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ON THE CLASSIFICATION

AND

GEOGRAPHICAL DISTRIBUTION

OF THE

MAMMALIA,

BEING THE
LECTURE ON SIR ROBERT READES FOUNDATION,
DELIVERED BEFORE THE

CAnibersity of Cambrivae, in the Senate-Wouse,
MAY to, 1859.

TO WHICH IS ADDED AN APPENDIX
“QN THE GORILLA,”

AND

“ON THE EXTINCTION AND TRANSMUTATION OF
SPECIES.” 7

BY

RICHARD OWEN, FE.RB:S.

READE’S LECTURER IN THE UNIVERSITY OF CAMBRIDGE, SUPERINTENDENT OF THE NATURAL
HISTORY DEPARTMENTS IN THE BRITISH MUSEUM, PRESIDENT OF THE BRITISH
ASSOCIATION FOR THE ADVANCEMENT OF SCIENCE, FOREIGN MEMBER OF
THE INSTITUTE OF FRANCE (ACADEMY OF SCIENCES), Xe.

LONDON:
JOHN W. PARKER AND SON, WEST STRAND.

M.DCCC.LIX.

CAMBRIDGE:
PRINTED BY C. J. CLAY, M.A.
AT THE UNIVERSITY PRESS.

2 ail

TO

THE REV. WILLIAM HENRY BATESON, D.D.,

MASTER OF ST JOHN’S COLLEGE,
AND

VICE-CHANCELLOR OF THE UNIVERSITY OF CAMBRIDGE.

My pear Sir,

I avait myself with pleasure of your permission to
dedicate to you the present Discourse, which owes its existence
principally to your favourable opinion of my ability to dis-
charge the trust which you have done me the honour of confiding
to me.

Believe me to be,
With the highest esteem and respect,

Your obliged and faithful Servant,

RICHARD OWEN.

ce 6, A
ome
oe a 3

wht

Mr Vicre-CHANCELLOR AND GENTLEMEN OF THE
UNIVERSITY,

My first impulse in availing myself of the privilege of
addressing you in this place, is, to give expression to
the deep sense which I entertain of the honour conferred
- on me by my appointment to ‘Sir Robert Reade’s Lecture-
ship,’ especially as it is the first which has been made since
the revival of that ancient foundation. Believe me, Sir, I truly
appreciate the favour of your choice, and am fully impressed
with the responsibilities which it involves. And if my ac-
knowledgments should seem curt or inadequate, I would be-
seech you to believe that this results from the wish not to tres-
pass too long on your most valuable time, but to devote to
the subject selected as much as may be of the period com-
monly allotted to an oral discourse.

In reviewing, for the choice of this subject, the field of
Natural Science in which I am a labourer, I desired to
‘select one that might be treated of with a certain degree
of completeness in a single Lecture, one that would enable
me to submit to you some of the more recent generalisa-
tions in Natural History, and at the same time exemplify
the applicability of that science, as a discipline, to the im-
provement of the intellect, and especially as a sharpener of the
faculties of observation and of methodical arrangement.

I trust that in the attempt to briefly unfold the Classi-
fication and Geographical Distribution of the Mammalia I
may attain the end I have in view.

The generalisation resulting in the idea of the natural
group of animals, so called, is one of ancient date. The
ZooToKA of Aristotle included the same outwardly diverse but
organically similar beings which constitute the MAMMALIA
of modern Naturalists. In that truly extraordinary compen-
dium of zoological and zootomical knowledge, the ‘ Tepi
Cwwy iotopias!,’ animals generally, and by implication the

1 Kd. Schneider, Leipzig, 1811, 4 Vols. Svo.

2

Zootoka, or air-breathing vivipara, are divided according to
the nature of their limbs into three sections:—1st, Dipoda;
2nd, Tetrapoda ; 3rd, Apoda. The first comprised the biped
human race, the second the hairy quadrupeds, the third the
whale-tribe, in which the limbs answering to the legs of man
are wanting. }

The second of these divisions, which includes the great
majority of mammals, and is commonly regarded as the class
itself, Aristotle subdivides into two great groups, according to
the modification of the extremities. In the first group the
foot is multifid, and a part of the digit—finger or toe—is left
free for the exercise of the faculty of touch, the hard nail or
claw being placed upon one side only; in the second group
the digits are inclosed in hoofs: these groups are recognised
in modern Zoology as the UNGUICULATA and UNGULATA.

Aristotle, in the generalised expressions of his observations
on the various conditions of the teeth, has indicated subdi-
visions of the UNGUICULATA according to characters of the
dental system. One subdivision includes those quadrupeds
which have the front teeth trenchant, and the back teeth
flattened, viz. the Pithecoitda or Ape-tribe. Another subdi-
vision includes the quadrupeds with diversified acuminated
front teeth and interlocking serrated back teeth, viz. the Kar-
charodonta, or Carnivora; whilst the animals now known as _
‘Rodents’ are indicated by a negative dental character.

With respect to the hoofed or Ungulate quadrupeds Aris-
totle in his generalisations on the organs of progressive motion
divides them into Dischide, or bisulcate quadrupeds, and
Aschide, or solidungulates, e. g. the horse and ass.

The term <Anepallacta, by which Aristotle signified the
animals in which the upper and lower teeth do not interlock,
is applicable to the herbivorous quadrupeds generally; in
which the Amphodonta, or those with teeth in both jaws,
e.g. the horse, are distinguished by him from those in
which the front teeth are wanting in the upper jaw, e. g.
the ox.

3

The bats were rightly recognised as true Zootoka, and the
genus was defined as Dermaptera.

The apodal Vivipara, which form the third of Aristotle’s
more comprehensive groups, embraces the Ketode, now called
Cetacea, and affords, by its position and co-ordinates in the
great philosopher’s zoological system, one of the most striking
examples of his sagacity and research. In generalising, how-
ever, on modes of reproduction Aristotle includes certain sharks
with the cetaceans, distinguishing the former by their gills, the
latter by their blow-hole.

I ought, also, to remark that, although Aristotle has ex-
emplified groups of animals which agree with many of the
modern Classes, Orders, and Genera, their relative value is not
so defined’; and his, in most respects, natural, assemblages
would have commanded greater attention and been earlier
and more generally recognised as the basis of later systems,
had its immortal author more technically expressed an ap-
preciation of the law of the subordination of characters; but
Aristotle applies to each of his groups the same denomination,
viz. yévos, genus; distinguishing, however, in some cases, the
greater from the less.

Centuries elapsed ere any advance was made in the science
of Zoology as it was bequeathed to the intellectual world by
the mind of Aristotle. Of no other branch of human know-
ledge does the history so strongly exemplify the fearful phe-
nomenon of the arrest of intellectual progress, resulting in the
‘dark ages.’ The well-lit torch which should have guided
to further explorations of the mighty maze of animated nature
was suffered to fall from the master-hand, and left to grow
dim and smoulder through many generations ere it was re-
sumed, fanned anew into brightness, and a clear view re-
gained both of the extent of ancient discovery and of the
right course to be pursued by modern research.

1 See the just and discerning remarks on this subject by Dr WHEWELL, in
his admirable History of the Inductive Sciences, 3rd ed., Vol. 11. p. 289.
B2

4

To Joun Ray, an ornament of this University, I would
ascribe the merit of proposing a classification of the Zootoka,
which first claims attention as in any respect an advance
upon that taught by the Father of Natural History. It is
given in a tabular form in Ray’s Synopsis Methodica Anima-
lium Quadrupedum, and is as follows :—(See p. 5).

In this Table the principle of the subordination of cha-
racters, or of their different values as applicable to groups of
different degrees of generalisation, is clearly exemplified ; and
herein perhaps is its chief value. But, in the exclusion of
the Dipoda and Apoda of Aristotle, Ray manifests a less
philosophical appreciation of the extent and essential nature
of the class Zootoka than his great predecessor. He is also
inferior in the discernment of the real significance of certain
modifications of zoological characters. Aristotle was not de-
ceived either by the claw-like shape of the hoofs of the camel,
or by the degree of subdivision of those of the elephant; he
knew that both quadrupeds were, nevertheless, essentially
Ungulate*.

Linn us first definitely and formally restored the great
natural class I am now treating of to its Aristotelian inte-
erity; and, applying to it that happy instinct of discernment
of significant outward characters which had enabled him to
effect so much for the sister Science of Botany, he proposed
for it the name MAMMALIA.

The active cultivation of the science of observation stimu-
lated by Ray, Linneus and Buffon, had brought to light
instances, €.g. in certain lizards, of viviparous quadrupeds
which differ in structures of classific importance from the
Zootoka tetrapoda of Aristotle. Certain forms of true fishes
were now known to bring forth their young alive, as well as
the fish-like Ketode. he term Zootoka ceased to be appli-
cable, exclusively, to the class of which Aristotle had sketched
out the bounds; and Naturalists gladly accepted and have
since retained the neat and appropriate and truly distinctive

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6

term proposed by Linnxus,—the term which was suggested
by the outward and visible part of that apparatus by which
the warm-blooded viviparous animals exclusively nourish
their new-born young".

Linneus, like Ray, founds his primary divisions of the
class MAMMALIA on the locomotive organs; but his second-
ary divisions or orders are taken chiefly from modifications of
the dentary system. The following is an abridged scheme of
his arrangement?:—

Front teeth, none in eitherjaw . . . Brura.

* Triguieulal Front teeth, cutters 2, laniarieso . . . GULIRES.
= JUCUMe \ Pront teeth, cutters 4, laniaries I. PRIMATES.
4

<j Front teeth, piercers (6, 2, 10), laniaries 1 FERS.

al Woihate Front teeth, i in both upper and lower j jaw. BELLU.
< IJ { Front teeth, none in the upper jaw... -PECORA.
= | Muticate Teeth variable. . . . Ses 3

On comparing the three preceding systems, it will be found
that the most important errors of arrangement have been com-
mitted, not by Aristotle, but by the modern naturalists. Both
Ray and Linneus have mistaken the character of the horny
parts enveloping the toes of the elephant, which do not defend
the upper part merely, as is the case with claws, but embrace
the under parts also, forming a complete case or hoof.

With respect to Linneeus, however, it must be observed,
that although he has followed Ray in placing the elephant in
the unguiculate group of quadrupeds, he has not overlooked
the great natural divisions which the latter naturalist adopted
from Aristotle; and his Ungulata is the more natural in the
degree in which it approaches the corresponding group in the
Aristotelian system.

I now proceed to the arrangement of the Mammalia pro-
posed by Cuvier in the last edition of his classical work
entitled ‘Le Régne Animal distribué @aprés son organisation.’

Adopting the same threefold primary division of the class
MAMMALIA as his predecessors, CUVIER subdivides it into

1 Aristotle knew that the Cetacea were mammiferous: ‘7a’ (6€ do pmév
fuacrovs) £8 évrds, womep derdls.’
2 From the Systema Nature, ed. x11. Holmiz, Tom. I. p. 24.

7

more naturally defined orders, according to various characters
afforded by the dental, osseous, generative and locomotive
systems, which his great anatomical knowledge had made
known to him.

That heterogencous order which Linnzeus—prepossessed
in favour of the easily recognisable outward character by
which he distinguished the class—had characterised by the
‘Mamme pectorales bine: dentes primores incisores: supe-
riores Iv paralleli*,’ was shewn, by the correlation of anatomi-
cal distinctions with the threefold modification of the limbs
of the Primates, to be divisible into as many distinct orders.
The hands on the upper limbs alone, and the lower limbs
destined to sustain the trunk erect, characterised the order
Bimana, the equivalent of the Linnean genus Homo. The
genus Sima of Linneeus, with hands on the four extremities,
became the order Quadrumana of Cuvier. The genus Ves-
pertio with the ‘manus palmate volitantes’ formed the
group Chetroptera, answerable to the Dermaptera of Aristotle.

Ray had pointed out certain viviparous quadrupeds with
a multifid foot as being ‘‘ anomalous species,’”’ instancing as
such “the tamandua, the armadillo, the sloth, the mole, the
shrew, the hedgehog, and the bat.’ The first three species
are associated with the scaly ant-eaters (Manis) of Asia and
Africa, with the Australian spiny ant-eaters (chidna), and
with the more strange duck-moles (Ornithorhynchus) of the
same part of the world, to form the order Hdentata ef Cuvier,
which answers to that called Bruta by Linneus, if the ele-
phant and walrus be removed from it. The rest of Ray’s
anomalous species exemplify the families Chetroptera and
Insectivora of the Cuvierian system, in which they are asso-
ciated with the true Carnivora in an order called ‘Carnas-
siers, answering to the Fere of Linneus.

Cuvier had early noticed the relation of the Austra-
lian pouched mammals, as a small collateral series, to the

1 Tom, cit. p. 24.

8

unguiculate mammals of the rest of the world; ‘some,’ he
writes, ‘corresponding with the Carnivora, some with the
Rodentia, and others again with the Hdentata, by their teeth
and the nature of their food.’ They formed a family of the
Carnassiers in the first edition of the ‘ Régne Animal’, but
were raised to the rank of an order under the name Marsupialia
in the second edition, where they terminate that series of the
Unguiculata, which possess the three kinds of teeth—incisors,
canines and molars.

The hoofed animals (UNGULATA, ‘animaux & sabots’) are
binarily divided into those that do, and those that do not,
chew the cud; the former constituting the order Pachyder-
mata, the latter that of Ruminantia.

The third primary group or subclass of Mammalia is indi-
cated, but without receiving any name distinct from that of
the single order Cetacea exemplifying it in the Cuvierian
system—an order which would be equivalent to the Mutica of
the Linnean system, save that the manatee which Linneus
placed in the same group as the elephant is associated with
the whale in the Régne Animal. 3

The Mammalian system of CUVIER is exemplified in the
subjoined Table :—(See p. 9).

Important as was the improvement it presented on previous
classifications, the progress of anatomical and physiological
knowledge, mainly stimulated by the writings and example
of Cuvier himself, soon began to make felt the defects of his
system. Shortly after its proposition, the zoological mind
began to be disagreeably impressed by the results of the ap-
plication of the characters employed by Cuvier in the forma-
tion of the primary and secondary groups of the class; the
sloth, for example, being placed above the horse, the mole
above the lynx, and the bat above the dog: even the Orni-
thorhynchus paradoxus—shewn by accurate anatomical scru-
tiny to be the most reptilian of the mammalian class—takes

1 8vo., 1816.

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10

precedence of the colossal and sagacious elephant in the Cu-
vierian scheme’.

The profound admiration and respect which I have always
entertained for my chief instructor in Zootomy and Zoology,
never blinded me to the necessity of much modification of his
arrangement of the Mammalia. 'The question, more especi-
ally, of the truly natural and equivalent primary groups of
the class, has been present to my mind whenever I have been
engaged in dissecting the rarer forms which have died at the
Zoological Gardens in London, or on other occasions. But I
propose first to submit to you, as briefly and clearly as I am
able, the results of this store of anatomical knowledge as ap-
plicable to the true organic characters of the class MAMMALIA.

Mammals are distinguished outwardly by an entire or
partial covering of hair’, and by having teats or mamme—
whence the name of the class.

All mammals possess mammary glands and suckle their
young: the embryo or foetus is developed ina womb. Their
leading anatomical character is, the highly vascular and mi-

oe eS OO

SSPE SSS SS Soaaeaenne ee

nutely cellular structure of the lungs, (fig. 1, 2) which are
freely suspended in a thoracic cavity separated by a musculo-
tendinous partition or ‘diaphragm’ from the abdomen, (ib. d.)

1 The modifications consequently proposed by Geoffroy St Hilaire, Illiger,
De Blainville, C. L. Bonaparte, J. E. Gray, Waterhouse, Milne Edwards,
Lesson, Wagner, Nilsson, Oken, Macleay, Sir E. Home, Gervais, and others,
have been cited and commented upon in my Papers communicated to the Lin-
nzan Society (Proceedings, 1857) and the Geological Society (Proceedings, Nov.
1847, pp. 135—140).

2 The foetal Cetacea shew tufts of hair on the muzzle.

11

Mammals, like Birds, have a heart composed of two ven-
tricles and two auricles, and have warm blood: they breathe
quickly ; but inspiration is performed chiefly by the agency
of the diaphragm ; and the inspired air acts only. on the capil-
laries of the pulmonary circulation.

The blood-discs are smaller than in Reptiles, and, save in
the Camel-tribe, are circular in form. The right auriculo-
ventricular valve is membranous, and the aorta bends over the
left bronchial tube.

The kidneys are relatively smaller and present a more
compact figure than in the other vertebrate classes; their
parenchyma is divided into a cortical and medullary portion,
and the secreting tubuli terminate in a dilatation of the excre-
tory duct, called the pelvis: they derive the material of their
secretion from the arterial system. Their veins are simple,
commencing by minute capillaries in the parenchyma and
terminating generally by a single trunk on each side in the
abdominal vena cava: they never anastomose with the mesen-
teric veins.

The liver is generally divided into a greater number of
lobes than in Birds. The portal system is formed by veins
derived exclusively from the spleen and chylopoietic viscera.
The cystic duct, when it exists, always joins the hepatic, and
does not enter the duodenum separately. The pancreatic
duct is commonly single.

The mouth is closed by soft flexible muscular lips: the
upper jaw is composed of palatine, maxillary and premaxillary
bones, and is fixed; the lower jaw consists of two side-halves,
or rami, which are simple or formed by one bony piece, and
are articulated by a convex (fig. 3, 6) or flat condyle to the
base of the zygomatic process, and not to the tympanic ele-
ment, of the temporal bone; the base of the coronoid process
(ib. c) generally extends along the space between the condy-
loid and the alveolar processes. The jaws of Mammals with
few exceptions are provided with teeth, which are arranged
in a single row; they are always lodged in sockets, and never

12

anchylosed with the substance of the jaw. The tongue is
fleshy, well-developed, with the apex more or less free. The
posterior nares are protected by a soft palate, and the larynx
by an epiglottis: the rings of the trachea are generally carti-
laginous and incomplete behind: there is no inferior larynx.
The cesophagus is continued without partial dilatations to the
stomach, which varies in its structure according to the nature
of the food, or the quantity of nutriment to be extracted there-
from.

The trunk-vertebree of Mammalia have their bodies ossified
from three centres, and present for a longer or shorter period
of life a discoid epiphysis at each extremity. They are arti-
culated by concentric ligaments with interposed glairy fluid
forming what are called the intervertebral substances; the
articulating surfaces are generally flattened, but sometimes, as
in the necks of certain Ruminants, they are concave behind
and convex in front. ‘The cervical vertebrae are seven in
number, neither more nor less. The lumbar vertebre are
more constant and usually more numerous than in other
classes of vertebrate animals. The atlas is articulated by
concave articular processes to two convex condyles, which
are developed from the ex-occipital elements, or neurapo-
physes, of the last cranial vertebra. ‘The tympanic element
of the temporal bone is restricted in function to the service
of the organ of hearing, and never enters into the articula-
tion of the lower jaw. ‘The olfactory nerves escape from
the cranial cavity through numerous foramina of a cribriform
plate. The optic foramina are always distinct from one
another.

The scapula is generally an expanded plate of bone; the
coracoid, with two (monotrematous) exceptions, appears as a
small process of the scapula. The sternum consists of a
narrow and usually simple series of bones: the sternal por-
tions of the ribs are generally cartilaginous and fixed to the
vertebral portions without the interposition of a distinct arti-
culation: there are no eristly or bony abdominal ribs or

13

abdominal sternum. The pubic and ischial arches are gene-
rally complete, and united together by bony confluence on the
sternal aspect, so that the interspace of the two pelvic arches
is converted into two holes, called ‘ foramina obturatoria.’

The sclerotic coat of the eye is a fibrous membrane, and
never contains bony plates. In the quantity of aqueous
humour and the convexity of the lens Mammals are generally
intermediate between Birds and Fishes. The organ of hearing
is characterized by the full development of the cochlea with a
lamina spiralis: there are three distinct ossicles in the tympa-
num ; the membrana tympani is generally concave externally ;
the meatus auditorius externus often commences with a com-
plicated external ear, having a distinct cartilaginous basis.
The external apertures of the organ of smell are provided
with moveable cartilages and muscles, and the extent of the
internal organ is increased by accessory cavities or sinuses
which communicate with the passages including the turbinated
bones.

There are few characters of the osseous system common,
and at the same time peculiar, to the class Mammalia. ‘The
following may be cited :—

1. Each half or ramus of the mandible consists of one
bony piece developed from a single centre: the condyle is
convex or flat, never concave. This has proved a valuable
character in the determination of fossils.

2. The second or distal bone, called ‘squamosal,’ in the
‘zygomatic’ bar continued backward from the maxillary arch,
is not only expanded, but is applied to the side-wall of the
cranium, and developes the articular surface for the mandible,
which surface is either concave or flat.

3. The presphenoid is developed from a centre distinct
from that of the basisphenoid.

In no other class of vertebrate animals are these osteo-
logical characters present.

The cancellous texture of mammalian bone is of a finer
and more delicate structure than in Reptiles, and forms a

14

closer network than in Birds. The microscopic radiating
cells are relatively smaller and approach more nearly to the
spheroid form.

The Mammalia, like Reptilia and Pisces, include a few
genera and species that are devoid of teeth; the true ant-
eaters (Myrmecophaga), the scaly anteaters or pangolins
(Manis), and the spiny monotrematous anteater (Zchidna),
are examples of strictly edentulous Mammals. The Orni-
thorhynchus has horny teeth, and the whales (Balena and
Balenoptera) have transitory embryonic calcified teeth, suc-
ceeded by whalebone substitutes in the upper jaw. The
female Narwhal seems to be edentulous, but has the germs
of two tusks in the substance of the upper jaw-bones; one
of these becomes developed into a large and conspicuous
weapon in the male Narwhal, whence the name of its genus
Monodon.

The examples of excessive number of teeth are presented,
in the order Bruta, by the priodont Armadillo, which has
ninety-eight teeth: and in the Cetaceous order by the Cacha-
lot, which has upwards of sixty teeth, though most of them
are confined to the lower jaw; by the common Porpoise, which
has between eighty and ninety teeth: by the Gangetic Dol-
phin, which has one hundred and twenty teeth; and by the
true Dolphins (Delphinus), which have from one hundred to _
one hundred and ninety teeth, yielding the maximum number
in the class Mammalia.

When the teeth are in excessive number, as in the Arma-
dillos and Dolphins above cited, they are small, equal, or
sub-equal, and usually of a simple conical form.

In most other mammals particular teeth have special forms
for special uses; thus, the front teeth, (figs. 2 and 3, ¢,) from
being commonly adapted to effect the first coarse division of
the food, have been called cutters or zncisors; and the back
teeth, (ib. m,) which complete its comminution, grinders or
molars; large conical pointed teeth situated behind the in-
cisors, and adapted, by being nearer the insertion of the biting

15

muscles, to act with greater force, are called holders, tearers,
laniaries, or more commonly canines, (ib. c,) from being well
developed in the Dog and other Carnivora.

It is peculiar to the class Mammalia to have teeth im-
planted in sockets by two or more fangs; but this can only
happen to teeth of limited growth, and generally characterizes
the molars and premolars: perpetually growing teeth require
the base to be kept simple and widely excavated for the per-
sistent pulp. In no mammiferous animal does anchylosis of
the tooth with the jaw constitute a normal mode of attachment.
Each tooth has its peculiar socket, to which it firmly adheres
by the close co-adaptation of their opposed surfaces, and by
the firm adhesion of the alveolar periosteum to the organized
cement which invests the fang or fangs of the tooth.

True teeth implanted in sockets are confined; in the Mam-
malian class, to the maxillary, premaxillary, and mandibular
or lower maxillary bones, and form a single row in each.

