UC-NRLF

B M 33T blS

A GUIDK

TO THE

ELEPHANTS

(RECENT AND FOSSIL)

EXHIBITED IN THE DEPARTMENT OF

GEOLOGY AND PALAEONTOLOGY

IN THE

BRITISH MUSEUM (NATURAL HISTORY),

CROMWELL ROAD, LONDON, S.W.7.

ILLUSTRATED BY 32 TEXT-FIGURES.

SECOND EDITION.

PRINTED BY ORDER OF THE TRUSTEES
OF THE BRITISH MUSEUM.

SOLD AT

THE BRITISH Mrskoi (NATrRAT, HISTORY), CROMWKI.I. ROAD, S.W. 7,

B. QUARITCH, LTD., 11 GRAKTOX STREET, NEW BOND STREET, W.I,
DVLAU & Co., LTD., 34-36 MARGARKT Sri;, RB, W. I,

. .. _ . ;, ; • • , • ; ',, • . ' '. : . \V!-.\ C«'R\! > 1 '

1922.

Sliillinq,

A GUIDB

TO THE

ELEPHANTS

(RECENT AND FOSSIL)

EXHIBITED IN THE DEPARTMENT OF

GEOLOGY AND PALAEONTOLOGY

THE

) BRITISH MUSEUM (NATURAL HISTORY),

CROMWELL ROAD, LONDON, S.W. 7.

PRESENTED

BY

The Trustees

OF

THE BRITISH MUSEUM

A GUIDE

TO THE

ELEPHANTS

(RECENT AND FOSSIL)

EXHIBITED IN THE DEPARTMENT OF

GEOLOGY AND PALAEONTOLOGY

) BRITISH MUSEUM (NATURAL HISTORY),

CROMWELL ROAD, LONDON, s.w.?.

ILLUSTRATED BY 32 TEXT-FIGURES.

SECOND EDITION.

LONDON :

PRINTED BY ORDER OF THE TRUSTEES
OF THE BRITISH MUSEUM.

SOLD AT

THE BRITISH MUSEUM (NATURAL HISTORY), CROMWELL ROAD, S.W. 7,

AND BY

B. QUARITCH, LTD., 11 GRAFTON STREET, NEW BOND STREET, W.I,
DULAU & Co., LTD., 34-36 MARGARKT STREET, CAVENDISH SQUARE, W. 1,
AND THE OXFORD UNIVERSITY PRESS, AMEN CORNER, E.C. 4.

1922.
(All rif/hts reserved.}

OU IDE

TO

ELEPHANTS (RECENT AND FOSSIL)

IN THE GALLERY OF FOSSIL MAMMALIA.

THE object of this Guide-book is to give a general account of
the Proboscidea or Elephants, referring especially to the various
stages they passed through in the course of their evolution
from the Eocene period to recent times.

Fortunately for the present purpose, the British Museum
possesses the most extensive series of Proboscidea to be
found anywhere, so that, except in very few instances, readers
can see in the S.E. Gallery of Geology the actual specimens,
or, at any rate, casts of the specimens, upon which the follow-
ing descriptions are based, and can to some extent check
the accuracy of the various statements for themselves.

Before proceeding to the description of the animals, it may
be advisable to refer to the geological horizons or periods of
the Earth's history during which they existed and in the rocks
of which their fossil remains are found; for it is necessary to
know the order in which the different forms appeared on the
Earth, just as in tracing the pedigree of a human family

2 ELEPHANTS.

the dates of the documents upon which it is founded must
be known. It will be seen from Table 1 that the history
of the life of the Earth falls into several great periods, to
which names have been given by geologists. The earliest of
these is called the Primary or Palaeozoic Period, and during
it the only backboned animals were fishes, amphibians (repre-
sented at the present time by newts, frogs, &c.), and, towards
the end, some reptiles. In the next great period, the Secondary
or Mesozoic, the reptiles were of the greatest importance : they
were very numerous and some were of gigantic size. They

TABLE 1.

f QUATERNARY f RECENT.

1 PLEISTOCENE. >
C^NOZOIC <{ f PLIOCENE.

LTEBTIABY.. - j Ml°CENE-

j OLIGOCENE,
I EOCENE.
f CRETACEOUS.
MESOZOIC or SECONDARY J jURASsic.

L TRIASSIC.
f PERMIAN.

CARBONIFEROUS.
PALAEOZOIC or PRIMARY DEVONIAN.

SILURIAN.

ORDOVICIAN.
^ CAMBRIAN.

became fitted for various modes of life, some inhabiting the
land, others the sea; some living on a vegetable diet, others
on flesh. During this period also the birds began to come into
existence, and remains of the most remarkable of the early
forms of birds, Archaopteryx, are shown in table-case 13. At
the same time the first of the warm-blooded mammals arose,
though they were as yet insignificant in size and numbers.
The third period, the Caenozoic, is that with which we
are chiefly concerned. During this the reptiles lost their
importance, while, on the other hand, the mammals took
their place, becoming extremely numerous, many of them

GEOLOGICAL SUCCESSION7. O

of great size and adapted to every kind of life and food. The
latter part of this period, sometimes called the Quaternary,
: extends till the present day, and during it the mammals
still continue to be the prominent backboned animals ; but
one of them, man, has become by far the most important
inhabitant of the world, and, instead of merely being slowly
fitted to new conditions of life, now to a large extent controls
(the conditions, and changes them to suit his own convenience.
At the present day the different main sub-divisions of the
Mammalia are, as a rule, very distinctly marked off from one

Fig. 1.

Upper and lower dentition of pig, showing the presence of the full set of
44 teeth and the low-crowned bunodont cheek-teeth adapted for a soft,
mainly vegetable diet, c., canine ; «'., incisors; m., molars; pm., pre-
molars.

another. For instance, the Carnivora (flesh-eaters, e. g.} tiger,
bear) are now widely different from the Ungulata (hoofed-
animals, e. y.y horse and ox), each of these groups being fitted
for some special manner of life, and particularly for living on
some special kind of food. Thus, the principal character of
most of the Carnivora is the possession of sharp claws and
teeth for killing and devouring other animals; while, on the
other hand, the vegetarian Ungulata have teeth fitted for

B2

4 ELEPHANTS.

grinding vegetable matter and feet adapted solely for moving
from place to place in search of pasturage. If, however, we
trace back through the earlier periods of the Earth's Geological
History the extinct animals from which these quite distinct
modern groups are descended, we find that, in nearly all cases
in which these earlier fossil forms are well known, there is a
tendency for them to become more and more alike. In the
early Eocene, indeed (see Table 1, p. 2), the Garni vora and
Ungulata are not always to be distinguished from one another

Fig. 2.

Skull and mandible of Striped Hyaena, showing the sharp cutting cheek-
teeth adapted for a flesh diet. Lettering as in fig. 1.

with certainty, so that the animals from which they and some
other mammals have descended may be placed in a single
group. Nearly all these early mammals have certain characters
in common: thus in most there are five toes on each foot and
forty-four comparatively simply constructed teeth. Characters
such as these are called " primitive/' and when as time goes
on they become gradually changed in different ways and
adapted for particular purposes, they are said to be more
"specialised." Thus in the horse the foot (fig. 3) is extremely
specialised, in that it has only a single complete toe instead of

SPECIALISATION.

the primitive number, five : its limbs are specially fitted for
swift movement over bard ground. To take another instance,
the teeth of the tiger are said to be highly " specialised/'
because there are only thirty of the original forty-four, and
these have become specially adapted for seizing living prey and
cutting and tearing its flesh. It must be added that all the
characters of a group of animals do not necessarily become
specialised, but that some may remain in the primitive condi-
tion. Thus man, in some ways the most highly specialised of
mammals, still retains the primitive number of five digits on
both limbs.

Fig. 3.

ii

Diagram showing the gradual loss of toes on the fore foot (a) and increase
of complexity in the grinding teeth (b) of successive horse-like Ungulata
from Europe, namely Hyracotherinm (Eocene) 4, Anchitherium (Mio-
cene) :3, HippaHon (Pliocene) 2, Equus (Pleistocene and Recent) 1.

