in the more ancient forms are unlike the molars, but like them in more
recent forms. The molars of the upper jaw have two crests parallel and
united by an outer crest. The fore-feet have four, the hind-feet three
toes.
The family is fully as ancient as that of the Equidae, but the
specialisation of the toes never advances so far. The modern
representatives of the order are, so far as the feet are concerned, in the
condition of very early representatives of the equine stock. Nor do the
teeth of the Tapirs ever reach the complicated pattern of that presented by
at least the modern Horses, or indeed of the Palaeotheres. Apart from this
it is not an easy matter to distinguish accurately between these several
families, including the Lophiodontidae, which, as already mentioned, is
placed nearer to the Tapiridae than to the Palaeotheriidae. Indeed the
differentiation of these two families, the Tapiridae and the
Lophiodontidae, seems to be a matter of the greatest difficulty. The
difficulty is well emphasised by the fact that naturalists disagree most
profoundly as to the relations of various genera of extinct Tapir-like
animals. For Mr. Lydekker the genus _Lophiodon_ includes also the American
genera _Isectolophus_ and _Systemodon_, which are placed by Zittel in the
sub-family Tapirinae as opposed to Lophiodontinae, which contains
_Lophiodon_ and _Helaletes_. The existing Tapirs can be differentiated from
the existing Horses with great ease, as the following account of the
existing genera will show.
{251} [Illustration]
FIG. 128.--American Tapir. _Tapirus terrestris._ x 1/10.
The genus _Tapirus_ is now met with only in South and Central America, and
in the Malay Peninsula and the islands of Java and Sumatra. This animal is
in many respects the most ancient of existing forms referable to the
Perissodactyle order. It has four toes on the front-feet, though only three
on the hind-feet. The number of teeth is 42--nearly the typical Eutherian
number. The Tapirs are always moderately-sized animals, entirely covered
with hair, and usually of a brownish-black colour. The Malayan Tapir is,
however, banded broadly with white--a single band; the young of the Tapir
is spotted, and striped with white. The nose and upper lip conjoined are
produced into a short trunk, precisely comparable with that of the
Elephant. As in the Rhinoceros--and in this both contrast with the other
existing Perissodactyle genus _Equus_--the temporal fossa is not separated
from the orbit by bone. Of existing Tapirs there are at any rate _T.
terrestris_,[163] _T. roulini_ (the "Tapir Pinchaque" of Cuvier), _T. dowi_
and _T. bairdi_ in America (the last two being sometimes separated into a
distinct genus, _Elasmognathus_, on account of the prolongation of the
ossified mesethmoid), and _T. indicus_ in the East. The tapir, probably _T.
terrestris_, is described by Buffon as "a dull and gloomy animal." It is
certainly mainly nocturnal in habit. The name _terrestris_ was given by
Linnaeus, who placed it in the same genus as {252} _Hippopotamus
amphibius_; hence the epithet applied to the Tapir. But as a matter of fact
it loves marshy neighbourhoods, and is in a way amphibious. This does not
of course apply to the Andesian _T. roulini_, which inhabits the cordillera
of Ecuador and Colombia. The distribution of existing Tapirs is, as is so
often the case, restricted when compared with that of their extinct
congeners and allies. In Europe the remains of the genus _Tapirus_ are
abundant from Pliocene strata, and its remains are there known from as far
back as the Miocene. The genus is thus one of the very oldest forms of
Mammalia at present inhabiting the earth.
[Illustration]
FIG. 129.--Malayan Tapir. _Tapirus indicus_, young. x 1/10. (From
_Nature_.)
The Malayan Tapir is to be distinguished from the American (_T.
terrestris_--the other species have not been dissected) by the greater
development of the valvulae conniventes in the intestine, the absence of a
moderator band in the heart, and the less elongated caecum, which is
sacculated by only three bands, there being four in _T. terrestris_.[164]
The animal frequents the most retired spots among the hill woods, by which
habit it seems {253} largely to escape the Tiger, its most formidable foe
in those regions of the world. Its quickness of senses enables it also to
slip away with rapidity. It can proceed at a great pace when disturbed, and
can readily push its way through obstacles. The young animal, like that of
the American species, is dark brown with yellowish spots. It is stated by
Mr. H. N. Ridley that the young animal lies during the hot part of the day
under bushes, in which situation "its coat is so exactly like a patch of
ground flecked with sunlight that it is quite invisible." It is interesting
to note that here, as with some other animals, it is the young that are
especially protected by such mechanisms. Moreover, some of the spots are
round and some are more elongated, so that the resemblance to spots of
sunlight which come in a direct and in a slanting direction is greatly
increased. Even the colours of the adult are not so conspicuous when it is
in its native haunts as might be supposed. The breaking up of the ground
colour into tracts of two different colours prevent it from striking the
eye so plainly as if it were of one colour throughout. "When lying down
during the day it exactly resembles a grey boulder, and as it often lives
near the rocky streams of the hill jungles, it is really nearly as
invisible then as it was when it was speckled."[165]
FAM. 3. RHINOCEROTIDAE.--This family is to be distinguished from the
preceding by a number of characters, which though not universal are
general. In the first place, there are commonly horns, or a horn,
consisting of what appears to be an agglomeration of hair-like structures
fixed upon a roughened patch of bone on the surface of the nasals. The
incisors are diminished or defective, and the upper canines are often
wanting. The molars and premolars are alike. The fore-feet are four- or
three-toed, but are functionally tridactyle; the hind-feet are three-toed.
The skeleton in this family is massive, and the limbs relatively short. The
skull, as in the Tapirs, has a confluent orbit and temporal fossa. The
upper lip is generally more or less prehensile; the body is as a rule--to
which the Pleistocene Hairy Rhinoceros is of course an exception--rather
sparsely covered with hair. In this feature the Rhinocerotidae contrast
both with the Tapiridae and the Equidae. The family in reality contains but
one existing genus, though three have been instituted, viz. {254}
_Rhinoceros_, _Ceratorhinus_, and _Atelodus_. As there are so few existing
species the subdivision of animals which agree in so many and such
highly-characteristic features seems to be an unnecessary procedure. The
existing Rhinoceroses are but a fragment of the total number of known forms
from past epochs. The family is very markedly on the wane.
The genus _Rhinoceros_ is characterised by its heavy build and thick,
almost smooth, skin--smooth, that is to say, so far as concerns the slight
development of hair--which is often thrown into folds. There is one or
there are two horns on the fore-part of the head, which are, as has already
been pointed out, structures _sui generis_, and not exactly comparable with
the horns of other living Ungulates. There are three nearly equal toes on
both fore- and hind-limbs. The canine teeth of existing species have
disappeared; the incisors are, or are not, present; the molars and
premolars are three and four in each half of each jaw.
The visceral anatomy of the Rhinoceros has been much investigated so far as
concerns the Asiatic forms. A curious feature, which serves to discriminate
some of the Asiatic species from others, is to be seen in the small
intestine. In _Rh. indicus_[166] this gut is furnished with numerous long
cylindrical narrow outgrowths "like tags of worsted"; in the allied _Rh.
sondaicus_ these tags are present, but are flatter and broader; while in
the two-horned _Rh. sumatrensis_ there are no tags at all, but only smooth
valve-like folds. Another mark by which these species can be distinguished
depends upon the variation in the presence or absence of certain glands
imbedded in the integument of the foot--the so-called "hoof glands." These
occur in _Rh. indicus_ and _Rh. sondaicus_, but are absent in _Rh.
sumatrensis_.
Sir W. Flower[167] studied some years since the skull features which serve
to differentiate the existing forms.
In _Rh. sumatrensis_ the two long downward processes of the squamosal bone,
termed respectively post-glenoid and post-tympanic, do not unite below the
auditory meatus. In this the species in question agrees with the African
forms but not with the one-horned Asiatic species, where the two processes
completely fuse. Again, another character, though perhaps less important,
{255} is the sloping backwards instead of forward of the occipital crest in
all two-horned species, whether African or Asiatic.
The Asiatic Rhinoceroses have, what the African animals have not,
functional incisor teeth throughout life. It has been proposed on these and
other grounds to separate generically the African and Asiatic forms.
[Illustration]
FIG. 130.--Indian Rhinoceros. _Rhinoceros indicus._ x 1/40.
The Asiatic Rhinoceroses include three well-differentiated species, in all
of which the skin is much thrown into folds. _Rh. indicus_ is the largest
form. It is one horned, and has enormous folds of skin at the neck and
hanging over the limbs. So like artificial armour is this thick plating,
that Albrecht Duerer may be excused for having given the beast the
appearance of being actually mail-plated in a sketch which he made of a
specimen sent over to the King of Portugal in 1513. This particular beast,
one of if not the first sent over to Europe, proved so intractable in
disposition that the king sent it as a present to the Pope. But "in an
access of fury it sunk the vessel on its passage"! The horn of this and of
other species was held until almost our times to have medicinal and other
more curious values. So recently as 1763 it was gravely asserted that a cup
made of its horn would fall to pieces if poison were poured into it. "When
wine is poured therein," wrote Dr. Brookes in the year referred to, "it
will rise, ferment, and seem to boil; but when {256} mixed with poison it
cleaves in two, which experiment has been seen by thousands of people."
John Evelyn also wrote of a well in Italy which was kept sweet by a
Rhinoceros' horn. This species seems to be long-lived, even in captivity; a
specimen now to be seen in the Zoological Society's Gardens has been there
since the year 1864.
_Rhinoceros sondaicus_, the Rhinoceros of the Sunderbunds, has a much wider
range than the last species or Indian Rhinoceros. This is unknown out of
India itself, and is there limited to a small region; the Sondaic form is
found in Bengal and in the Malayan Islands. It is a smaller species, and
the armour has a tesselated appearance. The female generally, if not
always, is hornless.
[Illustration]
FIG. 131.--Sumatran Rhinoceros. _Rhinoceros sumatrensis._ x 1/15. (From
_Nature_.)
The Sumatran species, _Rhinoceros sumatrensis_, is to be distinguished from
the last two by its two horns. It is also covered {257} by a much thicker
coat of hairs, which are sometimes blacker and sometimes redder. On account
of its two horns it has been proposed to separate it from the other
Oriental species into a distinct genus, _Ceratorhinus_. The animal has much
the same range as the last species, but extends to Borneo. A variety of
this species with hairy ears, from Assam, has been separated as a distinct
form, under the name of _Rh. lasiotis_, by Mr. Sclater. The animal upon
which that species was founded was until quite recently living in the
Zoological Society's Gardens.
[Illustration]
FIG. 132.--Hairy-eared Rhinoceros. _Rhinoceros lasiotis._ x 1/30.
There are only two certainly-known species of Rhinoceros in Africa. These
are the White Rhinoceros (_Rh. simus_) and the Black Rhinoceros (_Rh.
bicornis_). The origin of the names is not easy to understand, since the
"white" animal is, if anything, darker in colour than the Black Rhinoceros.
It is stated, however, that in past years the specimens of _Rh. simus_
found in the south-west of Cape Colony were "paler and whiter in colour
than those in the north-east." At present there are no grounds for
distinguishing the species by their colour characters. But they are plainly
distinguishable on other grounds. _Rhinoceros simus_ has a square upper
lip, and in relation to this crops the herbage upon the ground. _Rh.
bicornis_ has a prehensile upper lip projecting beyond the lower, and in a
corresponding fashion feeds principally upon the branches of shrubs, It has
been pointed out by Mr. {258} Coryndon[168] that the calf of _Rh. simus_
"always runs in front of the cow, while the calf of _Rh. bicornis_
invariably follows its mother." Both animals of course have two horns, and
upon the varying proportions of the horns a large number of "species" have
been made in the past. It is stated that the longest horn of the "White
Rhinoceros" known measures 56-1/2 inches; while that of _R. bicornis_ is
shorter, 40 inches being apparently the maximum. But the animal is smaller.
[Illustration]
FIG. 133.--Head of _Rhinoceros bicornis_.
The possible third African species of _Rhinoceros_[169] has been
provisionally named after Mr. Holmwood, and is based upon two horns 41 and
42 inches long, which may be abnormal horns of _Rh. bicornis_; but they are
thinner and have a smaller pedicel.
EXTINCT RHINOCEROTIDAE.--The existing Rhinoceroses are thus confined to
Africa, to certain parts of the continent of Asia, and to some of the large
islands lying to the south of that continent. But formerly the genus, and
allied genera, had a wider range. As far back as the Miocene we meet with
remains of Rhinoceroses closely allied to existing forms. The more ancient
forms have, as is natural, more ancient characters. Thus in _Rh.
schleiermacheri_ of the Miocene, canines appear to have been present. The
Miocene _Aceratherium_, primitive in the absence of horns as its {259} name
denotes,[170] had also canines and, in one species, six incisors in the
lower jaw. This _Aceratherium_ had, moreover, four toes in the fore-feet.
In the Miocene and later the Rhinoceros existed in Europe and America.
There was even a purely northern form, the _Rh. tichorhinus_, which
possessed a woolly covering and had the same range as the Mammoth. This
Rhinoceros was two-horned.