They may project only from the premaxillary bones, as in the
‘ Narwhal; or only from the lower maxillary bone, as in
Ziphius; or be limited to the superior and inferior maxillaries
and not present in the premaxillaries, as in the true Rum7-
nantia and most Bruta (Sloths, Armadillos, Orycteropes). In
most Mammals teeth are situated in all the bones above men-
tioned.

The teeth of the Mammalia usually consist of hard un-
vascular dentine, defended at the crown by an investment of
enamel, and everywhere surrounded by a coat of cement.

The coronal cement is of extreme tenuity in Man, Quad-
rumana and the terrestrial Carnivora; it is thicker in the
Herbivora, especially in the complex grinders of the Ele-
phant.

Vertical folds of enamel and cement penetrate the crown
of the tooth in the ruminating and many other Ungulata, and
in most Rodents, characterizing by their various forms the
genera of those orders.

No Mammal has more than two sets of teeth. In some

16

species the tooth-matrix does not develope the germ of a
second tooth, destined to sueceed the one into which the matrix
has been converted; such a tooth, therefore, when completed
and worn down, is not replaced. The Sperm Whales, Dol-
phins, and Porpoises are limited to this simple provision of
teeth. In the Armadillos and Sloths, the want of generative
power, as it may be called, in the matrix is compensated by
the persistence of the matrix, and by the uninterrupted growth
of the teeth.

In most other Mammalia, the matrix of the first-developed
tooth gives origin to the germ of a second tooth, which some-
times displaces the first, sometimes takes its place by the side
of the tooth from which it has originated.

All those teeth which are displaced by their progeny are
called ‘temporary,’ deciduous, or milk-teeth, (figs. 2 and 3,
d,1...4); the mode and direction in which they are displaced
and succeeded, viz. from above downwards in the upper, from
below upwards in the lower, jaw, in both jaws vertically—are
the same as in the Crocodile; but the process is never re-
peated more than once in any mammalian animal. A con-
siderable proportion of the dental series is thus changed; the
second or ‘ permanent’ teeth having a size and form as suitable
to the jaws of the adult, as the ‘ temporary’ teeth were adapted
to those of the young animal.

Those permanent teeth, which assume places not pre-
viously occupied by deciduous ones, are always the most pos-
terior in their position, and generally the most complex in
their form. The term ‘molar’ or ‘true molar’ is restricted
to these teeth (fig. 2 and 3, m). The teeth between them and
the canines are called ‘ premolars,’ (ib. 7); they push out the
milk-teeth, (ib. d,) and are usually of smaller size and simpler
form than the true molars.

Thus the class Mammalia, in regard to the times of form-
ation and the succession of the teeth, may be divided into
two groups, viz. Monophyodonts’ or those that generate a

1 udvos, once; dw, I generate ; ddovs, tooth.

17

single set of teeth, and the Diphyodonts* or those that generate
two sets of teeth. But this dental character is not so asso-
ciated with other organic characters as to indicate natural or
equivalent sub-classes. |

In the Mammalian orders with two sets of teeth, these
organs acquire individual characters, receive special denomi-
nations, and can be determinated from species to species.
This differentiation of the teeth is significative of the high
grade of organization of the animals manifesting it.

Originally, indeed, the names ‘incisors,’ ‘canines,’ and
‘molars,’ were given to the teeth, in Man and certain Mam-
mals, as in Reptiles and Fishes, in reference merely to the
shape and offices indicated by those names; but they are now
used as arbitrary signs, in a more fixed and determinate sense.
In some Carnivora, e.g., the front teeth have broad tubercu-
late summits adapted for nipping and bruising, while the
principal back-teeth are shaped for cutting and work upon
each other like the blades of scissors. The front-teeth in the
: Elephant project from the upper jaw, in the form, size and
direction of long pointed horns. Indeed, shape and size are
the least constant of dental characters in the Mammalia; and
the homologous teeth are determined, like other parts, by
their relative position, by their connexions, and by their
development.

Those teeth which are implanted in the premaxillary bones,
and in the corresponding part of the lower jaw, are called
‘incisors’ (fig. 2,7), whatever be their shape or size. The
tooth in the maxillary bone, which is situated at or near to
the suture with the premaxillary, is the ‘canine,’ as is also
that tooth in the lower jaw (ib. c), which, in opposing it,
passes in front of the upper one’s crown when the mouth
is closed. The other teeth of the first set are the ‘deciduous
molars’ (d. 1—3) ; the teeth which displace and succeed them
vertically are the ‘ premolars’ (p. 1—3) ; the more posterior

1 $is, twice ; d¥w and ddo’s. See Philosophical Transactions, 1850, p. 493.
fR.

18

teeth, which are not displaced by vertical successors, are the
‘molars’ properly so called (m. 1—4).

Hig? 2.

Lower Jaw of a young Opossum (Didelphys).

I have been led, chiefly by the state of the dentition in
most of the early forms of both carnivorous and herbivorous
Mammalia, which flourished during the eocene tertiary periods,
to regard 3 incisors, 1 canine, and 7 succeeding teeth, on each
side of both jaws, as the type formula of diphyodont dentition.

Three of the seven teeth may be ‘premolars’ (fig. 2, p.
1—3), and four may be true ‘ molars’ (ib. m. 1—4); or there
may be four premolars (fig. 3, p. 1—4), and three true mo-
lars (ib. m. 1—3). This difference forms a character of an

Lower Jaw of a young Pig (Sus.)

ordinal group in the mammalian class'. ‘The essential nature
of the distinction is as follows: true molars (ib. m.) are a
backward continuation of the first series of teeth (ib. d.); they
are developed in the same primary groove of the foetal gum;
they are ‘permanent’ because they are not pushed out by the
successional teeth (ib. p.), called ‘dents de remplacement’ by
Cuvier. Seven teeth developed in the primary groove is,

' Outlines of a Classification of the Mammalia, Trans. Zool. Soc. Vol. 1.
Pp. 330 (1839).

19

therefore, the typical number of first teeth, beyond the canines.
If, as in Didelphys (fig. 2), the anterior three develope tooth-
germs which come to perfection in a ‘ secondary groove,’ there
are then 3 deciduous teeth, 3 premolars, and 4 true molars:
if, as in Sus, fig. 3, the anterior four of the ‘ primary’ teeth
develope tooth-germs, which grow in a secondary groove,
there are then 4 deciduous teeth, 4 premolars, and 3 true mo-
lars. The first true molar of the marsupial (fig. 2, m. 1, d. 4), is
thus seen to be the homologue of the last milk-molar of the
placental (fig. 3, d. 4).

The Hog, the Mole, the Gymnure and the Opossum, are
among the few existing quadrupeds which retain the typical
number and kinds of teeth. In a young Hog of ten months
(fig. 3), the first premolar, p. 1, and the first molar, m. 1, are
in place and use together with the three deciduous molars,
d. 2, d. 3, and d.4; the second molar, m. 2, has just begun to
cut the gum; p. 2, p.3, and p.4, together with m. 3, are more
or less incomplete, and will be found concealed in their closed
alveoli’.

The last deciduous molar, d.4, has the same relative supe-
riority of size to d. 3 and d. 2, which m. 3 bears to m. 2 and
m.1; and the crowns of p. 3 and p. 4 are of a more simple
form than those of the milk-teeth, which they are destined to
succeed. When the milk-teeth are shed, and the permanent
ones are all in place, their kinds are indicated, in the genus
Sus, by the following formula :—

oe ae gee
3—3 1—1’ 4—4’

which signifies that there are on each side of both upper and

lower jaws 3 incisors, 1 canine, 4 premolars, and 3 molars,
making in all 44 teeth, each tooth bemg distinguished by its
appropriate symbol, viz. p.1 to p.4,m.1 to m.3. This number
of teeth is never surpassed in the placental diphyodont series.

1 IT recommend this easily acquired ‘subject’ to the young zoologist for a
demonstration of the most instructive peculiarities of the mammalian dentition.
He will see that the premolars must displace deciduous molars in order to rise
into place; the molars have no such relations.

C 2

20

When the premolars and the molars are below this typical
number, the absent teeth are missing from the back part of
the molar series, and usually from the fore part of the pre-
molar series. ‘The most constant teeth are the fourth premolar
and the first true molar. These being known by their order
and mode of development, the homologies of the remaining
molars and premolars are determined by counting the molars
from before backwards, e.g. ‘one,’ ‘two,’ ‘three,’ and the pre-
molars from behind forwards, ‘four,’ ‘three,’ ‘two,’ ‘one.’
The incisors are counted from the median line, commonly the
foremost part, of both upper and lower jaws, outwards and
backwards. The first incisor of the right side is the homo-
type, transversely, of the contiguous incisor of the left side in
the same jaw, and vertically, of its opposing tooth in the
opposite jaw; and so with regard to the canines, premolars,
and molars; just as the right arm is the homotype of the left
arm in its own segment, and also of the right leg of a suc-
ceeding segment. It suffices, therefore, to reckon and name
the teeth of one side of either jaw in a species with the typi-
cal number and kinds of teeth, e. g. the first, second, and
third incisors,—the first, second, third, and fourth premolars,
—the first, second, and third molars; and of one side of both
jaws In any case.

I have been induced to dwell thus long on the dental cha-
racters of the class Mammalia, because they have not been
rightly defined in any systematic or elementary work on
zoology, although an accurate formula and notation of the
teeth are of more use and value in characterizing genera in
this than in any other class of animals.

Mammals may. be surpassed in the rapidity with which
the blood circulates, in the extent and completeness of the
respiratory processes, in bodily temperature, in the concomi-
tant vigour of the muscular actions; all which superiorities,
in Birds, for example, result in those marvellous powers of
flight with which the feathered class is privileged. But in
their psychical phenomena the Mammalia, as a class, excel all

21

other animals. Let me exemplify this by reference to the
reproductive economy in the vertebrate series.

The instinctive sense of dependence upon another, mani-
fested by the impulse to seek out a mate,—which impulse,
even in fishes, is sometimes so irresistible that they throw
themselves on shore in the pursuit,—this first step in the
supercession of the lower and more general law of individual-
or self-preservation, although not first introduced at the ver-
tebrate stage of the animal series, is never departed from after
that stage has been gained. To this sexual relation is next
added a self-sacrificing impulse of a higher kind, viz. the
parental instinct. As we rise in the survey of vertebrate
phenomena, we see the entire devotion of self to offspring in
the patient incubation of the bird, in the unwearied exertions
of the Swift or the Hawk to obtain food for their callow brood
when hatched; in the bold demonstration which the Hen, at
other times so timid, will make to repel threatened attacks
against her cowering young.

Still closer becomes the link between the parent and off-
spring in the Mammalian class, by the substitution, for the
exclusion of a passive irresponsive ovum, of the birth of a
living young, making instinctive irresistible appeal, as soon
as born, to maternal sympathy; deriving nutriment immedi-
ately from the mother’s body, and both giving and receiving
pleasure by that act.

These beautiful foreshadowings of higher attributes are,
however, transitory in the brute creation, and the relations
cease, aS soon as the young quadruped can provide for itself.
Preservation of offsprmg has been superinduced on self-pre-
servation, but there is as yet no self-improvement: this is the
peculiar attribute of mankind. The human species is charac-
terised by the prolonged dependence of a slowly maturing off-
spring on parental cares and affections, in which are laid the
foundations of the social system, and time given for instilling
those principles on which Man’s best wisdom and truest hap-
piness are based, and by which he is prepared for another and

22

a higher sphere of existence. In this destination alone may
we discern an adequate end and purpose in the great organic
scheme developed upon our planet.

The progressive gradations in this scheme will be further
exemplified as I proceed to explain the principles and cha-
racters by which I have been guided in the formation of
the primary groups or divisions of the class Mammalia.

Prior to the year 1836 it was held by comparative ana-
tomists that the brain in Mammalia differed from that in all
other vertebrate animals by the presence of the large mass
of transverse white fibres, called ‘corpus callosum’ by the
anthropotomist; which fibres, overarching the ventricles and
diverging as they penetrate the substance of either hemisphere
of the cerebrum, bring every convolution of the one into com-
munication with those of the other hemisphere, whence the

?

other name of this part—the ‘ great commissure.’ In that year
I discovered that the brain of the kangaroo, the wombat, and
some other marsupial quadrupeds, wanted the ‘ great commis-
sure;’ and that the cerebral hemispheres were connected
together, as in birds, only by the ‘fornix’ and ‘anterior
commissure’.’ Soon afterward, I had the opportunity of
determining that the same deficiency of structure prevailed
in the Ornithorhynchus and Echidna’.

As many other modifications of structure, more or less
akin to those characterizing birds and reptiles, were found to
be associated with the above oviparous type of brain, together
with some remarkable peculiarities in the economy of repro-
duction *, I pointed out that the Mammalia might be divided
into ‘ placental’ and ‘ implacental*.’

Impressed, however, with the fact that such binary divi-
sion, like that which might be based upon the leading differ-
ences of dentition, was too unequal to be natural, the larger

1 See Philosophical Trans. for 1837, p. 87.

2 Art. MonoTREMATA, Cyclopedia of Anatomy and Physiology, Vol. 11. p.383.
3 Art. MARSUPIALIA, tom. cit. p. 257.

4 Art. MAMMALIA, tom. cit. p. 244.

23

group never presenting the same degree of correspondence of.
organic structure as the smaller moiety, I continued to pursue
investigations, with the view of gaining an insight into the
more natural and equivalent primary groups of the Mammalia ;
having my attention more especially directed to the cerebral
organ in this quest.

In 1842, I was able to demonstrate, in the ‘ Hunterian
Course of Lectures’ delivered at the Royal College of Sur-
geons, the leading modifications of the mammalian brain, and
their peculiar value in classification by reason of their asso-
ciation with concurrent modifications of other systems of
organs.

Nevertheless there were genera of Mammals, e.g. the
sloths, anteaters, armadillos, roussettes, giraffes, rhinoceroses,
&e. to which the cerebral test had to be applied. Fortunately
the rare species of these genera successively arrived at the
Zoological Gardens in London, and afforded me the means
_ of applying that test; so that, at length, having dissected
the brain in one species at least, of almost every genus or
natural family of the Mammalian class, I felt myself in a
position to submit to the judgment of my fellow-labourers in
zoology, at the Linnean Society, in 1857, the generalised
results of such dissections, comprising a fourfold primary divi-
sion of the MAMMALIA, based upon the four leading modi-
fications of cerebral structure in that class.

In some mammals the cerebral hemispheres are but feebly
and partially connected together by the ‘fornix’ and ‘ ante-
rior commissure:’ in the rest of the class the part called
‘corpus callosum’ is added, which completes the connecting
or ‘commissural’ apparatus.

With the absence of this great superadded commissure’ is
associated a remarkable modification of the mode of develop-
ment of the offspring, which involves many other modifica-
tions; amongst which are the presence of the bones called
‘marsupial,’ and the non-development of the deciduous body

1 On the Structure of the Brain in Marsupial Animals, Philos. Trans. 1837.

24

concerned in the nourishment of the progeny before birth,
called ‘placenta; the young in all this ‘implacental’ divi-
sion being brought forth prematurely, as compared with the
rest of the class.

This first and lowest primary group, or subclass, of Mam-
malia is termed, from its cerebral character, LyencepHaa’,
—signifying the comparatively loose or disconnected state of
the cerebral hemispheres. ‘The size of these hemispheres
(fig. 4, A) is so small that they leave exposed the olfactory gan-
glions (a), the cerebellum (c), and more or less of the optic
lobes (B); their surface is generally smooth; the anfractuosi-
ties, when present, are few and simple.

Brain of Opossum.

Brain of Beaver.

The next well marked stage in the development of the
brain is where the corpus callosum (indicated in fig. 5, by the
dotted lines d, d) is present, but connects cerebral hemispheres
as little advanced in bulk or outward character as in the pre-
ceding subclass; the cerebrum (A) leaving both the olfactory
lobes (a) and cerebellum (c) exposed, and being commonly

1 dw, to loose ; éyxédanos, brain.

25

smooth, or with few and simple convolutions in a very small
proportion, composed of the largest members, of the group.
The mammals so characterized constitute the subclass Lisszn-
CEPHALA’ (fig. 5).

The third leading modification of the Mammalian cere-
brum is such an increase in its relative size, that it extends
over more or less of the cerebellum; and generally more or
less over the olfactory lobes. Save in very few exceptional
cases of the smaller and inferior forms of Quadrumana (fig. 6),
the superficies is folded into more or less numerous gyri or
convolutions (fig. 7),—whence the name Grrzncepnara, Which
I propose for the third subclass of Mammalia’.

c

Brain of Marmoset Monkey.
(Nat. size).

Brain of Chimpanzee.
(Half nat. size),

In Man the brain presents an ascensive step in develop-
-ment, higher and more strongly marked than that by which
the preceding subclass was distinguished from the one below
it. Not only do the cerebral hemispheres overlap the olfac-
tory lobes and cerebellum, but they extend in advance of the

1 \ucods, smooth ; éyKépandos, brain.
, xy
2 yupdw, to wind about ; éyxédados, brain,

26

one and further back than the other (figs. 8& 9). Their pos-
terior development is so marked that anthropotomists have

assigned to that part the character and name of a‘ third lobe :’

it is peculiar and common to the genus Homo: equally pecu-

liar is the ‘posterior horn of the lateral ventricle’ and the

‘hippocampus minor,’ which characterize the hind lobe

of each hemisphere.
The superficial grey
matter of the cere-
brum, through the
number and depth
of the convolutions,
attains its maximum
of extent in Man.
Peculiar mental
powers are associ-
ated with this high-
estform of brain, and
their consequences
wonderfully illus-
trate the value of the
cerebral character ;
according to my es-
timate of which, I
am led to regard the
genus [Homo as not
merely a representa-
tive of a distinct or-
der, but of a distinct
subclass,of the Mam-
malia, for which I
propose the name

of ARCHENCEPHALA

(ages Oye:

Fig. 8.

Ib. Side view, one-third nat. size.

1 dpxw, to overrule ; éyxépados, brain.

20

With this preliminary definition of the organic characters,
which appear to guide to a conception of the most natural
primary groups of the class MAMMALIA, I next proceed to
define the groups of secondary importance, or the subdivisions
of the foregoing subclasses.

The Lyencephalous Mammalia are unguiculate: some
have the ‘optic lobes’ simple, others partly subdivided, or
complicated by accessory ganglions, the lobes being then
called ‘bigeminal bodies.’

The Lyenceruata with simple optic lobes are ‘ edentulous’
or without calcified teeth, and are devoid of external ears,
scrotum, nipples, and marsupial pouch: they are true ‘ testi-
conda:’ they have a coracoid bone extending from the
scapula to the sternum, and also an epicoracoid and epi-
sternum as in Lizards: they are unguiculate and pentadactyle,
with a supplementary tarsal bone supporting a perforated
spur in the male. The order so characterized is called
‘MONOTREMATA, in reference to the single excretory and
generative outlet, which, however, is by no means peculiar to
them among Mammalia. It includes two genera—LEchidna
and Ornithorhynchus. Of the first, the species are terrestrial,
insectivorous, chiefly myrmecophagous, having the beak-like
slender jaws, and long cylindrical tongue of the true anteaters ;
but they are covered, like the hedgehog, with spines. Of the
second genus, the species are aquatic, with a flattened beak,
like that of a duck, which is used in the anserine manner to
extract insects and worms from the mud: but they are clothed
with a close fine fur like that of a mole, whence the name
‘duck-mole’ by which these anomalous quadrupeds are com-
monly known to the colonists. Both genera of Monotremes
are strictly limited to Australia and Tasmania.

The Lyenceruera with divided optic lobes, forming the
‘corpora bigemina’ and ‘quadrigemina’ of anthropotomists,
have teeth, and with rare exceptions, the three kinds, viz.
incisors, canines, and molars. They are called MArsupia-
LIA, because they are distinguished by a peculiar pouch or

28

duplicature of the abdominal integument, which in the males
is everted, forming a pendulous bag, and in the females is
inverted, forming a hidden pouch containing the nipples and
usually sheltering the young for a certain period after their
birth: they have the marsupial bones in common with the
Monotremes; a much varied dentition, especially as regards
the number of incisors, but usually including 4 true molars ;
and never more than 3 premolars’ (fig. 2): the angle of the
lower jaw (ib. a) is more or less inverted’.

With the exception of one genus, Didelphys, which is Ame-
rican, and another genus Cuscus, which is Malayan, all the
known existing Marsupials belong to Australia, ‘Tasmania,
and New Guinea. ‘The grazing and browsing Kangaroos are
rarely seen-abroad in full daylight, save in dark rainy weather.
Most of the Marsupialia are nocturnal. Zoological wanderers
in Australia, viewing its plains and scanning its scrubs by
broad daylight, are struck by the seeming absence of mam-
malian life; but during the brief twilight and dawn, or by the
light of the moon, numerous forms are seen to emerge from
their hiding-places and illustrate the variety of marsupial life
with which many parts of the continent abound. We may
associate with their low position in the mammalian scale the
prevalent habit amongst the Marsupialia of limiting the exer-
cise of the faculties of active life to the period when they are
shielded by the obscurity of night.

The premature birth of the offspring, and its transference
to the tegumentary pouch, in which it remains suspended to
the nipple for a period answering to that of uterine life in
higher mammals, relate to the peculiarities of the climate of
Australia.

The adventurous and much-enduring explorers of that
continent bear uniform testimony to the want of water as the

1 Outlines of a Classification of the Marsupialia, Trans. Zool. Soc. Vol. 11. 1839.

2 For other Osteological and Dental characteristics of the Marsupialia, see
the paper above cited, and that On the Osteology of the Marsupialia, Trans.
Zool. Soc. Vol. 11. p. 379 (1838).

29

chief cause of their sufferings and danger. During the dry
season the rivers are converted into pools, ‘few and far be-
tween ;’ and the drought is sometimes continued so long as to
dry up these. An ordinary non-marsupial quadruped, such
as the wild cat or fox, having deposited her young in the nest
or burrow, would in such a climate, at the droughtiest period
of her existence, be compelled to travel a hundred, perhaps
two hundred miles, in order to quench her thirst. Before she
could return her blind and helpless litter would have perished.
By the marsupial modification the mother is enabled to carry
her offspring with her in the long migrations necessitated by
the scarcity of water.

With the climatal peculiarities of Australia, therefore, we
may connect the peculiar modifications of those members of
the mammalian class which are most widely distributed over
that continent. But the principle of final causes receives more
especial illustrations from the contingent particulars of the
marsupial organization. ‘The new-born Kangaroo is an inch
in length, naked, blind, with very rudimental limbs and tail:
in one which I examined the morning after the birth, I could
discern no act of sucking: it hung, like a germ, from the end
of the long nipple, and seemed unable to draw sustenance
therefrom by its own efforts. The mother, accordingly, is
provided with a peculiar adaptation of a muscle (cremaster) to
the mammary gland, by which she can inject the milk from
the nipple into the mouth of the pendulous embryo. Were
the larynx of the little creature like that of the parent, the
milk might—probably would—enter the windpipe and cause
suffocation: but the foetal larynx is cone-shaped, with the
opening at the apex, which projects, as in the whale-tribe,
into the back aperture of the nostrils, where it is closely em-
braced by the muscles of the ‘ soft palate.’ The air-passage
is thus completely separated from the fauces, and the injected
milk passes in a divided stream on either side the base of
the larynx into the cesophagus. These correlated modifica-
‘tions of maternal and fcetal structures, designed with especial

30

reference to the peculiar conditions of both mother and off-
spring, afford, as it seems to me, irrefragable evidence of
Creative foresight.

The Zissencepnata, or smooth-brained placental Mam-
malia, form a group which I consider as equivalent to
the Lyenceryata or Implacentals; and which includes the
following orders, Lodentia, Insectivora, Cheiroptera and
Bruta.