In order to trace back a modern specialised group of mammals
to its early primitive ancestors, a long series of fossil remains
from the successive geological periods is necessary. Unfor-
tunately, in many cases these fossils have yet to be found, but
every year further discoveries are made and gaps of more or
less importance are filled up. The series of changes undergone
by a group of mammals is perhaps best known in the horse

6 ELEPHANTS.

family *, but recent discoveries of remains of early forms of
the elephant group in the Eocene and Oligocene beds of the
Libyan Desert in Egypt have made it possible to trace
the history of the elephants also with considerable accuracy
and completeness.

In the fossil mammals it is the teeth that are of the greatest
importance in settling the relationships of different species to
one another, and in forming an opinion as to their food and
probable manner of life.

The reason for the importance of fossil teeth is that, while in
many ways they are very conservative, long retaining traces of
the earlier forms from which they have originated, nevertheless
they readily undergo change in accordance with the kind of
food they are called upon to seize and masticate. Moreover,
from their hardness they are more frequently preserved than
most other parts of the skeleton. It will be well, therefore,
before considering the teeth of the Proboscidea to give a short
account of mammalian teeth in general, so that the later
descriptions may be understood.

The tooth of a mammal (see "Mammalia" in Index collection
in Central Hall, Bay I.) consists of a root or roots embedded in
a socket in the jaw, and a crown which is exposed and is
adapted for the work it has to perform. In a section of a tooth
(fig. 4) it is seen that the greater part is composed of a hard
material called dentine, and that the crown of the tooth is usually
covered with a still harder substance known as enamel, while
in some cases the root, and in others (usually the more com-
plicated forms) both root and crown, may be coated with a softer
bony matter, called cement. The form of the crown differs
enormously in different animals and in different parts of the
mouth : it may be a simple cone like the canine or dog-tooth,
or it may form a large complicated grinding surface as in the
back teeth of the horse. The teeth are usually divided into
different series according to their particular position in the
mouth and the duties they have to perform (see figs. 1 & 2).
The front teeth implanted in the anterior part of the jaw are

* Casts of specimens showing the gradual " specialisation " of the teeth
and feet in the horses are shown in a case in the North Hall. See also
Pier-case 10 and Table-case 5 in Gallery of Fossil Mammals.

CHARACTERS OF TEETH.

7

mainly concerned in grasping and biting off the food: these are
called incisors. In mammals which, like the pig (fig. 1), possess
the primitive number of teeth, there are three of these on either
side in both upper and lower jaws. Behind these come the

Fig. 4.

Diagrammatic section of various teeth,

I. Section of tusk of elephant, a permanently growing tooth. II. Section
of a young human incisor still growing, the root not yet fully formed.

III. Section of human incisor fully formed, the root being complete.

IV. Section of human molar showing the low-crowned (brachyodont)
condition, the cusps being rounded tubercles (bunodont). V. Section of
the molar of an ox, showing the high (hypsodont) complexly folded
crown. In the figures the enamel is black, the pulp white, the
dentine represented by horizontal lines, the cement by dots.

canines, one on each side above and below ; these are generally
more or less pointed teeth, serving chiefly for fighting or defence.
Behind these again are the cheek-teeth, which have to do mainly

8 ELEPHANTS.

with the breaking up of the food before it is swallowed : in the
complete dentition there are seven of these on either side in both
jaws and they are divided into two groups, (1) four premolars
in front and (2) three molars behind: the premolars,, or at
least those posteriorly, replace the milk-molars of the young
animal, while the molars have no predecessors. It is in the
premolars and molars that the greatest variety of structure is
found, as might be expected, because it is these teeth that
are most affected by the nature of the food. Teeth suitable for
cutting up flesh would be quite unfitted for grinding hard vege-
table matter, and consequently in animals feeding on soft material
the teeth differ widely from those in which the food is hard and
requires much mastication. In the former the crowns of the
teeth are low and their cusps or tubercles are either sharp and
cutting in the case of flesh-feeders (e. g., lions and tigers) or
rounded (bunodont) in the case of animals feeding on a soft
vegetable or mixed diet (e.g., pigs and bears) . Teeth of this low-
crowned sort are called brachyodont (fig. 4, IV). In the case of
animals whose food is hard and requires much grinding, the
wear of the teeth is so great that simply-constructed low crowns
would be quickly worn out, and it must be remembered that
the length of an animal's life is largely dependent on the time
during which its teeth remain in good working order. To meet
the increased wear the crown of the tooth becomes higher and
moves up in the gum as wear takes place, sometimes through-
out the animal's life or only for a time ; teeth of this high-
crowned sort are called hypsodont (fig. 4, V) . With this increase
in height of the crown there is generally greater complication,
resulting from the infolding of the enamel in various ways and
the development of cement on the crown as well as on the roots.
One example of this gradual increase of height and complication
is about to be described in the case of the elephants ; another
extremely good instance is found in the gradual evolution of
the teeth in the horses, as excellently illustrated in the case in
the North Hall and in Table-case 5 (see fig. 3).

From the above account it will be seen that when the full
number of teeth is present there are three incisors, one canine,
four premolars, and three molars on each side in the upper jaw,

DENTITION.

and the same in the lower jaw. This is usually expressed shortly
by a formula, thus :— I.f, C. \, Pm.*, M.§ = ~, the letters
denoting the kind of teeth, the upper numbers the number
of each kind on one side in the upper jaw, the lower
numbers those in the lower jaw, so that in the example above
given there would be eleven teeth on each side in both the
upper and lower jaws, or forty-four in all. As has already been
mentioned, a greater or less number of these teeth may be
wanting in different animals and the formula will differ
accordingly: thus in man it is 1. 1, C.}, Pm.jj, M.y = g or
thirty-two in all. In the cat it is 1. 1, C. ~, Pm.|, M. \ = *
or thirty in all.

Among living mammals the elephants are perhaps the most
remarkable. Not only do they exceed in size all other living
land-animals, but they are further distinguished by the possession
of a mobile trunk or proboscis, which is at once a sensitive
organ of touch and a most efficient means of grasping objects,
both large and small. Furthermore, the structure of their teeth
reaches a degree of complication not to be found in any other
animals. At the same time, though in many respects so peculiar,
in others they retain primitive characters that have been lost in
most of the other Ungulata, with which they are usually classed.
The most notable of these primitive characters is the presence
of the original five toes on each foot,' while in most hoofed-
animals the feet have become " specialised " by the loss of one
or more of the digits.

It is now proposed to describe some of the principal stages
by which the elephants gradually came to be what they are at
the present day, and to show that the earliest-known forms
were much like other primitive hoofed-animals, a condition
to which the pigs and tapirs among living mammals perhaps
most nearly approach. It will be shown that the earliest-
known animal belonging to the Proboscidea or elephants was,
in fact, not unlike a large pig (see fig. 8), though in some
respects an even more primitive creature. From this beginning
we can trace a gradual increase in size in the later forms, a
gradual development of the trunk or proboscis, first as the

10 ELEPHANTS.

upper part of a long snout supported by the elongated lower
jaw, afterwards as the familiar movable organ so characteristic
of the modern elephants. We can also observe the gradual
increase in the size and degree of complication of the grinding-
teeth, accompanied by the complete loss of many of the teeth
found in the earlier forms. Finally, we have materials for
discussing the probable relationship of the elephants to some
other groups of animals.

The table on page 11 shows what are the chief forms
of Proboscideans living at the different periods and their
distribution over the world. It will be seen that the earliest
mammal which can be delinitely called a Proboscidean is
Moeritherium, a small tapir-like creature from the Upper
Eocene beds of the Fayum district of Egypt. This genus
existed also in the Lower Oligocene of the same region, but
was then accompanied by a larger and much more elephant-like
animal, Palaomastodon. At this time Africa was cut off from
the rest of the world to the north by a broad and deep sea which
extended from the Atlantic to the Pacific by way of Northern
India and Southern China, and the separation of Africa pre-
vented these early forms of elephant from wandering into other
regions till after the Lower Oligocene. After PalfPomas-
todon there is a large gap in the series, no fossil Proboscidea
having yet been found in the Upper Oligocene, though no doubt
their remains will be discovered somewhere in the fresh- water
deposits of that age in Egypt. Hitherto no Proboscidean
bones and teeth have been met with again till the Miocene, but
in rocks of that period they are found abundantly, not only in
Egypt, but in Europe, Asia, and North America. It is there-
fore clear that during the long lapse of time after the Eocene,
the deep sea above referred to must to some extent have been
replaced by land, over which the early elephants could spread
outwards from their home in Africa. The vast changes in the
distribution of land and water that took place in this region
will be apparent when it is understood that rocks crowded with
the shells that lived at the bottom of this ancient sea are to-day
found thousands of feet above sea-level in India and elsewhere.