The post-Pliocene and European _Elasmotherium_ was a colossal rhinocerotine
creature. This great beast had two horns and a body 15 feet long. Its limbs
are not known, and as the teeth are different from those of Rhinoceroses in
general, it may not have belonged to this group at all, though Osborn is
inclined to derive it from _Aceratherium_, admitting at the same time that
the evidence is "decidedly slender." The teeth in fact are like those of a
Horse in being hypselodont and prismatic in form. As to the two horns, they
were apparently not exactly like those of typical Rhinoceroses; there was
an enormous horn posteriorly, supported on a huge boss of bone, and in
front of this a roughened spot suggests a smaller or at least a much more
slender horn.
It is important to notice that fossil Rhinoceroses belonging to the
restricted genus _Rhinoceros_ were in Europe invariably two-horned; it is
only in India, where they still exist, that one-horned forms are met with
in a fossil state.
[Illustration]
FIG. 134.--Skeleton of _Hyracodon nebrascensis_. x 1/12 (After Scott.)
The Rhinoceroses of America were mostly hornless. _Diceratherium_ is an
exception; but in many cases it had two parallel not successive horns, and
these were, to judge from the slight prominences, but feeble in
development, and perhaps hardly exactly comparable with the formidable
weapons of the Old-World forms. _Aceratherium tridactylum_, with
indications of paired horns, may be ancestral to _Diceratherium_. The
American forms have weak and slender nasals in correspondence with the
absence of horns; the sagittal crest is retained in contradistinction to
the great flattened surface of the skull in the horned Rhinoceroses.
_Aceratherium_ of both divisions of the globe probably represents the
ancestral group of the horned and the hornless forms. This being the case
it is highly interesting to note a distinct convergence in the quite {260}
separate American genera towards the European horned genera. A genus
sometimes united with _Aceratherium_, but still differing from it in some
points, is _Aphelops_ (_Teleoceras_).[171] This animal is more nearly
approximated to "the modern standard" of Rhinoceroses than is its possible
ancestor _Aceratherium_. The skeleton in general is more robust, even
surpassing that of modern forms, and approaching the _Hippopotamus_. There
is a reduction in the upper incisors, which are limited to two pairs, and
the lower molars {262} are reduced to five. The lower incisors are only
two. The sagittal crest is less marked; the fifth digit is reduced to a
tiny nodule representing the metacarpus. It had a small nasal horn. There
are numerous other details of likeness to modern Rhinoceroses in this
creature, which has only community of descent with them from the older
hornless forms, such as _Aceratherium_ and _Caenopus_. In the genus
_Peraceras_ the upper incisors are as completely gone as in the living
African Rhinoceroses.
[Illustration]
FIG. 135.--Skeleton of _Aphelops (Teleoceras) fossiger_. x 1/15. (After
Osborn.)
The most ancient rhinocerotine types[172] are the Hyracodonts and the
Amynodonts. They both date from the Eocene, and became extinct in the
succeeding Oligocene. _Hyracodon_[173] (Fig. 134) was "an agile,
light-chested, and rather long-necked" type, resembling a Horse in build.
There were no horns present, but the hoofs were more like those of the
Horses than of the existing Rhinoceroses. These animals were apparently
plain dwellers and defenceless, which is held to account for their compact
hoofs and outward similarity to a Horse. The genus is Oligocene. The dental
formula is I 3/3 C 1/1 Pm 4/3 M 3/3.
It is surmised by Professor Scott that the number of dorso-lumbar vertebrae
was twenty-three or twenty-four. The radius and ulna are complete and
separate bones, but the latter is somewhat reduced. There are four
metacarpal bones, of which, however, the fifth is much reduced. The animal
is only three-fingered. The tibia and the fibula are distinct, and show no
tendencies towards fusion; but the fibula is much reduced. There are only
three metatarsals and three toes. Had this line, which is to be regarded as
a side branch of the Rhinoceros stem, not died out, it would probably have
resulted, thinks Professor Scott, in monodactyle--very Horse-like types. It
is later than the next genus to be described, _Hyrachyus_, of which it is
possibly a descendant. An intermediate type, _Triplopus_, appears to bind
together _Hyracodon_ and _Hyrachyus_.
In _Hyrachyus agrarius_ the skull is long and narrow, the facial region
being markedly longer than in existing Rhinoceroses. The mastoid portion of
the periotic bone is widely exposed upon the outer face of the skull, which
is, as has been said, not the case with the existing genus _Rhinoceros_.
The dentition is the complete Eutherian dentition of forty-four teeth. The
upper {263} molar teeth are strikingly like those of the genus
_Rhinoceros_. The fore-feet are pentadactyle, but functionally
tetradactyle; the hind-feet tridactyle. The ulna is less reduced than in
_Hyracodon_, and the dorso-lumbar vertebrae are twenty-five.
[Illustration]
FIG. 136.--Skeleton of _Metamynodon planifrons_. x 1/22. (After Osborn and
Wortman.)
The Amynodonts were short, heavy types, probably marsh-haunting in habit,
and possibly with a proboscis like that of the Tapir. The orbit is higher
than it is in the purely terrestrial {264} Hyracodonts, and it is suggested
that when swimming it was raised above the surface as with the
Hippopotamus. "This feature," observes Professor Osborn, "with the long
curved tusks, undoubtedly used in uprooting, suggests the resemblance
between the habits of these animals and those of the hippopotami." There
were no horns in the Amynodonts. The face is shorter than in the
Hyracodonts, and the mastoid is covered as in recent Rhinoceroses. The
canines are very strongly developed into tusks, but the incisors show signs
of disappearance. We know of the genera _Amynodon_, _Metamynodon_, and
_Cadurcotherium_. All except the last, which is European, are American in
range.
FAM. 4. TITANOTHERIIDAE.--These Oligocene Ungulates, often attaining to
large dimensions, are nearly peculiar, so far as is at present known, to
the North American Continent, and are at least most abundant in it.[174]
Many generic names, such as _Titanotherium_, _Brontotherium_, _Brontops_,
_Titanops_, and _Menodus_, have been given to them; but a recent study of
the entire material accessible for description or already described has led
Professor Osborn to the opinion that there was but a single genus, to which
the name _Titanotherium_ must be applied. Of this genus there are some
thirty well-characterised species, of which the gradual evolution can be
traced from the lowest strata of the White River beds where their remains
occur. An entire skeleton of _T. robustum_ enables us to understand the
osteology of these forms and to compare them with other Perissodactyles.
This animal was more than 13 feet long, standing some 7 feet 7 inches in
height. It seems to have presented during life the aspect of a Rhinoceros
with perhaps a touch of Elephant. The skull is not unlike that of a
Rhinoceros in general dimensions and shape; but there are a pair of
apparent horn cores anteriorly, which are smaller in the more ancient forms
and acquire a large size, a forward direction with a divergence of the two
in the later forms. A glance at the accompanying figures of skulls (Fig.
137) of early and later Titanotheres will exhibit the changes in this
particular which the skulls underwent in the lapse of time occupied by the
deposition of these Oligocene beds. The nasals are short in the later,
longer in the more early species, such as _T. heloceras_ and _T.
coloradense_. The zygomatic arch projects much, and is "shelf-like" in the
later forms, the skull thus getting an immense breadth, which, {266}
together with the long and divergent horn cores, must have given to the
living animal a most bizarre appearance. It is an interesting fact that
this animal, though a Perissodactyle, agrees with the Artiodactyla in the
nineteen dorso-lumbar vertebrae, of which seventeen bear ribs.
[Illustration]
FIG. 137.--Three figures showing the cranial evolution of _Titanotherium_.
Upper figure, _T. trigonoceras_; middle figure, _T. elatum_; lower figure,
_T. platyceras_. (After Osborn.)
The genus further agrees with the Artiodactyles in the structure of the
carpus. The toes of the fore-limb are four, those of the hind-limb three;
but while the hind-limb is undoubtedly Perissodactyle in the arrangement of
its component parts, the fore-limb shows a hint of an Artiodactyle mode of
structure. This limb is paraxonic, the axis of the limb passing between the
two middle digits. It may be that this genus represents more nearly than
any other Perissodactyle or Artiodactyle the primitive stem from which both
have diverged, though, of course, it is not old enough to be very near to
the actual ancestor. The molar dentition is the typical one; the incisors
seem to vary as to their presence or absence, and, if present, in their
numbers. In comparing the older with the more recent forms it is noteworthy
that there has been an increase of size exactly as there has been during
the evolution of the Camels and some other groups of Ungulates. As already
mentioned, the size of the horn cores also increases until it culminates in
the extraordinary species, _T. platyceras_ and _T. ramosum_, in which these
are half as long as the skull, flattened in form, and connected at their
bases by a "web" of bone. Arrived at this amount of specialisation the
genus _Titanotherium_ apparently exhausted its capacities for modification
and ceased to be. The many generic names may be explained by sexual
differences on the one hand and an incomplete knowledge of connecting links
on the other.[175]
_Palaeosyops_ is somewhat like a Tapir in build, the skull especially
resembling that of the Tapir. As in _Titanotherium_ the molar teeth,
instead of having an outer wall formed by fused cusps, have a [176]-shaped
outer wall on one side and two or one cusps on the opposite side. It is,
moreover, an Eocene form, and in correspondence with its greater age is
more primitive in some points of structure, for example, in the absence of
horns and in the full dental formula. The fore-limbs are four-toed, the
hind {267} three-toed. It was intermediate between a Tapir and a Rhinoceros
in size. It has been shown, too, from casts of the interior of the skull,
that the cerebral hemispheres are much less convoluted than were those of
_Titanotherium_.
Related to _Palaeosyops_ is another primitive Titanothere, the genus
_Telmatotherium_. This is also Eocene, from the Uinta Basin, the uppermost
of Eocene strata. The skull of these creatures was rather elongated, and
not unlike that of a Titanothere in general aspect. The dentition was
complete and the canines not very large. The horns, which acquire so
prodigious a development in the later Titanotheres, are just recognisable
in at any rate many species of this genus _Telmatotherium_, the name being
thus by no means an apt one. Better was that proposed by Dr. Wortman, of
_Manteoceras_ or "prophet horned." The horns are small elevations upon the
frontals just at the junction of these with the nasals, and, indeed, lying
partly upon the latter bones. In _T. cornutum_ the horns are chiefly borne
upon the very long nasals, whose size contrasts with the same bones in the
more highly-developed _Titanotherium_. It appears to be quite possible that
_Titanotherium_ was evolved from the genus _Telmatotherium_.[177]
SUB-ORDER 9. LITOPTERNA.
Whether the MACRAUCHENIIDAE should be considered as a separate group of
Ungulata is a matter of dispute. Cope placed them in a special order of
Ungulates which he called Litopterna. Zittel, on the other hand, regards
them as definitely Perissodactyles. One curious point of resemblance to
existing Horses is shown--that is the presence of a pit in the incisor
teeth. This matter seems to be so important as to need a placing of these
forms in the neighbourhood of the Perissodactyles, even of the Equidae; it
is so peculiar a character, and apparently so little related to any obvious
similarity in way of life, that it seems to mark a special affinity. Not so
the fact that in _Macrauchenia_ at any rate the orbit was entirely
surrounded by bone as in the Horse. We find that condition so frequently
acquired in many groups,--a development from an earlier condition where the
cavity for the lodgment of the eye is in continuity with the temporal {268}
fossa, that it cannot be regarded as anything more than a mark of
specialisation. It is, in fact, the case that the Macraucheniidae are in
many points specialised, while retaining many primitive features of
structure.
The chief primitive features are: the non-alternating positions of the
wrist- and ankle-bones; these, of course, interlock in the Perissodactyles
of to-day and in many extinct families. Then the absence of a diastema in
the tooth series, coupled with the presence in _Macrauchenia_ of a complete
dentition. The small brain may be referred to the same category.
_Macrauchenia_ must have been a strange-looking animal. It walked upon
three toes on each limb; the skull was Horse-like in general form, but the
nostrils are removed to a point about as far back as in the Whales or
nearly so, the nasal bones being correspondingly reduced. This it is
thought argues a proboscis. The humerus is particularly compared by
Burmeister[178] to that of a Horse. The radius and ulna though both well
developed are fused. The neck is long, and, as in the Camel, the vertebral
arteries run inside the neural arches. Since the fore-legs seem to have
been rather longer than the hind-legs, though only very slightly, and the
neck was long, the animal may have presented some likeness to the Giraffe.
It is interesting to note that in the proportions of humerus to ulna this
animal is more Lama-like than Horse-like. On the other hand, the
proportions of femur to tibia are more Horse-like. The remains of the
creature are limited to South America, and to quite superficial deposits.
It is evidently a specialised type, and has pursued a course parallel to
that of the Horse. Much nearer to the Horse however, but apparently by
convergence only, is the genus _Thoatherium_, usually placed in a separate
family, the PROTOROTHERIIDAE. In this creature, which has many archaic
characters, the toes are reduced to one in each foot. In an allied form,
_Protorotherium_, we have the two lateral toes diminishing just as in
_Anchitherium_.
* * * * *
{269}
CHAPTER XI
UNGULATA (_continued_)--ARTIODACTYLA (EVEN-TOED UNGULATES)--SIRENIA
SUB-ORDER 10. ARTIODACTYLA.
[Illustration]
FIG. 138.--Bones of the Manus--A, of Pig (_Sus scrofa_). x 1/3. B, of Red
Deer (_Cervus elaphus_). x 1/2. C, of Camel (_Camelus bactrianus_). x 1/8.