The RopEntrIA are characterized by two large and long
curved incisors in each jaw, separated by a wide interval
from the molars; the teeth being so constructed, and the
jaw so articulated, as to effect the reduction of the food
to small particles by acts of rapid and continued gnawing,
whence the name of the order. The orbits are not separated
from the temporal fosse. The male glands pass periodically
from the abdomen into a temporary scrotum, and are asso-
ciated with prostatic and vesicular glands. The placenta is
commonly discoid, but is sometimes a circular mass (Cavy),
or flattened and divided into three or more lobes (Lepus).
The Beaver and Capybara are the giants of the order, which
chiefly consists of small, numerous, prolific and diversified
unguiculate genera, subsisting wholly or in part on vegetable
food. Some Rodents, e.g. the Lemmings, perform remark-
able migrations, the impulse to which, unchecked by dangers
or any surmountable obstacles, seems to be mechanical.
Many Rodents build very artificial nests, and a few manifest
their constructive instinct in association. In all these inferior
psychical manifestations we are reminded of Birds. Many
Rodents hibernate like Reptiles. They are distributed over
all continents. About two-thirds of the known species of
Mammalia belong to the Rodent order.

The transition from the Marsupials to the Rodents is
made by the Wombats; and a transition from the Marsupials
is made, by an equally easy step, through the smaller Opos-
sums to the InsecTIVORA. ‘This term is given to the order of
small smooth-brained Mammals, the molar teeth of which are

51

bristled with cusps, and are associated with canines and
incisors: they are unguiculate, plantigrade, and pentadactyle,
and they have complete clavicles. Like Rodents, they are
temporary testiconda, and have large prostatic and vesicular
glands: like most other Lissencephala, the Insectivora have
a discoid or cup-shaped placenta. They do not exist in
South America and Australia; their office in these continents
is fulfilled by Marsupialia; but true Insectivora abound in
all the other continents and their contiguous islands.

The order CHEIROPTERA, with the exception of the modi-
fication of their digits for supporting the large webs that serve
as wings, repeat the chief characters of the Insectivora: a
few, however, of the larger species are frugivorous and have
corresponding modifications of the teeth and stomach. The
mammez are pectoral in position.

The most remarkable examples of periodically torpid
Mammals are to be found in the terrestrial and volant Insecti-
vora. The frugivorous Bats differ much in dentition from the
“true Cheiroptera, and would seem to conduct through the
Colugos or Flying Lemurs, directly to the Quadrumanous
order. ‘The Cheiroptera are cosmopolitan.

The order Bruta, called Hdentata by Cuvier, includes
two genera (Myrmecophaga and Manis) which are devoid of
teeth ; the rest possess those organs, which, however, have no
true enamel, are never displaced by a second series, and are
very rarely implanted in the premaxillary bones. All the _
species have very long and strong claws. The ischium as well
as the ilium unites with the sacrum; the orbit is not divided
from the temporal fossa. The Three-toed Sloths (Brady-
pus) manifest their affinity to the oviparous Vertebrata by the
supernumerary cervical vertebree supporting false ribs and by
the convolution of the wind-pipe in the thorax; and I may
add that the unusual number—three and twenty pairs—of
ribs, forming a very long dorsal, with a short lumbar, region
of the spine, in the Two-toed Sloth (Cholepus), recalls a
lacertine structure. The same tendency to an inferior type

32

is shown by the abdominal testes, the single cloacal outlet,
the low cerebral development, the absence of medullary canals
in the long bones in the Sloths, and by the great tenacity of
life and long-enduring irritability of the muscular fibre, in
both the Sloths and Anteaters’.

The order Bruta is but scantily represented at the present
period. One genus, Manis or Pangolin, is common to Asia
and Africa; the Orycteropus is peculiar to South Africa; the
rest of the order, consisting of the genera Myrmecophaga,
or true Anteaters, Dasypus or Armadillos, and Bradypus or
Sloths, are confined to South America.

Having defined the orders or subdivisions of the two fore-
going subclasses, | may remark that the LyencepHata cannot
be regarded as equivalent merely to one of the orders, say
Rodentia, of the ZissencepuHata, without undervaluing the
anatomical characters which are so remarkable and distinct
in the marsupial and monotrematous animals. The anato-
mical peculiarities of the edentulous Lyznceruata” appear to
me to be, at least, of ordinal importance. In these deduc-
tions I hold the mean between those who, with Geoffroy
St Hilaire, would make a distinct class of the MJonotremata,
and those who, with Cuvier, would make the Monotremes
a mere family of the Hdentata. In like manner, whilst I
regard the Zyencepuara as forming a group of higher rank
than an order, I do not consider it as forming an equivalent
primary group to that formed by all the placental Mammalia.

The true value of the Zyencepuata is that of one of four

1 This latter vital character attracted the notice of the earliest observers of
these animals. Thus Marcgrave and Piso narrate of the Sloth :—‘Cor motum
suum validissimé retinebat, postquam exemptum erat e corpore per semiho-
rium :—exempto corde ceteris visceribus multo post se movebat et pedes lenté
contrahebat sicut dormituriens solet.’ Buffon, who quotes the above from the
Historia Naturalis Brasilie, p. 322, well remarks, ‘ Par ces rapports, ce quad-
rupede se rapproche non seulement de la tortue, dont il a la lenteur, mais en-
core des autres reptiles et de tous ceux qui n’ont pas un centre du sentiment
unique et bien distinct. —/Hist. Naturelle, 4to, Tom. XII. p. 45.

2 See my article Monotremata, in the Cyclopedia of Anatomy, part xxvi.

IS4T.

33

primary divisions or subclasses of the Mammalia; its true
equivalency is with the Lissencepuaa, and all its analogical
relations are to be found more directly in that smooth-brained
subclass than in the Placentalia at large.

The following Table exemplifies the correspondence of the
groups in the Lyencephalous and Lissencephalous series :—

LYENCEPHALA, D1ssENCEPHALA.
OME ODNAGO, ines ssthalyotn eben Burrowing Rodentia.
OCMIOGO: scivces axtgncsadacha. Dipodide and Leporide.
LE UGMFUS! Sodvid ines cde oaesaves Pteromys.
ERGIGNGUUAG. .cccsesecseavs . Sciuride and prehensile-tailed

arboreal Rodents. —

TE ASCOLATCLOS cnvunecasivea-a +> Bradypus.
Perameles and Myrmecobius. Krinaceide.
COTO DUS <i swiaias dy ocsodiane o« Macroscelis.
Didelphys and Phascogale . Soricide.
DEERE sa nlite ab ves lcncoee's Centetes, Gymnura.
NGO) iva ca tecuss ewen'es tess Manis.

Besides the more general characters by which the Lisszn-
CEPHALA, in common with the Lyencepuara, resemble Birds
and Reptiles, there are many other remarkable indications of
their affinity to the Oviparous Vertebrata in particular orders
or genera of the subclass. Such, e.g., are the cloaca, con-
voluted trachea, supernumerary cervical vertebre and their
floating ribs, in the three-toed Sloth ; the numerous trunk-ribs
in the two-toed Sloth ; the irritability of the muscular fibre, and
persistence of contractile power in the Sloths and some other
Bruta; the long, slender, beak-like edentulous jaws and
gizzard of the Anteaters ; the imbricated scales of the equally
edentulous Pangolins, which have both gizzard and gastric
glands like the proventricular ones in birds; the dermal bony
armour of the Armadillos like that of loricated Saurians; the
quills of the Porcupine and Hedgehog; the brilliant iridescent
colours of the fur of the Cape-mole (Chrysochlora aurea) ;
the proventriculus of the Dormouse and Beaver; the pre-

1 On the Classification of the Marsupialia, Trans. of the Zool, Soc, Vol, 11.

p- 315, 1830.
D

o4

valence of disproportionate development of the hind limbs in
the Rodentia, coupled, in the Jerboa, with confluence of the
three chief metatarsals into one bone, as in birds; the keeled
sternum and wings of the Bats; the aptitude of the Chetrop-
tera, Insectivora, and certain Rodentia to fall, like Reptiles,
into a state of true torpidity, associated with a corresponding
faculty of the heart to circulate carbonized or black blood :—
these, and the like indications of coaffinity with the Lyzn-
cepHaLa to the Oviparous air-breathing Vertebrata, have
mainly prevailed with me against an acquiescence in the
elevation of different groups of the Dissencupuara to a higher
place in the Mammalian series, and in their respective associa-
tion, through some single character, with better-brained orders,
according to Mammalogical systems which, at different times,
have been proposed by zoologists of deserved reputation.
Such, e.g., as the association of the long-clawed Bruta with
the Ungulata’, and of the shorter-clawed Shrews, Moles and
Hedgehogs, as well as the Bats, with the Carnivora* ; of the
Sloths with the Quadrumana’; of the Bats with the same
high order*; and of the Jnsectivora and Rodentia in immediate
sequence after the Linnean ‘ Primates,’ as in the latest pub-
lished ‘System of Mammalogy,’ from a distinguished French
author’.

So far as their ordinal affinities are known, the most
ancient Mammals, the fossil remains of which have been found
in secondary strata, are either ly- or liss- encephalous, and belong
either to the Marsupialia or the Insectivora. (Appendix A).

In the G'yrencepuazta we look in vain for those marks of
affinity to the oviparous vertebrate animals which have been in-
stanced in the preceding subclasses; although, it is true, that
when we proceed to consider the subdivisions of the Gyren-

1 Macleay, Linn. Trans. Vol. xvi. (1833) ; Gray, Dr. J. E., Mammalia in
the British Museum, 12mo, 1843, p. xii.

2 Cuvier, Régne Animal, 1829, p. 110.

3 De Blainville, Ostéographie, 4to, Fasc. 1. p. 47 (1839).

4 Linneus, Systema Nature, Ed. 12, Tom. 1. p. 26.

> Prof. Gervais, Zoologie et Paléontologie Francais, 4to, 1852, p. 194.

35

chPHALA, We seem at first to descend in the scale by finding in
that wave-brained subclass a group of animals, having the
form of Fishes: but a high grade of mammalian organization
is masked beneath this form.

The Gyrenceruata are primarily subdivided, according to
modifications of the locomotive organs, into three series, for
which the Linnean terms may well be retained ; viz. Mutilata,
Ungulata and Unguiculata, the maimed, the hoofed, and the
clawed series.

These limb-characters can only be rightly applied to the
gyrencephalous subclass; they do not indicate natural groups,
save in that section of the Mammalia. ‘To associate the
Livencepuara and Lissencepuata with the unguiculate Gyren-
CEPHALA into one great primary group, as in the Mammalian
systems of Ray, Linneus and Cuvier, is a misapplication of a
solitary character akin to that which would have founded a
primary division on the discoid placenta or the diphyodont
dentition. No one has proposed to associate the unguiculate
Bird or Lizard with the unguiculate Ape; and it is but a
little less violation of natural affinities to associate the Mono-
tremes with the Quadrumanes in the same primary (unguicu-
late) division of the Mammalian class.

The three primary divisions of the G'yrencerHaza are of
higher value than the ordinal divisions of the Lisszncepwava ;
just as those orders are of higher value than the representative
families of the LyzncepHaua,

_The Mutilata, or the maimed Mammals with folded brains,
are so called because their hind limbs seem, as it were, to have
been amputated; they possess only the pectoral pair of limbs,
and these in the form of fins: the hind end of the trunk
expands into a broad, horizontally flattened, caudal fin. They
have large brains with many and deep convolutions, are
naked, and have neither neck, scrotum, nor external ears.

The first order, called CETACEA, in this division are either
edentulous or monophyodont, and the latter have teeth of one
kind and usually of simple form. They are ‘testiconda,’ and

D2

36

have no ‘vesicule seminales.. The mamme are pudendal ;
the placenta is diffused; the external nostrils—single or
double—are on the top of the head, and called spiracles or
‘blow-holes.’ They are marine, and, for the most part,
range the unfathomable ocean; though with certain geogra-
phical limits as respects species. The ‘right whale’ of the
northern hemisphere (Balena mysticetus) is represented by a
distinct species (Balena australis) in the southern hemi-
sphere: the high temperature of the waters at the equatorial
zone bars the migration of either from one pole to the other.
True Cetacea feed on fishes or marine animals.

The second order, called Strenta, have teeth of different
kinds, incisors which are preceded by milk-teeth, and molars
with flattened or ridged crowns, adapted for vegetable food.
The nostrils are two, situated at the upper part of the snout ;
the lips are beset with stiff bristles ; the mamme are pectoral ;
they are ‘testiconda,’ but have ‘vesicule seminales.’ The
Sirenia exist near coasts or ascend large rivers; browsing on
fuci, water plants, or the grass of the shore. There is much
in the organization of this order that indicates its nearer
affinity to members of the succeeding division, than to the
cetaceous order.

The Dugongs (Halicore) inhabit the Red sea, the Ma-
layan Archipelago, and the soundings of the Australian
coasts: the Manatees (Manatus) frequent the shores of tropi-
cal America and Africa.

In the Ungulata the four limbs are present, but that por-
tion of the toe which touches the ground is incased in a hoof,
which blunts its sensibility and deprives the foot of prehen-
sile power. With the limbs restricted to support and loco-
motion, the Ungulates have no clavicles; the two bones of the
fore leg are fixed together in the position anatomists call
‘prone;’ as a general rule hoofed quadrupeds feed on vegetables.

A particular order, or suborder, of this group is indicated
by fossil remains of certain South American genera, e. g.
Toxodon and Nesodon, with long, curved, rootless teeth,

37

having a partial investment of enamel, and with certain pecu-
larities of cranial structure: the name TOXODONTIA is pro-
posed for this order, all the representatives of which are
extinct.

A second remarkable order, most of the members of which
have also passed away, is characterized by two incisors in
the form of long tusks; in one genus (Dinotherium) projecting
from the under jaw, in another genus (Hlephas) from the
upper jaw, and in some of the species of a third genus (Masto-
don) from both jaws. There are no canines: the molars are
few, large and transversely ridged ; the ridges sometimes few
and mammillate, often numerous and with every intermediate
gradation. The nose is prolonged into a cylindrical trunk,
flexible in all directions, highly sensitive, and terminated by
a prehensile appendage like a finger: from this peculiar organ
is derived the name Proposcipra given to the order. The feet
are pentadactyle, but the toes are indicated only by divisions of
the hoof; the placenta is annular; the mamme are pectoral.

Elephants are dependent chiefly upon trees for food. One
species now finds the conditions of its existence in the rich
forests of tropical Asia; a second species in those of tropical
Africa. Why, we may ask, should not a third be living at
the expense of the still more luxuriant vegetation watered by
the Oronoko, the Essequibo, the Amazon, and the La Plata,
in tropical America? Geology tells us that at least two kinds
of Elephant (Mastodon Andium and Mast. Humboldtit) for-
merly did derive their subsistence, along with the great Mega-
therioid beasts, from that abundant source: two other kinds
of Elephant (Mastodon ohioticus and Elephas textanus) existed
in the warm and temperate latitudes of North America. Twice
as many species of Mastodon and Elephant, distinct from all
the others, roamed in pliocene times in the same latitudes of
Europe. At a later or pleistocene period, a huge elephant,
clothed with wool and hair, obtained its food from hardy trees,
such as now grow in the 65th degree of north latitude; and

1 Philosophical Transactions, 1853, p. 291.

38

abundant remains of this Hlephas primigentus (as it has been
prematurely called, since it was the last of our British ele-
phants) have been found in temperate and high northern lati-
tudes in Europe, Asia and America. This, like other Arctic
animals, was peculiar in its family for its range in longitude.
The Musk Buffalo was its contemporary in England and Eu-
rope, and still lingers in the northernmost parts of America.

I have received evidences of Elephantine species from
China and Australia, proving the proboscidian pachyderms to
have once been the most cosmopolitan of hoofed herbivorous
quadrupeds.

Both the proboscidian and toxodontal orders of Uneunara
may be called aberrant: the dentition of the latter, and several
particulars of the organization of the Elephant, indicate an
affinity to the Rodentia; the cranium of the Toxodon, like
that of the Dinothere, resembles that of the Sirenda in its re-
markable modifications.

The typical Ungulate quadrupeds are divided, according
to the odd or even number of the toes, into PERISSODACTYLA
and ARTIODACTYLA!: the single hoof of the horse, the triple
hoof of the tapir, exemplify the first: the double hoof of the
camel, the quadruple hoof of the hippopotamus, exemplify the
second. In the perissodactyle or odd-toed Uneutara, the dorso-
lumbar vertebree differ in number in different species, but are
never fewer than twenty-two; the femur has a third trochanter,
and the medullary artery does not penetrate the fore part of
its shaft. The fore part of the astragalus is divided into two
very unequal facets. The os magnum and the digitus medius
which it supports are large, in some disproportionately so, and
the digit is symmetrical: the same applies to the ectocunel-
form and the digit which it supports in the hind foot. If the
species be horned, as the Rhinoceros, the horn is single; or,
if there be two, they are placed on the median line of the
head, one behind the other, each being thus an odd horn.

1 From mepiccoddkrudos, qui digitos habet impares numero; and dptuvos,
par, ddxrvdos, digitus.—Quarterly Journal of the Geological Society, No. 14,
May, 1848.

39

There is a well-developed post-tympanic process which is
separated by the true mastoid from the paroccipital in the
Horse, but unites with the lower part of the paroccipital in
the T'apir, and seems to take the place of the mastoid in the
Rhinoceros and Hyrax. The hinder half, or a‘larger propor-
tion, of the palatines enters into the formation of the posterior
nares, the oblique aperture of which commences in advance
either of the last molar, or, as in most, of the penultimate one.
The pterygoid process has a broad and thick base and is per-
forated lengthwise by the ectocarotid. The crown of from one
to three of the hinder premolars is as complete as those of the
molars: that of the last lower milk-molar is commonly bi-
lobed. 'To these osteological and dental characters may be
added some important modifications of internal structure, as,
e. g., the simple form of the stomach and the capacious and
sacculated cecum, which equally evince the mutual affinities
of the odd-toed or perissodactyle quadrupeds with hoofs, and
. their claims to be regarded as a natural group of the Uneurara.
Many extinct genera, e. g. Coryphodon, Pliolophus, Lophiodon,
Taptrotherium, Paleotherium, Ancitherium, Hipparion, Acero-
therium, Elasmotherium, &c., have been discovered, which
once linked together the now broken series of Perissodactyles,
represented by the existing genera Lhinoceros, Hyrax, Ta-
pirus, and Equus. The placenta is replaced by a diffused
vascular villosity of the chorion in all the recent genera of this
order, excepting the little /yrax, in which there is a localised
annular placenta, as in the Elephant. But the diffused pla-
centa occurs in some genera of the next group, shewing the
inapplicability of that character to exact classification.

In the even-toed or ‘artiodactyle’ Ungulates, the dorso-
lumbar vertebre are the same in number, as a general rule,
in all the species, being nineteen. The recognition of this
important character appears to have been impeded by the
variable number of moveable ribs in different species of the
Artiodactyles, the dorsal vertebree, which those ribs charac-
terize, being fifteen in the Hippopotamus and twelve in the

40

Camel. And the value of this distinction has been exag-
gerated owing to the common conception of the ribs as special
bones distinct from the vertebre, and their non-recognition as
parts of a vertebra equivalent to the neurapophyses and other
autogenous elements.

The vertebral formule of the Artiodactyle skeletons shew
that the difference in the number of the so-called dorsal and
lumbar vertebre does not affect the number of the entire
dorso-lumbar series: thus the Indian Wild Boar has d. 13,
1.6 =19; the Domestic Hog and the Peccari have d. 14, 1.
5 =19; the Hippopotamus has d. 15, 1. 4=19; the Gnu and
Aurochs have d. 14, 7.5 =19; the Ox and most of the true
Ruminants have d.13, 7.6 =19; the aberrant Ruminants have
d.12,1.7=19. The natural character and true affinities of
the Artiodactyle group are further illustrated by the absence
of the third trochanter in the femur, and by the place of per-
foration of the medullary artery at the fore and upper part of
the shaft, as in the Hippopotamus, the Hog, and most of the
Ruminants. The fore part of the astragalus is divided into
two equal or subequal facets: the os magnum does not exceed,
or is less than, the unciforme in size, in the carpus; and the
ectocuneiform is less, or not larger, than the cuboid, in the
tarsus. The digit answering to the third in the pentadactyle
foot is unsymmetrical, and forms, with that answering to the
fourth, a symmetrical pair. If the species be horned, the
horns form one pair, as in most Ruminants, or two pairs, as
in the four-horned Antelope and Sivathere; they are never
developed singly, of symmetrical form, from the median line.
The post-tympanic does not project downward distinctly from
the mastoid, nor supersede it in any Artiodactyle; and the
paroccipital always exceeds both those processes in length.
The bony palate extends further back than in the Perisso-
dactyles ; the hinder aperture of the nasal passages is more
vertical and commences posterior to the last molar tooth. The
base of the pterygoid process is not perforated by the ecto-
carotid artery. ‘The crowns of the premolars are smaller and

41

less complex than those of the true molars, usually represent-
ing half of such crown. The last milk-molar is trilobed.

To these osteological and dental characters may be added
some important modifications of internal structure, as, e. g., the
complex form of the stomach in the Hippopotamus, Peccari,
and Ruminants; the comparatively small and simple cecum
and the spirally folded colon in all Artiodactyles, which equally
indicate the mutual affinities of the even-toed hoofed quad-
rupeds, and their claims to be regarded as a natural group of
the Unevutata. The placenta is diffused in the Camel-tribe
and non-ruminants; is cotyledonal in the true Ruminants.
Many extinct genera, e.g. Chwropotamus, Anthracotherium,
Hyopotamus, Entelodon, Dichodon, Merycopotamus, Xiphodon,
Dichobune, Anoplotherium, Microtherium, &c., have been dis-
covered, which once linked together the now broken series
of Artiodactyles, represented by the existing genera, Hippo-
potamus, Sus, Dicotyles, Camelus, Auchenia, Moschus, Camelo-
. pardalis, Cervus, Antilope, Ovis, and Bos.

A well-marked, and at the present day very extensive
subordinate group of the Artiodactyles, is called Ruminantia,
in reference to the second mastication to which the food is
subject after having been swallowed; the act of rumination
requiring a peculiarly complicated form of stomach. The
Ruminants have the ‘cloven foot,’ 1.e. two hoofed digits on
each foot forming a symmetrical pair, as by the cleavage of a
single hoof; in most species there is added a pair of small
supplementary hoofed toes. The metacarpals of the two func-
tional toes coalesce to form a single ‘ cannon-bone,’ as do the
corresponding metatarsals. The Camel-tribe have the upper
incisors reduced to a single pair; in the rest of the Ruminants
the upper incisors are replaced by a callous pad. The lower
canines are contiguous to the six lower incisors, and, save in
the Camel-tribe, are similar to them, forming part of the same
terminal series of eight teeth, between which and the molar
series there is a wide interval. The true molars have their
erinding surface marked by two double crescents, the con-

42

vexity of which is turned inwards in the upper and outwards
in the under jaw.

Many fossil Artiodactyles, with similar molars, appear to
have differed from the Ruminants chiefly by retaining struc-
tures which are transitory and embryonic in most existing
Ruminants, as, e.g. upper incisors and canines, first pre-
molars, and separate metacarpal and metatarsal bones; these
are among the lost links that once connected more intimately
the Ruminants with the Hog and Hippopotamus.

The Pachyderms in the Cuvierian system included all the
non-ruminant hoofed beasts; they were divided by the great
French anatomist into the Proboscidia, Solidungula, and Pachy-
dermata ordinaria, the latter again being subdivided according
to the odd or even number of the hoofs. I have on another
occasion! adduced evidence to shew that the right progression
of the affinities of the Unevnara was broken by the interpo-
sition of the Horse and other Perissodactyles between the
non-ruminant or omnivorous and the ruminant Artiodactyles ;
and that too high a value had been assigned to the Rumi-
nantia by making them equivalent to all the other Ungulates
collectively.