From the Miocene period onwards we meet with elephant-like

GEOLOGICAL SUCCESSION.

il

H

12 ELEPHANTS.

animals in great variety all over the Northern Hemisphere,
"wherever suitable deposits for the preservation of their remains
occur. At the end of the Pliocene period the group also
spread into South America, but at the present day it is totally
wanting in the whole Western Hemisphere.

During the later Miocene and Pliocene periods the head-
quarters of these animals seem to have been in India, for it is
there that we meet with the greatest number and variety of
forms, showing all grades of structure between the Miocene
types above referred to and elephants almost like those now
existing. From the end of the Pliocene to the beginning o'f
the Quaternary Period may be regarded as the time during
which the elephants reached their most flourishing condition,
both in the number of kinds that existed and in the wide
range over which they were spread. After this a gradual
decline in the group took place, till, at the present day, it is
represented by two species only, the African elephant confined to
Tropical Africa, and the Indian elephant found in India, Ceylon,
Burma, the Malay Peninsula, and some of the neighbouring
islands. But for restrictions placed upon their slaughter,
even these la-t remnants of one -of the oldest, and in many
ways the most remarkable, groups of mammals would soon
disappear, just as has happened, for instance, in the case of
the great ground-sloths of South America, the giant lemurs
of Madagascar, and the giant marsupials such as Diprotodon
in Australia.

A more detailed account of the changes that the Proboscidea
have passed through will now be given, the following animals
being selected for description as representing six of the most
important of the successive stages at present known : —

1. Moeritherium, Upper Eocene and Lower Oligocene.

2. PaltPomastodon, Lower Oligocene.

3. Tetrabelodon angustidens , Miocene.

4. ,, longirostris, Lower Pliocene.

5. Stegodon insigms, Pliocene.

6. Elephas, Pliocene. Pleistocene, and Recent.

Some reference will also be made to other types, such as
Dinotherium and Mastodon.

ANCESTRY OF ELEPHANTS.

1 6.

13

Recent
Pleistocene
Upper Plioce fie

ELEPHAS
(shortchin)

Lower Pliocene TETRABELODON

[LONGIROSTRIS STAGE
Uflfier Miocene (shortening chin)

Middle Miocene TETRABELODON

[ANGUSTIDENS STAGE]
Lower Niocene (longchin)

UnnerOligocene ™ 9™*™ from Africa
into Lurone -

Lower Olyocene PAL AEO MASTODON

(lengthening chin)
ditto ditto \

n. [ MOERITHERIUM

Upper Eocene y (short cflm)

Lower Eocene •

Diagram showing- some stages in the gradual increase in size, and alteration
in form, of the skull and mandible occurring in the Proboscidea from
the Eocene to the present day.

14 ELEPHANTS.

MOERITHERIUM.

Moeritherium (figs. 6, 7,8) (Wall-case 43 ; Table-case 24) was
an animal about the size of the tapir, which it must have
much resembled in general appearance. It was common in the
region that is now known as the Fayum in Lower Egypt,
where its fossil remains occur in considerable quantities in
the Upper Eocene beds, intermingled with bones of toothed
whales (Zeuglodori), sea-cows (Eosiren), marine turtles (Pse-
phophorus and Thalassochelys), and snakes (Pterosphenus) , as
well as skeletons of fishes. From this mixture of land and
aquatic animals it may be concluded that Moeritherium lived
near the shore, probably in swamps at the mouth of a great
river, where the remains of both marine and of drowned land-
animals would be mingled and entombed together in the muds
and clays, which accumulated in the estuary and now make up
much of the strata found in this locality. In the Lower
Oligocene beds, overlying those just described, remains of
Moeritherium are also found; here, however, there is no inter-
mingling of marine animals, but instead we find remains of
many remarkable land-mammals, crocodiles, and immense
quantities of trunks of fossil trees embedded in the sands and
gravels of a great river. Probably both the animals and the
tree-trunks were swept away by floods, their remains piled up
in shallows and places where the current was slack, and buried
in the mud and sand carried down by the stream.

The skull of Moeritherium (see fig. 6) differs in no very
marked manner from that of other primitive hoofed-animals,
and shows scarcely any trace of the peculiarities of the
skulls of the later Proboscidea. The most important feature
is the large nasal opening not quite at the end of the
snout, the nasal bones being short ; this indicates that probably
there was already a short proboscis, something like that of the
tapir. Another interesting point is that some of the bones at
the back of the skull are thickened and contain air-chambers ;
in the later elephants this development of air-cells is carried to
such an extent that the whole form of the skull, particularly the

MOKKITIIKKIUM. 15

.posterior portion, is entirely altered by it (see the broken
skull of the Indian elephant in Gallery). The reason for
this swelling of the bones is that, as the head becomes
heavier, owing in great part to the development of the trunk
and tusks, a' larger surface for the attachment of the muscles
which support the head is necessary, and even in the small
Eocene Moeritherium change in this direction had begun.

In Moeritherium the dental formula (see p. 9) is I. ^, C, J,
Pm. |, M. |=£ or thirty-six in all (figs. 5 & 7). From the
formula it will be seen that in the upper jaw only one premolar
is wanting to complete the primitive number, while in the

Fig. 6.

Skull and lower jaw of Moeritherium from the Upper Eocene of the Fay urn,

Egypt. 4 nat. size.

ant.orb., antorbital foramen ; c., canine ; e.r.oc., exoccipital ; />'.,
frontal; i. 1-3, incisors; ju., jugal; m. 1-3, molars; mx., maxilla;
n., nasal; p.a., parietal; par., paroccipital ; pm. 2-4, premolars;
p.mx., premaxilla; pt., post-tympanic process of squamosal ; s.oc., supra-
occipital ; sq., squamosal.

lower jaw an incisor and the canine are missing on each side
in addition to one premolar.

Of the upper incisors the second pair (i. 2) are greatly
enlarged and form strong downwardly directed tusks, the
beginning of the great tusks so characteristic of the later
elephants. The canine (c.) seems to have been quite small and
unimportant, being on the way to disappearance. The pre-
molars (pm.) are separated from the canine by a short interval,
and, as already mentioned, the anterior one of the full denti-
tion is wanting. The remaining premolars are all simpler in

16

ELEPHANTS.

structure than the molars behind them, and consist of three main
cusps only, the two front cusps being arranged in a transverse
line in the third and fourth premolars. These teeth are preceded
by milk-teeth which they displace from above as in the ordinary
mammals ; in the later elephants we shall see that this usual
replacement of milk-teeth by premolars is gradually lost.

Fig. 7.

Upper and lower teeth of Moeritherium.

A. Upper teeth. B. PremaxiHa, large tusk-like second incisor. C. Man-
dible from other side, c., socket of canine ; i. 1-3, incisors ; m. 1-3,
molars ; pm. 2-4, premolars. \ nat. size.

The molars (fig. 7. A, m.) are the most interesting and im-
portant of the teeth, because it is in them that the most nearly
complete series of gradually more and more complicated forms
can be traced. In Moeritherium the crown of each upper molar is
composed of two transversely arranged pairs of knobs, giving
rise to a pair of transverse crests ; there are also in many cases
small posterior knobs — the first trace of the tendency to increase
the number of transverse crests by additions to the back
of the tooth, which is characteristic of the whole group.

MOERITHERIUM.