_c_, Cuneiform; _l_, lunar; _m_, magnum; _m_^2, _m_^5, second and fifth
metacarpals; _R_, radius; _s_, scaphoid; _td_, trapezoid; _u_, unciform;
_U_, ulna; _II-V_, second to fifth fingers. (From Flower's _Osteology_.)
The Artiodactyle or "Even-toed" Ungulates are to be distinguished from the
Perissodactyla, and from other Ungulate groups, by a number of trenchant
characters. The most salient {270} of these, and that which has given its
name to the group, concerns the arrangement of the digits. Instead of there
being but one prevailing digit--the third, in the hand and foot, through
which the axis of the foot passes, there are two, numbers three and four,
between which the same axis passes, and which are perfectly symmetrical
with each other. This type of foot has been termed "paraxonic," as opposed
to the "mesaxonic" Perissodactyle foot (see Fig. 121 B, p. 235). It has
been attempted to prove that the single prevailing digit of the Horse's
foot is a fused pair of digits, and the state of affairs which
characterises the Camel, where the two metacarpals or metatarsals are to an
almost complete extent united, has been urged in proof; so, too, certain
abnormalities, such as those called "solid-hoofed pigs."[179] These latter
are simply Pigs in which the two central metacarpals and the terminal hoofs
are completely fused with one another. In some of such cases there is not
the slightest trace of the union of the separate metacarpals and phalanges.
Even the sesamoid bones, attached behind to the toes, are two in number
instead of four. And, furthermore, the tendon supplying the bones is
single, though showing traces of its double origin. Such Pigs often show
the abnormality from generation to generation, and they proved convenient
for those whose scruples would not allow them to eat the flesh of a beast
"dividing the hoof" and not chewing the cud. More singular still, as
showing a pathological approach from another side to the Perissodactyle
condition in an Artiodactyle, is a calf, where the foot ended in three
equi-sized digits, of which the middle one lay in the longitudinal axis of
the limb. From the opposite side cases are known of a Horse with a split
hoof and phalanges, thus presenting the most striking likeness to a Camel.
There is, furthermore, in certain groups of Artiodactyles (_e.g._ the
Tragulidae) a tendency for the two middle metacarpals to unite, quite apart
from such "sports" as those illustrated by the cases just set forth. And,
as already mentioned, the union of the two middle metacarpals culminates in
the Camel, Ox, etc. There is, however, absolutely no trace of such a fusion
in the series of Perissodactyle animals known to us; and it would be by
fusion rather than dismemberment that, as it would appear on this theory,
the modern Ungulate foot has been arrived at. Of course {271} the facts of
Ungulate descent are absolutely destructive of any such comparisons.
As is the case with the Perissodactyles, the Artiodactyles show a
historical series, the primitive five-toed condition being almost preserved
in _Oreodon_, up to the most modern modification exemplified by the Ox,
Sheep, etc., in which animals there are not even vestiges of the fourth and
fifth toes. It has been stated, however, that the foetal Sheep has traces
of those rudiments. The so-called cannon bone (the fused third and fourth
metapodia) is accompanied in its fusion by an increase in length. At the
same time the functional middle metacarpals push aside the rudiments and,
forming a broad surface for that purpose, articulate with the magnum and
unciform bones to the exclusion of the rudiments. This has been termed an
"adaptive reduction." In the "inadaptive reduction" there is the same
reduction of the metacarpals, but the rudiments still articulate as in the
primitive Artiodactyle foot, _i.e._ Mc II with trapezium, trapezoid, and
magnum; Mc III with magnum and unciform; Mc IV and V with unciform. This
would appear to give greater solidity and consequently greater strength to
the foot.
[Illustration]
FIG. 139.--Dorsal surface of right tarsus of Red Deer (_Cervus elaphus_). x
1/3. _a_, Astragalus; _c_, calcaneum; _c_^3, cuneiform; _cb_, cuboid;
_mIII_, _mIV_, metatarsals; _n_, navicular. (From Flower's _Osteology_.)
The carpal bones of the Artiodactyla alternate in their articulation; the
primitive state of affairs[180] is not retained even in the earliest types.
The femur has no third trochanter, so prevalent in the Perissodactyles. In
the hind-foot the calcaneum has an articular facet for the fibula, which is
not characteristic of the Perissodactyla. In the more modern forms, _e.g._
the Cervidae, the navicular and cuboid become fused into one bone; and
there are even further fusions which will be referred to later as
characteristic features of different groups. It is interesting to notice
that the reduction begins earlier and is clearer in the hind-foot than in
the fore. One {272} can see how this may be purely adaptive, the push of
the hind-legs in running needing a firmer support. In _Hyomoschus_ this is
the case. The hind-limbs are provided with a cannon bone, while the
metacarpals of the fore-feet are still free.
The number of dorso-lumbar vertebrae is less in the Artiodactyle than in
the Perissodactyle Ungulates. Whereas the former have but nineteen, the
latter have, as a rule, twenty-three such vertebrae.[181] The number of
ribs varies from twelve (_Camelus_, _Hydropotes_) through thirteen
(_Cervus_, _Gazella_) to fourteen in _Dicotyles_, _Giraffa_, etc.
The curious form of teeth known as "selenodont" is characteristic of the
Artiodactyla, though only found well developed in the modern forms, and of
those only in the Pecora. The more primitive forms had "bunodont" teeth
with typically four tubercles (if we except the tritubercular and but
little-known _Pantolestes_); and the intermediate "buno-selenodont" type
characterises such groups as the Anthracotheriidae.
While the stomach of the Perissodactyles is always a simple sac, it is
complicated, or shows signs of complication, in the Artiodactyles. That of
the Hippopotamus is divided into two chambers; there are three in
_Tragulus_, and four in the typical Ruminants such as _Cervus_, _Ovis_,
etc.
Had we to deal only with the still living genera of Artiodactyles, it would
be easy to sort them into two groups on the characters of the teeth; for
the Pigs and Hippopotamus are provided with tubercular molars; they are
bunodont. The Deer, Camels, Oxen, Giraffes, etc., have selenodont molars.
Besides, the latter are "Ruminants," and have a more complicated stomach.
The existing Chevrotains forbid a more trenchant division, since they are,
as will be pointed out in due course, somewhat intermediate in structure;
the feet are more Pig-like, and the stomach is not so typically Ruminant.
In any case such a division is prevented by certain extinct families which
are perhaps ancestral to both. They have teeth which are not quite bunodont
and not quite selenodont. These teeth have been termed buno-selenodont or
buno-lophodont.
The distribution of the living Artiodactyles presents us with some
interesting facts. The vast preponderance of species occurs in the Old
World--34 in America as against over 250 species {273} in Europe, Asia, and
Africa. The Neotropical region has no Oxen, or Sheep, or Antelopes. The
latter are confined to Africa, Asia, and certain parts of the Palaearctic
region; they are vastly more prevalent in Africa, where they take the place
of the totally absent Deer. The Pig tribe is almost entirely Oriental and
Ethiopian in distribution, only one form, the European Wild Boar, ranging
into the Palaearctic region; and the two species of Peccary are found in
both North and South America. Broadly speaking, the Ethiopian region is the
headquarters of the Artiodactyla. But the great island of Madagascar has
but one form of Artiodactyle, a Pig of the genus _Potamochoerus_.[182]
GROUP I.--_SUINA._
FAM. 1. HIPPOPOTAMIDAE.--The family Hippopotamidae contains of existing
genera only _Hippopotamus_, for the Liberian dwarf Hippopotamus is not now
regarded, as it was formerly, as the type of another genus, _Choeropsis_.
The reasons for its former separation were the loss of the outer pair of
incisors and the different proportions of various parts of the skull. This
little Liberian animal has, however, been shown by Sir W. Flower[183] to
possess the missing incisors occasionally; and as to the proportions of the
skull, it is exceedingly common for small animals to vary from larger
relatives in this way. Hence, considering the characteristic features of
the Hippopotamus and the fewness of species, it seems unnecessary to divide
it up further. We shall therefore only recognise one genus.
The Hippopotamus at present is African in range, and confined to that
continent. But quite recently it inhabited Madagascar; and further back
still in time the existing African species, _H. amphibius_, ranged into
Europe; there were also Indian forms, which were contemporary with the
Stone-age man. The Common Hippopotamus is a great thick-skinned beast with
but few hairs. It has four toes on each foot, a complex stomach, but no
caecum. The strong incisors continue growing through life, as do the great
canines. The number of incisors is two on each side of each jaw. Some of
the extinct species had six in each {274} jaw, and they were distinguished
as a genus _Hexaprotodon_, contrasting with _Tetraprotodon_, until
intermediate conditions were observed. _Choeropsis_, as already observed,
was a still further reduction of the tetraprotodont type. The molars (the
formula is Pm 4/4 M 3/3) when worn show a double trefoil pattern. The
orbital cavity is encircled by bone. As with many other aquatic mammals the
kidneys are lobulated.
[Illustration]
FIG. 140.--Hippopotamus. _Hippopotamus amphibius._ x 1/40.
A very singular fact about the Hippopotamus is the production of a "bloody
sweat," a carmine-coloured secretion, containing small crystals and
corpuscles, from the skin. This coloured fluid has of course nothing to do
with blood.[184]
The animal grows to a length of at any rate 14 feet. The limbs and the tail
are short. Like other aquatic animals the nostrils are on the surface of
the head, and can be closed when the animal is under water. When it reaches
the surface of the water after a prolonged immersion, it spouts like a
Whale. Sir Samuel Baker says that ten minutes is the longest time that the
Hippopotamus can remain below the water. It is frequently a dangerous
animal to encounter, as it will capsize boats, and even bite large pieces
out of their bottoms; with its huge teeth it {275} can and does attack and
destroy human beings. The Hippopotamus not only swims, but can walk along
the bottom of a river with great rapidity. It occasionally puts out to sea
from the mouths of rivers frequented by it; and it is supposed that in this
way Madagascar was populated with Hippopotamuses, whose remains are now
found in swamps in that island.
[Illustration]
FIG. 141.--Wild Boar. _Sus scrofa._ x 1/12.
FAM. 2. SUIDAE.--The Pig family, Suidae, differ from the last in their
smaller size, in the terminal nostrils and mobile snout, which is not
grooved, except faintly as in _Babirusa_. They are generally hairy, but the
Babyroussa is an exception, while _Phacochoerus_ is but slightly haired.
Though there are four digits, as in the Hippopotamus, only two reach the
ground in walking. The stomach, furthermore, is simple, and (except in
_Dicotyles_) there is a caecum. The kidneys are smooth, and the liver is
more lobate than in _Hippopotamus_. The orbital cavity is confluent with
the temporal fossa. The typical genus, _Sus_, is distributed over Europe,
Asia, and the islands of the Malay Archipelago, reaching as far as Borneo
and Celebes. The dentition[185] is complete. A single species, the
so-called _S. sennaariensis_, is from Ethiopian Africa, but it is not
certain how far this animal may be an escaped species introduced by man. A
very large number of "species" of _Sus_ have been described, but Dr.
Forsyth {276} Major is disposed to reduce them to four if not to fewer
species. He allows the widely-ranging _S. scrofa_, _S. vittatus_, and the
eastern Malayan _S. verrucosus_ and _S. barbatus_.
[Illustration]
FIG. 142.--Pygmy Hog (from _Nature_). _Sus salvania._ x 1/6.
The Pygmy Hog of the Bhotans seems to be not entitled to specific rank,
certainly not to generic (in the opinion of some), though it has been
termed _Porcula salvania_.[186] The Wild Boar of Europe is _Sus scrofa_. It
was formerly quite abundant in this country; not merely are its remains
exhumed from fens, caves and peat bogs, but there is ample evidence of its
continuance down to a comparatively late historic period. Enactments are on
record as to the hunting of these animals; there are places, such as
Boarstall, whose names are clearly derived from the name of the animal,
presumably once a native of the locality; and various documents all show
the presence of the Wild Boar {277} in this country down to so late a
period as the end of the sixteenth century.
[Illustration]
FIG. 143.--Wart Hog. _Phacochoerus aethiopicus._ x 1/6.
The African Wart Hog, genus _Phacochoerus_, is usually regarded as the type
of a distinct genus of Pigs. This animal, "superlatively ugly" with its
huge tusks and great protuberances upon the face, is chiefly to be
distinguished from the genus _Sus_ by these characters, and by the
complexity of the last molar, which, with the tusks, are sometimes in aged
animals the only teeth left. The complete formula is Pm 2/2 M 3/3. There
are two species of this genus, _P. aethiopicus_ and _P. africanus_. When
enraged the Wart Hog is said to carry its tail directly up, and to present
a ludicrous as well as ferocious appearance.
[Illustration]
FIG. 144.--Head of Wart Hog.
The Celebesian Babyroussa, genus _Babirusa_, is an almost hairless hog with
enormously upturned tusks in both jaws of the {278} male. In the Wild Boar
there is a hint of this, which is carried still further in _Phacochoerus_;
but in _Babirusa_ the upper tusks turn upwards before they leave the
substance of the jaw, for which reason they appear to arise on its dorsal
surface; the lower tusks are nearly as long. It has been found that the
young of this Pig are not striped as are those of other Pigs. By means of
the curved upper tusks this animal has been said by old writers to suspend
itself to branches of trees, just as does by his downwardly-projecting
tusks the male Chevrotain! There is but one species, _B. alfurus_.