It is interesting, in relation to the needs of mankind, to find
that, whilst some groups of Uneuxara, e. g. the Perissodactyles
and omnivorous Artiodactyles, have been gradually dying out,
other groups, e.g. the Ruminants, have been augmenting in
genera and species. Most interesting also is it to observe,
that in existing Ungulates there is a more specialized struc-
ture, a further departure from the general type, than in their
representatives of the miocene and eocene tertiary periods:
such later and less typical Mammalia do more effective work
by virtue of their adaptively modified structures.

The Ruminants, e. g., more effectually digest and assimi-
late grass, and form out of it a more nutritive and sapid kind
of meat, than did the antecedent more typical and less spe-
cialized non-ruminant Herbivora.

1 Proceedings of the Geological Society, November 3, 1847, p. 135.

43

The monodactyle Horse is a better and swifter beast of
draught and burthen than its tridactyle predecessor the mio-
cene [Hipparion could have been. The nearer to a Tapir ora
Rhinoceros in structure, the further would an equine quadruped
be left from the goal in contending with a modern Racer.

With respect to the geographical distribution of the hoofed
Mammalia, I may first remark that the order Ruminantia is
principally represented by Old World species, of which 162
have been defined; only 24 species have been discovered in
the New World, and none in Australia, New Guinea, New
Zealand, or the Polynesian Isles.

The Camelopard is now peculiar to Africa; the Musk-deer
to Africa and Asia: out of about 50 defined species of Ante-
lope, only one is known in America, and none in the central
and southern divisions of the New World. The Bison of
- North America is distinct from the Bison of Europe. The
Musk-ox, peculiar for its limitation to high northern latitudes,
is the sole bovine species that roams over the arctic coasts of
both Asia and America. The Deer-tribe are more widely distri-
buted. The Camels and Dromedaries of the Old World are
represented by the Llamas and Vicugnas of the New. As, in
regard to a former (tertiary) zoological period, the fossil
Camelide of Asia are of the genus Camelus, so those of
America are of the genus Auchenia. This geographical
restriction ruled prior to any evidence of man’s existence.

Paleontology has expanded our knowledge of the range of
the Giraffe ; during miocene or old pliocene periods, species of
Camelopardalis roamed in Asia and Europe. Passing to the
non-ruminant Artiodactyles, geology has also taught us that
the Hippopotamus was not always confined, as now, to African
rivers, but bathed, during pliocene times, in those of Asia and
Europe. But no evidence has yet been had that the Girafte
or Hippopotamus were ever other than Old-World forms of
Ungulata.

With respect to the Hog-tribe, we find that the true Swine
(Sus) of the Old World are represented by Peccaries (Dico-

44

tyles) in the New; and geology has recently shewn that
tertiary species of Dicotyles existed in North as well as South
America. But no true Sus has been found fossil im either
division of the New World, nor has any Décotyles been found
fossil in the Old World of the geographer. Phacocherus
(Wart-hogs) is a genus of the Hog-tribe at present peculiar
to Africa.

The Rhinoceros is a genus now represented only in Asia
and Africa; the species being distinct in the two continents.
The islands of Java and of Sumatra have each their peculiar
species ; that of the latter being two-horned, as all the African
Rhinoceroses are. Three or more species of two-horned Rhi-
noceros formerly inhabited Europe’, one of which we know to
have been warmly clad and adapted for a cold climate; but
no fossil remains of the genus have been met with save in the
Old World of the geographer. One of the earliest forms of
European Rhinoceros was devoid of the nasal weapon: it has
long been extinct.

Geology has given a wider prospect of the range of the
Horse and Elephant, than was open to the student of living
species only. The existing Eqguide and Elephantide properly
belong, or are limited to, the Old World; and the Elephants
to Asia and Africa, the species of the two continents being
quite distinct. The horse, as Buffon remarked, carried terror
to the eye of the indigenous Americans, viewing the animal
for the first time, as it proudly bore their Spanish conqueror.
But species of Hquus, like species of Mastodon, coexisted with
the Megatheritum and Megalonyx in both South and North
America, and perished with them, apparently before the
human period.

The third division of the Gyrenceruata enjoy a higher
degree of the sense of touch than the Ungulates through the
ereater number and mobility of the digits and the smaller
extent to which they are covered by horny matter. This
substance forms a single plate, in the shape of a claw or nail,

1 See my History of British Fossil Mammals, 8vo, p. 350.

45

which is applied to only one of the surfaces of the extremity
of the digit, leaving the other, usually the lower, surface pos-
sessed of its tactile faculty; whence the name Uneuicurara,
applied to this group, which, however, is here more restricted
and natural than the group to which Linneus extended the
term. All the species are ‘diphyodont,’ and the teeth have
a simple investment of enamel. 3
The first order, CARNIVORA, includes the beasts of prey,
properly so called. With the exception of a few Seals the

aes Be 3—3 . ‘ 1—1
incisors are [— in number; the canines ees

than the other teeth, and usually exhibiting a full and perfect
development as lethal weapons; the molars graduate from a

, always longer

trenchant to a tuberculate form, in proportion as the diet
deviates from one strictly of flesh, to one of a more miscella-
neous kind. The clavicle is rudimental or absent; the
innermost digit is often rudimental or absent; they have no

vesicule seminales; the teats are abdominal; the placenta is
~ zonular.

The Carnivora are divided, according to modifications of
the limbs, into ‘ pinnigrade,’ ‘ plantigrade,’ and ‘ digitigrade’
tribes. In the Pinnigrades (Walrus, Seal-tribe) both fore
and hind feet are short, and expanded into broad, webbed
paddles for swimming, the hinder ones being fettered by con-
tinuation of integument to the tail. In the Plantigrades
(Bear-tribe) the whole or nearly the whole of the hind foot
forms a sole, and rests on the ground. In the Digitigrades
(Cat-tribe, Dog-tribe, &c.) only the toes touch the ground,
the heel being much raised.

It has been usual to place the Plantigrades at the head of
the Carnivora, apparently because the higher order, Quadru-
mana, can put the heel to the ground: but the affinities of the
Bear, as evidenced by internal structure, e.g. the renal and
genital organs, are closer to the Seal-tribe; the broader and
flatter pentadactyle foot of the plantigrade is nearer in form to
the flipper of the seal than is the digitigrade, retractile-clawed,

46

long and narrow hind foot of the feline quadruped, which is
the highest and most typical of the Carnivora.

With the exception of the Dingo no true Carnivore exists
in Australia, and that wild dog may have as little claim to
be considered an autochthon as the low variety of Man, with
whom it is sometimes associated in a half-tamed state.

The genus Ursus is represented by species indigenous to
Europe, Asia, Africa, and America; but those of the temper-
ate and warmer latitudes of the New World are distinct from
the species of the Old World. Certain plantigrade genera,
e. g. Procyon (Racoons), Nasuwa (Coati-mondis) and Cercolep-
tes (Kinkajous) are peculiarly American: other plantigrade
genera, e.g. Mydaus, Ailurus, and Arctictis, are peculiarly
Asian.

The genus Hyeena is limited to the Old World, and one
species (H. crocuta) to Southern Africa.

The Skunks (Mephitis) are peculiar to America; the viver-
rine Carnivores to the Old World.

The great fulvous felines (Leo) of Africa and Asia are
represented in America by the smaller Pumas: the Old
World spotted felines by the Jaguars: the great striped
felines (Z%gris) are now restricted to Asia.

The principle of the more specialized character of actual
organisations receives illustration in the genetic history of the
present order. |

The genera Felis and Machairodus, with their curtailed
and otherwise modified dentition and their strong short jaws,
become, thereby, more powerfully and effectively destructive
than the eocene Hycnodons and miocene Pterodons, with
their numerically typical dentition and their three carnassial
teeth on each side of the concomitantly prolonged jaws, could
have been.

In the most strictly carnivorous Glyrencepnaa the paw is
perfected as an instrument for retaining and lacerating a strug-
eling prey by the superadded elastic structures for retracting
the claws and maintaining them sharp. We next find in

47

‘the unguiculate limb such a modification in the size, shape,
position, and direction of the innermost digit that it can be op-
posed, as a thumb, to the other digits, thus constituting what is
properly termed a ‘hand.’ Those Unguiculates which have
both fore and hind limbs so modified, form the order QUAD-

RUMANA. They have

= incisors, and — broad tuber-
culate molars; perfect clavicles; pectoral mamme; vesicular
and prostatic glands; a discoid, sometimes double, placenta.
The Quadrumana have a well-marked threefold geographical
as well as structural division.

The Strepsirhines are those with curved or twisted termi-

a

nal nostrils, with much modified incisors, commonly isc ¢

remolars ——— or
P B= 3 9 9

tubercles: the second digit of the hind limb has a claw.
This group includes the Galagos, Pottos, Loris, Aye-Ayes,
. “Indris, and the true Lemurs; the three latter genera being
restricted to Madagascar, whence the group diverges in one
direction to the continent of Africa, in the other to the Indian

Archipelago.

in number, and molars with sharp

The Platyrhines are those with. the nostrils subterminal
ee :
53 m number, the molars with
blunt tubercles; the thumbs of the fore-hands not opposable
or wanting; the tail in most prehensile; they are peculiar to

South America.
The Catarhines have the nostrils oblique and approximated
below, and opening above and behind the muzzle: the pre-

and wide apart; premolars

—_
molars are =

a :
5 in number; the thumb of the fore-hand is

opposable. They are restricted to the Old World, and, save
a single species on the rock of Gibraltar, to Africa and Asia.
The highest organized family of Catarhines is tailless, and

offers in the Orang, Chimpanzee, and Gorilla, the nearest
approach to the human type.

48

The Catarhine monkeys include the Macaques, most of
which are Asiatic, a few are African, and one European; the
Cercopitheques, most of which are African, and a few Asiatic;
and other genera which characterize one or other continent
exclusively. Thus the true Baboons (Papio) are African,
as are the thumbless Monkeys (Colobus) and the Chimpanzees
(Troglodytes). The Semnopitheques, Gibbons (Hylobates),
and Orangs (Pithecus) are peculiarly Asiatic. Paleontology
has shown that a Macaque, a Gibbon and an Orang existed
during the older tertiary times in Europe; and that a Sem-
nopitheque existed in miocene times in India. But all the
fossil remains of Quadrumana in the Old World belong to
the family Catarhina, which is still exclusively confined to
that great division of dry land. The tailless Macaque (Jnwus
sylvanus) of Gibraltar may have existed in that part of the
Old World before Europe was separated by the Straits of
Gibraltar from Africa. Fossil remains of Quadrumana have
been discovered in South America; they indicate Platyrhine
forms: a species, for example, allied to the Howlers (I/ycetes),
but larger than any now known to exist, has left its remains
in Brazil.

Whilst adverting to the geographical distribution of
Quadrumana, [ would contrast the peculiarly limited range of
the Orangs and Chimpanzees with the cosmopolitan powers
of mankind. The two species of Orang (Pithecus) are con-
fined to Borneo and Sumatra; the two species of Chimpanzee
(Troglodytes) are limited to an intertropical tract of the western
part of Africa. ‘They appear to be inexorably bound to their
localities by climatal influences regulating the assemblage
of certain trees and the production of certain fruits. With
all our care, in regard to choice of food, clothing, and contri-
vances for artificially maintaining the chief physical condi-
tions of their existence, the healthiest specimens of Orang or
Chimpanzee, brought over in the vigour of youth, perish
within a period never exceeding three years, and usually
much shorter, in our climate. By what metamorphoses, we

49

may ask, has the alleged humanized Chimpanzee or Orang
been brought to endure all climates? The advocates of
‘transmutation’ have failed to explain them. Certain it is
that those physical differences in cerebral, dental, and osteo-
logical structure, which place, in my estimate of them, the
genus Homo in a distinct group of the Mammalian class, zoo-
logically of higher value than the ‘order,’ are associated with
equally contrasted powers of endurance of different climates,
whereby Man has become a denizen of every part of the globe
from the torrid to the arctic zones.

Climate rigidly limits the range of the Quadrumana in
latitude: creational and geographical causes limit their range
in longitude. Distinct genera represent each other in the same
latitudes of the New and Old Worlds; and also, in a great
degree, in Africa and Asia. But the development of an Orang
out of a Chimpanzee, or reciprocally, is physiologically incon-
ceivable. (Appendix B).

The sole representative of the ArcyencepHaxa, is the ge-
nus Homo. His structural modifications, more especially of
the lower limb, by which the erect stature. and bipedal gait
are maintained, are such as to claim for Man ordinal distinc-
tion on merely external zoological characters. But, as I have
already argued, his mental powers, in association with his
extraordinarily developed brain, entitle the group which he
represents to equivalent rank with the other primary divi-
sions of the class Mammalia founded on cerebral characters.
In this primary group Man forms but one genus, Homo, and
that genus but one order, called BImMANA, on account of the
opposable thumb being restricted to the upper pair of limbs.
The mamme are pectoral. The placenta is a single, sub-
circular, cellulo-vascular, discoid body.

Man has only a partial covering of hair, which is not
merely protective of the head, but is ornamental and distinc-
tive of sex. The dentition of the genus Homo is reduced to
thirty-two teeth by the suppression of the outer incisor and

E

50

the first two premolars of the typical series on each side of
both jaws, the dental formula being :—

.2-72. 1-1 2—2 | 8—3 _ ao
y EIS NG Se ONE Be ot eat | eee Nh ap

All the teeth are of equal length, and there is no break in the
series ; they are subservient in Man not only to alimentation,
but to beauty and to speech.

The human foot is broad, plantigrade, with the sole, not
inverted as in Quadrumana, but applied flat to the ground;
the leg bears vertically on the foot; the heel is expanded be-
neath; the toes are short, but with the innermost longer and
much larger than the rest, forming a ‘ hallux’ or great toe,
which is placed on the same line with, and cannot be opposed
to, the other toes; the pelvis is short, broad, and wide, keep-
ing the thighs well apart; and the neck of the femur is long,
and forms an open angle with the shaft, increasing the basis
of support for the trunk. The whole vertebral column, with
its slight alternate curves, and the well-poised, short, but
capacious subglobular skull, are in like harmony with the
requirements of the erect position. The widely-separated
shoulders, with broad scapule and complete clavicles, give a
favourable position to the upper limbs, now liberated from the
service of locomotion, with complex joints for rotatory as well
as flexile movements, and términated by a hand of matchless
perfection of structure, the fit instrument for executing the
behests of a rational intelligence and a free will. Hereby,
though naked, Man can clothe himself, and rival all natural
vestments in warmth and beauty; though defenceless, Man
can arm himself with every variety of weapon, and become
the most terribly destructive of animals. Thus he fulfils his
destiny as the master of this earth, and of the lower Creation.

Such are the dominating powers with which we, and we
alone, are gifted! I say gifted, for the surpassing organisa-
tion was no work of ours. It is He that hath made us; not

1 Vid. p. 19, for the type-formula and explanation of its symbols.

51

we ourselves. This frame is a temporary trust, for the uses
of which we are responsible to the Maker.

Oh! you who possess it in all the supple vigour of lusty
youth, think well what it is that He has committed to your
keeping. Waste not its energies; dull them not by sloth:
spoil them not by pleasures! The supreme work of Creation
has been accomplished that you might possess a body—the
sole erect—of all animal bodies the most free—and for what ?
for the service of the soul.

Strive to realise the conditions of the possession of this
wondrous structure. Think what it may become—the T'emple of
the Holy Spirit! Defile it not. Seek, rather, to adorn it with
all meet and becoming gifts, with that fair furniture, moral
and intellectual, which it is your inestimable privilege to ac-
quire through the teachings and examples and ministrations
of this Seat of Sound Learning and Religious Education.

Such, Sir, are the sentiments that have naturally flowed
from the contemplation of the highest of the gradations of
Mammalian structure of which we have now completed the
ascensive survey: and I know of no topic more fitting to the
words in which, with a grateful sense of the most kind re- —
ception and attention accorded to me, I conclude the duty of
this Chair. :

Ba iyi

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APPENDIX A.
ON THE EXTINCTION OF SPECIES.

Being the Conclusion of the Fullerian Course of Lectures on
Physiology, for 1859.

In a Report to the British Association for the Advancement of
Science, On the Hatinct Mammals of Australia, published in the
Volume of Reports for 1844, evidence is adduced in proof of the
law, that with extinct as with existing mammalia particular forms
were assigned to particular provinces, and that the same forms
were restricted to the same provinces at a former geological period
as they are at the present day. That period, however, was the
more recent tertiary one.

In carrying back the retrospective comparison of existing and
extinct mammals to those of the eocene and oolitic strata, in rela-
tion to their local distribution, we obtain indications of extensive
changes in the relative position of sea and land during those epochs,
through the degree of incongruity between the generic forms of the
mammalia which then existed in Europe, and any that actually
exist on the great natural continent of which Europe now forms
part. It would seem, indeed, that the further we penetrate into
time for the recovery of extinct mammalia, the further we must
go into space to find their existing analogues. To match the eo-
cene palzeotheres and lophiodons we must bring tapirs from Suma-
tra or South America; and we must travel to the antipodes for
myrmecobians, the nearest living analogue to the amphitheres and
spalacotheres of our oolitic strata.

On the problem of the extinction of species I have little to say ;
and of the more mysterious subject of their coming into being,
nothing profitable or to the purpose. As a cause of extinction in
times anterior to man, it is most reasonable to assign the chief
weight to those gradual changes in the conditions affecting a due
supply of sustenance to animals in a state of nature which must
have accompanied the slow alternations of land and sea brought

56

about in the eons of geological time. Yet this reasoning is appli-
cable only to land-animals; for it is scarcely conceivable that such .
operations can have affected sea-fishes.

There are characters in land-animals rendering them more ob-
noxious to extirpating influences, which may explain why so many
of the larger species of particular groups have become extinct,
whilst smaller species of equal antiquity have survived. In pro-
portion to its bulk is the difficulty of the contest which the animal
has to maintain against the surrounding agencies that are ever
tending to dissolve the vital bond, and subjugate the living matter
to the ordinary chemical and physical forces. Any changes, there-
fore, in such external agencies as a species may have been origi-
nally adapted to exist in, will militate against that existence in a
degree proportionate to the size which may characterise the spe-
cies. If a dry season be gradually prolonged, the large mammal
will suffer from the drought sooner than the small one; if such
alteration of climate affect the quantity of vegetable food, the bulky
herbivore will first feel the effects of stinted nourishment; if new
enemies be introduced, the large and conspicuous animal will fall a
prey while the smaller kinds conceal themselves and escape. Small
quadrupeds, moreover, are more prolific than large ones. Those
of the bulk of the mastodons, megatheria, glyptodons, and dipro-
todons, are uniparous. The actual presence, therefore, of small spe-
cies of animals in countries where larger species of the same na-
tural families formerly existed, is not the consequence of degenera-
tion—of any gradual diminution of the size—of such species, but
is the result of circumstances which may be illustrated by the
fable of the ‘Oak and the Reed;’ the smaller and feebler animals
have bent and accommodated themselves to changes to which the
larger species have succumbed.

That species should become extinct appears, from the abundant
evidence of the fact of extinction, to be a law of their existence;
whether, however, it be inherent in their own nature, or be rela-
tive and dependent on inevitable changes in the conditions and
theatre of their existence, is the main subject for consideration.
But, admitting extinction as a natural law which has operated
from the beginning of life on this planet, it might be expected
that some evidence of it should occur in our own time, or within
the historical period. Reference has been made to several in-
stances of the extirpation of species, certainly, probably, or pos-
sibly, due to the direct agency of man; but this cause avails not in

OT

the question of the extinction of species at periods prior to any evi-
dence of human existence; it does not help us in the explanation
of the majority of extinctions; as of the races of aquatic inverte-
brata which have successively passed away.

Within the last century academicians of St. Petersburg and
good naturalists have described and given figures of the bony and
the perishable parts, including the alimentary canal, of a large and
peculiar fucivorous Sirenian—an amphibious animal like the Ma-
natee, which Cuvier classified with his herbivorous Cetacea, and
called Stelleria, after its discoverer. This animal inhabited the
Siberian shores and the mouths of the great rivers there disem-
boguing. It is now believed to be extinct, and this extinction
seems not to have been due to any special quest and persecution
by man. We may discern, in this fact, the operation of changes
in physical geography which have, at length, so affected the con-
ditions of existence of the Stelleria as to have caused its extinction.
Such changes had operated, at an earlier period, to the extinction
of the Siberian elephant and rhinoceros of the same regions and
latitudes. A future generation of zoologists may have to record

the final disappearance of the Arctic buffalo (Ovibos moschatus).
- Fossil remains of Ovibos and Stelleria shew that they were con-
temporaries of Llephas primigenius and Rhinoceros tichorrhinus.

The Great Auk (Alca impennis, L.) seems to be rapidly
verging to extinction. It has not been specially hunted down,
like the dodo and dinornis, but by degrees has become more scarce.
Some of the geological changes affecting circumstances favourable
to the well-being of the Alca impennis, have been matters of ob-
servation. A friend’, who last year visited Iceland, informs me
that the last great auks, known with anything like certainty to
have been there seen, were two which were taken in 1844 during
a. visit made to the high rock called ‘Eldey,’ or ‘Meelsoekten,’
lying off Cape Reykianes, the 8. W. point of Iceland. This is one
of three principal rocky islets formerly existing in that direction,
of which the one, specially named from this rare bird ‘ Geirfugla
Sker,’ sank to the level of the surface of the sea during a volcanic
disturbance in or about the year 1830. Such disappearance of the
fit and favourable breeding-places of the Alcea impennis must form
an important element in its decline towards extinction. The
numbers of the bones of Alea impennis on the shores of Iceland,
Greenland, and Denmark, attest the abundance of the bird in

1 John Wolley, jun., Esq. F.Z.S.

58

former times. A consideration of such instances of modern partial
or total extinctions may best throw light on, and suggest the truest
notions of, the causes of ancient extinctions.

As to the successions, or coming in, of new species, one might
speculate on the gradual modifiability of the individual; on the
tendency of certain varieties to survive local changes, and thus
progressively diverge from an older type; on the production and
fertility of monstrous offspring; on the possibility, e.g. of a variety
of auk being occasionally hatched with a somewhat longer winglet,
and a dwarfed stature; on the probability of such a variety better
adapting itself to the changing climate or other conditions than
the old type—of such an origin of Alca torda, e.g. ;—but to what
purpose? Past experience of the chance aims of human fancy,
unchecked and unguided by observed facts, shews how widely they
have ever glanced away from the gold centre of truth.

The sum of the evidence which has been obtained appears to
prove that the successive extinction of Amphitheria, Spalacotheria,
Triconodons, and other mesozoic forms of mammals, has been
followed by the introduction of much more numerous, varied, and
higher-organised forms of the class, during the tertiary periods.

There are, however, geologists who maintain that this is an
assumption, based upon a partial knowledge of the facts. Mere
negative evidence, they allege, can never satisfactorily establish
the proposition that the mammalian class is of late introduction,
nor prevent the conjecture that it may have been as richly repre-
sented in secondary as in tertiary times, could we but get evidence
of the terrestrial fauna of the oolitic continent. To this objection
I have to reply : in the paleozoic strata, which, from their extent
and depth, indicate, in the earth’s existence as a seat of organic
life, a period as prolonged as that which has followed their depo-
sition, no trace of mammals has been observed. It may be con-
ceded that, were mammals peculiar to dry land, such negative
evidence would weigh little in producing conviction of their non-
existence during the Silurian and Devonian eons, because the ex-
plored parts of such strata have been deposited from an ocean, and
the chance of finding a terrestrial and air-breathing creature’s re-
mains in oceanic deposits is very remote. But, in the present
state of the warm-blooded, air-breathing, viviparous class, no genera
and species are represented by such numerous and widely dispersed
individuals, as those of the order Cetacea, which, under the guise
of fishes, dwell, and can only live, in the ocean.