17

In the lower jaw (figs. 6 & 7 C) the middle incisors (i 1) are
small, the second pair (/ 2) large and tusk-like ; both are
directed forward and their upper surface continues forward the

oo

surface of the spout-like anterior portion of the jaw. The third
incisors, the canine, and the first premolar of the full dentition
are wanting. The remaining three premolars (pm.), which re-
place milk-molars, are simpler than the molars, and only in the

c

18 ELEPHANTS.

third ami fourth is there any '.arrangement of the anterior
cusps to form a transverse ridge. The first and second molars
(fig. 7 C), like those of the upper jaw, consist of two transversely-
arranged pairs of knobs and a posterior knob which is larger
than in the upper teeth. The last lower molar (fig. 10 A) has
a third ridge forming a sort of heel or talon, as it is called ; this
tooth, though much smaller, is remarkably similar to the molars
of some of the earlier forms oE Mastodon.

The skeleton of Moeritherium is not well known, but some of
the most important points in its structure are: — (1) the neck
is proportionated longer than in the later elephants in which it
is much shortened, (2) the hip-bones are narrow, while in the
later forms, owing to the great increase in size and weight, they
are much expanded. The humerus also is rather different,
particularly at its lower end, the great supinator ridge, so
characteristic of the larger Proboscidea, being scarcely developed.
On the other hand, the thigh -bone is extremely like that of a
very small elephant.

To sum up the primitive characters of Moeritherium :—

1 . The skull differs little from that of an ordinary ungulate.

2. There is a full set of three pairs of incisors in the upper,

and only one pair wanting in the lower jaw.

3. There is a canine on each side in the upper jaw.

4. There are three premolars in the upper and lower jaws

replacing milk-molars.

5. The molars have only two transverse ridges and a small

hind lobe ; each transverse ridge is composed of two
distinct cusps ; the third lower molar has also a heel
or talon.

Some of the more important of the Proboscidean characters
are : —

1. The large size of the external nasal opening, its some-

what backward situation, and the, small size of the
nasal bones.

2. The commencement of the development of air-cells in

the bones of the back of the skull.

PALjEOMASTODON. 19

3. The enlargement of the second incisors in both jaws to

form tusks.

4. The transversely ridged character of the molars.

5. The spout-like anterior portion of the lower jaw.

All these characters become much more marked in the next
stage, which is represented by Palaomastodon from the Upper
Eocene of JEgypt.

PAL^OMASTODON.

The genus Paiteomastodon (figs. 9-11, 13 A) (Wall-case 43 ;
Table -case 24) is represented by species varying in size from
an animal little larger than Moeritherium to one nearly as
large as a small elephant, so that in size alone there is a great
advance in the direction of the modern elephants. In the
structure of the skull and teeth, as well as in the rest of the
skeleton, so far as known, the advance is likewise very striking.

In the skull (see fig. 9) the opening of the nostril (nar.) has

Fig. 9.

Skull aud lower jaw of Palcco mastodon, showing the elongated chin with
a pair .of terminal incisors (Li.), from the Lower Oligocene of the
Fayum, Egypt. T^ nat. size.

nar., position of opening of nose ; u.i., upper second i.icisoi* or tusk.

shifted far back from near the end of the snout, though it is
still in front of the orbit of the eye. The nasal bones are still
shorter and smaller than in Moeritherium. At the back of the
-skull the development of air-cells in some of the bones has

20

ELEPHANTS.

enormously increased, but lias not yet invaded the root' of the
skull, so that the sides are only separated by a sharp median
crest. The posterior surface of the skull slopes forward above
the condyles, and there is a deep pit in the middle line for the
attachment of the muscles necessary to support the increasingly
heavy head.

Of the incisor teeth in the upper jaw only the second pair
now remains, and these have been still further enlarged, forming-
Fig. 10.

Posterior lower molars (m2,ma) of (A) Moerithenum^ (B) Palvomastodon,
showing- the increase in the number of ridges of Palceomastodon.
I nat. size.

downwardly directed, curved, somewhat flattened tusks with a
broad band of enamel along their outer sides only. The
canine has disappeared, but there are still three premolars
replacing milk-molars. The anterior premolar is a simple
cone, while the crown of the posterior one consists of two

PAL^OMASTODON. 21

transverse ridges, these teeth being subjected to two distinct
influences, namely, the tendency to the reduction in the
front of the series and the tendency to become more like the
molars at the back. The three molars themselves show a
distinct advance, the crown of each consisting of three transverse
ridges, each ridge composed primarily of t\\ o main cusps which
may, however, show small traces of sub-division into secondary
cusps.

The mandible (figs. 9 & 13 A) differs from that of Moeri-
therium in the much greater prolongation of the spout-like
anterior portion ; this now projects a considerable distance
in front of the skull and is prolonged still further forward by
the single remaining (second) pair of incisor teeth, which meet
in the middle line and from a sort of shovel-shaped extension ;
the edges are worn both 011 the upper and lower surface, so that
these teeth were probably used for grubbing about in the ground
to procure food, and the upper surface must have been further
worn by working against the lower surface of the trunk or
elongated upper lip.

There are only two premolars in the lower jaw, the anterior
of the three milk-molars (figs. 9 & 13 A) falling out without
being replaced from below ; the molars are three in number,
the two anterior with three transverse ridges, the third some-
times having in addition a small heel (fig. 10 B). It should be
noticed that in the full-grown animal all the molars and pre-
molars are in position and use at the same time ; it will be seen
that in the later forms of elephant-ancestors this is not so.

The skeleton, so far as known, is almost exactly like that of
a small elephant, the only important differences being that the
neck is longer and the limbs most likely less massive. The animal,
as a whole (fig. 11), must have been very like a small elephant,
but would be distinguishable by the longer head and neck, and
by the fact that instead of possessing a flexible trunk it had a
long snout, the lower portion consisting of the elongated lower
jaw,, the upper without bony support and probably extending
beyond the lower ; the projecting portion most likely was more
or less flexible and capable of seizing objects, and was the
beginning of the prehensile trunk.

22

ELEPHANTS.

TETRABELODON. 23

The chief steps taken by Palaomastodon in advance of Moeri-
therium towards greater likeness to the later elephants are : —

1. Considerable increase in size.

2. Lengthening of the snout, as shown by the mandible.

3. Loss of canines and all the incisors except the second

pair in both jaws.

4. Three-ridged molars.

5. Greater development of air-cells at back of skull.

6. Shifting further back of the nose-opening and smaller

size of the nasal bones.

7. Greater similarity of the bones of the skeleton to those

of ordinary elephants.

TETRABELODON.

The next stage is found in Tetrabelodon angustidens (figs. 12-
14, 16) (Pier-cases 41, 42; Table-case 23), from the Lower

Fig. 12.

Skull and mandible of Tetrabelodon angustidem, showing the greatly elongated
chin with a pair of terminal cutting incisors (/.£.), from the Lower and
Middle Miocene, France. ^ nat. size.

nar.f position of opening of nose ; u.i.j upper incisors.

Miocene of Northern Africa, Europe, and probably Asia. This
animal is as large as a medium-sized elephant, and its teeth and

24 ELEPHANTS.

skull are much more elephant-like than in Palteomastodon.
Thus the nostrils have shifted still further back, and the great
development of air-cells in the bones at the back of the skull
has led to the disappearance of the ridge along the middle
of the roof, which is now flat. Further, the upper tusks,
which were quite small and flattened in Palaomastodon, are
now large and round, and differ from those of a modern
elephant only in curving downward instead of upward, and in
having a band of enamel along the outer side relatively much

Fig. 13.

jimi

sym.

rn.2

Lower milk-dentition of (A) Palcsomastodon, (B) Tetrabelodon anyustidens.
Showing some of the milk-teeth in situ, with the germs of replacing
premolars. The germs of the 2nd molars are shown at the back of the
jaw. About I nat. size.

i., incisor; m. 1-2, permanent molars ; mm. 3-4, third and fourth
milk-molars ; mm., socket of second milk-molar ; pin. 3-4, premolars ;
sym., symphysis of mandible.

narrower than in Palcsomastodon, in which nearly the whole
outer face of the tooth is enamel-clad. As the tusks increase in
size, the dentine of which they are composed acquires a peculiar
structure, which is shown in transverse sections of the tusks (see

TETRABELODON. 25

Table-case 24), and appears as a series of crossing lines curving
out from the middle of the tooth and giving a pattern like the
engine-turning on the case of a watch. This is quite peculiar to
elephant-tusks, and by it even small pieces of elephant ivory
can be at once distinguished. This structure depends on the
frequent bending of the tubules which make up the dentine, and
one result of it is, that true ivory is one of the most perfectly
elastic of substances, and is therefore specially suitable for making:
billiard balls. This form of ivory is not found in either the upper
or lower tusks of Moeritherium and Palaomastodon or in the
lower tusks of Dinotherium, and appears for the first time in
the large upper tusks of Tetrabelodon angmtidens. In some
of the American Tetrabelodonts in which the lower tusks are
very large, this structure seems to occur.