From _Sus_ proper the African and Malagasy _Potamochoerus_, including the
Red River Hog, is barely separable generically. Their principal claim to
generic distinction lies in the existence of a horny outgrowth arising from
a bony apophysis above the canine in the male. These have been compared to
the osseous "horn cores" in the extinct Dinocerata. But the Javan _Sus
verrucosus_ shows at least the beginning of a similar modification. The
popular name of the animal is derived from the fine rufous colour of its
pelage, not seen, however, in all the species. Dr. Forsyth Major[187]
recognises five species, of which only one is from Madagascar.
[Illustration]
FIG. 145.--Peccary. _Dicotyles tajacu._ x 1/6.
FAM. 3. DICOTYLIDAE.--The Peccaries are generally placed in a different
family from that of the other Pigs. This family, {279} Dicotylidae,
contains but one genus, _Dicotyles_, with at most two species. The name of
the animal is connected with the dorsal gland; the animal thus appeared to
possess two navels. The Peccaries, exclusively confined to the New World,
differ from the Old-World Pigs in one or more important characters. They
have only three toes on the hind-feet, and the stomach is complicated.
Though the Peccaries have but small tusks they hunt in packs and are very
dangerous animals to meet with. They owe, too, their safety from many foes
to their sociable habits. Being nocturnal animals they are liable to the
attacks of the Jaguar, which will speedily overpower and devour a Peccary
that has strayed from its herd.
FOSSIL SWINE.--The existing genera of the Pig tribe are also known in a
fossil condition. _Sus_ itself goes back as far as the Upper Miocene. _Sus
erymanthius_, the Erymanthine Boar, is known from beds of that age in
Greece, England, and Germany. This genus is not known to have had a wider
distribution in the past than it has in the present. _Dicotyles_ occurs in
the Pleistocene of both North and South America, the regions which it
inhabits at the present day. The genus _Listriodon_, also Miocene, is
remarkable for having lophodont instead of bunodont teeth, that is so far
as concerns the molars, which resemble those of the Tapir. It was European
and Indian in range. A number of genera, more remote from the existing Pigs
than those which have just been dealt with, are placed together in a
special sub-family, Achaenodontinae. The type genus, _Achaenodon_, had a
somewhat short skull for a Pig; and it is in general aspect and in the
characters of the canine teeth highly suggestive of that of a Carnivore.
The bunodont molars, however, are Suine, as is the form of the lower jaw
with a rounded angle. This is an Eocene animal found in Wyoming.
_Elotherium_[188] occurs chiefly in the Miocene of both North America and
Europe; but _E. uintense_ is Eocene. The orbits are completely encircled by
bone in the more modern forms; this is not the case in the last-described
genus, with which _E. uintense_ agrees. The skull is also longer and more
Pig-like. The zygomatic arch is powerful, with sometimes a large descending
process, such as is found in _Diprotodon_, more faintly in Kangaroos, and
in Sloths and certain extinct Edentates. The lower jaw has a pair {280} of
dependent processes near the symphysis, which suggest processes occupying a
corresponding position in _Dinoceras_. The skull and body are heavy, but
the two-toed limbs are slender. There is a smaller pair of toes behind
these. The dentition is complete, and the canines are not inordinately
developed. The brain is very diminutive. Perhaps _E. uintense_ should be
separated as a distinct genus, _Protelotherium_.[189]
_Hyotherium_ (which is regarded as identical with _Palaeochoerus_) has a
sharp sagittal crest; the orbit is nearly but not quite closed. The canines
are not strongly developed. The upper canines have double fangs as in
_Triconodon_ among extinct mammals, and as in the Hedgehog and other forms
among living Mammalia. The premolars have the cutting and serrated edge of
those of some other Pigs, a feature which gives them a curious resemblance
to the "grinding" teeth of Seals. The molars are tuberculate, and like
those of living Pigs. It is European and Indian in range, and Miocene.
The genus _Choeropotamus_ has a complete dental formula save for the loss
of a premolar in the lower jaw. Though it has lost this tooth, it is from
an older stratum than some of those forms which have retained that
premolar; it has been found in the Upper Eocene of the Isle of Wight and of
the neighbourhood of Paris.
The American and Miocene _Chaenohyus_ has lost the corresponding teeth of
the upper jaw.
_Homacodon_[190] is a genus consisting of several species, which has a
bunodont and complete dentition. The molars are sextubercular in the upper
jaw. _H. vagans_ was of about the size of a Rabbit, and it appears to have
had a curved neck. The limbs had five digits, as is so generally the case
with Eocene Ungulates. It is known from the Middle Eocene of Wyoming.
GROUP II.--_RUMINANTIA._
The Selenodontia or Ruminantia form the second division of existing
Artiodactyles. The characters of the teeth, which give them their name,
have already been referred to. They also differ in that there are never
more than a single pair of incisors {281} in the upper jaw, and very
usually there are none. As a general rule the third and fourth metacarpals
and metatarsals become united to form a cannon bone. To this there is but
one exception, the African _Hyomoschus_. Moreover, the second and fifth
digits are nearly always rudimentary, and may practically disappear
altogether. Here again the Tragulidae are an exception. The Ruminantia are
so-called on account of the fact that they "ruminate," that is, after the
food has been rapidly swallowed, it is forced back up the gullet and more
thoroughly masticated. Associated with this is a complex stomach, which is
divided into several compartments. This stomach has at least three
compartments, as in the Tragulidae; but it has usually four. Its characters
are illustrated in Fig. 146. The majority of the Selenodontia possess
horns, which are partly formed of solid protuberances of the frontal bones.
In the Giraffe they are somewhat different.
[Illustration]
FIG. 146.--Stomach of Ruminant opened to show the internal structure. _a_,
Oesophagus; _b_, rumen; _c_, reticulum; _d_, psalterium; _e_, abomasum;
_f_, duodenum. (After Flower and Lydekker.)
This group may be divided into--A. TRAGULINA, Chevrotains; B. TYLOPODA,
Camels, Lamas; and C. PECORA, Deer, Antelopes, Oxen, Giraffes, Goats,
Sheep.
A. TRAGULINA.
As the Tragulina are undoubtedly the most ancient of the Selenodontia it
will be logical to commence with an account of them. {282}
FAM. 4. TRAGULIDAE.--This family comprises a number of small Deer-like
animals, which are really in many points more related to the Pigs than to
the true Deer. They are known as Chevrotains; and the term "Deerlet,"
introduced by Professor Garrod, is certainly appropriate, since they have
the aspect of very small and hornless Deer. If it were not for their
Artiodactyle feet one might at a glance confuse these creatures with some
Marsupial type. The family is Oriental and West African in range. The two
genera (whose individual peculiarities will be considered later) differ
from other Artiodactyles in a number of rather important characters.
[Illustration]
FIG. 147.--Indian Chevrotain. _Tragulus meminna._ x 1/4.
They are absolutely hornless in both sexes. The canines are present in both
jaws, and are especially well developed in the upper jaw. The dental
formula is I 0/3 C 1/1 Pm 3/3 M 3/3. In the skull the tympanic bulla is
usually, as in the non-ruminating Artiodactyles, filled with loose bony
tissue. The feet (usually) have the four toes of the Suina, and are
therefore in a more primitive condition than in Deer and Antelopes. But as
the middle metacarpals are fused in _Tragulus_ (though separate in
_Hyomoschus_) they are a stage further than are the Pigs, in the direction
of the typical Ruminants.
The stomach is comparatively simple, thus offering {283} intermediate
characters between the Pigs and the Ruminants; there are but three separate
compartments. A highly-interesting character is afforded by the placenta.
This is in the present family of the diffuse kind, not presenting the
separated and tufted cotyledons of the Ruminant placenta. This we may
fairly assume is a further proof of the less-specialised characters of this
group[191] as compared with the Ruminantia, a view, however, which is not
universally accepted. While the molars have the selenodont character of
other Pecora, the premolars are more adapted for cutting, with sharp edges.
The genus _Tragulus_ consists of several species (e.g. _T. stanleyanus_,
_T. napu_, etc.), which have been aptly compared in external appearance to
certain Rodents such as the Agoutis. The legs are delicate and slender,
hardly "thicker than an ordinary cedar pencil." These creatures have got
among the Malays a considerable reputation for astuteness, embodied in the
saying, "Cunning as a _kanchil_." The male has tusks, which greatly
contributed to the confusion of this creature with the totally different
Musk Deer, _Moschus moschiferus_. It is even said to suspend itself by
their aid to the branches of trees, and so avoid danger.
_Hyomoschus_ (or _Dorcatherium_ as it should properly be called) is West
African. Its rich brown colour, with spots and stripes, is much like that
of the Chevrotains, but it has shorter limbs. The only species is _D.
aquaticum_, which is sometimes called, on account of its frequenting the
banks of streams, the Water Chevrotain. Remains of this genus occur in
Miocene and Pliocene strata of Europe.
The separate metacarpals, comparatively simple stomach, absence of horns,
diffuse placenta, and spotted pelage are features which argue the primitive
position of these animals among existing Artiodactyles.
Besides the two existing genera which have just been treated of, there are
a number of extinct genera undoubtedly belonging to the same group.
_Gelocus_ (Eocene and Oligocene in range) is a European genus known from
France. It differs from the living members of the group by the fact that
the second and fifth toes on both hind- and fore-feet are represented, as
in certain Deer, {284} by rudiments at the upper and at the lower end only;
they are deficient in the middle. The middle large metacarpals, though
closely applied, are not fused. The metatarsals, on the other hand, are, or
are not fused, according to the species. A later form is the genus
_Leptomeryx_ from the Miocene of North America. This genus departs from the
typical Traguline structure in more than one point. The tympanic bulla is
hollow instead of being filled with cancellated bone; the cuneiform is not
fused with the cuboid and navicular, though the latter are with each other;
the lateral digits of the hind-feet are rudimentary. The magnum and
trapezoid, however, are fused. In the fore-feet the middle metacarpals are
separate, and the lateral less perfect metacarpals have toes. The
metatarsals are fused.
Not definitely referable to the Tragulidae, but coming near to them, are
the PROTOCERATIDAE. Of this family there is but one well-known genus,
_Protoceras_,[192] from the Miocene of North America.
The skull is singularly reminiscent of _Dinoceras_, with which this quite
Artiodactyle genus has, of course, nothing to do. It merely exemplifies the
phenomenon of "parallelism." In general form it is peculiarly long and low.
There are three pairs of bony protuberances: one, the largest, pair are
situated on the maxillae rising up just behind the implantation of the
canine teeth; the parietals have a second pair; and a third much more
diminutive pair of bosses are upon the frontals, near their junction with
the nasals. This description refers to the male; the female has only traces
of the parietal bosses. These were all possibly tipped or sheathed with
horn or roughened skin. The dentition of this genus is precisely that of
the Tragulidae, _i.e._ I 0/3 C 1/1 Pm 4/4 M 3/3. The orbit is completely
encircled by bone; the auditory bulla is not swollen; the premaxillae are
small.
The nasal cavity is very large and open, the end of the nasal bones
anteriorly being situated at about the middle of the skull; this would seem
to indicate at least a flexible and long nose like that of the Saiga
Antelope, if not a trunk.
The brain was of good size, and quite well convoluted.
The limbs are constituted on the Traguline plan; in the fore-limbs the
middle metacarpals are quite free from each other, and the more diminutive
lateral digits are complete. The {285} metatarsals are free, but with a
tendency to fusion; the lateral toes are only represented at the upper
extremity. The carpal bones are separated.
This animal, which was about the size of a Sheep, though of more delicate
proportions, was allied not only to the Tragulidae but to the Giraffidae;
it is impossible to refer it definitely to either family.
B. TYLOPODA.
FAM. 5. CAMELIDAE.--This small group of Selenodonts includes only the
Camels and Lamas. The limbs are long and have no traces of the second and
fifth toes. The fused metacarpals and metatarsals diverge somewhat at their
distal ends. In the upper jaw is a single pair of incisors. The stomach
differs from that of the typical Ruminants. The rumen has smooth and not
papillose walls, and from it are developed the "water cells," diverticula
with narrow mouths provided with a closing sphincter muscle. The psalterium
is reduced to a mere vestige, and so the stomach has, as in the Tragulina,
but three chambers. This, so far ancient, character in the structure of the
Camel tribe is associated with another, also seen in the more primitive
Ungulates, viz. the diffuse character of the placenta. A very singular
peculiarity of this group is the fact that the blood corpuscles instead of
showing the ordinary mammalian round contours are elliptical.
[Illustration]
FIG. 148.--Bactrian Camel. _Camelus bactrianus._ x 1/30.