59

In all cetacea the skeleton is well ossified, and the vertebre
are very numerous: the smallest cetaceans would be deemed large
amongst land-mammals; the largest surpass in bulk any creatures
of which we have yet gained cognizance: the hugest ichthyosaur,
iguanodon, megalosaur, mammoth, or megathere, is a dwarf in com-
parison with the modern whale of a hundred feet in length.

During the period in which we have proof that Cetacea have
existed, the evidence in the shape of bones and teeth, which latter
enduring characteristics in most of the species are peculiar for their
great number in the same individual, must have been abundantly
deposited at the bottom of the sea; and as cachalots, grampuses,
dolphins, and porpoises are seen gambolling in shoals in deep
oceans, far from land, their remains will form the most charac-
teristic evidences of vertebrate life in the strata now in course of
formation at the bottom of such oceans. Accordingly, it consists
with the known characteristics of the cetacean class to find the
marine deposits which fell from seas tenanted, as now, with verte-
brates of that high grade, containing the fossil evidences of the
order in vast abundance.

The red crag of our eastern counties contains petrified frag-

*” ments of the skeletons and teeth of various Cetacea, in such quanti-

ties as to constitute a great part of that source of phosphate of lime
for which the red crag is worked for the manufacture of artificial
manure. The scanty and dubious evidence of Cefacea in newer
secondary beds’ seems to indicate a similar period for their begin-
ning as for the soft-scaled cycloid and ctenoid fishes which have
superseded the ganoid orders of mesozoic times.

We cannot doubt but that had the genera [chthyosaurus, Plio-
saurus, or Plesiosaurus, been represented by species in the same
ocean that was tempested by the Baleenodons and Dioplodons of
the miocene age, the bones and teeth of those marine reptiles
would have testified to their existence as abundantly as they do at
a previous epoch in the earth’s history. But no fossil relic of an
enaliosaur has been found in tertiary strata, and no living enalio-
saur has been detected in the present seas: and they are conse-
quently held by competent naturalists to be extinct.

Tn like manner does such negative evidence weigh with me in
proof of the non-existence of marine mammals in the liassic and
oolitic times. In the marine deposits of those secondary or meso-

1 See ‘Introduction’ to Owen’s History of British Fossil Mammalia, 8vo.,
1846, p. xv.

60

zoic epochs, the evidence of vertebrates governing the ocean, and
preying on inferior marine vertebrates, is as abundant as that of
air-breathing vertebrates in the tertiary strata; but in the one the
fossils are exclusively of the cold-blooded reptilian class, in the
other, of the warm-blooded mammalian class. The Hnaliosauria,
Cetiosauria, and Crocodilia, played the same part and fulfilled
similar offices in the seas from which the lias and oolites were
precipitated, as the Delphinide and Balenide did in the tertiary,
and still do in the present, seas. The unbiassed conclusion from
both negative and positive evidence in this matter is, that the
Cetacea succeeded and superseded the Lnaliosauria. To the mind
that will not accept such conclusion, the stratified oolitic rocks
must cease to be monuments or trustworthy records of the con-
dition of life on the earth at that period.

So far, however, as any general conclusion can be deduced from
the large sum of evidence above referred to, and contrasted, it is
against the doctrine of the Uniformitarian. Organic remains,
traced from their earliest known graves, are succeeded, one series by
another, to the present period, and never re-appear when once lost
sight of in the ascending search. As well might we expect a
living Ichthyosaur in the Pacific, as a fossil whale in the Lias : the
rule governs as strongly in the retrospect as the prospect. And
not only as respects the Vertebrata, but the sum of the animal
species at each successive geological period has been distinct and
peculiar to such period.

Not that the extinction of such forms or species was sudden or
simultaneous : the evidences so interpreted have been but local :
over the wider field of life at any given epoch, the change has been
gradual ; and, as it would. seem, obedient to some general, but as
yet, ill-comprehended law. In regard to animal life, and its as-
signed work on this planet, there has, however, plainly been ‘an
ascent and progress in the main.’

Although the mammalia, in regard to the plenary development
of the characteristic orders, belong to the Tertiary division of geo-
logical time, just as ‘Hchini are most common in the superior
strata, Ammonites in those beneath, and Producti with numerous
Encrim in the lowest’* of the secondary strata, yet the beginnings
of the class manifest themselves in the formations of the earlier
preceding division of geological time.

No one, save a prepossessed Uniformitarian, would infer from

1 A generalisation of WILLIAM SMITH’s.

61

the Lucina of the permian, and the Opis of the trias, that the
Lamellibranchiate Mollusks existed in the same rich variety of
development at these periods as during the tertiary and present
times; and no prepossession can close the eyes to the fact that the
Lamellibranchiate have superseded the Palliobranchiate bivalves.

On negative evidence Orthisina, Theca, Producta, or Spirifer are
believed not to exist in the present seas: neither are the existing
genera of siphonated bivalves and univalves deemed to have
abounded in permian, triassic or oolitic times. To suspect that
they may have then existed, but have hitherto escaped observation,
because certain Lamellibranchs with an open mantle, and some
holostomatous and asiphonate Gastropods, have left their remains
in secondary strata, is not more reasonable, as it seems to me, than
to conclude that the proportion of mammalian life may have been
as great in secondary as in tertiary strata, because a few small
forms of the lowest orders have made their appearance in triassic
and oolitic beds.

Turning from a retrospect into past time for the prospect of
time to come,—and I have received more than one inquiry into
the amount of prophetic insight imparted by Paleeontology—I may
crave indulgence for a few words, of more sound, perhaps, than
significance. But the reflective mind cannot evade or resist the
tendency to speculate on the future course and ultimate fate of
vital phenomena in this planet.

There seems to have been a time when life was not; there may,
therefore, be a period when it will cease to be.

Our most soaring speculations still shew a kinship to our
nature : we see the element of finality in so much that we have
cognizance of, that it must needs mingle with our thoughts, and
bias our conclusions on many things.

The end of the world has been presented to man’s mind under
divers aspects :—as a general conflagration ; as the same, preceded
by a millennial exaltation of the world to a Paradisiacal state,—
the abode ofa higher and blessed race of intelligences.

If the guide-post of Paleontology may seem to point to a
course ascending to the condition of the latter speculation, it
points but a very short way, and in leaving it we find ourselves
in a wilderness of conjecture, where to try to advance is to find
ourselves ‘in wandering mazes lost.’

With much more satisfaction do I return to the legitimate
deductions from the phenomena we have had under review.

62

In the survey which I have taken in the present course of
lectures of the genesis, succession, geographical distribution, affini-
ties, and osteology of the mammalian class, if I have succeeded in
demonstrating the perfect adaptation of each varying form to the
exigencies, and habits, and well-being of the species, I have ful-
filled one object which I had in view, viz. to set forth the bene-
ficence and intelligence of the Creative Power.

If Ihave been able to demonstrate a uniform plan pervading
the osteological structure of so many diversified animated beings,
I must have enforced, were that necessary, as strong a conviction |
of the unity of the Creative Cause.

If, in all the striking changes of form and proportion which
have passed under review, we could discern only the results of
minor modifications of the same few osseous elements,—surely we
must be the more strikingly impressed with the wisdom and power
of that Cause which could produce so much variety, and at the
same time such perfect adaptations and endowments, out of means
so simple.

For, in what have those mechanical instruments,—the hands
of the ape, the hoofs of the horse, the fins of the whale, the trowels
of the mole, the wings of the bat,—so variously formed to obey
the behests of volition in denizens of different elements—in what,
I say, have they differed from the artificial instruments which we
ourselves plan with foresight and calculation for analogous uses,
save in their greater complexity, in their perfection, and in the
unity and simplicity of the elements which are modified to con-
stitute these several locomotive organs ?

Everywhere in organic nature we see the means not only sub-
servient to an end, but that end accomplished by the simplest
means. Hence we are compelled to regard the Great Cause of
all, not like certain philosophic ancients, as a uniform and quies-
cent mind, as an all pervading anima mundi, but as an active and
anticipating intelligence.

By applying the laws of comparative anatomy to the relics of
extinct races of animals contained in and characterizing the dif-
ferent strata of the earth’s crust, and corresponding with as many
epochs in the earth’s history, we make an important step in
advance of all preceding philosophies, and are able to demonstrate
that the same pervading, active, and beneficent intelligence which
manifests His power in our times, has also manifested His power in
times long anterior to the records of our existence.

63

But we likewise, by these investigations, gain a still more im-
portant truth, viz. that the phenomena of the world do not succeed
each other with the mechanical sameness attributed to them in the
cycles of the Epicurean philosophy ; for we are able to demonstrate
that the different epochs of the earth were attended with corre-
sponding changes of organic structure; and that, in all these in-
stances of change, the organs, as far as we could comprehend their
use, were exactly those best suited to the functions of the being.
Hence we not only show intelligence evoking means adapted to
the end; but, at successive times and periods, producing a change
of mechanism adapted to a change in external conditions. Thus
the highest generalizations in the science of organic bodies, like the
Newtonian laws of universal matter, lead to the unequivocal con-
viction of a great First Cause, which is certainly not mechanical.

Unfettered by narrow restrictions,—unchecked by the timid

and unworthy fears of mistrustful minds, clinging, in regard to mere
physical questions, to beliefs, for which the Author of all truth has
been pleased to substitute knowledge,—our science becomes con-
nected with the loftiest of moral speculations.
If I believed,—to use the language of a gifted contemporary,—
‘that the imagination, the feelings, the active intellectual powers,
bearing on the business of life, and the highest capacities of our
nature, were blunted and impaired by the study of physiological
and paleontological phenomena, I should then regard our science
as little better than a moral sepulchre, in which, like the strong
man, we were burying ourselves and those around us in ruins of
our own creating.

But surely we must all believe too firmly in the immutable
attributes of that Being, in whom all truth, of whatever kind, finds
its proper resting-place, to think that the principles of physical and
moral truth can ever be in lasting collision’.

1 Sedgwick, Address to the Geological Society, 1831.

64

APPENDIX B.
ON THE ORANG, CHIMPANZEE, AND GORILLA,
With reference to the ‘ Transmutation of Species.’

For about two centuries, naturalists have been cognizant of a
small ape, tailless, without cheek-pouches, and without the ischial
callosities, clothed with black hair, with a facial angle of about 60°,
and of a physiognomy milder and more human-like than in the
ordinary race of monkeys, less capricious, less impulsive in its
habits, more staid and docile. This species, brought from the West
Coast of Africa, is that which our anatomist, Tyson, dissected: he
described the main features of its organisation in his work pub-
lished in 1699°. He called it the Homo Sylvestris, or pigmy. It
is noted by Linnzeus, in some editions of his Systema Nature, as
the Homo Troglodytes. Blumenbach, giving a truer value to the
condition of the innermost digit of the hind foot, which was like a
thumb, called it the Simia T'roglodytes,; it afterwards became more
commonly known as the ‘ Chimpanzee.’

At a later period, naturalists became acquainted with a similar
kind of ape, of quiet docile disposition, with the same sad, human-
like expression of features. It was brought from Borneo or Suma-
tra; where it is known by the name of Orang, which, in the language
of the natives of Borneo, signifies ‘man,’ with the distinctive
addition of Outan, meaning ‘Wood-man,’ or ‘Wild Man of the
Woods.’ This creature differed from the pigmy, or Simia T'ro-
glodytes of Africa, by being covered with hair of a reddish-brown
colour, and by having the anterior, or upper limbs, much Jonger in
proportion, and the thumb upon the hind feet somewhat less. It
was entered in the zoological catalogue as the Simia Satyrus. A
governor of Batavia, Baron Wurmb, had transmitted to Holland,
in 1780, the skeleton of a large kind of ape, tailless, like this small
species from Borneo, but with a much-developed face, and large
canine teeth, and bearing thick callosities upon the cheeks, giving
it, upon the whole, a very baboon-like physiognomy ; and he called
it the Pongo.

At the time when Cuvier revised his summary of our knowledge
of the animal kingdom, in the second edition of his ‘Régne Animal,’

1“ Orang-Outang, sive Homo sylvestris ; or the Anatomie of a Pygmie, com-
pared with that of a Monkey, an Ape and a Man,’ 4to, 1699.

65

1829, the knowledge of the anthropoid apes was limited to these
three forms. It had been suspected that the pongo might be the
adult form of the orang; but Cuvier, pointing to its distinctive
characters, suggested that it could hardly be the same species. The
facial angle of the small red orang of Borneo, and of the small -
black chimpanzee of Africa, brought them, from the predominant
cranium, and small size of the jaws and small teeth, nearer than
any other known mammalian animal to the human species, par-
ticularly to the lower, or negro forms. It was evident, from the
examination of these small chimpanzees and orangs, that they
were the young of some large species of ape. The small size and
number of their teeth, (there being, in some of the smaller speci-
mens, only twenty, like the number of deciduous teeth in the
human species,) and the intervals between those teeth, all showed
them to be of the first or deciduous series. In 1835 I availed
myself of the rich materials in regard to these animals collected
about that time by the Zoological Society, to investigate the state
of dentition, especially that of the permanent teeth which might be
hidden in the substance of the jaws, of both the immature orang-
outang and the chimpanzee, and I found that the germs of those
‘teeth in the orang-outang agreed in size with the permanent teeth
that were developed in the jaws of a species of the pongo of Wurmb,
which Sir Stamford Raffles had presented to the museum of the
College of Surgeons some years before. Specimens of orangs since
acquired, of an intermediate age, have shown the progressive
change of the dentition.

In the substance of the jaw were found the germs of the great
canines, and of large bicuspid teeth ; foreshowing the changes that
must take place when the jaw is sufficiently enlarged to receive
permanent teeth of this kind; and, when the rest of the cranium is
modified, concomitantly, for the attachment of muscles to work the
jaw so armed, denoting that all these changes must result in the
acquisition of characters such as are presented by the skulls of the
large pongo, or Bornean baboon-like ape. The specific identity of
the pongo with certain of the young orang-outangs, was thus
satisfactorily made out, and is now admitted by all naturalists.
With regard to the chimpanzee, the germs of similarly propor-
tioned large teeth were also discovered in the jaws, indicating, in
like manner, that it must be the young of a much larger species
of ape.

The principal osteological characters of the chimpanzee and

EF

66

orang, commencing from the vertebral column, are as follows :—
The vertebral column describes only one curve, inclining forward,
where it supports the head with its large jaws and teeth. The
‘vertebre in the neck, seven in number as usual in the mammalia,
are chiefly remarkable for the great length of the simple spinous
processes developed more than in most of the inferior apes, in
relation to the necessities of the muscular masses that are to sus-
tain and balance the head that preponderates so much forward on
the neck. The vertebree maintain a much closer correspondence
in size, from the cervical to the dorsal and lumbar region, than in
the human skeleton. With regard to the.dorsal vertebree, or those
to which moveable ribs are articulated, there are twelve pairs in the
orang; seven of them send cartilages to join the sternum, which is
more like the sternum in man than in any of the inferior quadru-
mana: it is shorter and broader. In the smaller long-armed apes
(Hylobates), which make the first step in the transition from the
ordinary quadrumana to the man-like apes, the sternum is remark-
ably broad and short. The lumbar vertebree are, originally, five
in number in the orang; but one or two may coalesce with the
sacrum. The sacrum is broader than in the lower quadrumana,
but it is still narrow in comparison with its proportions in man.
The pelvis is longer. The iliac bones are more expanded than
in the lower quadrumana, but on the same plane, and are flat-
tened and long. The tuberosities of the ischia are remarkably
developed, and project outward. All these conditions of the ver-
tebral column indicate an animal capable only of a semi-erect
position, and present a modification of the trunk much better
adapted for a creature destined for a life in trees, than one that, is
to walk habitually erect upon the surface of the ground. But
that adaptation of the skeleton is still more strikingly shown in
the unusual development of the upper prehensile extremities. The
scapula is broad, with a well-developed spine and acromion; there
is a complete clavicle; the bone of the arm (Awmerus) is of remark-
able length, in proportion to the trunk; the radius and the ulna
are also very long, and unusually diverging, to give increased sur-
face of attachment to muscles; the hand is remarkable for the
length of the metacarpus, and of the phalanges, which are slightly
bent towards the palm; the thumb is less developed than the cor-
responding digit in the foot; the whole hand is admirably adapted
for retaining a firm grasp of the boughs of trees. In the structure
of the carpus, there is a well-marked difference from the human

67

subject, and a retention of the character met with in the lower
quadrumana ; the scaphoid bone being divided in the orang-outang.
In the chimpanzee the bones of the carpus are eight, as in the
human subject, but differ somewhat in form. If the upper ex-
tremities are so extraordinary for their disproportionate length, the
lower ones are equally remarkable for their disproportionate small-
ness in comparison with the trunk, in the orang. The femur is
short and straight, and the neck of the thigh-bone comparatively
short. The head of the thigh-bone in this animal, which requires the
use of these lower prehensible organs to grasp the branches of trees,
and to move freely in many directions, is free from that ligament
which strengthens the hip-joint in man; the head of the femur in
the orang is quite smooth, without any indication of that attach-
ment. Here, again, the chimpanzee manifests a nearer approach
to man, for the ligamentum teres is present in it in accordance
with the stronger and better development of the whole hind-limb.
This approximation, also, is more especially marked in the larger
development of the innermost of the five digits of the foot in the
chimpanzee, which is associated with a tendency to move more fre-
* quently upon the ground, to maintain a more erect position than
the orang-outang, and to walk further without the assistance of a
stick. The foot, in both these species of anthropoid orangs, is
characterized by the backward position of the ankle-joint surface
presented by the astragalus to the tibia, which serves for the trans-
ference of the superincumbent weight upon the foot; by the
comparatively feeble development of the backward projecting pro-
cess of the calcaneum; by the obliquity of the articular surface of
the astragalus, which tends to incline the foot a little inwards,
taking away from the plantigrade character of the creatures and
from their capacity to support themselves in an erect position, and
giving them an equivalent power of applying their prehensile
feet to the branches of the trees in which they live.

In both the orang and chimpanzee the skull is articulated to
the spine by condyles, which are placed far back on its under surface.
The cranium is small, characterised by well-developed occipital and
sagittal ridges; the occipital ridges in reference to the muscles sus-
taining the head; and the sagittal ones in reference to an increased
extent of the temporal muscles. The zygomatic arches are strong,
and well arched outwards. The lower jaw is of great depth, and
has powerful ascending rami, but the chin is wanting. The facial
angle is about 50° to 55° in the full-grown Simia satyrus, and

F2

68

55° to 60° in the Zroglodytes niger. The difference in the facial
angle between the young and adult apes, (which, in the young
chimpanzee, approaches 60° to 65°,) depends upon those changes
consequent upon the shedding of the deciduous teeth and the con-
comitant development of the jaws and intermuscular processes of
the cranium.

But the knowledge of the species of these anthropoid apes has
been further increased since the acquisition of a distinct and pre-
cise cognisance of the characters of the adults of the orang and
chimpanzee. Jirst, in reference to the orangs of Borneo, speci-
mens have reached this country which show that there is a smaller
species in that island, the Simia Morio, in which the canines are
less developed, in which the bony cristw are never raised above
the level of the ordinary convexity of the cranium, and in which
the callosities upon the cheeks are absent, associated with other
characteristics plainly indicating a specific distinction. The Rajah
Brooke has confirmed the fact of the existence in the island of
Borneo of two distinct species of red orangs; one of a smaller size
and somewhat more anthropoid; and the larger species presenting
the baboon-like cranium. 7

In reference to the black chimpanzee of Africa also, another
very important addition has been, recently, made to our knowledge
of those forms of highly developed quadrumana. In 1847 I received
a letter from Dr Savage, a church-missionary at Gaboon, on the
west coast of tropical Africa, enclosing sketches of the crania of an
ape, which he described as much larger than the chimpanzee,
ferocious in its habits, and dreaded by the negro natives more
than they dread the lion or any other wild beast of the forest.
These sketches showed plainly one cranial characteristic by which
the chimpanzee differs in a marked degree from the orangs; viz.
that produced by the prominence of the super-orbital ridge, which
is wanting in the adult and immature of the orangs. That ridge
was strongly marked in the sketches transmitted. At a later
period in the same year, were transmitted to me from Bristol two
skulls of the same large species of chimpanzee as that notified in
Dr Savage's letter ; they were obtained from the same locality in
Africa, and brought clearly to light evidence of the existence in
Africa of a second larger and more powerful ape,—the Tvroglodytes
gorilla. They are described and figured in the third Volume
of the Transactions of the Zoological Society, 1848.

The additional facts, subsequently ascertained respecting the

69

gorilla, although they prove its nearer approach to man than any
other tailless ape, have not in any degree affected or invalidated
the conclusions at which I then arrived.

Since the date of that memoir, skeletons and the entire carcase
preserved in spirits of the gorilla have successively reached the
Museums of Paris, Vienna, and London; and have formed the
subjects of several memoirs, the results of the recorded observa-
tions differing only in regard to the interpretation of the facts.

Dr Wyman, the accomplished anatomical professor at Boston,
U.S., agrees with the writer in referring the gorilla to the same
genus as the chimpanzee (Zroglodytes), but he regards the latter as
more nearly allied to the human kind.

Professors Duvernoy and Isidore Geoffroy St Hilaire consider
_the differences in the osteology, dentition, and outward character
of the gorilla to be of generic importance; and they enter the
species in the zoological catalogues as Gorilla gina, the trivial
name being that by which the animal is called by the natives of
Gaboon ; the French naturalists also concur with the American in
placing the gorilla below the chimpanzee in the zoological scale ;
and some have more lately been disposed to place both below the
slamangs, gibbons or long-armed apes (Hy/lobates).

The following are the principal external characters of the
Gorilla exhibited by the specimen preserved in spirits which was
received in 1858, at the British Museum, and is now mounted
and exhibited in the Mammalian Gallery. My attention was first
attracted by the shortness, almost absence, of neck, due to the back-
ward position of the junction of the head to the trunk, to the great
length of the cervical spines, causing the ‘nape’ to project beyond
the ‘occiput,’ to the great size and elevation of the scapule, and
to the oblique rising of the clavicles from their sternal attachments
to above the level of the angles of the jaw. The brain-case, low
and narrow, and the lofty ridges of the skull, make the cranial
profile pass in almost a straight line from the occiput to the super-
orbital ridge, the prominence of which gives the most forbidding
feature to the physiognomy of the gorilla; the thick integument
overlapping that ridge forming a scowling pent-house over the
eyes. The nose is more prominent than in the chimpanzee or
orang-utan, not only at its lower expanded part, but at its upper
half, where a slight prominence corresponds with that which the
author had previously pointed out in the nasal bones. The mouth
is very wide, the lips large, of uniform thickness, the upper one

70

with a straight, as if incised margin, not showing the coloured
lining membrane when the mouth is shut. The chin is short
and receding, the muzzle very prominent. The eyelids with eye-
lashes, the eyes wider apart than in the orang or chimpanzee;
no defined eyebrows; but the hairy scalp continued to the super-
orbital ridge. The ears are smaller in proportion than in man,
much smaller than in the chimpanzee; but the structure of the
auricle is more like that of man. On a direct front view of the
face, the ears are on the same parallel with the eyes’. The huge
canines in the male give a most formidable aspect to the beast:
they were not fully developed in the younger and entire specimen,
now mounted. The profile of the trunk describes a slight con-
vexity from the nape to the sacrum,—there being no inbending at
the loins, which seem wanting, the thirteenth pair of ribs being
close to the ‘labrum ili.’ The chest is of great capacity; the
shoulders very wide across; the pectoral regions are slightly
marked, and shew a pair of nipples placed as in the chimpanzee
and human species. The abdomen is somewhat prominent, both
before and at the sides. The pelvis relatively broader than in
other apes.