In the cheek-teeth of Tetrabelodon angustidens (fig. 16 A)
great changes have also taken place. The milk-molars are still
replaced by premolars, but these are quickly dropped out
(fig. 13 B). This is chiefly the result of the great increase in
size of the true molars, particularly of the third, which is not
accompanied by a corresponding increase in the length of the
tooth-bearing portion of the jaws, so that there is not room for
the premolars and molars to remain in position at the same
time. The consequence of this is. that as the posterior molars
are cut they move forward to take up their position in the jaw,
thrusting out the teeth in front of them so that in the adult
only the two large back molars remain on each side in both
jaws, and in old individuals perhaps only the last is left. This
mode of replacement is shown in fig. 13 B. In this species,
while the first and second molars still have only three ridges,
as in Palceomastodon, the last commonly has five, and all are
proportionately very large.

In the mandible (fig. 12) the anterior portion is extremely long
and projects much further beyond the skull than it does in Palao-
mastodon ; and, indeed, it is in this species that we have the
greatest degree of lengthening of the lower ja\v. The two lower
incisors, as in Palaomastodon, help to add to the length, and
were no doubt used for grubbing in the earth. The remarks

xJ(> ELEPHANTS.

made about the upper cheek-teeth are equally true of the
lower. The neck seems to have been a little longer and more
flexible than in the modern elephant, but the limbs and other
parts were much the same. This animal when living (fig. 14)
must have been still more like an elephant than Palteo-
mastodon, and the most noticeable difference would be that
here also, instead of the flexible trunk, there was a long stiff

Fig. 14.

Restoration of Tetrabelodon aiigustidens.

snout, which was supported by the elongated front of the lower
jaw. Probably the end of the upper lip and nose was free and
movable, and may even have been able to grasp objects to some
extent, but the whole arrangement seems to have been rather
clumsy. In most groups of animals as size increases the length
of the neck becomes greater in proportion, so that the animal can
still reach the ground; but in these early elephants, in spite of

TETRABELODON.

27

the great increase in size, the neck actually shortened, audit was
only this extraordinary lengthening of the snout that enabled
the animals to reach the ground. It seems certain that all the
sub-divisions of the Proboscidea must have passed through this
iongirostrine stage.

The next stage in this strange history is found in Tetra-
belodon longirostris (fig. 15, 16) (Pier-cases 41, 42 ; Table-case

Skull and mandible of Tetrabelodon longirostris, from the Lower

Pliocene, Eppelsheim, Hesse-Darmstadt.
»., lower incisor; m. 2-3, second and third molars. About J^ nat. size.

23), aji elephant of which the remains are common in the Lower
Pliocene of Eppelsheim in Germany and other localities. In

4O ELEPHANTS.

this animal the skull, so far as known, does not differ to any
great extent from that of Tetrabelodon angustidens. The teeth,
however, have advanced considerably in size and complication.
The first and second molars may have four or five trans-
verse ridges, while in the last there may be as many as six

Fig. 10.

Second and third lower molars of (A) Tetrabelodon angustidens and
(-B) Tetrabelodon longirostris. £ nat. size.

ridges (fig. 16 B). Only one of the milk- molars is now
replaced by a premolar, and both this and the other milk-
molars are early pushed out by the forward growth of the
large molars, only two of which at most on each side remain

TET11ABELODON.

29

in position in old animals. It is in the lower jaw, how-
ever, that the chief changes have taken place. Here the

Fig. 17.

Mandible of Tetrabelodon (Rhynchotheriuni) dinotherioides, from the
Loup Fork Beds, North-Western Kansas.

A. Eight side of mandible. iV nat. size.

B. Upper view of entire mandible. TV nat. size.

C. Upper view of third right lower molar, g- nat. size.

alv.j alveolus of tusk ; cond., condyle ; cor., coronoid process ;
m3j third lower molars.

elongated anterior part (fig. 15), so striking in the last type, has
become shortened till it projects but little in advance of the

30 ELEPHANTS.

skull, and, although its upper surface is still deeply grooved and
spout-like as in the earlier forms, the lower incisors no longer
meet in the middle line and prolong the spout, but are rounded,
directed downward, and separated from one another. In this
animal it is clear that the lower jaw was shortening np and could
no longer reach the ground, but doubtless the fleshy upper lip
and nose, now freed from their bony support for at least part
of their length, became flexible and better adapted for grasping
the animal's food. In fact, this species must have looked much
the same as a modern elephant, except that it had a longer chin
bearing a pair of small downwardly directed tusks.

In some of the American Tetrabelodons of about the same
age as T. longirostris, the lower tusks, instead of undergoing
reduction, seem to have become greatly enlarged, and at the
same time the symphysial portion of the mandible is slightly
deflected, so that the mandible with its tusks is to some degree
similar to that of Dinotherium. An example of this form of
jaw is seen in the case of the mandible of a Tetrabelodon
(T. dinotherioides) from the Loup Fork Beds (Upper Miocene)
of Kansas, exhibited in Pier-case 42 (fig. ] 7) .

DINOTHERIUM.

Tetrabelodon dinotherioides has no near relationship with
Dinotherium (fig, 18) (Wall-case 43 ; Case C), which forms a
side branch of the Proboscidea, and is widely different from
all the other members of the group. The earliest-known
member of the genus is Dinotherium cuvieri, a comparatively
small animal, which is found in the same deposits as the earliest-
known remains of Tetrabelodon angustidens, and, as in the case
of that species, its ancestors probably lived in Africa, though up
to the present no traces of them have been discovered. In the
later Miocene beds occur a number of species, some of enormous
size (e. g., D, gigantissimum from Roumania). The genus finally
disappears in Lower Pliocene times. The chief peculiarity
of these animals is that the front part of their lower jaw is
turned sharply downward and bears two large tusks (fig. 18,
Case C) ; probably there were no tusks in the upper jaw. The
skull js remarkable for the great expansion of the occipital

DIXOTIIERIUM.

31

surface, which is strongly inclined forward : there seems to
have been scarcely any development of spongy bone in the
occipital region. The teeth are of a much simpler character
than those found in the Mastodon-elephant line. In the upper
jaw of the adult there are two premolars, both simpler than
the molars; of these molars the anterior one consists of three
transverse crests, the other two of two crests only, a notable
peculiarity, since in the other Proboscidea it is the hindermost

Fig. 18.

Skull of Dinothcrium giganteum from the Lower Pliocene of Eppelsheiin,
Hesse-Darmstadt. jV nat. size.

molar which is the most complex. The posterior milk-molar
also has three crests. In the lower jaw in addition to the
down-turned tusks there are two premolars and three molars.
As in the upper jaw, the premolars are simpler than the molars :
the anterior molar has three crests, the second and third two
only, though in the last there may be a trace of a third. In
the young there are three milk-teeth, the hinder one having

32 ELEPHANTS.

three crests, like the anterior true molar. All the molars and
premolars remain in use throughout the animal's life, a condition
already lost in the earliest Tetrabelodons. It has been suggested
that Dmotherium was an aquatic animal, but there is nothing

Fig. 19.

Mandible of a young individual of Mastodon americanus, showing the small
remnants of the lower incisors. In the adult these are lost. From a
Pleistocene deposit, North America. About % nat. size.

in the structure of the limbs to give any support to this idea>
though the lowness of the crowns of the teeth probably
indicates that it lived on soft herbaceous vegetation, such as
may have grown in swampy places.

MASTODON. 33

MASTODON.

In Tetrabelodon longirostris the main characteristics of the
modern elephants are already established, and the later changes
of importance include, (1) the still further reduction of the
inandibular symphysis and the loss of the lower incisors, and
(2) the great increase in the size and complication of the
cheek-teeth. It will be convenient to consider these changes
separately.