The genus _Camelus_, confined to the Old World, is made up of two quite
distinct species, the Bactrian Camel, _C. bactrianus_, with two humps, and
the Dromedary, _C. dromedarius_, with only one. The former species is
Asiatic. It is a singular fact that neither of the species is known to
occur in a genuinely wild condition. The so-called "wild" Camels appear to
be invariably feral. The two species will interbreed; and there is at the
Zoological Society's Gardens such a hybrid, which has the general
appearance and shaggy brown hair[193] of the Bactrian animal, but the one
hump of the Dromedary. It may be that the Bactrian Camels of Lob-nor are
really wild; but the desert contains so many remains of cities destroyed by
sand-storms that these reputed wild {286} Camels may be the descendants of
animals belonging to the inhabitants of those cities. A strayed herd of
Camels has established itself in a feral state in Spain. Otherwise the
genus does not occur in Europe. The Camels are also represented in the New
World. The genus _Lama_ (_Auchenia_ of many authors) belongs to this
family. These Camels differ from their allies in the Old World by their
smaller size, by the absence of the characteristic hump, and by the
dropping of one premolar, the dental formula being otherwise similar. A
variety of names, Lama, Alpaca, Huanaco, Vicuna, have been applied to these
animals; but it appears that the names are in excess of the number of the
species. Mr. Thomas, who has lately inquired into the matter, will only
allow two, the Huanaco, _Lama huanacos_, of which there are two domestic
races, the Llama and the Alpaca, and the Vicuna, _Lama vicugna_. They are
both South American in range. Not only is there a herd of escaped Camels in
Spain, but the Spaniards attempted to introduce and acclimatise the useful
Lama. The first Lama ever seen in Europe was brought in the year 1558 to
{287} the town of Middelburg in Holland; it was purchased and presented to
the Emperor of Germany. Gesner gives a curious figure of it, representing
the animal as a comparatively colossal beast submitting itself to the
guidance of a dwarfish man. The habit of "spitting" of the Lama is well
known. Augustin de Zarate and Buffon speak of the Lama as having no
protection save this habit, which is more than a mere ejection of saliva:
the contents of the stomach are forcibly shot at the object of its
annoyance. It can also kick and bite. In the intestines (as in those of
some other mammals) are found Bezoar stones, or Bezards as they are
variously spelt. These were once valued in medicine, and even so lately as
1847 were, according to Gay, the historian of Chili, in vogue; these
concretions, comparable to the ambergris of the Whales, were supposed to be
an antidote to poison.
[Illustration]
FIG. 149.--Lama. _Lama huanacos._ x 1/12.
EXTINCT CAMELS.--The earliest cameloid type is the genus _Protylopus_,[194]
of which we are acquainted with an imperfect skull {288} and the greater
part of a radius and ulna belonging to one individual, and most portions of
the hind-limbs in other specimens. The one species, _P. petersoni_, was
about the size of a "jack rabbit," and is late Eocene (Uinta formation) and
American in range. The teeth of this mammal are the typical forty-four, and
the canines are not pronounced, being incisiform in shape. In the skull the
nasals overhang, as in the genus _Poebrotherium_. The orbit is not closed
by bone. There is in this ancient Camel a trace of the supra-orbital notch
so characteristic of the Camel tribe. "The vertebrae resemble those of the
modern Lamas closely in their general proportions." The lumbars have the
usually Cameloid formula of 7. This genus has but two functional toes on
the hind-feet, the second and fifth being reduced to vestiges. It is
interesting to note that the radius and ulna appear to remain distinct,
except in very old animals, in which they come to be co-ossified in the
middle only, thus foreshadowing their complete union in the next genus,
_Poebrotherium_. The present genus, moreover, as well as _Poebrotherium_,
was distinctly unguligrade; it has not acquired the characteristic
phalangigrade mode of progression of the modern types of Camels.
[Illustration]
FIG. 150.--Skull of _Poebrotherium wilsoni_. _i_^1, _i_^2, _i_^3, Incisors
1-3. x 1/2. (After Wortman.)
The American and Oligocene _Poebrotherium_ has been recently and
exhaustively studied by Professor Scott.[195] It was considerably smaller
than a Lama. Its neck was long as compared with other Artiodactyles, but
still shorter than that of the Lama. It was a lightly-built, graceful
creature, with apparently some external likeness to a Lama. It is an
important fact to notice that at this {289} period, and for a long time
after, there were no types referable to the Camelidae in the Old World.
Though a Camel in many features of its organisation, _Poebrotherium_ was
"generalised" in many ways. Thus the metacarpals and metatarsals were not
fused to form a cannon bone, and the two lateral digits were represented by
splint rudiments of metacarpals and metatarsals. The dentition was
complete. The skull though distinctly Tylopodan, also shows more
generalised characters. Thus the orbit is not quite, though nearly,
completed by bone. In the Camel it is quite closed. The nasal bones are
much longer, reaching nearly to the end of the snout. The odontoid process
of the axis vertebra is not spout-like as in existing forms, but
cylindrical, though slightly flattened upon the upper surface. The scapula
is described as being more like that of the Lama than of the Camel, though
variations occur which approximate to the Camel. The brain, judging of
course from casts, has those sulci "which are common to the whole series of
Ungulates, and closely resemble those of a foetal Sheep."
[Illustration]
FIG. 151.--Anterior surface of axis of Red Deer, x 2/3. _o_, Odontoid
process; _pz_, posterior zygapophysis; _sn_, foramen for second spinal
nerve. (From Flower's _Osteology_.)
Later in historical sequence than _Poebrotherium_, and structurally
intermediate between it and _Protolabis_, is the Miocene genus
_Gomphotherium_. It shows an advance in structure upon _Poebrotherium_, in
that the orbit is completely encircled by bone, though the posterior wall
is thin; the lower canines instead of being incisiform are curved back as
in later Camels, and separated by a wide diastema from the preceding and
the succeeding teeth.
Later in age than _Poebrotherium_ is _Protolabis_, a _Tylopod_ in which the
full number of teeth is still retained; its skull presents no particular
changes from the Poebrotherine type; the nasals, however, are somewhat
shortened.
Later still in point of time is _Procamelus_. In this form we have
apparently an ancestral stock, whence both Camels and Lamas were derived.
The upper incisors are as in existing forms, but the first and second
persist for a somewhat longer time. The skull shows two well-marked types
of structure; in _P. occidentalis_ {290} there are more points of likeness
to the Lama, in _P. angustidens_ to the Camel. In both, the orbits are
completely encircled by bone. The nasals are much shortened. The odontoid
process of the axis is still more concave than in _Poebrotherium_, but not
spout-like as in existing forms. This fact shows that the spout-like
character of the Camels' odontoid process is not a point of affinity to
other Artiodactyles--in fact the occurrence of the same form of odontoid
process in Perissodactyles is enough proof of this. We must come to the
conclusion that the form is adaptive in all cases. If we were not obliged
on palaeontological evidence to come to this conclusion, the structure in
question is just one which would be fastened upon as evidence of genetic
affinity; for it is a resemblance in a small though distinctive point of
structure having no obvious relation to utility. The metacarpals and
metatarsals have coalesced to form the cannon bones, though a rudiment of
one metacarpal seems to remain. The genera referred to appear to be on the
direct line of descent of the modern representatives of the family. But
there are other forms which are offshoots of the main stem. Such are
_Homocamelus_, _Eschatia_, and _Holomeniscus_. The last two are Pliocene
and American; the teeth are much reduced.
C. PECORA.
The Pecora are a group which possess so many characters in common that it
is not an easy task further to subdivide them.
In all there are but two functional digits on the feet, and the metacarpals
and metatarsals of these are fused. There are no upper incisors, and
canines in the upper jaw are not universal, and generally small. Horns are
confined to this group of the Selenodontia.[196] The premolar teeth are of
a simpler form than the molars. The stomach has four chambers, of which two
may be regarded as belonging to its cardiac half and two to the pyloric.
The former are, in the first place, a large paunch or rumen, followed by a
smaller reticulum, so called on account of the network arrangement of the
folds of its lining membrane. Connected with the latter, and constituting
the first part of the pyloric half of the stomach, is the psalterium or
"manyplies," so called on account of the longitudinal folds, like the
leaves of a {291} book, into which its lining membrane is raised. Finally
there is the abomasum, out of which proceeds the small intestine. Garrod
has observed that the chamber of the stomach which varies most among the
Pecora is the psalterium. This chamber is specially large in _Bos_, and
particularly small in the Antelopes _Nannotragus_ and _Cephalophus_. But
its variation relates more especially to the folds of its mucous membrane.
These folds are of varying lengths and have a definite arrangement There
may be as many as five sets of laminae of regular depths. The most simple
psalterium is that of _Cephalophus_, where there are only two sets of
laminae of different sizes, a deeper set and a very much shallower set;
this form is termed by Garrod "duplicate." Most common is the
"quadruplicate" arrangement, with four sets of laminae of differing depths.
In all Pecora the liver is but little divided by fissures.
FAM. 6. CERVIDAE.--The Deer tribe is a very extensive one, and, with the
exception of Africa and Australia, world-wide in distribution.[197]
The Deer are absolutely distinguished from all other Ruminant animals by
the existence of antlers, which are invariably present in the male sex,
save in the aberrant genera _Moschus_ and _Hydropotes_; in the Reindeer
alone are antlers present in both sexes. The general characters of these
appendages have been dealt with on a former page (p. 200), where they are
compared to, or rather contrasted with, the horns of the Bovidae. These
antlers, so characteristic of the Cervidae, are very variously developed
among the members of the family. Thus in _Elaphodus_ the antlers are very
small and entirely unbranched. In the Muntjacs, _Cervulus_, the antlers are
hardly larger, but they have a small anterior branch arising from near the
pedicel, the "brow tine." In _Cariacus antisiensis_ only one branch, the
brow tine, is present, but it is nearly as long as the main stem of the
antler, the "beam." In _Capreolus capraea_ the beam bears two tines; in
_Cervus sika_ three; in _C. duvauceli_ two of the three tines present bear
secondary branches. There are other complications (some of which are
illustrated in Figs. 152-157) of the simple antler which culminate in the
complex antlers with their expanded "palms" of the Elk and the Fallow Deer.
{292}
Another highly-interesting fact concerning these same antlers is their
gradual increase in complexity of tines and palm from the Miocene _Cervus
matheroni_ to the great Irish Elk of post-Tertiary times.
Beyond the antlers there seems to be no character of universal
applicability which distinguishes the Cervidae from the nearly-related
Antelopes. There are, however, a number of structural features which are
_nearly_ universally characteristic. Excepting _Moschus_ (which Professor
Garrod would not allow to be a "Deer"), no Cervine has a gall-bladder[198]
to its liver. All Bovidae (including Antelopes) have, with the exception of
_Cephalophus_.
A small but constant character of the Deer is the existence of two orifices
to the lachrymal duct. The genus _Tragelaphus_ alone among Antelopes shows
this character.
So far as is known the placenta of the Deer has but few cotyledons, that of
the Bovidae many. But not many types are known.
The navicular, cuboid and ectocuneiform are often united. This is never the
case in the Bovidae.
The first and second phalanges of the lateral (imperfectly developed)
digits are always present except in the Muntjacs; they are never found in
Bovidae. The Deer always present a light brown to a darker brown
coloration. _Elaphodus michianus_ is almost black. There is commonly white
on the under parts and beneath the short tail. Some Deer, such as the
Fallow Deer, are spotted; and the young of others that are uniformly
coloured when adult are spotted. In some cases a winter coat, darker than
the summer coat, is developed.
Altogether some sixty species of Deer are known, of which the preponderance
are Old-World forms. The Deer of the Old World are distributed among the
genera[199] _Cervus_ (all Europe and Asia); _Cervulus_, the Muntjacs
(India, Burmah, China, etc.); _Hydropotes_ (Eastern China); _Capreolus_
(Europe and Central Asia); _Elaphodus_ (Eastern China); there is one
American _Cervus_, the Wapiti. The American genera are _Cariacus_ and
_Pudua_. The Elk (_Alces_) and the Reindeer (_Rangifer_) are circumpolar.
The principal structural modification which occurs within {293} the family
Cervidae concerns the rudimentary fifth and second toes. In _Capreolus_,
_Hydropotes_, _Moschus_, _Alces_, _Rangifer_, and _Pudua_ there are
considerable remains of the lower parts of metacarpals II. and V.; in the
other genera smaller traces of the upper ends of the same bones.
The two most abnormal genera are _Moschus_ and _Hydropotes_, more
particularly the former, which neither Sir V. Brooke nor Professor Garrod
allow to be members of the family at all. _Moschus_ is usually placed in a
special sub-family by itself, Moschinae, the remaining Deer being referred
to another sub-family, Cervinae.
SUB-FAM. 1. CERVINAE.--The genus _Cervus_ comprises rather over twenty
existing species, which, except the Wapiti (_C. canadensis_), are
exclusively Old World in distribution. The principal features of variation
in the genus, in accordance with which it has been divided up into
sub-genera, are (1) palmated (Fallow Deer, _Dama_) or non-palmated antlers;
(2) adults spotted with white at all ages and seasons (_Axis_), or in
summer only (_Pseudaxis_), or not at all; (3) spotted or unspotted young;
(4) existence or absence of rudimentary canines in the upper jaw.
Among the members of this genus, _Cervus (Elaphurus) davidianus_ is
interesting as having been first observed by the missionary Pere David in a
park belonging to the Emperor of China near Pekin. Its horns are remarkable
for dividing early into two branches of equal length, of which the anterior
again branches into two. Specimens of this Deer were ultimately obtained
for the Zoological Society's Gardens.
The species of _Cervus_ are fairly distributed between the Palaearctic and
the Indian regions. The Palaearctic species, such as Luehdorff's Deer (Fig.