The chief deviations from the human structure are seen in the
limbs, which are of great power, the upper ones prodigiously strong.
The arm from below the short deltoid prominence preserves its
thickness to the condyles; a uniform circumference prevails in the
fore-arm ; the leg increases in thickness from below the knee to the
ankle. There is no ‘calf’ of the leg. These characters of the limbs
are due to the general absence of those partial muscular enlarge-
ments which impart the graceful varying curves to the outlines of
the limbs in man. Yet they depend rather on excess, than defect,
of development of the carneous as compared with the tendinous
parts of the limb-muscles, which thus continue of almost the same
size from their origin to their insertion, with a proportionate gain
of strength to the beast.

The difference in the length of the upper limbs between the
gorilla and man is but little in comparison with the trunk; it
appears greater through the arrest of development of the lower
limbs. Very significant of the closer anthropoid affinities of the
gorilla is the superior length of the arm (humerus) to the fore-
arm, as compared with the proportions of those parts in the chim

1 On the Anthropoid Apes; Proceedings, R. I, Vol. 11. (1855) p. 26, and in
the Transactions of the Zoological Society, 1848,

71

panzee. The hair of the arm inclines downward, that of the
fore-arm upward, as in the chimpanzee. The thumb extends a
little beyond the base of the proximal phalanx of the fore-finger ;
it does not reach to the end of the metacarpal bone in the chim-
panzee or any other ape: the thumb of the siamang is still shorter
in proportion to the length of the fingers of the same hand: the
philosophical zoologist will see great significance in this fact. In
man the thumb extends to, or beyond, the middle of the first
phalanx of the fore-finger.

The fore-arm in the gorilla passes into the hand with very
slight evidence, by constriction, of the wrist; the circumference of
which, without the hair, is fourteen inches, that of a strong man
averaging eight inches. The hand is remarkable for its breadth
and thickness, and for the great length of the palm, occasioned
both by the length of the metacarpus and the greater extent of
undivided integument between the digits than in man; these only
begin to be free opposite the middle of the proximal or first
phalanges in the gorilla. The digits are thus short, and appear as
if swollen and gouty; and are conical in shape after the first joint,
«by tapering to nails, which, being not larger or longer than those
of man, are relatively to the fingers much smaller. The circum-
ference of the middle digit at the first joint in the gorilla is 54
inches; in man, at the same part, it averages 23 inches. The skin
covering the middle phalanx is thick and callous on the backs of
the fingers, and there is little outward appearance of the second
joint. The habit of the animal to apply those parts to the ground,
in occasional progression, is manifested by these callosities. The
back of the hand is hairy as far as the divisions of the fingers; the
palm is naked and callous. The thumb, besides its shortness,
according to the standard of the human hand, is scarcely half so
thick as the fore-finger. The nail of the thumb did not extend to
the end of that digit; in the fingers the nail projected a little
beyond the end, but with a slightly convex worn margin, resem-
bling the human nails in shape, but relatively less.

In the hind-limbs, chiefly noticeable was that first appearance
in the quadrumanous series of a muscular development of the
gluteus, causing a small buttock to project over each tuber ischii.
This structure, with the peculiar expanse (in Quadrumana) of the
iliac bones, leads to an inference that the gorilla must naturally
and with more ease resort occasionally to station and Die
on the lower limbs than any other ape.

(2

The same cause as in the arm, viz. a continuance of a large
proportion of fleshy fibres to the lower end of the muscles, co-
extensive with the thigh, gives a great circumference to that
segment of the limb above the knee-joint, and a more uniform size to
it than in man. The relative shortness of the thigh, its bone being
only eight-ninths the length of the humerus (in man the humerus
averages five-sixths the length of the femur), adds to the appearance
of its superior relative thickness. Absolutely the thigh is not of
greater circumference at its middle than is the same part in man.

The chief difference in the leg, after its relative shortness, is the
absence of a ‘calf, due to the non-existence of the partial accumu-
lation of carneous fibres in the gastrocnemii muscles, causing that
prominence in the type-races of mankind. In the gorilla the
tendo-achillis not only continues to receive the ‘penniform’ fibres
to the heel, but the fleshy parts of the muscles of the foot receive
accessions of fibres at the lower third of the leg, to which the
greater thickness of that part is due, the proportions in this
respect being the reverse of those in man. The leg expands at
once into the foot, which has a peculiar and characteristic form,
owing to the modifications favouring bipedal motion being super-
induced upon an essentially prehensile, quadrumanous type. The
heel makes a more decided backward projection than in the chim-
panzee; the heel-bone is relatively thicker, deeper, more expanded
vertically at its hind end, besides being fully as long as in the
chimpanzee. This bone, so characteristic of anthropoid affinities,
is shaped and proportioned more like the human calcaneum than
in any other ape. The malleoli do not make such well-marked
projections as in man; they are marked more by the thickness of
the fleshy and tendinous parts of the muscles that pass near them,
on their way to be inserted into parts of the foot. Although the
foot be articulated to the leg with a slight inversion of the sole, it
is more nearly plantigrade than in the chimpanzee or any other
ape. The hallux (great toe, thumb of the foot), though not rela-
tively longer than in the chimpanzee, is stronger; the bones are
thicker in proportion to their length, especially the last phalanx,
which in shape and breadth much resembles that in the human
foot. The hallux in its natural position diverges from the other
toes at an angle of 60 deg. from the axis of the foot; its base is
large, swelling into a kind of ball below, upon which the thick
callous epiderm of the sole is continued. The transverse indents
and wrinkles show the frequency and freedom of the flexile move-

73

ments of the two joints of the hallux; the nail is small, flat and
short. The sole of the foot gradually expands from the heel
forward to the divergence of the hallux, and seems to be here cleft,
and almost equally, between the base of the hallux and the common
base of the other four digits. These are small and slender in pro-
portion, and their beginnings are enveloped in a common tegumen-
- tary sheath as far as the base of the second phalanx. A longitudinal
indent at the middle of the sole, bifurcating—one channel defining
the ball of the hallux, the other running towards the interspace
between the second and third digit,—indicates the action of op-
posing the whole thumb (which seems rather like an inner lobe
or division of the sole), to the outer division terminated by the four
short toes. What is termed the ‘instep’ in man is very high in
the gorilla, owing to the thickness of the carneo-tendinous parts of
the muscles as they pass from the leg to the foot over this region.
The mid-toe (third) is a little longer than the second and fourth;
the fifth, as in man, is proportionally shorter than the fourth, and
is divided from it by a somewhat deeper cleft. The whole sole is
wider than in man—relatively to its length much wider—and in
that respect, as well as by the off-set of the hallux, and the defini-
tion of its basal ball, more like a hand, but a hand of huge dimen-
sions and of portentous power of grasp.

The hairy integument is continued along the dorsum of the
foot to the-clefts of the toes, and upon the first phalanx of the
hallux: the whole sole is bare.

In regard to the outward coloration of the gorilla, only from
the examination of the living animal could the precise shades of
colour of the naked parts of the skin be truly described. Much
of the epiderm had peeled off the subject of the present descrip-
tion; but fortunately in large patches, and the texture of these
had acquired a certain firmness, apparently by the action of the
alcohol upon the albuminous basis. The parts of the epiderm
remaining upon the face indicated the skin there to be chiefly of a
deep leaden hue; it is everywhere finely wrinkled, and was some-
what less dark at the prominent parts of the supraciliary roll and
the prominent margins of the nasal ‘ale, the soles and palms
were also of a lighter colour.

Although the general colour of the hair appears, at first sight,
and when moist, to be almost black, it is not so, but is rather of
a dusky grey: it is decidedly of a less deep tint than in the
chimpanzee (Z'rogl. niger): this is due to an admixture of a few

74

reddish, and of more greyish, hairs with the dusky coloured ones
which chiefly constitute the ‘pelage:’ and the above admixture
varies at different parts of the body. The reddish hairs are so
numerous on the scalp, especially along the upper middle region, as
to make their tint rather predominate there; they blend in a less
degree with the long hairs upon the sides of the face. The greyish
hairs are found mixed with the dusky upon the dorsal, deltoidal
and anterior femoral, regions; but on the limbs, not in such pro-
portion as to affect the impression of the general dark colour, at
first view. Near the margin of the vent are a few short whitish
hairs, as in the chimpanzee. The epiderm of the back shewed the
effects of habitual resting, with that part against the trunk or
branch of a tree, occasioning the hair to be more or less rubbed
off: the epithelium was here very thick and tough.

It is most probable, from the degree of admixture of different
coloured hairs above described, that a living gorilla seen in bright
sunlight, would in some positions reflect from its surface a colour
much more different from that of the chimpanzee than appears by
a comparison of the skin of a dead specimen sent home in spirits.
It can hardly be doubted, also, that age will make an appreciable
difference in the general coloration of the Z’roglodytes gorilla.

The adult male gorilla measures five feet six inches from the
sole to the top of the head, the breadth across the shoulders is
nearly three feet, the length of the upper limb is three feet four
inches, that of the lower limb is two feet four inches; the length
of the head and trunk is three feet six inches, whilst the same
dimension in man does not average three feet.

In the foregoing remarks are given the results of direct obser-
vations made on the first and only entire specimen of the gorilla
which has reached England. A more important labour, however,
remains, The accurate record of facts in natural history is one
and a good aim; the deduction of their true consequences is a
better. I proceed, therefore, to reconsider the conclusions from
which my experienced French and American fellow-labourers in
natural history differ from me.

The first—it may be called the supreme—question in regard to
the gorilla is, its place in the scale of nature, and its true and
precise affinities.

Is it or not the nearest of kin to human kind? Does it form,
like the chimpanzee and orang, a distinct genus in the anthropoid
or knuckle-walking group of apes? Are these apes, or are the

i)

long-armed gibbons, more nearly related to the genus Homo? O
the broad-breast-boned quadrumana, are the knuckle-walkergs or
the brachiators, i.e. the long-armed gibbons, most nearly and essen-
tially related to the human subject?

At the first aspect, whether of the entire animal or of the
skeleton, the gorilla strikes the observer as being a much more
bestial and brutish animal than the chimpanzee. All the features
that relate to the wielding of the strong jaws and large canines are
exaggerated ; the evidence of brain is less; its proper cavity is
more masked by the outgrowth of the strong occipital and other
cranial ridges. But then the impression so made that the gorilla
is less like man, is the same which is derived from comparing a
young with an adult chimpanzee, or some small tailless monkey
with a full-grown male orang or chimpanzee. Taking the cha-
racters that cause that impression at a first inspection of the gorilla,
most of the small South American monkeys are more anthropoid ;
they have a proportionally larger and more human-shaped cranium,
much less prominent jaws, with more equable teeth.

On comparing the skeletons of the adult males of the gorilla,
chimpanzee, orang, and gibbon, the globular cranium of the last,
and its superior size compared with the jaws and teeth, seemed to
shew the gibbons to be more nearly akin to man than any of the
larger tailless apes. And this conclusion had been formed by
a distinguished French paleontologist, M. Lartet, and accepted
by a high geological authority at home’. The experienced Professor
of Human Anatomy at Amsterdam had been also cited as supporting
this view; but I have failed to find any statement of the grounds
upon which it was sustained. In the art. Quadrumana of Todd’s
Cyclopedia, cited by Lartet*, Professor Vrolik. briefly treats of the
osteology of the Quadrumana according to their natural families.
In ‘a first genus, Sima proper, or ape,’ he includes the chimpanzee
or orang, noticing some of the chief points by which these apes
approach the nearest to man. He next goes to the second genus,
the gibbon (Hylobates), notices their ischial callosities, and the
nearer approach of their molars, in their rounded form, to the
teeth of carnivora than the molars of the genus Simia. Then,
comparing the siamang with other species of Hylobates, Vrolik says,
‘its skeleton approaches most to that of man,’ which may be true

1 Sir C. Lyell, Supplement to the 5th Edition of a Manual of Elementary

Geology, 1859, p. 15.
2 Comptes Rendus de V Académie des Sciences, Juillet 28, 1856.

76

in comparison with other gibbons, but certainly is not so as respects
the higher Simiw. No details are given to illustrate the proposition
even in its more limited application; but the minor length of the
arms in the siamang, as compared with ylobates lar, was apa
the obvious character in Vrolik’s mind.

The appearance of superior cerebral development in the siamang
and other long-armed apes is due to their small size and the con-
comitant feeble development of their jaws and teeth. The same
appearance makes the small platyrrhine monkeys of South America
equally anthropoid in their facial physiognomy, and much more
human-like than are the great orangs and chimpanzees. It is an
appearance which depends upon the precocious growth of the brain,
as dependent on the law of its development. In all quadrumana
the brain has reached its full size before the second set of teeth is
acquired, almost before the first set is shed. If a young gorilla,
chimpanzee, or orang, be compared with a young siamang, of cor-
responding age, the absolutely larger size and better shape of brain,
the deeper and more numerous convolutions of the cerebrum, and
the more completely covered cerebellum, unequivocally demonstrate
the higher organization of the shorter-armed apes; ‘in the structure
of the brain,’ writes Vrolik', in accordance with all other com-
parative anatomists, ‘they’ (chimpanzee and orang-utan) ‘approach
the nearest to man.’ The degree to which the chimpanzee and
orang so resembled the human type seemed much closer to Cuvier,
who knew those great apes only in their immaturity, with their
small milk-teeth and precociously developed brain. Accordingly, the
anthropoid characters of the Simia satyrus and Sina troglodytes, as
deduced from the facial angle and dentition, are proportionally
exaggerated in the Régne Animal *. As growth proceeds, the
milk-teeth are shed, the jaws expand, the great canines succeed
their diminutive representatives, the biting muscles gain a propor-
tional increase of carneous fibres, their bony fulcra respond to the
call for increased surface of attachment, and the sagittal and occipital
crests begin to rise: but the brain grows no more ; its cranial box
retains the size it shewed in immaturity ; it finally becomes masked
by the superinduced osseous developments in those apes which
attain the largest stature and wield the most formidably armed
jaws. Yet under this disguise of physical force, the brain is still
the better and the larger than is that of the little long-armed ape,

1 Art. Quadrumana, Cyclopedia of Anatomy, Vol. IV. p. 195.
2 Ed..1829, pp..87, 89.

17

which retains throughout life so much more of the characters of
immaturity, especially in the structure of the skull.

The siamang and other gibbons have smaller lower but longer
upper canines, relatively, than in the orangs and chimpanzees; the
permanent ones more quickly attain their full size, and are sooner
in their place in the jaws; consequently the last molar teeth—what
we call the ‘ wisdom-teeth ’—come last into place as they do in the
human species. But, if this be interpreted as of importance in
determining the relative affinity of the longer-armed and shorter-
armed apes to man, it is a character in which, as in their seeming
superior cerebral development, the Hylobates agree with some much
lower Quadrumana with still smaller canines.. The comparative
anatomist, pursuing this most interesting comparison with clear
knowledge of the true conditions and significance of a globular
cranium and small jaws within the quadrumanous order, turns
his attention to the true distinctive characters of the human
organization.

In -respect to the brain, he would look not so much for its
relative size to the body, as for its relative size in the species
compared one with another in the same natural group. He would
inquire what quadrumanous animal shews absolutely the biggest
brain ? what species shows the deepest and most numerous and
winding conyolutions? in which is the cerebrum largest, as com-
pared with the cerebellum? If he found all these characters
highest in the gorilla, he would not be diverted from the just
inference because the great size and surpassing physical power
attained in that species masked the true data from obvious view.

The comparative anatomist would look to the cecum and the
ischial integument: if he found in one subject of his comparisons
(Troglodytes) a long ‘appendix vermiformis ceci, as in man, but
no ‘callosities, and in another subject (Hylobates) the ischial
callosities but only a short rudiment of the cecal appendix, he
would know which of the two tailless apes were to be placed next
‘the monkeys with ischial callosities and no vermiform appendix,’
and which formed the closer link toward man. He would find that
the anthropoid intestinal and dermal characters were associated with
the absolutely larger and better developed brain in the gorilla,
chimpanzee, and orang ; whilst the lower quadrumanous characters
exhibited by the cecum and nates were exhibited by the smaller-
brained and longer-armed tailless gibbons.

Pursuing the comparison through the complexities of the bony

78

framework, the comparative anatomist would first glance at the
more obvious characters; and such, indeed, as would be given by
the entire animal. The characteristics of the limbs in man are
their near equality of length, but the lower limbs are the longest.
The arms in man reach to below the middle of the thigh ; in the
gorilla they nearly attain the knee; in the chimpanzee they reach
below the knee ; in the orang they reach the ankle; in the siamang
they reach the sole; in most gibbons the whole palm can be applied
to the ground without the trunk being bent forward beyond its
naturally inclined position on the legs. These gradational dif-
ferences coincide with other characters determining the relative
proximity of the apes compared with man. In no quadrumana
does the humerus exceed the ulna so much in length as in man ;
only in the very highest and most anthropoid, viz. the gorilla and
chimpanzee, does it exceed the ulna at all in length; in all the rest,
as in the lower quadrupeds, the fore-arm is longer than the arm.

The humerus, in the gorilla, though less long, compared with
the ulna, than in man, is longer than in the chimpanzee; in the
orang it is shorter than the ulna; in the siamang and other gibbons
it is much shorter, the peculiar length of arm in those ‘long-armed
apes’ is chiefly due to the excessive length of the antibrachial bones.

The difference in the length of the upper limbs, as compared
with the trunk, is but little between man and the gorilla. The
elbow-joint in the gorilla, as the arm hangs down, is opposite the
‘labrum ili,’ the wrist opposite the ‘tuber ischii;’ it is rather lower
down in the chimpanzee; is opposite the knee-joint in the orang;
and opposite the ankle-joint in the siamang.

Man’s perfect hand is one of his peculiar physical characters ;
that perfection is mainly due to the: extreme differentiation of the
first from the other four digits, and its concomitant power of
opposing them as a perfect thumb. An opposable thumb is present
in the hand of most Quadrumana, but is usually a small appendage
compared with that of man. It is relatively largest in the gorilla.
In this ape the thumb reaches to a little beyond the base of the
first phalanx of the fore-finger; it does not reach to the end of the
metacarpal bone of the fore-finger in the chimpanzee, orang, or
gibbon; it is relatively smallest in the last tailless ape. In man
the thumb extends to or beyond the middle of the first phalanx of
the fore-finger. The philosophical zoologist will see great signi-
ficance in the results of this comparison. Only in the gorilla and
chimpanzee are the carpal bones eight in number, as in man; in

79

the orangs and gibbons they are nine in number, as in the tailed
monkeys.

The scapulee are broader in the gorilla than in the chimpanzee,
orang, or long-armed apes; they come nearer to the proportions of
that bone in man. But a more decisive resemblance to the human
structure is presented by the iliac bones. In no other ape than the
gorilla do they bend forward, so as to produce a pelvic concavity ;
nor are they so broad in proportion to their length im any ape as
in the gorilla. In both the chimpanzee and orang the iliac bones
are flat, or present a concavity rather at the back than at the
forepart. In the siamang they are not only flat, but are narrower
and longer, resembling the iliac bones of tailed monkeys and
ordinary quadrupeds.

The lower limbs, though characteristically short in the gorilla,

are longer in proportion to the upper limbs, and also to the entire
trunk, than in the chimpanzee; they are much longer in both
proportions and more robust than in the orangs or gibbons. But
the guiding points of comparisons here are the heel and the hallux
(great toe or thumb of the foot).
_ The heel in the gorilla makes a more decided backward projection
than in the chimpanzee; the heel-bone is relatively thicker, deeper,
more expanded vertically at its hind end, besides being fully as long
as in the chimpanzee: it is in the gorilla shaped and proportioned
more like the human calcaneum than in any other ape. Among
all the tailless apes the calcaneum in the siamang and other gibbons
least resembles in its shape or proportional size that of man.

Although the foot be articulated to the leg with a slight inversion
of the sole it is more nearly plantigrade in the gorilla than in the
chimpanzee. The orang departs far, and the gibbons farther, from
the human type in the inverted position of the foot.

_ The great toe which forms the fulcrum in standing or walking
is perhaps the most characteristic peculiarity in the human structure;
it is that modification which differentiates the foot from the hand,
and gives the character to his order (Limana). In the degree of
its approach to this development of the hallux the quadrumanous
animal makes a true step in affinity to man.

The orang-utan and the siamang, tried by this test, descend far
and abruptly below the chimpanzee and gorilla in the scale. In the
orang the hallux does not reach to the end of the metacarpal of the
second toe; in the chimpanzee and gorilla it reaches to the end of
the first phalanx of the second toe; but in the gorilla the hallux

80

is thicker and stronger than in the chimpanzee. In both, however,
it is a true thumb, by position, diverging from the other toes, in the
gorilla, at an angle of 60° from the axis of the foot.

Man has 12 pairs of ribs, the gorilla and chimpanzee have 13
pairs, the orangs have 12 pairs, the gibbons have 13 pairs. Were
the naturalist to trust to this single character, as some have trusted
to the cranio-facial one, and in equal ignorance of the real condition
and value of both, he might think that the orangs (Pithecus) were
nearer akin to man than the chimpanzees (Zroglodytes) are. But
man has sometimes a thirteenth pair of ribs; and what we term
‘ribs’ are but vertebral elements or appendages common to nearly
all the true vertebrze in man, and only so called, when they become
long and free. The genera Homo, T'roglodytes, and Pithecus, have
precisely the same number of vertebre: if Zvroglodytes, by the
development and mobility of the pleurapophyses of the 20th ver-
tebree from the occiput seem to have an additional thoracic vertebra,
it has one vertebra less in the lumbar region. So, if there be, as
has been observed in the same genus, a difference in the number of
sacral vertebre, it is merely due to a last lumbar having coalesced
with what we reckon the first sacral vertebra in man.

The thirteen pairs of ribs, therefore, in the gorilla and chim-
panzee are of no weight, as against the really important characters
significative of affinity with the human type. But, supposing the
fact of any real value, how do the advocates of the superior resem-
blance of the gibbon’s skeleton to that of man dispose of the
thirteenth pair of ribs ?

In applying the characters of the skull to the determination of
the important question at issue those had first to be ascertained by
which the genus Homo trenchantly differs from the genus Sima, of
Linneus. To determine these osteal distinctions I have compared
the skulls of many individuals of different varieties of the human
race together with those of the male, female, and young of species
of Troglodytes, Pithecus, and Hylobates; the detailed results of
which comparisons will be found in the Catalogue of the Osteo-
logical Series in the Museum of the Royal College of Surgeons,
4to, 1853. In the present Appendix, I restrict myself to a few of
these results.