In the shortening of the chin the next stage is found in
Pliocene forms like Mastodon atticus ( Pier-case 37) from the
Lower Pliocene of Pikermi. In this the symphysis, though
much shortened, is still more or less spout-like, and in very
young individuals the incisors may be present, though they are
soon shed ; .the absence of these teeth in the adult is the chief
character distinguishing the genus Mastodon (fig. 19) from
Tetrabelodon. One of the best-known species belonging to this
stage of development is Mastodon arvernensis (Pier-cases 37—40),
which is found in Europe in Pliocene deposits; in it the lower
incisors seem to be entirely wanting and the anterior molars
have four transverse ridges. A nearly allied species, M . sivalensis
(fig. 21), from the Pliocene of India, is notable as showing a
tendency to acquire five ridges to its molars.

The region in which the passage from the Mastodons to the
true elephants occurred seems to have been Southern Asia,
where, in a succession of Pliocene and Pleistocene deposits,
there is a complete series of forms passing from the Mastodon
up to the recent Indian Elephant (Pier-cases 34-37). How
far these changes may have gone on in the rest of the Northern
Hemisphere is not known, but the history of the Mastodons
in America is rather different from that of the Old World
forms. The Proboscidea make their first appearance in North
America in the Miocene, and are there represented by a great
variety of forms. Some (see pages 29—30) have the lower tusks
much enlarged and borne in a down-turned symphysis ; these
are regarded by Professor Osborn as belonging to a distinct
genus Uhynchotherium ; one of them is shown in figure 17.
Others are straigl it-jawed Tetiabelodonts, some with compara-
tively short symphyses (e. g., T. productus\ some with the

34 ELEPHANTS.

symphysis greatly elongated (e. g., T. giganteus). These with the
later Mastodons probably represent several waves of immigration
from Asia. Somewhere about the beginning of the Pliocene
some of these North American forms, probably short-jawed
Tetrabelodonts, migrated into South America, where several
peculiar species are found which persisted with little change till
the Pleistocene. Probably the reason why these Mastodons,
as well as M. americanus (Stand B) of North America, con-
tinued in a comparatively primitive condition is to be found
in their isolation and freedom from competition. One Soutli
American species, M. andium (Pier-cases 39, 40), is par-
ticularly interesting on account of its variability, especially in
the length, of the chin and in the presence or absence of the
lower tusks. Some of these differences are due to sex and,.

Fig. 20.

Vertical longitudinal section of a molar tooth of a Mastodon, showing the
low crown, the open valleys between the cross-ridg'es, and the thick
enamel (I), c., dentine. | nat. size.

perhaps age, but the variations are no doubt mainly due
to the fact that the symphysis and lower tusks were undergoing
reduction and that the latter were about to disappear altogether.
Neither in North nor in South America does it appear that the
Mastodons gave rise to more advanced types, arid the presence in
the Pleistocene of North America of true elephants (E. columbi
and E. primigenius) is due to immigration from Asia. No-
species of Elephas reached South America.

To return to the scries of stages of development found in
Southern Asia, the first species that need be considered is
Mastodon cautleyi (Table-case 23), which in the character of its

ELEPHAS. 35

teeth is nearly related to Tetrabelodon Ion giro stris, but the ridges
of the molars are comparatively higher. There is, however, as
yet no cement in the valleys (fig. 20), which are more or less
obstructed by small tubercles, and some of the cusps wear into a
trefoil pattern, as in Tetrabelodon angustidens and T. longirostris.
While, however, the anterior molars are almost identical with
those of T. longirostris , the posterior lower molar is very similar

Fig. 21.

Grinding-surface of a lower molar tooth of Mastodon sivalensis.
From the Lower Pliocene of the Siwalik Hills, India, f- nat. size.

to that of M. laiidens, which in its turn approaches Elephas
(Stegodon) clifti (fig. 22) very closely. In M. latidens there are,
as a rule, five transverse ridges in the second upper molar and
six in the last.

ELEPHAS.

The next stage is represented by Elephas (Stegodon) clifti
(fig. 22) (Pier-cases 35, 36 ; Table-case 24, Stand K). With
this species we reach the true elephants, though the molars
have much lower crowns and fewer transverse ridges than in
the modern species of » Elephas ; and, in fact, these earlier
forms are sometimes separated into another genus called
Stegodon. Since, however, no real line can be drawn between
them and the later types it is perhaps best to call all Elephas , but
distinguish these earlier forms by adding the name Stegodon in
brackets as above. In all this group the lower incisors have
entirely disappeared, the anterior elongation of the chin at
the same time being reduced to a mere peg-like process
(figs. 24 & 31), and a greater or less amount of cement (see

D2

36 ELEPHANTS.

above, p. 8) fills the transverse valleys in the crowns of the
molars (fig. 23). In Elephas (Steyodori) clifti the number of
ridges is greater than in Mastodon latidens. In order to express
briefly the number of ridges in the molars of this and other species,
a formula is used, thus : —in E. (Steyodon) clifti the formula
M 1 ~, M 2 °, M 3 J=|, means that in the first true molar (M 1)
there are in the upper jaw 6-7 ridges, while in this case the
number in the lower is not known. In the second molar (M 2)
there are 6 in the upper, the lower being uncertain, while in
the last molar (M 3) there are 7-8 ridges in both the upper and
lower jaw. It will be seen, therefore, that the numerator of the

Fig-. 22.

G rinding-surface of an upper molar tuoth of Elephas (Stegodon) clifti. From
Lower Pliocene, Siwalik Hills, India. Showing six transverse ridges.

fractions represents the number of ridges in the upper teeth,
the denominator the number in the lower. When the two
numbers are given thus, 6-8, it means that the number of
ridges varies between them. This formula will be employed
below in describing the molars, and a table showing the gradual
increase in the number of ridges will be given at the end.

In E. clifti premolars are still developed, but must have
been pushed out almost as soon as the milk-molars they replace.
The crowns of the molars are quite low, and there is only a
small quantity of cement in the valleys between the ridges.

In Elephas (Steyodori) bombifrons andE. (£.) insiynis (fig. 23),

ELEPHAS (STEGODON).

37

which represent the next stage, the transverse ridges are some-
what more numerous and at the same time are higher, and the
valleys are filled with cement to a greater degree. Nevertheless,

Vertical longitudinal section of molar tooth of Elephas (Steyodori) insignis.
From Lower Pliocene, Siwalik Hills, India. Showing the wide valleys
between the cross-ridges filled with cement (a), the dark band marked
b being the enamel and beneath that the dentine (c). £ nat. size.

these differences are not very marked, and in the case of indi-
vidual teeth it is often difficult to be sure to which of these

Fig. 24.

Skull. and mandible of Elephas yanesa, showing the immense upper tusks
and the shortened chin. From Lower Pliocene, Siwalik Hills, India.
About -g-V nat. size.

species they belong, In some of the Stegodont elephants the
tusks attain an enormous size : for instance, in a skull of
Etephas (Stegodon] ganesa (fig. 24), exhibited in the gallery

38

ELEPHANTS.

(Stand K),the tusks project for a distance o£ 9 ft. 9 in. beyond
the sockets.

* In the next stage we pass from the low-crowned Stegodont
group to animals in which the ridges are considerably higher

Fig-. L>5.

Grinding-surface of an incomplete upper molar of Elephas planifrons.
From Lower Pliocene, Siwalik Hills, India. § nat. size.

and the valleys completely filled with cement ; this is called
the Loxodont group. The most primitive member is Elephas
planifrons (figs. 25 & 26) (Pier-case 34 ; Table-case 24), in which

Fig. 26.

Vertical longitudinal section of molar tooth of Elephas planifrons, showing
the deeper and narrower valleys completely filled with cement («). The
enamel-layer is marked b, the dentine c. •§• nat, size.

the posterior molars may have as many as twelve ridges. This
is the last of the elephants in which premolars have been

ELEPHAS. O1J

observed ; these teeth are small and closely crowded up beneath
the milk-molars, so that when those teeth are greatly worn the
premolars are exposed to view, and are no doubt shed as soon
as, or even before, the posterior part of the niilk-molars they
should replace.