152), are mainly Asiatic. _Cervus elaphus_ and _Cervus dama_ alone are
European and British. The former of course is the Red Deer, the latter the
Fallow Deer. The Red Deer is reddish-brown in summer and greyish-brown in
winter, with the white patch on the rump so common in the Deer tribe. The
Red Deer is genuinely wild in Scotland, in certain parts of Devonshire and
Westmoreland, and in the New Forest. At the beginning of the last century,
according to Gilbert White, there were 500 head of deer in Wolmer Forest,
which were inspected by Queen Anne. The antlers may have as many as
forty-eight points; and a stag with more than the three anterior tines is
termed a "Royal Hart." The Fallow Deer has {294} palmated antlers, and is
usually spotted. It seems to be an introduced species, common report
pointing to the Romans as the introducers. It would be more correct to say
"re-introduced," for fossil remains of this Deer have been met with.
[Illustration]
FIG. 152.--Luehdorff's Deer. _Cervus luehdorffi._ x 1/15. (From _Nature_.)
_Elaphodus_[200] contains probably two species, _E. cephalophus_ of
Milne-Edwards and _E michianus_ of Swinhoe, both from China. The antlers
are small and unbranched; the canines in the male are massive; it differs
from _Cervulus_, to which it is closely allied, principally in the absence
of frontal glands. The second {295} species has a dark iron-grey pelage,
and the late Mr. Consul Swinhoe described it as very Goat-like in aspect.
_Capreolus._--The Roe Deer has fairly complex antlers. It is a small Deer
and has spotted young. The common Roe Deer, _C. capraea_, is a native of
this country. It is the smallest of our Deer, and its antlers only have
three tines in stags of the third year. It is a singular fact about this
Deer that though the pairing season is in July and August, the young are
not born until the following May or June, a period which does not represent
that of gestation. The germ remains dormant for some time before
developing.
[Illustration]
FIG. 153.--Mule Deer. _Cariacus macrotis._ x 1/15. (From _Nature_.)
The Muntjacs, _Cervulus_, form a distinct generic type confined to the
Indian and the South-Eastern Palaearctic region. They are small Deer with
spotted young, and short one-branched antlers placed upon pedicels as long
as themselves. The canines are strongly developed in the males. There are
about half-a-dozen species.
_Cariacus_ is exclusively American in range, and contains about twenty
species. There are or are not upper canines. The young {296} are spotted.
The antlers are occasionally very simple; in _C. rufus_ and a few allies
(placed in a special sub-genus _Coassus_) they are simple spikes without
branches. In this genus, and in the nearly allied and also New-World
_Pudua_, the vomer is prolonged backwards and divides the posterior nares
into two. The bulk of the species are South American.
[Illustration]
FIG. 154.--Chilian Deer. _Cariacus chilensis._ x 1/12. (From _Nature_.)
_Pudua_, just mentioned, comes from the Chilian Andes. It is a small Deer
without canines and with minute antlers. Other generic names have been
proposed for various species of American deer.
_Hydropotes inermis_ is a small perfectly hornless Deer, living on the
islands of the Yang-tse-kiang. The male has tusks; the young are spotted.
Though, like other deer, _Hydropotes_ has no gall-bladder, both Mr.
Garrod[201] and Mr. Forbes[202] found the {297} rudiments of one in the
shape of a white ligamentous cord. Mr. Forbes has especially dwelt upon the
likeness of the brain to that of _Capreolus_. The female has four teats,
and produces three to six young at a time.
[Illustration]
FIG. 155.--Water Deer. _Hydropotes inermis._ x 1/10. (From _Nature_.)
_Alces machlis_, the Elk or Moose, is a circumpolar species with palmated
antlers and is of large size. The young are unspotted. This animal is the
largest of the Deer tribe. The aspect of this creature is by no means that
of a Deer, the long, thick, and rather prehensile upper lip not by any
means suggesting the family to which it belongs; the legs, too, are
ungainly through their unusual length. The Moose has a curious method of
protecting himself from Wolves. Instead of moving about during heavy
snowstorms, and being thus on the heavy ground an easy prey for these agile
enemies, the animal forms what is known as a "Moose yard." An area of
ground is kept well {298} trampled down, and the animal contents itself
with browsing upon the adjacent stems. The well-trampled ground gives an
easy footing, and by his powerful horns the great stag is able to keep his
enemies at bay.
[Illustration]
FIG. 156.--Moose. _Alces machlis._ x 1/20.
_Rangifer tarandus_, the Reindeer, is unique among Deer by reason of the
fact that both sexes wear antlers. These antlers are palmated. The brow
tine and the next or bez tine are also palmated and are directed forwards
and a little downwards. The young are unspotted. The pelage alters in
winter. Like the Moose, the Reindeer is circumpolar. As is well known,
during the Pleistocene period the Reindeer extended its range as far as the
South of France. Even in the historic period it is said to have been hunted
in Caithness.
Reindeer, like so many other particularly Arctic animals, have regular
migrations. In Spitzbergen, for instance, the animal migrates in the summer
to the inland region of the island, and in {299} the autumn back again to
the sea coast to browse upon the seaweed. These migrating herds have been
stated to be led by a large female.
[Illustration]
FIG. 157.--Reindeer. _Rangifer tarandus._ x 1/15.
SUB-FAM. 2. MOSCHINAE.--_Moschus moschiferus_[203] is a native of the
Asiatic Highlands. It is 3 feet or so high, perfectly hornless, and with
very large canines in the male. It is noteworthy that in _Hydropotes_,
where the canines are also very large, horns are absent. These are
examples, perhaps, of correlation. The musk sac (whence the name) is
present on the abdomen of the male only. There is no crumen or suborbital
gland, which is so generally (though by no means universally) present in
Cervidae. But the male has, in addition to the musk glands, glands near the
tail and on the outside of the thigh. Unlike other Deer, the lachrymal bone
of _Moschus_ bears but one orifice. The feet, so far as concerns the
preservation of the outer rudimentary {300} metacarpals, are of the more
ancient type represented in _Alces_, _Hydropotes_, etc. A gall-bladder is
present. The young, as in so many Cervidae are spotted; but the adult is of
a greyish-brown colour.
[Illustration]
FIG. 158.--Musk Deer. _Moschus moschiferus._ x 1/6. (From _Nature_.)
There is no doubt that _Moschus_ is more nearly related to the Cervidae
than to any other Ruminants. It is regarded by Sir W. Flower as "an
undeveloped deer--an animal which in most points (absence of horns, smooth
brain, retention of gall-bladder, etc.) has ceased to progress with the
rest of the group, while in some few (musk gland, mobile feet) it has taken
a special line of advance of its own."
The musk itself, which gives its name to the creature, is found in a gland
on the belly, about the size of a hen's egg. The whole gland is cut out and
sold in this condition. Such quantities of musk deer have been and are
killed for this purpose that the rarity of the animal is increasing. In the
seventeenth {301} century it was so common that the traveller Tavernier
purchased 7673 musk "pods" in one journey, or, according to Buffon, 1663.
The tusks, which recall those of _Hydropotes_, to which _Moschus_ is not
nearly allied, and of _Tragulus_, with which it has of course still less
connexion, are said to be used for the digging up of roots. Its feet, in
relation to its mountain-ranging habits, are very mobile.
EXTINCT SPECIES OF DEER.--It has been already mentioned that the most
primitive kinds of Deer had no horns at all, resembling in this the modern
_Moschus_ and _Hydropotes_, and that with lapse of time went hand in hand
an increasing complexity of antler; the facts of palaeontology harmonising
in the most striking manner with the facts of individual development from
year to year. The oldest forms seem to be more nearly akin to the living
Muntjacs, and their remains occur in the lowest Miocene beds of both Europe
and America. At present the group is confined to the warmer parts of Asia
and some of the islands belonging to that continent.
One of the oldest types is _Amphitragulus_. This genus, which consists of
several species, inhabited Europe, and differed from living Muntjacs in
being totally hornless in both sexes; the skull had no lachrymal fossa or
deficient lateral ossification.
Nearly allied is _Dremotherium_ of similar age and range.
The Middle Miocene has furnished the remains of the genus _Dicroceras_.
This is the earliest Deer in which horns have been found. The horns are, as
the name of the genus implies, bifid, and have, like those of the living
Muntjac, a very long pedicel. This is also a European genus like the last.
From this period we come across true Deer, which commence in the Upper
Miocene and have branched horns. Moreover they belong, at least for the
most part, to the existing genera. One of the most remarkable forms is
_Cervus sedgwicki_ (sometimes placed in a separate genus, _Polycladus_)
from the Forest Bed of Norfolk and from the Upper Pliocene of the Val
d'Arno. This creature was remarkable for its multitudinously-branched
antlers. These end in no less than twelve points. No Deer exists or has
existed in which the horns are so completely branched. They are like those
of a Red Deer exaggerated.
[Illustration]
FIG. 159.--Giraffe. _Giraffa camelopardalis._ x 1/40.
FAM. 7. GIRAFFIDAE.--Undoubtedly the type of a distinct family, Giraffidae,
is the genus _Giraffa_. It is characterised by {302} the long neck, which,
nevertheless, consists of only the normal seven vertebrae, and by the
"horns" which differ from those of all other Ruminants; they are small bony
prominences of the frontal bones, which become fused with the skull, and
which are covered with unmodified skin. They are not shed. Between them is
a median prominence. This cranial armature is present in the female as well
as in the male, and is well developed even in the {303} new-born young. The
orbits are completely encircled by bone, and there is no lachrymal fossa,
so common in Deer and Antelopes. There are no canines above; but these are
present in the lower jaw. The rudimentary digits of other Ruminants have
disappeared in this genus. There are fourteen pairs of ribs as in many
other Artiodactyla. The liver of the Giraffe[204] is, as in many, but not
all, Ruminants, devoid of a gall-bladder; neither has it a caudate or a
Spigelian lobe. The caecum is actually largish (2-1/2 feet in length), but
is relatively very small, as the small and the large intestines measure 196
and 75 feet in length respectively. The Giraffe has a well-marked
"ileo-caecal" gland, found in many Ruminants; its appearance in _Giraffa_
is especially compared by Garrod with its appearance in _Alces_.
Considered by itself, _Giraffa_ forms a very isolated type of Ruminant. But
after we have dealt with certain facts concerning extinct forms clearly
allied to _Giraffa_, the isolation of the family will be found to be less
marked.
The Giraffe ("one who walks swiftly," the word means in Arabic) is, as
every one knows, limited in its range to the African continent. It is not,
however, so familiar a fact that there are two quite distinct species of
Giraffe, one a northern form from Somaliland, and the other South African.
The distinctness of these two, _G. camelopardalis_ and _G. australis_, has
been lately worked out in some detail by Mr. de Winton.[205] The principal
point of difference between them consists in the large size of the median
horn in the Cape species, which is represented by the merest excrescence in
the other species. The Giraffe of West Africa is held to differ from the
northern and southern species, coming nearer to the former. It appears in
the first place to be a larger animal, and slight differences in the skull
have been pointed out. This series of peculiarities may be expressed, for
those who do not object to trinomial nomenclature, by calling this novel
western form _Giraffa camelopardalis peralta_. The existence of the three
horns covered with unaltered skin is the main characteristic of this
Ungulate. But the Giraffe also differs from other Artiodactyles by its
enormously long neck, which enables it to browse upon trees inaccessible to
the common herd {304} of Ruminants. The neck is often supposed to have some
relation to this method of feeding. But a more ingenious explanation of its
inordinate length is that it serves as a watch-tower. The long grass of the
districts inhabited by the animal swarms with Lions and Leopards, which
must be foes. The long neck allows of a wide look out being kept, and it is
noteworthy that the Ostrich, living under similar conditions, is also
renowned for its length of neck. It is the spots upon the Giraffe which
have given it its name of Cameleopard; these spots present in the southern
form a series of chocolate-coloured areas, sharply marked off by white
spaces. Of these spots it is asserted that they serve as a means of
concealing their possessor. Sir Samuel Baker[206] wrote of it in the
following words: "The red-barked mimosa, which is its favourite food,
seldom grows higher than 14 or 15 feet. Many woods are almost entirely
composed of these trees, upon the flat heads of which the giraffe can feed
when looking downwards. I have frequently been mistaken when remarking some
particular dead tree-stem at a distance that appeared like a decayed relic
of the forest, until upon nearer approach I have been struck by the
peculiar inclination of the trunk; suddenly it has started into movement
and disappeared."
The Giraffe, remarked Pliny, "is as quiet as a sheep." The Roman public, to
whom the first Giraffe ever brought into Europe was exhibited, expected
from its name "to find in it a combination of the size of the camel and the
ferocity of a panther." As a matter of fact, Giraffes in captivity are not
always sheep-like in temper. They will kick with viciousness and vigour,
and will even initiate an attack upon their keeper. At the same time they
are singularly nervous creatures, and have been known to die from a shock.
In moving, the Giraffe uses the fore- and hind-limb of each side
simultaneously; this gives to its gait a peculiar rocking motion, the
singularity of which is heightened by the curving movements of the long
neck, which even describes now and then a figure of eight in the air.
_Giraffa camelopardalis_ and the species (?) already referred to are the
only existing Giraffes (of the genus _Giraffa_), and they are not found out
of Africa. Sir Harry Johnston has lately given a brief account of a larger
and more brilliantly coloured species from Uganda {305} which will probably
prove to belong to a distinct genus. It has five horns, the additional pair
being placed above the ears.