The first and most obvious differential character is the glo-
bular form of the brain-case, and its superior relative size to the
face, especially the jaws, in man. But this, for the reasons al-
ready assigned, is not an instructive or decisive character, when

81

comparing quadrumanous species, in reference to the question at
issue. It is exaggerated in the human child, owing to the acquisi-
tion of its full, or nearly full size, by the brain, before the jaws
have expanded to lodge the second set of teeth. It is an anthropoid
character in which the quadrumana resemble man in proportion
to the diminution of their general bulk. If a gorilla, with milk-
teeth, have a somewhat larger brain and brain-case than a chimpanzee
at the same immature age, the acquisition of greater bulk by the
gorilla, and of amore formidable physical development of the skull,
in reference to the great canines in the male, will give to the chim-
panzee the appearance of a more anthropoid character, which really
does not belong to it; which could be as little depended upon in a
question of precise affinity as the like more anthropoid characters of
the female, as compared with the male, gorilla or chimpanzee.
Much more important and significant are the following cha-
‘racters of the human skull :—the position and plane of the occipital
foramen; the proportion and size of the condyloid and petrous
processes; the mastoid processes, which relate to balancing the
head upon the trunk in the erect attitude; the small premaxillaries
and concomitant small size of the incisor teeth, as compared with
the molar teeth. These characters relate to the superiority of the
psychical over the physical powers in man. They govern the feature
in which man recedes from the brute; and to them may be added
the prominence of the nasal bones in most, and in all the typical,
races of man. The somewhat angular form of the bony orbits,
tending to a square, with the corners rounded off, is, likewise, a
good human character of the skull; which is difficult to compre-
hend as an adaptive one, and therefore the better in the present
inquiry. The same may be said of the production of the floor of
the tympanic or auditory tube into the plate called ‘vaginal.’
Believing the foregoing to be sufficient to test the respective
degrees of affinity to man within the limited group of quadrumana
to which it is now proposed to apply them, I forbear to cite the
characters of minor importance. The question at issue is, as
between the anthropoid apes and man. Cuvier deemed the orang
(Pithecus) to be nearer akin to man than the chimpanzee (7’roglo-
dytes) is. That belief has long ceased to be entertained. I pro-
ceed, therefore, to compare the gorilla, chimpanzee, and gibbon, in
reference to their human affinities.
Most naturalists entering upon this question would first look
to the premaxillary bones, or, owing to the early confluence of
: G

82

those bones with the maxillaries in the gorilla and chimpanzee, to
the part of the upper jaw containing the incisive teeth, on the
development of which depends the prognathic or brutish character -
of a skull. Now the extent of the premaxillaries below the nostril
is not only relatively but absolutely less in the gorilla, and con-
sequently the profile of the skull is less convex at this part, or less
‘prognathic, than in the chimpanzee. Notwithstanding the degree
in which the skull of the gorilla surpasses in size that of the chim-
panzee, especially when the two are compared on a front view, the
breadth of the premaxillaries and of the four incisive teeth is the
same in both. In the relative degree, therefore, in which these
bones are smaller than in the chimpanzee, the gorilla, in this most
important character, comes nearer to man. In the gibbons the
incisors are relatively smaller than in the gorilla, but the pre-
maxillaries bear the same proportional size, in the adult male
slamang.

Next, as regards the nasal bones. In the chimpanzee, as in the
orangs and gibbons, they are as flat to the face as in any of the
lower Simiw. In the gorilla, the median coalesced margins of the
upper half of the nasal bones are produced forwards ; in a slight
degree it is true, but affording a most significant evidence of
nearer resemblance to man. In the same degree they impress
that anthropic feature upon the face of the living gorilla. In some
pig-faced baboons there are ridges and prominences in the naso-
facial part of the skull; but they do not really affect the question
as between the gorilla and chimpanzee. All naturalists know that
the semnopithéques of Borneo have long noses; but the proboscidi-
form appendage which gives so ludicrous a mask to those monkeys
is scarcely the homologue of the human nose, and is unaccompanied
by any such modification of the nose-bones as gives the true
anthropoid character to the human skull, and to which only the
gorilla, in the ape tribe, makes any approximation.

No orang, chimpanzee, or gibbon shews any rudiment of mas-
toid processes; but they are present in the gorilla, smaller indeed
than in man, but unmistakeable; they are, as in man, cellular, and
with a thin outer plate of bone. This fact led me to express,
when in respect to the gorilla, only the skull had reached me, the
following inference, viz.: ‘from the nearer approach which the
gorilla makes to man in comparison with the chimpanzee, or orang,
in regard to the mastoid processes, that it assumed more nearly
and more habitually the upright attitude than those inferior anthro-

83

poid apes do.” This inference has been fully borne out by the rest
of the skeleton of the gorilla, subsequently acquired.

In the chimpanzee, as in the orangs, gibbons, and inferior
Simi, the lower surface of the long tympanic or auditory process
is more or less flat and smooth, developing in the chimpanzee
only a slight tubercle, anterior to the stylohyal pit. In the gorilla
the auditory process is more or less convex below, and developes a
ridge, answering to the vaginal process, on the outer side of the
carotid canal. The processes posterior and internal to the glenoid
articular surface, are better developed, especially the internal one,
in the gorilla than in the chimpanzee; the ridge which extends
from the ectopterygoid along the inner border of the foramen ovale,
terminates in the gorilla by an angle or process answering to that
called ‘styliform’ or ‘spinous’ in man, but of which there is no
trace in the chimpanzee, orang, or gibbon.

The orbits have a full oval form in the orang; they are almost
circular in the chimpanzee and siamang; more nearly circular,
and with a more prominent rim in the smaller gibbons; in the
gorilla alone do they present the form which used to be deemed
peculiar to man. There is not much physiological significance in
some of the latter characters; but, on that very account, I
deem them more instructive and guiding in the actual com-
parison. The occipital foramen is nearer the back part of the
cranium, and its plane is more sloping, less horizontal, in the
siamang, than in the chimpanzee and gorilla. Considering the less
relative prominence of the fore part of the jaws in the siamang, as
compared with the chimpanzee, the occipital character of that
gibbon and of-other species of Hylobates indicates well their
inferior position in the quadrumanous scale.

In the greater relative size of the molars, compared with the
incisors, the gorilla makes an “important closer step towards man
than does the chimpanzee. The molar teeth are relatively so
small in the siamang, that notwithstanding the small size of the
incisors, the proportion of those teeth to the molars is only the
same as in the gorilla: in other gibbons (Hylobates lar), the four
lower incisors occupy an extent equal to that of the first four
molars, in the chimpanzee equal to that of the first three molars,
in the siamang equal to that of the first two molars and rather
more than half of the third, in man equal to the first two molars
and half of the third: in this comparison the term molar is applied
to the bicuspids.

G2

84

The proportion of the ascending ramus to the length of the
lower jaw tests the relative affinity of the tailless apes to man.

In a profile of the lower jaw, compare the line drawn vertically
from the top of the coronoid process to the horizontal length along
the alveoli. In man and the gorilla it is about 7-10ths, in the
chimpanzee 6-10ths, in the siamang it is only 4-10ths. The
siamang further differs in the shape and production of the angle of
the jaw, and in the shape of the coronoid process, approaching the
lower simiz in both these characters. In the size of the post-
glenoid process, in the shape of the glenoid cavity which is almost
flat, in the proportional size of the petrous bone, and in the position
of the foramen caroticum, the siamang departs further from the
human type and approaches nearer that of the tailed simiz than
the gorilla does, and in a marked degree.

Every legitimate deduction from a comparison of cranial cha-
racters makes the tailless Quadrumana recede from the human
type in the following order,—gorilla, chimpanzee, orangs, gibbons ;
and the last-named in a greater and more decided degree.

Those comparisons have of late been invested with additional
interest from the discoveries of remains of quadrumanous species in
different members of the tertiary formations.

The first quadrumanous fossil, the discovery of which by Lieuts.
Baker and Durand is recorded in the Jowrnal of the Asiatic So-
ciety of Bengal, for November, 1836, has proved to belong, like
subsequently discovered quadrumanous fossils in the Sewalik (pro-
bably miocene) tertiaries, to the Indian genus Semnopithecus. The
quadrumanous fossils discovered in 1839, in the eocene deposits of
Suffolk, belong to a genus (Hopithecus) having its nearest affinities
gvith Macacus. The monkey’s molar tooth from the pliocene beds
of Essex is most closely allied to the Macacus sinicus. The
remains of the large monkey, 4 feet in height, discovered in 1839
by Dr Lund in a limestone cavern in Brazil was shewn by its
molar dentition (» = m 53) to belong to the platyrrhine
family now peculiar to South America. The lower jaw and teeth —
of the small quadrumane discovered by M. Lartet in a miocene
bed of the south of France, and described by him and De
Bainville, is so closely allied to the gibbons as to scarcely justify
the generic separation which has been made for it under the name
Pliopithecus.

Finally, a portion of a lower jaw with teeth and the shaft of a

85

humerus of a quadrumanous animal (Dryopithecus), equalling the
size of those bones in man, have been discovered by M. Fontan,
of Saint-Gaudens, in a marly bed of upper miocene age, forming
the base of the plateau on which that town is built. The molar
teeth present the type of grinding surface of those of the gibbons
(Hylobates), and as in that genus the second true molar is larger
than the first, not of equal size, as in the human subject and chim-
panzee. The premolars have a greater antero-posterior extent,
relatively, than in the chimpanzee; and in this respect agree more
with those in the siamang. The first premolar has the outer cusp
raised to double the height of that of the second; its inner lobe
appears from M. Lartet’s figure to be less developed than in the
gorilla, certainly less than in the chimpanzee. The posterior talon
of the second premolar is more developed, and consequently the
fore and aft extent of the tooth is greater than in the chimpanzee ;
thereby the second premolar of Dryopithecus more resembles that
in Hylobates, and departs further from the human type.

The canine, judging from the figures published by M. Lartet’,
seems to he less developed than in the male chimpanzee, gorilla, or
‘orang. In which character the fossil, if it belonged to a male,
makes a nearer approach to the human type; but it is one which
many of the inferior monkeys also exhibit, and is by no means to
be trusted as significant of true affinity, supposing even the sex of
the fossil to be known as being male.

The shaft of the humerus, found with the jaw, is peculiarly
rounded, as it is in the gibbons and sloths, and offers none of those
angularities and ridges which make the same bone in the chim-
panzee and orang come so much nearer in shape to the humerus of
the human subject. The fore part of the jaw, as in the siamang, is
more nearly vertical than in the gorilla or chimpanzee, but whe-
ther the back part of the jaw may not have departed in a greater
degree from the human type than the fore part approaches it, as is
the case in the siamang, the state of the fossil does not allow of
determining. One significant character is, however, present,—the
shape of the fore part of the coronoid process. It is slightly con-
vex forwards, which causes the angle it forms with the alveolar
border to be less open. The same character is present in the
gibbons. The fore part of the lower half of the coronoid process
in man is concave, as it is likewise in the gorilla and chimpanzee.
I am acquainted with this interesting fossil, referred to a genus

1 Comptes Rendus de UV’ Académie des Sciences, Paris, Vol, XLII.

86

called Dryopithecus, only by the figures published in the 43rd
volume of the Comptes Rendus de 0 Académie des Sciences. From
these it appears that the canine, two premolars, and first and
second true molars are in place. The socket of the third molar is
empty, but widely open above; from which I conclude that the
third molar had also cut the gum, the crown being completed, but
not the fangs. If the last molar had existed as a mere germ, it
would have been preserved in the substance of the jaw.

In a young siamang, with the points of the permanent canines
just protruding from the socket, the crown of the last molar is
complete, and on a level with the base of that of the penultimate
molar, whence I infer that the last molar would have cut the gum
as soon as, if not before, the crown of the canine had been com-
pletely extricated. This dental character, the conformation and
relative size of the grinding teeth, especially the fore-and-aft
extent of the premolars, all indicate the close affinity of the
Dryopithecus with the Pliopithecus and existing gibbons; and
this, the sole legitimate deduction from the maxillary and dental
fossils, is corroborated by the fossil humerus, fig. 9, in the above-
cited plate.

There is no law of correlation by which, from the portion of
jaw with teeth of the Dryopithecus, can be deduced the shape of
the nasal bones and orbits, the position and plane of the occipital
foramen, the presence of mastoid and vaginal processes, or other
cranial characters determinative of affinity to man; much less any
ground for inferring the proportions of the upper to the lower
limbs, of the humerus to the ulna, of the pollex to the manus, or
the shape and development of the iliac bones. All those charac-
ters which do determine the closer resemblance and affinity of the
genus Troglodytes to man, and of the genus Hylobates to the tailed
monkeys, are at present unknown in respect of the Dryopithecus.
A glance at fig. 5 (Gordla), and fig. 7 (Dryopithecus), of the plate
of M. Lartet?s memoir, would suffice to teach their difference of
bulk, the gorilla being fully one-third larger. The statement that
the parts of the skeleton of the Dryopithecus as yet known, viz.,
the two branches of the lower jaw and the humerus, ‘are sufficient
to shew that in anatomical structure, as well as stature, 1t came
nearer to man than any quadrumanous species, living or fossil,
before known to zoologists'’, is without the support of any ade-

1 Sir Chas. Lyell, Supplement to the Fifth Edition of a Manual of Ele-
mentary Geology, 8vo., 1859, p. 14.

87

quate fact, and in contravention of most of those to be deduced
from M. Lartet’s figures of the fossils. Those parts of the Dryo-
pithecus merely shew—and the humerus in a striking manner—its
nearer approach to the gibbons. The most probable conjecture
being that it bore to them, in regard to size, the like relations
which Dr Lund’s Protopithecus bore to the existing Mycetes.
Whether, therefore, strata of such high antiquity as the miocene
may reveal to us ‘forms in any degree intermediate between the
chimpanzee and man’ awaits an answer from discoveries yet to
be made; and the anticipation that the fossil world ‘may here-
after supply new osteological links between man and the highest
known quadrumana’ may be kept in abeyance until that world
has furnished us with the proofs that a species did formerly exist
which came as near to man as does the orang, the chimpanzee, or
the gorilla.

Of the nature and habits of the last-named species, which really
offers the nearest approach to man of any known ape, recent or
fossi], the lecturer had received many statements from individuals
resident at or visitors to the Gaboon, from which he selected the
« following as most probable, or least questionable.

Gorilla-land is a richly wooded extent of the western part of
Africa, traversed by the rivers Danger and Gaboon, and extending
from the equator to the 10th or 15th degree of south latitude.
The part where the gorilla has been most frequently met with
presents a succession of hill and dale, the heights crowned with
lofty trees, the valleys covered by coarse grass, with partial scrub
or scattered shrubs. Fruit trees of various kinds abound both on
the hills and in the valleys; some that are crude and uncared for
by the negroes are sought out and greedily eaten by the gorillas,
and as different kinds come to maturity at different seasons, they
afford the great denizen of the woods a successive and unfailing
supply of these indigenous fruit trees. [I am able through the
contributions of kind and zealous correspondents to specify the
following :—

The palm-nut (Zlais guiniensis) of which the gorillas greatly
affect the fruit and upper part of the stipe, called the ‘ cabbage.’
The negroes of the Gaboon have a tradition that their forefathers
first learnt to eat the ‘cabbage, from seeing the gorilla eat it,
concluding that what was good for him must be good for man.

The ‘ginger-bread tree’ (Parinariwm excelsum), which bears a
plum-lke fruit.

88

The papau tree (Carica papaya).

The banana (Musa sapientivm), and another species (Jusa
paradisiaca).

The Amomum Afzelii and Am. grandiflorum.

A tree, with a shelled fruit, like a walnut, which the gorilla
breaks open with the blow of a stone.

A tree, also botanically unknown, with a fruit like a cherry.

Such fruits and other rich and nutritious productions of the
vegetable kingdom, constitute the staple food of the gorilla, as they
do of the chimpanzee. The molar teeth, which alone truly indicate
the diet of an animal, accord with the statements as to the
frugivorous character of the gorilla: but they also sufficiently
answer to an. omnivorous habit to suggest that the eggs and callow
brood of nests discovered in the trees frequented by the gorilla
might not be unacceptable.

The gorilla makes a sleeping place like a hammock, connecting
the branches ofa sheltered and thickly leaved part of a tree by means
of the long tough slender stems of parasitic plants, and lining it
with the broad dried fronds of palms, or with long grass. , This
hammock-like abode may be seen at different heights, from 10 feet to
40 feet from the ground, but there is never more than one such
nest in a tree.

They avoid the abodes of man, but are most commonly seen in
the months of September, October, and November, after the negroes
have gathered their outlying rice crops, and have returned from the
‘bush’ to the village. So observed, they are described to be usually
in pairs; or, if more, the addition consists of a few young ones, of
different ages, and apparently of one family. The gorilla is not
gregarious. The parents may be seen sitting on a branch, resting
the back against the tree-trunk—the hair being generally rubbed off
the back of the old gorilla from that habit—perhaps munching their
fruits, whilst the young gorillas are at play, leaping and swinging
from branch to branch, with hoots or harsh cries of boisterous
mirth.

If the old male be seen alone, or when in quest of food, he is
usually armed with a stout stick, which the negroes aver to be the
weapon with which he attacks his chief enemy the elephant. Not
that the elephant directly or intentionally injures the gorilla, but,
deriving its subsistence from the same substances, the ape regards
the great proboscidian as a hostile intruder. When therefore he
discerns the elephant pulling down and wrenching off the branches

89 ’

of a favourite tree, the gorilla, stealing along the bough, strikes
the sensitive proboscis of the elephant with a violent blow of his
club, and drives off the startled giant trumpeting shrilly with
rage and pain. |

In passing along the ground from one detached tree to another
the gorilla is said to walk semi-erect, with the aid of his club, but
with a waddling awkward gait; when without a stick, he has been
seen to walk as a biped, with his hands clasped across the back of
his head, instinctively so counterpoising its forward projection. If
the gorilla be surprised and approached while on the ground, he
drops his stick, betakes himself to all-fours, applying the back part of
the bent knuckles of his fore-hands to the ground, and makes his
way rapidly, with an oblique swinging kind of gallop, to the nearest
tree. There he awaits his pursuer, especially if his family be near,
and requiring his defence. No negro willingly approaches the tree
in which the male gorilla keeps guard. Even with a gun the negro
does not rashly make the attack, but reserves his fire in self-defence.
The enmity of the gorilla to the whole negro race, male and female,
is uniformly testified to. The young men of the Gaboon tribe make
armed excursions into the forests, in quest of ivory. The enemy
they most dread on these occasions is the gorilla. If they have
come unawares too near him with his family, he does not, like the
lion, sulkily retreat, but comes rapidly to the attack, swinging
down to the lower branches, and clutching at-the nearest foe. The
hideous aspect of the animal, with his green eyes flashing with rage, »
is heightened by the skin over the prominent roof of the orbits being
drawn rapidly backward and forward, the hair erected, and causing
a horrible and fiendish scowl. If fired at and not mortally hit, the
gorilla closes at once upon his assailant and inflicts most dangerous,
if not deadly, wounds with his sharp and powerful tusks. The
commander of a Bristol trader informed me that he had seen a
negro at the Gaboon frightfully mutilated by the bite of the
gorilla, from which he had recovered. Another negro exhibited
to the same voyager a gun-barrel bent and partly flattened by the
bite of a wounded gorilla, in its death-struggle.

Negroes when stealing through the gloomy shades of the tropical
forest become sometimes aware of the proximity of one of these
frightfully formidable apes by the sudden disappearance of one of
their companions, who is hoisted up into the tree, uttering, perhaps,
a short choking cry. In a few minutes he falls to the ground a
strangled corpse. The gorilla, watching his opportunity, has let

90

down his huge hind-hand, seized the passing negro by the neck,
with vice-like grip, has drawn him up to higher branches, and
dropped him when his struggles had ceased.

The strength of the gorilla is such as to make him a match for
a lion, whose tusks his own almost rival. Over the leopard,
invading the lower branches of the gorilla’s dwelling tree, he will
gain an easier victory; and the huge canines, with which only the
male gorilla is furnished, doubtless have been assigned to him for
defending his mate and offspring.

The skeleton of the old male gorilla obtained for the British
Museum in 1857, shews an extensive fracture, badly united, of the
left arm-bone, which has been shortened, and gives evidence of long
suffering from abscess and partial exfoliation of bone. The upper
canines have been wrenched out or shed, some time before death,
for their sockets have become absorbed.

The redeeming quality in this fragmentary history of the gorilla
is the male’s care of his family, and the female’s devotion to her
young.

It is reported that a French natural-history collector, accom-
panying a party of the Gaboon negroes into the gorilla woods,
surprised a female with two young ones on a large boabdad
(Adansonia), which stood some distance from the nearest clump.
She descended the tree, with the youngest clinging to her neck,
and made off rapidly on all-fours to the forest, and escaped. The
deserted young one on seeing the approach of the men began to
utter piercing cries: the mother, having disposed of her infant
in safety, returned to rescue the older offspring, but before she
could descend with it her retreat was cut off. Seeing one of the
negroes level his musket at her, she, clasping her young with one
arm, waved the other, as if deprecating the shot; the ball passed
through her heart, and she fell with her young one clinging to her.
It was a male, and survived the voyage to Havre, where it died on
arriving. I have examined the skeleton of this young gorilla in
the museum of natural history at Caen, and am indebted to Professor
Deslongchamps, Dean of the Faculty of Sciences in that town, for
drawings of this rare specimen.

There might be more difficulty in obtaining a young gorilla for
exhibition than a young chimpanzee. But as no full-grown chim-
panzee has ever been captured, we cannot expect the larger and
much more powerful adult gorilla to be ever taken alive.

A bold negro, the leader of an elephant-hunting expedition,

91

being offered a hundred dollars if he would bring back a live
gorilla, replied, ‘If you gave me the weight of yonder hill in gold
coins, I could not do it!’ : ;

All the terms of the aborigines in respect to the gorilla imply
their opinion of his close kinship to themselves. But they have a
low opinion of his intelligence. They say that during the rainy
season he builds a house without a roof. The natives on their
hunting excursions light fires for their comfort and protection by
night; when they have gone away, they affirm that the gorilla will
come down and warm himself at the smouldering embers, but has
not wit enough to throw on more wood, out of the surrounding
abundance, to keep the fire burning,—‘ the stupid old man!’

Every account of the habits of a wild animal obtained at
second hand from the reports of aborigines has its proportion of
‘apocrypha.’ I have restricted myself to the statements that
have most probability and are in accordance with the ascertained
structures and powers of the animal, and would only add the
averment and belief of the Gaboon negroes that when a gorilla
dies, his fellows cover the corpse with a heap of leaves and loose
earth collected and scraped up for the purpose.

A most singular phenomenon in natural history, if one reflects
on the relations of things, is this gorilla! Limited as it is in its
numbers and geographical range, one discerns that the very peculiar
conditions of its existence—abundance of wild fruit—needs must
be restricted in space; but, concurring in a certain part of Africa,
there lives the creature to enjoy them.

The like conditions exist in Borneo and Sumatra, and there
also a correlative human-like ape, of similar stature, tooth-armour,
and force, exists at their expense. Neither orangs nor gorillas,
however, minister to man’s use directly or indirectly. Were they
to become extinct, no sign of the change or break in the links of
life would remain. What may be their real significance ?

In regard to the ancient notices which may relate to the great
anthropoid ape of Africa, I may quote the following passage from
the ‘Periplus, or Voyage of Hanno, which has been supposed to
refer to the species in question :—‘ On the third day, having sailed
from thence, passing the streams of fire, we came to a bay called the
Horn of the South. In the recess there was an island like the first,
having a lake, and in this there was another island full of wild men.
But much the greater part of them were women, with hairy bodies,
whom the interpreters called “ gorillas.” But, pursuing them, we

92

were not able to take the men; they all escaped, being able to climb
the precipices, and defended themselves with pieces of rock. But
three females, who bit and scratched those who led them, were
not willing to follow. However, having killed them, we flayed
them, and conveyed the skins to Carthage. For we did not sail
any further, as provisions began to fail.’ This encounter indicates,
therefore, the southernmost point on the west coast of Africa
reached by the Carthaginian navigator.