The species E. meridionalis (fig. 27) (Pier-case 33; Table-
case 20), of which remains are found in the Pliocene of Middle
and South Europe, seems to be closely related to E. planifrons,
and is in about the same stage of evolution, or perhaps a little
more advanced. The last molars may have 13-14 ridges,
and in some cases approach the condition seen in E. hysudricus.
E. meridionalis attained enormous dimensions, some individuals

Fig. 27.

Grinding-surface of upper molar of Elephas meridionalis.
Forest Bed, Norfolk. About £ nat. size.

probably standing about 15 feet in height at the shoulder.
Remains of this species occur in the Forest Bed of Norfolk
(see Table-case 20) and in the Upper Pliocene deposits of the
Val d'Arno and the Auvergne. According to Leith Adams
(' British Fossil Elephants/ p. 232) , the molars are distinguished
by the following characters: — " Crowns very broad ; columns
short as compared with E. antiquus, and generally as com-
pared with E. primigenius ; the enamel of the discs thick and
rarely crimped, but usually uneven, looped or channelled ;
plates wide apart, with thick wedges of cement." Numerous
teeth of this species are shown in Table-case 20.

In this species when the plates come into wear the lateral

40

ELEPHANTS.

columns tend to fuse into laminae while the central column
remains for a time distinct and more or less circular in section
(see fig. 27) ; in E. antiquus the lateral columns tend to remain
annular, while the medials unite into a lamella (fig. 28).

Returning to the Indian series, the next stage may be taken
as represented by Elephas hysudricus (Pier-case 33) , in which
there is a considerable increase in the height of the teeth and
in the number of the ridges. The skull is in many respects
similar to that of E. maximus (E. indicus), the modern Indian
elephant, which may have been its direct descendant. E. hysu-
dricus, like E. meridionalis, sometimes attained a very large
size. This species occurs at the end of the Pliocene and

Fisr. 28.

Grinding-surface of second lower molar of Elephas antiquus from the
Pleistocene of Grays, Essex. £ nat. size.

perhaps in the early Pleistocene beds of the Narbada. Con-
temporary with it was E. antiquus (fig. 28) (Pier-case 33 ;
Table-cases 19, 19 A), an elephant found in the late Pliocene and
early Pleistocene of Europe. This species, in which the molar
teeth are relatively small and tusks nearly straight, sometimes-
attained a gigantic size. A skeleton recently found at Upnor,
near Rochester (Pier-case 31), indicates an animal standing
about fifteen feet high at the shoulder, the humerus being
4 ft. 4 in. long (1/322 meters), a foot longer than the humerus.'
of an African Elephant said to have been lift. 4 in. high at
the shoulder. A variety of this species, E. antiquus recki from
East Africa, likewise attained a gigantic size. E. zulu from
Zululand is probably a related form. In India E. antiquus is

ELEPHAS. 41

represented by E. namadicus ; in this species the skull is peculiar
from the development of a sort of overfolded ridge on its-
frontal portion,, forming an overhanging fold on the fore-
head (see Pier-case 34). In the changes that took place
in the distribution of land and water at the end of the
Pliocene and the beginning of the Pleistocene, portions of the
regions inhabited by Elephas antiquus and its varieties became
isolated as islands, and in these restricted habitats the species
became dwarfed, and the dwarf forms in the different islands
at the same time became specifically distinct from one another.
Instances of these small forms are Elephas melitensis, E. mnai-
driensis, E. Cypriotes, and E. creticus (Table-cases 17 A,
21, 21 A). E. melitensis and E. mnaidriensis are found
in Malta. Of the first-named species a small form, sometimes
called E. falconeri, did not stand more than about three
feet high at the shoulder. The ridge-formula of the molar
teeth is :— M 1 g, M 2 £, M 3 jgf. A large collection of
remains of these species, obtained by Admiral Spratt and
Professor Leith Adams, is shown in Table-cases 21 & 21 A.
E. mnaidriensis is also found in Sicily ; probably this species,
which is larger than E. melitensis , represents the intermediate
stage between it and E. antiquus. E. Cypriotes from Cyprus
and E. creticus from Crete were both discovered and described
by Miss D. M. A. Bate, who collected the specimens shown
in Table-case 17 A.

Turning again to the main line, we find that Elephas hysu-
dricus probably passed into some such species as E. armeniacus
(Table-case 17), which in many respects is intermediate between
the Mammoth (E. primiyenius) (Pier-cases 30-32 ; Table-cases
17-19) and the living Indian Elephant (E. maximus). The
Mammoth seems to represent the highest pitch of evolution
attained in the Elephantidse, being in some respects in advance
even of the Indian Elephant. It is here that we meet
with the greatest number of ridges in the molars (fig. 29), the
formula being M 1 ££, M 3 JJlJt M 3 £^. These teeth
represent the culmination of the long series of changes above
described, all tending to increase the efficiency of the molars

42

ELEPHANTS.

as grinding organs. The great size, and especially height, of
the crown gives them a prolonged period of wear, while the
numerous alternating plates of enamel, dentine, and cement,
of different degrees of hardness, ensure that the grinding-
surface will remain sufficiently rough for its purpose through-
out the period during which the tooth remains in use. The
Mammoth was a very widely-distributed form, being found all
over Northern Europe, Asia, and America, and it seems to
have been particularly abundant in Siberia and the islands to the
north, where remains occur in great abundance, and whence
the tusks are actually exported for commercial purposes. The

Fig. 29.

Grinding-surface of molar tooth of the Mammoth (Elephas primigeniui),
showing some still unworn posterior plates. About £ nat. size.

extinction of the Mammoth appears to have been a compa-
ratively recent event, and in Siberia portions of carcases with
the skin and flesh in good preservation are found in the frozen
tundras. An instance of this kind is illustrated by drawings
and photographs on the pillar between Pier-cases 31—32. In
this instance the animal seems to have fallen into a hole and to
have died in its efforts to scramble out. The mouth was found
:still filled with the grass on which the animal was browsing at
the time when it met with the accident. This individual,
restored and mounted in the attitude in which it was found,
is now exhibited in the Academy of Sciences at Petrograd,
and shows with many others that the Mammoth was covered
with a reddish-brown wool and long dark hair, while the tail
ended in a large tuft of hair. A piece of the skin with its

ELEPHAS.

woolly covering and some of the long hair are shown in Pier-
case 31. Further confirmation of this peculiarity of the
Mammoth is found in the rude sketches scratched on ivory
by early Man, who was evidently quite familiar with the
animal. A reproduction of one of these early drawings is
shown near Wall-case 1.

The finest Mammoth skull hitherto collected in England
is shown in. Case M : this specimen was obtained from the

Fig. 30.

Grinding-surface of upper molars of (A) the Asiatic and (B) the
African Elephant. About -| nat. size.

Pleistocene brick-earth at Ilford in Essex. Most of the skeleton
seems to have been found with it, but the bones were
unfortunately destroyed before their interest was recognised.
The tusks in this skull are 10 feet 6 inches in length beyond
the sockets.

In North America several species of Mammoth occur, such
as JZ. imperator and E. jeffersoni ; some of these attained a

44 ELEPHANTS.

much larger size than E. primigenius, and possessed teeth of a
more primitive character, approximating to those of the
probably ancestral E. meridionalis and E. hysudricus.

The Indian elephant, E. maximus (fig. 30 A) (Stands G, J)r
one of the two surviving species of the suborder, is found in India,
Ceylon, and the Malay Peninsula to'Sumatra. The chief pecu-
liarities of the species, distinguishing it from the African elephant,
are the flatness of the forehead, the comparatively small ears,
the presence of a single finger-like process at the front of the
end of the trunk (fig. 31 A). As might be expected from the
wide range of this species, different local forms can be dis-
tinguished, and in some cases these have even been regarded as-
specifically distinct, as, for instance, the elephant of Sumatra,

Fig. 31.

Skull of the African Elephant (Elephas africanus). About j-8 nat. size.

which has been called E. sumafranus. It seems, however, that
all are merely geographical races of the same animal. The
Ceylon form is said to be, as a rule, tuskless, and, although tusk-
bearing forms do occur in the island, they may be either animals
imported from the mainland of India or the result of former
interbreeding with such. In India, also, some individuals, called
Muchnas, are tuskless or have very small tusks (see mounted
skin, Stand G). The Sumatran type differs in being rather
more slightly built, and in possessing a rather longer trunk and
more expanded end to its tail. The elephants from Further
India and the Malay Peninsula are probably also a distinct

ELEPHAS. 45

race which seems especially liable to produce albino forms,
the white elephants of Siam and Burma being well known.

