4, C 19. Limbs rather
long, especially the anterior pair, digitigrade, four subequal toes on
each, with stout non-retractile claws. Pollex and hallux only represented
by rudimentary metacarpal and metatarsal bones. Tail rather short. A
large post-anal median glandular pouch, into which the largely developed
anal scent glands pour their secretion.
[Illustration: FIG. 245.—Outer (_A_) and palatal (_B_) aspects of the
right upper carnassial tooth of the Striped Hyæna (_Hyæna striata_). From
the _Quart. Journ. Geol. Soc._]
The three existing species of Hyæna are divisible into two sections, to
which some zoologists assign generic rank, but fossil forms show such
a transition between these two types as to render any such division
impracticable.
The typical or _Euhyænine_ group presents the following distinctive
features. Upper molar moderately developed and three-rooted. An inner
cusp and hind talon more or less developed on the lower molar. Ears
large, pointed. Hair long, forming a mane on the back and shoulders. _H.
striata_, the Striped Hyæna (Fig. 246) of Northern Africa and Southern
Asia. _H. brunnea_, of South Africa, in some respects intermediate
between this and the next group.
The Striped Hyæna is dirty gray in colour, with narrow transverse
tawny or blackish stripes on the body and legs; the length of the
head and body is 3½ feet, and that of the tail, with its hair, 1½
feet. It occurs throughout peninsular India, where it is most common
in open hilly districts, and in North Africa. Mr. Blanford[469] gives
the following account of its habits: “It is a nocturnal animal, and
although an occasional individual may be met with returning to its den
in the early morning, its rambles are usually commenced after sunset
and ended before sunrise. During the night it roams far and wide, and
no tracks of wild animals are more common in the countries where it is
found than its unmistakable footprints, very like a dog’s in shape, but
with the marks of the hind feet conspicuously smaller than those of the
fore feet. Unlike the Spotted Hyæna, the Striped species appears to
be solitary in its habits, and it is rare to meet with more than two
together. The principal food of the Hyæna consists of the carcases of
animals that have died of disease or been killed by beasts of prey, and
very often it carries off portions of the body to its den. I once shot
one that was carrying away the hind leg of a Nilghai. The powerful jaws
and large teeth are admirably adapted for crushing bones, which are
consumed by Hyænas, after the flesh has been picked off by vultures and
jackals. Occasionally sheep or goats, and more often dogs, are carried
off by Hyænas, and the latter at all events are often taken alive to the
animal’s den.” The Striped Hyæna is essentially a cowardly animal, and
one that is much more silent than _H. crocuta_. Remains of _H. striata_
are found in the cavern-deposits of the south of France, and also in the
Upper Pliocene of the Val d’Arno in Tuscany, and in the English Red Crag.
[Illustration: FIG. 246.—The Striped Hyæna (_Hyæna striata_).]
The _Crocutine_ group presents the following characters. Upper molar
extremely small, two- or one-rooted, often deciduous. Lower molar
without trace of inner cusp, and with an extremely small talon. Ears
moderate, rounded. Hair not elongated to form a mane. _H. crocuta_, the
Spotted Hyæna (Fig. 247), from Africa south of the Sahara. In dental
characters as well as in its visceral anatomy, especially as regards the
reproductive organs of the female,[470] this species may be considered
as by far the more specialised form. The Spotted Hyæna is a larger and
bolder animal than the Striped species, hunting in packs, and uttering
very frequently its unearthly cry. The coloration consists of dark brown
spots on a yellowish ground. It was formerly very common at the Cape.
Remains of a large race of this species are exceedingly common in the
cavern-deposits of Europe, where they were first described under the
name of _Hyæna spelæa_; teeth have also been met with in the Norfolk
Forest-bed, and in cavern-deposits in Madras—the latter locality being
exceedingly interesting from a distributional point of view.
[Illustration: FIG. 247.—The Spotted Hyæna (_Hyæna crocuta_).]
In addition to the remains of existing species, to which reference has
been already made, there were numerous extinct forms of _Hyæna_ in the
upper Tertiaries of Europe, from the horizon of the Lower Pliocene
Pikermi beds of Greece upwards. In the Crocutine group _H. colvini_ of
the Pliocene of India (Fig. 248), and _H. robusta_ of that of Italy,
appear to have been ancestral forms allied to _H. crocuta_; the former
being distinguished by the loss of the first upper premolar. _H. eximia_,
of the Pikermi beds, is a more generalised form, in which the first lower
premolar (lost in existing forms) is retained. In the typical group, _H.
arvernensis_ and _H. perrieri_, of the Upper Pliocene of the Continent,
approximate to _H. brunnea_; although _H. perrieri_ makes a farther step
towards the Crocutine group by the loss of the inner cusp in the lower
carnassial. The extinct _Hyænictine_ group, as represented by the Indian
_H. sivalensis_ and the Grecian _H. græca_, connects _H. striata_ with
_Palhyæna_. Both are characterised by the presence of a small second
lower molar behind the carnassial; while _H. græca_ also has four lower
premolars. Still more generalised is the _Lychyænine_ group; comprising
_H. macrostoma_ of India and _H. chæretis_ of the Pikermi beds; in these
forms the muzzle was longer, and the premolars much more compressed than
in the existing species, thus making a very decided approach to the
_Viverridæ_. There were four lower premolars; the lower carnassial had
an inner cusp, and it is probable that there was a second lower molar;
while the first upper molar was placed partially behind the carnassial.
The Lower Pliocene _Palhyæna hipparionum_, in which the dental formula
is _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ²⁄₂, is a smaller form with long jaws
and compressed premolars which approaches so closely to the Viverroid
genus _Ictitherium_ as to show pretty clearly how the Hyænas have been
gradually modified from that stock.
[Illustration: FIG. 248.—Outer view of part of the right ramus of the
mandible of _Hyæna colvini_, showing the third and fourth premolars and
the carnassial. (From the _Palæontologia Indica_.)]
_Section_ CYNOIDEA.
_Family_ CANIDÆ.
This section contains the single family of the _Canidæ_, or Dog-like
animals, which appear to hold an intermediate position between the other
two sections, retaining also many of the more generalised characters of
the ancient members of the order. The structure of the auditory bulla and
adjacent parts of the bones of the skull is intermediate between that
of the Æluroid and Arctoid forms. In the number and arrangement of the
teeth they more nearly approach the primitive heterodont type than any
other existing Carnivora. A cæcum is always present, sometimes short and
simple, but when long it is folded upon itself in a characteristic manner.
[Illustration: FIG. 249.—Right lateral aspect of the skull of the Dog
(_Canis familiaris_).]
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.
[Illustration: FIG. 250.—Cæcum of the Arctic Fox (_Canis lagopus_). _i_,
Ileum; _c_, colon. In the natural position the colon is uppermost.]
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.
[Illustration: FIG. 251.—The last four left upper teeth of an extinct
Wolf (_Canis cautleyi_). From the _Palæontologia Indica_.]
_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.