Sir Harry Johnston has quite recently made known another genus of
Giraffidae living in the Semliki forest, Belgian Congo district. The skin
and two skulls, as well as the bones of the feet, are known from specimens
sent by Sir Harry Johnston to the Natural History Museum, and briefly
described to the Zoological Society by Professor Ray Lankester.[207] This
creature, of which the native name is "Okapi," is proposed to be called
_Ocapia johnstoni_. The first actual specimens which reached this country
were two bandoliers made from the skin of the flanks, which were striped
black and white, and were not unnaturally held to be portions of the skin
of a new species of Zebra. The animal is of about the size of a Sable
Antelope, and the back and sides are of a rich brown colour; it is only the
fore- and hind-limbs which are striped, the striping being longitudinal,
_i.e._ parallel with the long axis of the body. The head is Giraffe-like,
but there are no external horns; wisps of curled hairs seem to represent
the vestiges of the horns of other Giraffes. The tail is rather short, and
the neck is rather thick and short. The skull is clearly Giraffine. The
basicranial axis is straight, and the fontanelle in the lachrymal region is
very large. Upon the frontal bones near their parietal border is a large
boss on either side, which presumably represents the horn core or "os
cornu." On the mandible the great length of the diastema between the
incisors and premolars is a Giraffine characteristic. The Okapi lives in
pairs in the deepest recesses of the forest.
We are acquainted with a few extinct forms, belonging to _Giraffa_, which
are extra-African in range. _G. sivalensis_ is from the Pliocene of the
Siwalik Hills in India, _G. attica_ from Greece. These remains, however, do
not include the top of the skull, so that it is doubtful whether their
horns were as in _G. camelopardalis_.
A closely-allied genus is the extinct _Samotherium_. This flourished in
Miocene times, and its remains have been found in the Greek island of
Samos. The neck and limbs are shorter than in the Giraffe, and the horns,
longer than in _Giraffa_, are placed just above the orbit upon the frontal
bones alone, instead of upon the boundary line of frontals and parietals as
in _Giraffa_. In several ways, therefore, the existing Giraffe is a more
modified or {306} specialised animal than its forerunner of the Miocene. In
the latter, the male alone carried horns, and in neither sex does the
unpaired median bony excrescence appear. The remains of this genus
(probably even the same species, _S. boissieri_) also occur in Persia.
_Helladotherium_ (there is but one species, _H. duvernoyi_) has its four
limbs of nearly the same length; the skull of the only known example is
hornless; the neck is shorter than in _Giraffa_. It is known from the
Miocene deposits of Pikermi in Greece.
_Palaeotragus_ is a genus which is not referred to the Giraffidae by all
systematists. Its very name, given to it by the eminent French
palaeontologist M. Gaudry, indicates his opinion as to its Antelopine
affinities. The chief and indeed (according to Forsyth Major[208]) the only
reason for placing this Ruminant with the Antelopes is the large size of
the horns. They undoubtedly suggest the horn cores of Antelopes. But they
are placed wider apart than in those animals. It is thought that the
hornless _Camelopardalis parva_ is the female of this species, which is
from Pikermi.
Rather more different from _Giraffa_ is the extinct genus _Sivatherium_,
from the Siwalik deposits of India. Here again there has been some
discussion as to its affinities. Some place it in the neighbourhood of
_Antilocapra_, but most palaeontologists now regard it as a Giraffe. The
main peculiarity of this large beast was the existence of two pairs of horn
cores; the larger are upon the parietal bones, and are of a palmated form,
with a few short tines, which are highly suggestive of those of the Elk
(_Alces_). The shorter anterior pair are upon the frontal bones. The neck
is short, the limbs of equal length, and there are no additional toes upon
the limbs. _Sivatherium_ was almost as large as an Elephant, and in
restorations it is depicted as having a fleshy dilated nose like the Saiga
Antelope; this view is based upon the position and size of the nasal bones.
Hornless skulls have been identified as the female of _Sivatherium_.
_Vishnutherium_, _Hydraspotherium_, and _Bramatherium_ are allied genera.
FAM. 8. ANTILOCAPRIDAE.--This family contains but one genus and species,
the N. American "Pronghorn," _Antilocapra americana_. This animal deserves
a family to itself on account of the singular structure of the horns, which
are intermediate in character {307} between those of the Deer and those of
the Antelopes. They are unquestionably "hollow-horned" Ruminants, in that
there is an osseous horn core, upon which lies the actual horn. This,
however, is softer than in Bovidae, and is semicorneous. It is, indeed,
more like the velvet of the stag's horn. Moreover the horn is branched, and
there are sometimes even three prongs. Furthermore, it is now certainly
known that the Pronghorn sheds its horns not merely occasionally, but with
definite annual periodicity. It so far resembles the Deer. But it must be
borne in mind that in the Deer the horn shedding is a twofold process.
There is first of all the stripping off of the velvet, and secondly the
shedding of a portion of the horn core down to the burr. What happens in
the Prongbuck is the shedding of the true horn only ( = the shedding of the
velvet), _not_ of the horn core. It appears, however, that occasionally
(once in their lifetime?) certain undoubted Antelopes may cast their
horns.[209] Another external character of this animal is the total absence
of "false hoofs," the last vestiges of the second and fifth digits. The
Pronghorn is a gregarious creature running in bands of six up to hundreds.
FAM. 9. BOVIDAE.--This family, more extensive than that of the Cervidae,
contains not only the Oxen, Sheep, and Goats, but also the Antelopes, save
only _Antilocapra_, which must be placed in a family by itself. The only
two points which distinguish all Bovidae from all Cervidae[210] are the
nature of the horns already described, and the polycotyledonary condition
of the placenta. Moreover the horns are usually present in both sexes,
though there are exceptions, such as the Sheep and Goats, and various
genera of Antelopes (_Tragelaphus_, _Tetraceros_, etc.). There are never
the first two phalanges belonging to the rudimentary digits II., V., as
there are in all Deer excepting _Cervulus_. There is as a rule but one
orifice to the lachrymal duct. There are never persistent upper canine
teeth in either sex.
It is exceedingly difficult to separate the Antelopes from the Sheep, Oxen,
and Goats. Their inclusion along with these creatures in one family,
Bovidae, shows that no differences of an important character exist. The
term Antelope is rather of popular than {308} of zoological significance.
As a rule there are horns in both sexes; but this rule is not without
exceptions, of which one is the genus _Strepsiceros_, the Koodoo. Many
other Bovidae are horned in the males only, e.g. _Saiga_, _Tragelaphus_.
The Antelopes further differ from the true Oxen in their more graceful
build, and in the fact that the horns, if they curve at all, generally
curve backwards towards the neck. In the Oxen, on the other hand, the build
is stouter, and the horns usually curve outwards. The same remarks apply to
the Sheep. Such an Antelope, however, as the Eland (_Orias_) is very
Ox-like in habit. Another feature which may be remarked upon, though not of
absolute differential value, is that while the Antelopes are as a rule
smooth and sleek in their skins, the Oxen tend to be rough and shaggy. The
Zebu, however, in this, in its hump, and in general aspect, is far from
being unlike an Eland. But then the Zebu is a domestic race, and we do not
know what the wild stock was like. It is perhaps with the Goats that the
Antelopes have the nearest affinities, and it is difficult to place such a
form as _Nemorrhaedus_, and indeed some others. In the Antelopes as a rule
the middle lower incisors are larger than the lateral ones; in the Sheep
and Goats they are alike in size. The parietal bones, too, in the Antelopes
are moderately large and are much shortened in the remaining Cavicornia,
especially in the Oxen. As the Antelopes are the oldest, so far as we know,
of all bovine animals, one would expect to find them combining the
characters of the rest. But they do this so effectually that a
disentanglement is really impossible. They date from the Miocene. Antelopes
are now limited to Europe, Asia, and Africa; they have always had the same
range, though more abundant in former times in Europe. They preponderate
now in tropical Africa, and abound in genera and species. Messrs. Sclater
and Thomas[211] allow altogether thirty-five genera, of which twenty-four
are exclusively Ethiopian in range.
In the following summary of the group Messrs. Sclater and Thomas's work is
followed. They commence with a section or sub-family of which the type is
the Hartebeest.
_Bubalis_, or _Alcelaphus_ as it is sometimes called, is an African genus,
ranging however into Arabia. These Antelopes are characterised by the long
skull and the doubly-curved horns. There are eight species of the genus, of
which _B. caama_ is the {309} best known; this is the animal known as the
Hartebeest. The Bontebok and Blessbok belong to a closely-allied genus,
_Damaliscus_, distinguished mainly by the fact that the bony base of the
horn cores is not extended upwards, and therefore the parietal bones are
visible when the skull is viewed from in front, which is not the case in
_Bubalis_.
[Illustration]
FIG. 160.--Brindled Gnu. _Connochaetes taurinus._ x 1/20.
The Gnus, _Connochaetes_, are familiar owing to their curious aspect. The
hairy face, and rump and tail like those of a pony are highly
characteristic. The horns are bovine in appearance, standing outwards and
then curving upwards.[212] There are three species of Gnu, all from South
Africa. They are _C. gnu_, _C. taurinus_, and _C. albogulatus_.
Of the Cephalophine section there are two genera:--
_Cephalophus_ is an African genus. These animals are known as Duikerboks;
they are small, and have short non-curved horns in the male sex only. Their
general aspect is not unlike that of certain Deer with simple horns, such
as _Cervulus_. Messrs. Sclater and Thomas allow thirty-eight species. The
{310} smallest species do not exceed the dimensions of a Hare. None are
really large.
_Tetraceros_ is an Indian genus characterised, as its name denotes, by the
fact that it possesses four horns. It is the posterior pair which
correspond to the single pair of _Cephalophus_. The anterior pair, which
are much smaller and are sometimes absent, are a new pair. The female of
this Antelope is hornless. Sheep are occasionally four-horned, and there is
indeed a breed of such in Kashmir. A four-horned Chamois was described by
the late Mr. Alston.
The Klipspringer, _Oreotragus saltator_, is the first type of a third
section; as its name denotes, it is an Antelope with Goat-like habits,
being found particularly among rocks. The horns are short and straight.
This, the only species of the genus, is African in range, of which its
Dutch name gives evidence. A specimen in the Zoological Society's Gardens
(as has been pointed out to me by Mr. Mercer) had the habit of depositing
the secretion of the tear gland upon a mass of concrete in its enclosure,
the secretion thus exuded forming a pointed heap of hardish matter. It may
be that the object of this is to guide its fellows to its whereabouts.
_Ourebia_ is a less-known genus, larger in size, but with horns of the same
character, though longer.
The Grysbok and the Steinbok, genus _Raphiceros_, have similar horns. This
as well as the last two genera have horns in the male only.
One of the smallest of Antelopes belongs to an allied genus; this is
_Neotragus pygmaeus_. It is known as the Royal Antelope, a name apparently
derived from Bosman's statement that the negroes called it "the king of the
harts." Its horns are very small. The height of the animal is only 10
inches. Horns are present in the male alone. The last three genera are
African.
The Cervicaprine series, which is also African, includes the Waterbucks and
Reedbucks, so called on account of their water-loving propensities. As in
the last series, from which they are separated by Sclater and Thomas, but
with which they are united by Flower, there are horns in the male only.
These horns, though not twisted, are long. The typical genus is _Cobus_, of
which there are eleven species. The Waterbuck, _C. ellipsiprymnus_, and the
Sing-sing, _C. unctuosus_, are perhaps the best-known species; the former
is {311} blackish grey, the latter browner in colour. In _C. maria_ and one
or two other species the horns are more curved backwards and again forwards
than in some of the others, where their form is sublyrate.
The Reedbucks, _Cervicapra_, are closely allied to _Cobus_; they are,
however, of smaller size. Here, as in that genus, the females are hornless,
and the horns of the males are of medium size. Five species are referred to
the genus. They are all of a brownish fawn colour. A genus _Pelea_, with
but one species, _P. capreolus_, has been separated on account of the fact
that the horns are nearly straight and that there is no naked patch of skin
beneath the ears. This animal has received its name on account of its
resemblance to the Roebuck.
The Antilopine section includes a number of genera.
The genus _Antilope_ is Indian in range. It includes but one species, _A.
cervicapra_. This Antelope is of medium size, with a brown pelage getting
blacker with years; it is thus known as the Black-buck. The female, which
is hornless, is lighter brown. The horns are long, spirally twisted, and
closely ringed.
_Aepyceros_, with two species, is African. The Palla (_Ae. melampus_) is a
large Antelope, with longish lyrate horns in the male, which are
half-ringed.
The Saiga Antelope, genus _Saiga_, is one of the most remarkable types of
Antelope in its outward appearance. Its nose is very large and inflated,
the two nostrils being quite widely separated, a depression indeed lying
between them dorsally. The horns are lyrate in the male, absent in the
female. The "ovine expression" of this bovine animal is more pronounced in
the female. Corresponding with the clumsy nose are very short nostrils, the
commencement of the narial aperture being therefore very far back. It is
almost suggestive of _Macrauchenia_ in this respect. The fleece is also
Sheep-like. The genus occurred in this country during the Pleistocene. It
is now an inhabitant of Eastern Europe and Western Asia. The only species
is _S. tartarica_.