The origin of the other living species of elephant, Elephas
ajricanus (fig. 31), is not very clearly known, owing to the want
of fossil remains. Several closely-allied species, e.g. E. atlariticus,
have been described from the Pleistocene of Algeria, but 110
form that can be regarded as ancestral has been found in the
Tertiary Beds of Africa. It is now generally supposed that
E. africanus arose from some at present unknown Stegodont,
and not as prevously suggested from an E. antiquus-like animal
in which the molar teeth were already more complex.

The African Elephant (Central Hall) to-day ranges very widely
over Africa south of the Sahara, but fossil remains have been
found in Northern Africa and in the south of Europe. It is
distinguished by its convex forehead (Stands H, I), its very

Fig. 32.

A

Tip of trunk of (A) the Asiatic and (B) the African Elephant.

large ears, and by the presence of two finger-like processes on
the tip of the trunk (see fig. 32 B) . The molar teeth are con-
siderably simpler than those of the Indian species (see fig. 30 B),
the ridges being fewer in number and widening out in the
middle in a peculiar manner; the teeth also are relatively
smaller than in other elephants.

It has lately been shown that, although there is only a single
species of African Elephant, nevertheless, in different parts of
the continent, there are different local races which may perhaps
be regarded as subspecies, and are in fact species in the making.
Differences in the form of the skull of these different races can
be detected, but the most striking characters distinguishing
them from one another are the size and shape of the ears.

46 ELEPHANTS.

According to Mr. Lydekker the distinctive features of these
local races are : —

I. In the Addo Bush, or East Cape, Elephant (Elephas
ofricanus capensis) the ears are rather small, somewhat square
in shape, with rounded corners, and a small, sharply pointed,
angular lappet at the lower angle. The forehead falls away
towards the temples, so as to appear highly arched. Several
young individuals of this variety are now shown in the Central
Hall.

II. The West Cape Elephant (E. a. toxotis) has the ears much
larger (4 ft. 5 in. in a female 8 ft. 8 in. high), longer, and
semi-oval in shape ; although, as in the preceding race, sharply
inflected at the lappet. (Later regarded by Lydekker as a
synonym of I.)

III. In the Matabeleland race (E. a. selousi) the ear is much
less elliptical than in the West Cape Elephant, and approaches
more to that of the Camerim race, but agrees with that of the
former in that the lappet underhangs the jaw and chin.

IV. In the West African Elephant (E. a. cyclotis), typically
from South Cameruns, the ears are very large, but of quite
different shape, the contour being oval and the lappet in the
form of half-ellipse. The skin has a mosaic-like appearance,
and its colour is paler grey than in most other races. The
Congo Elephant, which comes very close to this type, has long
and slender tusks.

V. The Masai Elephant (E. a. knochenhaueri] , typically from
German East Africa, has small triangular ears, with the lappet
angulated and pointed. The exhibited specimen (which stands
11 ft. 4 in. in height, with ears measuring 4 ft. 2J in. by 3ft.
5 in.) may belong to this race.

VI. In the Aberdare Elephant (E. a. peeli), typically from
the Aberdare Mountains, British East Africa, the ears are pear-
shaped, with the lappet very long, although somewhat rounded
at the tip.

VII. The Lake Rudolf Elephant (E. a. cavendishi) is nearly
allied to the last, but has broader ears, in which the lappet is
shorter.

VIII. In the Abyssinian, or Sudan, Elephant (E. a. oxyotis)
the ears form an elongated triangle, with the upper border

RELATIONSHIPS OF THE TltOBOSCIDEA. 47

rounded and the lappet very sharply pointed and angular. This
elephant attains very large dimensions.

IX. The N. Somali Elephant (E. a. orleansi), on the other
hand, is small,, with the upper border of the small ears straight
and the lappet short and distinctly defined.

X. In the West Sudan Elephant (E. a. rothschildi) the ears are
in some respects intermediate between those of the Abyssinian
and those of the West African race, although approximating to
the former in the shape of the lappet.

XI. The Albert Nyanza Elephant has been separated as E. #..
albertensis and is characterised by the unusually short and broad
skull.

A dwarf elephant (Elephas pumilio) is known from the Congo-
region, and another species (E/ep/ias fransseni), the so-called
Water Elephant, from the neighbourhood of Lake Leopold IK,
Congo, may also have been a small form. Specimens of the
tusks of this last species are shown in the Central Hall and the
mounted skin is in the East Corridor.

RELATIONSHIPS OP THE PROBOSCIDEA.

The discovery of the Eocene Proboscidea proves that, although
the elephants are no doubt rightly included among the Ungu-
lata or hoofed-animals, they are at the same time very widely
separated from the other members of that group, or at least
from the existing members of it, and seem to have formed an
independent series from the earliest Tertiary times, when they
probably arose from some quite generalised form of the primitive
group called the Condylarthra. A very early side-branch from
the Proboscidean stem is probably represented by the Sirenia
or Sea-cows, aquatic animals which, though now as unlike
elephants as possible, still possess a number of anatomical
peculiarities in common with them, so that this relationship
was recognised long ago. Recently a number of Sirenians have
been found in the Eocene strata of Egypt (Pier-case 30), one
in the same horizon as Moeritherium. These early Sirenians
are much less specialised than the existing forms, having
the full series of teeth and a complete pelvis, and probably a

48

ELEPHANTS.

functional hind limb. In many points they resemble Moeri-
therium, as, for instance, in the form of the brain, teeth, and
pelvis ; and it seems likely that both they and Moeritherium
had a common ancestor in Lower Eocene times. It was pro-
bably a swamp-living creature, some of whose descendants
became gradually more and more exclusively aquatic in their
habits, thus giving rise to the Sirenia, while others became
exclusively terrestrial and gave rise to the elephants as described
above.

TABLE SHOWING THE RIDGE-FORMULA OF THE TRUE MOLARS IN THE
APPROXIMATE LlNE OF DESCENT OF THE ELEPHANTS FROM
Moeritherium.

Ml.

M2.

M3.

Moeritherium

2
2

2
2

2
3

Pal&omastodon

3
3

3

3

3

3

Tetrabelodon angustidens . . .

3

3

8
3

4—5
4=5

„ longirostris .

4
4

4
4

ft— 6
5=0

Mastodon cautleyi

4
4

4
4

5

5

„ latidens

4
4

4—5
4—5

5—6
5—6

Elephas (Stegodon) ciifti . . .

6-7
P

6

7—8
7—8

,, „ bombifrons .

6
7

6—7
7—8

8—9
8—9

insignis . -

7—8
7—10

7—8
8—12

9—11
9—13

Elephas planifrons

7
7

8—9
8—9

10-12
10—13

„ hysudricus

9—12'
9—12

10—12
12—13

13—17
14—18

12

16

24

,, maximus

12

16

24—27

„ primigenius ....

9—15
9—15

14—16
14—16

18-27
18=27

A series of specimens illustrating the gradual increase in the
number of ridges on the molars is shown in Table-case 24.

Printed by TAYLOR and FRANCIS, Red Lion Court, Fleet Street.

BRITISH MUSEUM

(ffATUJLAL HISTORY),

Cromwell Road, London, S.W. 7.

DATS AND HOURS OF ADMISSION.

The Museum is open to the Public, free daily — on WEEK-
DAYS, throughout the year from 10A.M., in

January and February ... ... to 5 P.M.

March to September (inclusive) „ 6 ,,

October, November, and December ., 5 „

on SUNDAYS, throughout the year from 2.30 to 6 P.M.

On Sundays the back galleries in the east wing are closed,
namely : — Fossil Fishes, Fossil Cephalopods, Fossil Shells,
Starfishes &c.? Fossil Corals, Fossil Plants, and Special Fossil
Collections.

The Museum is closed on Good Friday and Christmas Day.

BY ORDER OF THE TRUSTEES,

i

S. F. HARMEB,

I

Director.

\

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