The Chiru, _Pantholops_, is allied to the Saiga. The horns of the male are
long and nearly straight; they are ringed in front. The muzzle is swollen
in the male; the nostrils are large, and provided with extensive sacs
internally. The colour of this animal, which is exclusively Thibetan in
range, is a pale fawn. The hair, in accord with its habitat, is very
woolly. No living specimens have ever been brought to Europe. This creature
has accumulated much {312} legend. Its blood is believed by the Mongols to
possess virtues, and by means of the rings on the horns fortunes are told.
Naturally the animal is on these grounds hard to stalk and shoot.
[Illustration]
FIG. 161.--Loder's Gazelle. _Gazella loderi._ x 1/10.
The Gazelles, genus _Gazella_, are fairly numerous in species, which are
both Palaearctic and Ethiopian. There are altogether twenty-five of them.
The genus as a whole is characterised by the small or moderate size, the
sandy coloration with white belly, the presence of dark and light stripes
on the face and on the flanks. These streaks, however, are not always
present, and their presence or absence serves to differentiate some of the
species. The horns are usually present in both sexes. The horns are of fair
length, ringed, and of lyrate form.
The Springbok is separated from the rest of the Gazelles, to which genus it
is clearly most nearly related, as a genus _Antidorcas_. This genus differs
from _Gazella_ by having only two lower premolars as in _Saiga_. Otherwise
it resembles the Gazelles; there is but a single species, _A. euchore_,
which is African. {313}
_Ammodorcas_ is closely allied to the Gazelles, but differs from them in
having an elongated neck and also a long tail. _A. clarkei_, the only
species, is limited to Somaliland.
_Lithocranius_, not unlike the last, has a still longer neck, which makes
it almost Giraffe-like; its tail, however, is short. The scientific name is
derived from the "solid stony character of the cranium." In running, this
Gazelle carries the head forward in a straight line with the body. It is
African.
_Dorcotragus_ with one species, _D. megalotis_, is a pigmy Gazelle
restricted to Somaliland. Its likeness, on account of size and in some
other superficial features, to the Klipspringer, led to its original
confusion with that genus (_Oreotragus_).
[Illustration]
FIG. 162.--Sable Antelope. _Hippotragus niger._ x 1/20. The horns of the
specimen figured have not nearly reached their full dimensions.
A sub-family Hippotraginae, or Hippotragine section, includes a number of
Antelopes which agree in the possession of four mammae, and of molars more
like those of the true Oxen, of {314} horns of some length, present in both
sexes, and of a longish tail. They are all African in range.
The type genus _Hippotragus_ has its horns placed above the orbits; they
are not twisted, but curved backwards. There are three species in the
genus. Of these the best known is _H. niger_, the beautiful Sable Antelope.
Its general colour is a rich, dark, glossy brown with white stripes on the
face, and with a white belly. The other species are the Roan Antelope, _H.
equinus_, and the Blaaubok, _H. leucophaeus_, of which the last specimen
was probably killed in 1799.[213]
[Illustration]
FIG. 163.--Beatrix Antelope. _Oryx beatrix._ x 1/16. (From _Nature_.)
The genus _Oryx_ (chiefly African, but also Arabian and Syrian) also
contains a number of species, which are fairly familiar through the fact
that several of them are always on view in the Zoological Society's
Gardens. The genus differs from _Hippotragus_ in that the horns, present in
both sexes, are placed behind the orbits, and slant backwards in a line
with the face. They are annulated. The Leucoryx (_O. leucoryx_) is of a
pale colour, but {315} this is not so marked as in _O. beatrix_, which is
largely white with, however, brown legs. The Gemsbok is a handsome creature
with greyish tawny colour, much darker on the legs, and with a
Gazelle-like, dark, side stripe. It has received its vernacular name on
account of its supposed likeness to the Chamois ("Gemse"), just as the
Rehbok was so-called from its supposed likeness to the Roe Deer, and the
Eland to the Elk. The Beisa (_O. beisa_) is of a similar tawny colour to
the last, and also with darker stripes.
The Addax (_Addax_) of North Africa, Arabia and Syria, has but one species
(_A. nasomaculatus_). The horns are spirally twisted.
[Illustration]
FIG. 164.--Speke's Antelope. _Tragelaphus spekii_ ([female]). x 1/16.
The Tragelaphine section includes the Kudus, Elands, Nilgais, and Harnessed
Antelopes. They are all long-horned (when the horns are present in both
sexes), the horns being twisted; the nose is naked with a slight median
groove, and all are Ethiopian or Oriental in range.
The genus _Tragelaphus_ includes the Harnessed Antelopes, so called on
account of the direction of the stripes suggesting harness. The females are
hornless, and the colours of the two sexes are different. The hoofs are
long and the toes rather unusually separable, which state of affairs is in
accord with the {316} swampy country affected by many. _T. gratus_ and _T.
spekei_ are larger forms; the Boschbok, _T. sylvaticus_, is smaller.
The Kudus, genus _Strepsiceros_, have more markedly twisted horns, which
are absent in the female. The body is vertically striped with white. The
largest species is _S. kudu_; a smaller form, _S. imberbis_, is from
Somaliland.
[Illustration]
FIG. 165.--Eland. _Orias canna._ x 1/25.
The last genus of this section or sub-family is the African Eland, genus
_Oreas_[214] (which it appears should be spelt _Orias_). The Elands are
perhaps more Ox-like in appearance than the other members of this group,
and in both sexes have horns, in which the spiral twisting is more close.
_Orias canna_ is the name of the common Eland. _O. livingstonii_ has been
applied to an East African variety, which has thin and faint lateral
stripes like the other members of the group to which it belongs.
The genus _Boselaphus_ includes only _B. tragocamelus_, the Nilgai, which
is purely Indian in range. The female is hornless, and the horns of the
male are smooth and not long. {317}
The members of the Bovine section or Oxen are to be distinguished from
other hollow-horned Ruminants by their stouter build and by the fact that
the horns stand out from the sides of the skull and are simply curved, not
twisted; and smooth, not annulate like those of other Ruminants. The muffle
is naked, broad, and moist. The Oxen are widely distributed; but are
entirely absent from the Australian region and from South America and
Madagascar.
The true Oxen are perhaps best considered to form but a single genus,
_Bos_. They have, however, been divided into a number of genera. Even the
supposed aberrant _Anoa depressicornis_ of Celebes hardly differs
sufficiently to warrant its separation. In favour of this view, too, is the
extraordinary ease with which different "genera" will cross with each other
and produce fertile offspring. The following is the pedigree of an animal
lately living in the Zoological Society's Gardens. The female offspring of
a male Zebu and a female Gayal was mated with a male Bison. The female calf
was again mated with a Bison and produced a calf, also a female, which
contained therefore the three species, _Bos indicus_, _Bibos frontalis_,
and _Bison americanus_. It is clearly unwise in view of this fact to insist
too much upon generic distinctions in any of those types.[215]
Of this genus the Oriental Gaur (_Bos gaurus_), the Gayal (_B. frontalis_),
and the Banteng (_B. sondaicus_) form a well-marked section, characterised
by their dark coloration and by the somewhat flattened horns.
The Gaur, _Bos gaurus_, has a more concave forehead than its allies; the
horns are less curved than those of the Banteng, and less so than the horns
of the Gayal (_Bos frontalis_). It inhabits the Indian Peninsula; and
extends through Burmah to the extremity of the Malay Peninsula. The Malay
name of this animal is Sakiutan, which simply means wild cattle. It chiefly
frequents wooded hills and is an excellent mountain climber.
_Bos frontalis_, the Indian Gayal, has a white caudal disc like the last
species, but the forehead is flat and the horns curve but little. It is
chiefly known as a tame animal, and its occurrence in the wild state has
been doubted. It has furthermore been suggested that it is merely a tame
race of the Gaur altered {318} slightly through domestication. It is,
however, said not to cross in a state of nature with the Gaur.[216]
[Illustration]
FIG. 166.--Gayal. _Bos frontalis._ x 1/20.
The Banteng, _B. sondaicus_, is distributed through Chittagong, Tenasserim,
and the Malay Peninsula to Java and Borneo. There are apparently two races
of this animal. The species differs from the others by the fact that the
horns are smaller and more curved; there is a white caudal disc; the
forehead is narrower and the skull longer than in the others.
The American Bison and the European Aurochs form another section; they are
indeed extremely alike, specific differences being hardly recognisable. The
Bison of America, formerly present in such numbers that the prairies were
black with countless herds, has now diminished to about a thousand head.
One of the largest of existing Bovidae is the Aurochs, Wisent, or European
Bison, _Bos bonasus_ (or _Bison europaeus_). It is exceedingly like its
American relative. Formerly the animal was much more widely spread than it
is now, extending its range from Europe into North America. It is now
limited to certain districts on the Urals, in the Caucasus, and a herd of
them are kept up through the fostering care of the Emperor of Russia in the
forest of Bielovege in Lithuania. The term "Aurochs" should not really be
applied to this species but to the Wild Cattle, _Bos taurus_. It is,
however, so generally used for the Wisent (which is the German name) that
it {319} is not necessary to change it. The Sclavonic name is Zubr or
Suber. It is a great beast, standing 6 feet or so in height at the
shoulder. It ranged further over Europe well within the historic period. In
the days of Charlemagne it was spread over Germany and was a beast of the
chase. In the year 1848 the Emperor of Russia presented a pair of these
Oxen to the Zoological Society of London. At the time of their presentation
an interesting communication was made to the Society by M. Dolmatoff, on
the method of the capture of these two examples. The creature is not easy
to capture and is alarming to confront. "The eyes," says an old writer,
"are red and fiery; the looks are furious and commanding." It has of course
the shaggy mane and hump of the American animal. The herd in Lithuania was
said to be 1900 in the year 1856. Mr. E. N. Buxton,[217] who has lately
visited the forest, quotes M. Neverli to the effect that at present the
numbers are not more than 700.
[Illustration]
FIG. 167.--Bison. _Bison americanus._ x 1/25.
Allied to this animal, and apparently still nearer to the American Bison,
is the extinct _B. priscus_ of Europe. The Pleistocene Bisons of North
America, _B. antiquus_ and _B. latifrons_, are not remote from the living
forms. Finally, the Miocene _B. sivalensis_ from India, and the Pliocene
_B. ferox_ and _B. alleni_ of North America, take back this group to as
remote a period as any other genus of Oxen. {320}
[Illustration]
FIG. 168.--Yak. _Bos grunniens._ x 1/15.
[Illustration]
FIG. 169.--British Wild Ox. _Bos taurus._ From Vaynol Park, Bangor. x 1/20.
The Yak, _Bos grunniens_, is a long-haired peculiar type, confined to the
Thibetan plateau. _B._ (_Anoa_) _depressicornis_ of Celebes is
characterised by its straight horns; allied to it is _B. mindorensis_
(Philippine Islands), supposed, however, to be a hybrid between {321} it
and some other species. Africa has at least two Buffaloes. We may finally
mention the Wild Ox of Europe, _B. primigenius_, the supposed progenitor of
our domestic cattle, believed to be still surviving in the herds at
Chillingham, Chartley, and elsewhere. This animal is sometimes called the
Aurochs. The Romans spoke of it as the Urus, and it appears to have
formerly attained to more gigantic proportions than at present. It is the
small size of the present race that is the chief objection to tracing them
back to the large Oxen existing near London in 1174, and found sub-fossil
in the Cambridgeshire fens.
[Illustration]
FIG. 170.--Punjab Wild Sheep. _Ovis vignei._ x 1/10.
Of the true sheep, genus _Ovis_, there are a considerable number of
species. The Sheep are to be distinguished from the Goats by their rather
stouter build and by the absence of the beard in the male. The horns are
developed in both sexes, and are usually twisted and often of large size.
The Sheep are almost entirely Palaearctic and Nearctic. They only just get
into the Oriental region. One of the finest species is the great Pamir
Sheep, _O. poli_, whose length reaches 6 feet {322} 7 inches, and height 3
feet 10 inches. The horns of this fine Sheep may measure more than five
feet round the curves. The Rocky Mountain Bighorn (_O. montana_) is a Sheep
ranging along the Rockies as far south as New Mexico, and also to the far
north; they are not confined to the chain of mountains mentioned, but occur
also on the mountains of British Columbia down to those of California. The
horns are not quite as large as those of the last species, but measurements
give a length (along the curve) of 32 to 40 inches.
[Illustration]
FIG. 171.--Himalayan Burrhel Sheep. _Ovis burrhel._ x 1/12. (From
_Nature_.)
[Illustration]
FIG. 172.--Blanford's Sheep. _Ovis blanfordi._ x 1/10. (From _Nature_.)
Just as the Goats are often limited to islands and small stretches of
country, so are the Sheep. Thus Cyprus has a species, _O. ophion_, peculiar
to itself. This, which is known as the Cyprus Mouflon, is limited to a
range of mountains, the Troodos, in that island. In 1878 it was believed
that the animal was nearly exterminated, a flock of twenty-five members
alone surviving. They have, however, since increased. Confined {324} to the
Thibetan plateau are _O. hodgsoni_ and _O. nahura_. Corsica has the
Mouflon, _O. musimon_; and the Barbary Sheep or Arui, _O. tragelaphus_, is
found only in Northern Africa. _Ovis burrhel_ and _O. blanfordi_ are Indian
forms.
[Illustration]
FIG. 173.--Barbary Sheep. _Ovis tragelaphus._ x 1/10.
[Illustration]