To an inquiry by an eminent Greek scholar, how far the
newly-discovered great ape of Africa bore upon the question of
the authenticity of the Periplus? I have replied:—‘The size and
form of the great ape, now called “gorilla,” would suggest to
Hanno and his crew no other idea of its nature than that of a
kind of human being; but the climbing faculty, the hairy body,
and the skinning of the dead specimens, strongly suggest that
they were large anthropoid apes. The fact that such apes, having
the closest observed resemblance to the negro, being of human
stature and with hairy bodies, do still exist on the west coast of
Africa, renders it highly probable that such were the creatures
which Hanno saw, captured, and called “Gorullai.” ’

The brief observation made by Battell in West tropical Africa,
1590, recorded in Purchas’s Pilgrimages, or Relations of the World,
1748, of the nature and habits of the large human-like ape which
he calls ‘ pongo,’ more decidedly refers to the gorilla. Other notices,
as by Nieremberg and Bosman, applied by Buffon to Battell’s pongo, .
were deemed valueless by Cuvier, who altogether rejected the
conclusions of his great predecessor as to the existence of any such
ape. ‘This name of pongo or boggo, given in Africa to the chim-
panzee or to the mandril, has been applied,’ writes Cuvier, ‘by
Buffon to a pretended great species of ourang-utan, which was
nothing more than the imaginary product of his combinations.’
After the publication of Cuvier’s Régne Animal, the supposed
species was, by the high authority of its author, banished from
natural history; it has only been authentically reintroduced since
the intelligent attention of Dr Savage was directed to the skull,
which he first saw at the Gaboon in 1847, and took my opinion
upon.

Having premised the foregoing account of the mature characters
of the different species of orangs and chimpanzees, in regard to their
relative proximity to the human species, I next proceed to shew how
their structure contrasts with-that of man. With regard to the

93

dentition of these anthropoid apes, the number and kinds of the teeth,
like those of all the quadrumana of the old world, correspond with
those in the human subject; but all these apes differ in the larger
proportionate size of the canine teeth, which necessitates a certain
break in the series, in order that the prolonged points of the canine
teeth may pass into their place when the mouth is completely
closed. In addition to the larger proportionate size of the incisors
and canines, the bicuspids in both jaws are implanted by three
distinct fangs—two external and one internal: in the human
species, the bicuspids are implanted by one external and one
internal fang: in the highest races of man these two fangs are
often connate ; very rarely is the external fang divided, as it con-
stantly is in all the species of the orang and the chimpanzee.

With regard to the catarrhine, or old-world quadrumana, the
number of milk teeth is twenty, as in the human subject. But
both chimpanzees and orangs differ from man in the order of de-
velopment of the permanent series of teeth: the second true molar
comes into place before either of the bicuspids have cut the gum,
and the last molar is acquired before the permanent canine. We
may well suppose that the larger grinders are earlier required by
the frugivorous apes than by the omnivorous human race; and
one condition of the earlier development of the canines and
bicuspids in man, may be their smaller relative size as compared
with the apes. The great difference is the predominant develop-
ment of the permanent canine teeth, at least in the males of the
orangs and chimpanzees; for this is a sexual distinction, the canines
in the females never presenting the same large proportion. In man,
the dental system, although the formula is the same as in the
apes, is peculiar for the equal length of the teeth, arranged in
an uninterrupted series, and shews no sexual distinctions. The
characteristics of man are exhibited in a still more important
degree in the parts of the skeleton. His whole framework pro-
claims his destiny to carry himself erect ; the anterior extremities
are liberated from any service in the mere act of locomotion.

With regard to the foot, I have shewn in my work On the
Nature of Limbs, that in tracing the manifold and progressive
changes of the feet in the mammalian series, in those forms where
it is normally composed of five digits, the middle is usually the
largest; and this is the most constant one. The modifications in
the hind foot, in reference to the number of digits, are, first, the
reduction and then the removal, of the innermost one; then the

94.

corresponding reduction and removal of the outer one; next, of
the second and fourth digits, until it is reduced to the middle
digit, as in the horse.

The innermost toe, the first to dwindle and disappear in the
brute series, is, in Man, developed to a maximum size, becoming
emphatically the ‘great toe,’ one of the most essential charac-
teristics of the human frame. It is made the powerful fulcrum
for that lever of the second kind, which has its resistance in the
tibio-astragalar joint, and the power applied to the projecting heel-
bone: the superincumbent weight is carried further forward upon
the foot, by the more advanced position of the astragalus, than in
the ape tribe; and the heel-bone is much stronger, and projects
more backwards.

The arrangement of the powerfully-developed tarsal and meta-
tarsal bones is such as to form, in Man, a bony arch, of which the
two piers rest upon the proximal joint of the great toe and the end
of the heel. Well-developed cuneiform bones combine with the
cuboid to form a second arch, transverse to the first. There are no
such modifications in the gorilla or orang, in which the arch, or
rather the bend of the long and narrow sole, extends to the extreme
end of the long and curved digits, indicating a capacity for grasping.
Upon these two arches the superincumbent weight of man is solidly
and sufficiently maintained, as upon a low dome, with this further
advantage, that the different joints, cartilages, coverings, and
synovial membranes, give a certain elasticity to the dome, so that
in leaping, running, or dropping from a height, the jar is diffused
and broken before it can be transmitted to affect the enormous
brain-expanded cranium. ‘The lower limbs in man are longer in
proportion to the trunk than in any other known mammalian
animal. The kangaroo might seem to be an exception, but if the
hind limbs of the kangaroo are measured in relation to the trunk,
they are shorter than in the human subject. In no animal is the
femur solong in proportion to the leg as in man. In none does
the tibia expand so much at its upper end. Here it presents two
broad, shallow cavities, for the reception of the condyles of the
femur. Of these condyles, in man only is the innermost longer
than the outermost; so that the shaft of the bone inclines a little
outwards to its upper end, and joinsa ‘neck’ longer than in other
animals, and set on at a very open angle. The weight of the body,
received by the round heads of the thigh bones, is thus transferred
to a broader base, and its support in the upright posture facilitated.

-

. 95

There is also the collateral advantage of giving more space to those
powerful adductor muscles that assist in fixing the pelvis and trunk
upon the hind limbs. With regard to the form of the pelvis, the
iliac bones, compared with those in the gorilla, are short and
broad : they are more bent forwards, the better to receive and
sustain the abdominal viscera, and are more expanded behind to
give adequate attachment to the powerful glutei muscles, which are
developed to a maximum in the human species, in order to give a
firm hold of the trunk upon the limbs, and a corresponding power
of moving the limbs upon the trunk. The tuberosities of the
ischium are rounded, not angular, and not inclined outwards, as
in the gorilla and the rest of the ape tribe. The symphysis
pubis is shorter than in the apes. The tail is reduced to three or
four stunted vertebre, anchylosed to form the bone called ‘os
coccygis.’ The true vertebre, as they are called in human anatomy,
correspond in number with those of the chimpanzee and the orang,
and in their divisions with the latter species, there being twelve
thoracic, five lumbar, and seven cervical. This movable part of the
column is distinguished by a beautiful series of sigmoid curves, con-
vex forwards in the loins, concave in the back, and again slightly
convex forwards in the neck. The cervical vertebrae, instead of
having long spinous processes, have short processes, usually more
or less bifurcated. The bodies of the true vertebrze increase in size
from the upper dorsal to the last lumbar, which rests upon the
base of the broad wedge-shaped sacrum, fixed obliquely between
the sacro-iliac articulations. All these curves of the vertebral
column, and the interposed elastic cushions, have relation to the
libration of the head and upper limbs, and the diffusion and the
prevention of the ill effects from shocks in many modes of loco-
motion which man, thus organised for an erect position, is capable
of performing. The arms of man are brought into more symme-
trical proportions with the lower limbs; and their bony framework
shews all the perfections that have been superinduced upon it in
the mammalian series, viz., a complete clavicle, the antibrachial
bones so adjusted as to permit the rotary movements of pronation
and supination, as well as of flexion and extension; manifesting
those characters which adapt them for the manifold application of
that most perfect and beautiful of prehensile instruments, the hand.
The scapula is broad, with the glenoid articulation turned out-
wards ; the clavicles are bent ina slight sigmoid flexure; the
humerus exceeds in length the bones of the fore-arm. The carpal

96 :

‘bones are eight in number. The thumb is developed far beyond
any degree exhibited by the highest quadrumana, and is the most
perfect opposing digit in the animal creation.

The skull is distinguished by the enormous expansion of the
brain-case; by the restricted growth of the bones of the face,
especially of the jaws, in relation to the small, equally-developed
teeth; and by the early obliteration of the maxillo-intermaxillary
suture. To balance the head upon the neck-bone, we find the
condyles of the occiput brought forward almost to the centre of
the base of the skull, resting upon the two cups of the atlas,
so that there is but a slight tendency to incline forwards when
the balancing action of the muscle ceases, as when the head nods
during sleep, in an upright posture. Instead of the strongly
developed occipital crest, we find a great development of true
mastoid processes advanced nearer to the middle of the sides of
the basis cranil, and of which there is only the rudiment in the
gorilla. The upper convexity of the cranium is not interrupted
by any sagittal or parietal criste. The departure from the arche-
type, in the human skull, is most conspicuous, in the vast expanse
of the neural spines of the three chief cranial vertebra, viz.
occipital, parietal, and frontal.

‘To what extent,’ it may next be asked, ‘does man depart from
the typical character of his species?’ With regard to the kind and
amount of variety in mankind, we find, propagable and character-
istic of race, a difference of stature, a difference in regard to colour
of skin, difference in both colour and texture of the hair, and cer-
tain differences in the osseous framework.

As to stature, the Bushmen of South Africa and the natives of
Lapland exhibit the extreme of diminution, ranging from four to
five feet. Some of the Germanic races and the Patagonian Indians
exhibit the opposite extreme, ranging from six to seven feet. The
medium size prevails generally throughout the races of mankind.

With reference to the characteristics of colour, which are ex-
treme, we have now opportunities of knowing how much that
character is the result of the influence of climate. We know it
more particularly by that most valuable mode of testing such influ-
ences which we derive from the peculiarity of the Jewish race. For
1800 years that race has been dispersed in different latitudes and
climates, and they have preserved themselves distinct from inter-
mixture with other races of mankind. There are some Jews still
lingering in the valleys of the Jordan, having been oppressed by the

vv er. ee

97

successive conquerors of Syria for ages,—a low race of people, and
described by trustworthy travellers as being as black as any of the
Ethiopian races. Others of the Jewish people, participating in
European civilization, and dwelling in the northern nations, shew
instances of the light complexion, the blue eyes, and light hair of
the Scandinavian families. The condition of the Hebrews, since
their dispersion, has not been such as to admit of much admixture
by the proselytism of household slaves. Weare thus led to account
for the differences in colour, by the influence of climate, without
having to refer them to original or specific distinctions.

As to the difference in size in mankind, it is slight in com-
parison with what we observe in the races of the domestic dog,
where the extremes of size are much greater than can be found
in any races of the human species.

With reference to the modifications of the bony structure, as
characteristic of the races of mankind, they are almost confined
to the pelvis and the cranium. In the pelvis the difference is a
slight, yet apparently a constant one. The pelvis of the adult
negro may sometimes be distinguished from that of the European
. by the greater proportional length and less proportional breadth of
the iliac bones ; but how trifling is this difference compared with
that marked distinction in the pelvis which the gorilla and orang-
outang present !

With regard to the cranial differences, 1 have selected for com-
parison three extreme specimens of skulls characteristic of race: one
of an aboriginal of Van Diemen’s Land (the lowest of the Melanian
or dark-coloured family), a well-marked Mongolian, and a well-
formed European skull. The differences are chiefly these. In
the low, uneducated, uncivilised races, the brain is rather smaller
than in the higher, more civilised, and more educated races;
consequently the cranium rises and expands in a less degree.
Concomitant with this contraction of the brain-case is a greater
projection of the fore part of the face; whether it may be from a
longer exercise of the practice of suckling, or a more habitual ap-
plication of the teeth in the premaxillary part of the jaw, and in
the corresponding part of the lower jaw, in biting and gnawing
tough, raw, uncooked substances,—the anterior alveolar part of the
jaws does project more in those lower races; but still to an insigni-
ficant degree compared with the prominence of that part of the
skull in the large apes. And while alluding to them, I may again
advert to the distinction between them and the lowest of the

H

98

human races, which is afforded by the pre-maxillary bone, already
referred to. In the young orang-utan, even when the change of
dentition has begun, the suture between that bone and the
maxillary is present; and it is not until the large canine teeth are
developed, that the stimulus of the vascular system, in the con-
comitant expansion and growth of the alveoli, tends to obliterate
the suture. In the young chimpanzee, the maxillary suture dis-
appears earlier, at least on the facial surface of the upper jaw. In
the human subject those traces disappear still earlier, and in regard
to the exterior alveolar plates, the inter-maxillary and maxillary
bones are connate. But there may be always traced in the human
foetus the indications of the palatal and nasal portions of the
maxillo-intermaxillary suture, of which the poet Goethe was the
first to appreciate the full significance.

In the Mongolian skull there is a peculiar development of the
cheek-bones, giving great breadth and flatness to the face, a broad
cranium, with a low forehead, and often with the sides sloping away
from the median sagittal tract, something like a roof; whereas, in
the European, there is combined, with greater capacity of the
cranium, a more regular and beautiful oval form, a loftier and
more expanded brow, a minor prominence of the malars, and a less
projection of the upper and lower jaws. All these characteristics
necessarily occasion slight differences in the facial angle. On a
comparison of the basis cranil, the strictly bimanous characteristics
in the position of the foramen magnum and occipital condyles, and
of the zygomatic arches, are as well displayed in the lowest as in the
highest varieties of the human species.

With regard to the value to be assigned to the above defined
distinctions of race:—in consequence of not any of these differences
being equivalent to those characteristics of the skeleton, or other
parts of the frame, upon which specific differences are founded by
naturalists in reference to the rest of the animal creation, I have
come to the conclusion that Man forms one species, and that these
differences are but indicative of varieties. As to the number of
these varieties :—from the very well marked and natural character
of the species, just as in the case of the similarly natural and
circumscribed class of birds, scarcely any two ethnologists agree as to
the number of the divisions, or ag to the characters upon which those
varieties are to be defined and circumscribed. In the subdivision
of the class of birds, the ornithological systems vary from two
orders to thirty orders; so with man there are classifications of

99

races varying from thirty to the three predominant ones which
Blumenbach first clearly pointed out,—the Ethiopian, the Mon-
golian, and the Caucasian or Indo-European. These varieties
merge into one another by easy gradations. The Malay and the
Polynesian link the Mongolian and the Indian varieties; and the
Indian is linked by the Esquimaux again to the Mongolian. The
inhabitants of the Andaman Islands, New Caledonia, New Guinea,
and Australia, in a minor degree seem to fill up the hiatus between
the Malayan and the Ethiopian varieties ; and in no case can a well
marked definite line be drawn between the. physical characteristics
of allied varieties, these merging more or less gradationally the one
into the other.

In considering the import and value of the osteological differ-
ences between the gorilla—the most anthropoid of all known brutes
—and man, in reference to the hypothesis of the origination of
species of animals by gradual transmutation of specific characters,
and that in the ascending direction :—it may be admitted that the
skeleton is modifiable to a certain extent by the action of the
muscles to which it is subservient, and that in domesticated races
. the size of the animal may be brought to deviate in both directions
from the specific standard. By the development of the processes,
ridges, and crests, and also by the general proportions of the bones
themselves, especially those of the limbs, the human anatomist
judges of the muscular power of the individual to whom a skeleton
under comparison has appertained.

The influence of muscular actions in the growth of bone is
more strikingly displayed in the change of form which the cranium
of the young carnivore or the sternum of the young bird undergoes
in the progress of maturity; not more so, however, than is mani-
fested in the progress of the development of the cranium of the
gorilla itself, which results in a change of character so great, as
almost to be called a metamorphosis.

In some of the races of the domestic dog, the tendency to the
development of parietal and occipital cristze is lost, and the cranial
dome continues smooth and round from one generation of the smaller
spaniel, or dwarf pug, e.g. to another; while, in the large deer-
hound, those bony cristz are as strongly developed as in the wolf,
Such modifications, however, are unaccompanied by any change in
the connexions, that is, in the disposition of the sutures, of the
cranial bones; they are due chiefly to arrests of development, to
retention of more or less of the characters of immaturity: even

100

the large proportional size of the brain in the smaller varieties of
house-dog is in a great degree due to the rapid acquisition by the
cerebral organ of its specific size, agreeably with the general law
of its development, but which is attended in the varieties cited by
an arrest of the general growth of the body, as well as of the particu-
lar developments of the skull in relation to the muscles of the jaws.

No species of animal has been subject to such decisive experi-
ments, continued through so many generations, as to the influence
of different degrees of exercise of the muscular system, difference
in regard to food, association with man, and the concomitant stimu-
lus to the development of intelligence, as the dog; and no domestic
animal manifests so great a range of variety in regard to general
size, to the colour and character of the hair, and to the form of the
head, as it is affected by different proportions of the cranium and
face, and by the intermuscular crests superadded to the cranial
parietes. Yet, under the extremest mask of variety so superin-
duced, the naturalist detects in the dental formula and in the
construction of the cranium the unmistakeable generic and specific
characters of the Canis familiaris.

This and every other analogy applicable to the present question
justifies the conclusion that the range of variety allotted to the
gorilla, chimpanzee, and orang-utan, under the operation of ex-
ternal circumstances favourable to their higher development, would
be restricted to differences of size, of colour, and other characters of
the hair, and of the shape of the head, in so far as this is influenced
by the arrest of general growth after the acquisition by the brain of
its mature proportions, and by the development, or otherwise, of
processes, crests, and ridges for the attachment of muscles. The
most striking deviations from the form of the human cranium which
that part presents in the great orangs and chimpanzees result from
the latter acknowledged modifiable characters, and might be simi-
larly produced; but not every deviation from the cranial struc-
ture of man, nor any of the important ones upon which the
naturalist relies for the determination of the genera Tvroglodytes
and Pithecus, have such an origin or dependent relation. The
gorilla, indeed, differs specifically from both the orang and man
in one cranial character, which no difference of diet, habit, or
muscular exertion can be conceived to affect.

The prominent superorbital ridge, for example, is not the con-
sequence or concomitant of muscular development; there are no
muscles attached to it that could have excited its growth. It is a

101

characteristic of the cranium of the genus Z’roglodytes from the time
of birth to extreme old age; by the prominent superorbital ridge,
for example, the skull of the young gorilla or chimpanzee with de-
ciduous teeth may be distinguished at a glance from the skull of an
orang at the same immature age; the genus Pithecus, Geoffr., being
as well recognised by the absence, as the genus 7’roglodytes is by the
presence, of this character. We have no grounds, from observation
or experiment, to believe the absence or the presence of a prominent
superorbital ridge to be a modifiable character, or one to be gained
or lost through the operations of external causes, inducing par-
ticular habits through successive generations of a species. It
may be concluded, therefore, that such feeble indication of the
superorbital ridge, aided by the expansion of the frontal sinuses,
as exists in man, is as much a specific peculiarity of the human
skull, in the present comparison, as the exaggeration of this
ridge is characteristic of the chimpanzees and its suppression of
the orangs.

The equable length of the human teeth, the concomitant absence
’ of any diastema or break in the series, and of any sexual difference
in the development of particular teeth, are to be viewed by the
light of actual knowledge, as being primitive and unalterable spe-
cific peculiarities of man.

Teeth, at least such as consist of the ordinary dentine of mam-
mals, are not organised so as to be influenced in their growth by
the action of neighbouring muscles; pressure upon their bony
sockets may affect the direction of their growth after they are pro-
truded, but not the specific proportions and forms of the crowns of
teeth of limited and determinate growth. The crown of the great
canine tooth of the male Zroglodytes gorilla began to be calcitied
when its diet was precisely the same as in the female, when both
sexes derived their sustenance from the mother’s milk. Its growth
proceeded and was almost completed before the sexual development
had advanced so as to establish those differences of habits, of force,
of muscular exercise, which afterwards characterise the two sexes.
The whole crown of the great canine is, in fact, calcified before it
cuts the gum or displaces its small deciduous predecessor; the
weapon is prepared prior to the development of the forces by which
it is to be wielded; it is therefore a structure fore-ordained, a
predetermined character of the great ape, by which that creature
is made physically superior to man; and one can as little conceive
the development of the canine tooth to be a result of external

102

stimulus, or as being influenced by the muscular actions, as the
development of the stomach, or of any internal gland.

The two external divergent fangs of the premolar teeth, and
the slighter modifications of the crowns of the molars and _ pre-
molars, appear likewise from the actual results of observation to be
equally predetermined and non-modifiable characters.

No known cause of change productive of varieties of mammalian
species could operate in altering the size, the shape, or the con-
nexions of the premaxillary bones, which so remarkably distinguish
the Troglodytes gorilla, not from man only, but from all other
anthropoid apes. We know as little the conditions which protract
the period of the obliteration of the sutures of the premaxillary
bones in the 7'r. gorilla beyond the period at which they disappear
in the 77. niger, as we do those that cause them to disappear in
man earlier than they do even in the smaller species of chimpanzee.

There is not, in fact, any other character than those founded
upon the developments of bone for the attachment of muscles, which
is known to be subject to change through the operation of external
causes; nine-tenths, therefore, of the differences, especially those
very striking ones manifested by the pelvis and pelvic extremities,
which I have cited in the memoirs on the subject, published in the
Zoological Transactions, as distinguishing the gorilla and chimpanzee
from the human species, must stand in contravention of the hypo-
thesis of transmutation and progressive development, until the
supporters of that hypothesis are enabled to adduce the facts and
cases which demonstrate the conditions of the modifications of such .
characters.

If the consideration of the cranial and dental characters of the
Troglodytes gorilla has led legitimately to the conclusion that it is
specifically distinct from the Z'roglodytes niger, the hiatus is still
greater that divides it from the human species, between the ex-
tremest varieties of which there is no osteological and dental
distinction which can be compared to that manifested by the shorter
premaxillaries and larger incisors of the Z'roglodytes niger as com-
pared with the 7’. gorilla.

The analogy which the establishment of the second and more
formidable species of chimpanzee in Africa has brought to light
between the representation of the genus Z'roglodytes in that con-
tinent, and that of the genus Pithecus in the great islands of the
Indian Archipelago, is very close and interesting. As the Z'roglo-
dytes gorilla parallels the Pithecus Wurmbii, so the Troglodytes

103

niger parallels the Pithecus morio, and an unexpected illustration
has thus been gained of the soundness of the interpretation
of the specific distinction of that smaller and more anthropoid
orang.

Tt is not without interest to observe, that as the generic forms
of the Quadrumana approach the Limanous order, they are repre-
sented by fewer species. The gibbons (Hy/lobates) scarcely number
more than half-a-dozen species; the orangs (Pithecus) have but two
species, or at most three; the chimpanzees (7’roglodytes) are repre-
sented by two species.

The unity of the human species is demonstrated by the con-
stancy of those osteological and dental characters to which the
attention is more particularly directed in the investigation of the
corresponding characters in the higher Quadrumana.

Man is the sole species of his genus, the sole representative of
his order and subclass.

Thus I trust has been furnished the confutation of the notion
of a transformation of the ape into man, which appears from a
favourite old author to have been entertained by some in his day.

“ And of a truth, vile epicurism and sensuality will make the
soul of man so degenerate and blind, that he will not only be con-
tent to slide into brutish immorality, but please himself in this very
opinion that he is areal brute already, an ape, satyre or baboon;
and that the best of men are no better, saving that civilising of
them and industrious education has made them appear in a more
refined shape, and long inculcate precepts have been mistaken for
connate principles of honesty and natural knowledge; otherwise
there be no indispensable grounds of religion and virtue, but what
has happened to be taken up by over-ruling custom. Which things,
I dare say, are as easily confutable, as any conclusion in mathe-
matics is demonstrable. But as many as are thus sottish, let them
enjoy their own wildness and ignorance; it is sufficient for a good
man that he is conscious unto himself that he is more nobly de-
scended, better bred and born, and more skilfully taught by the

purged faculties of his own minde’,.”

1 Henry More’s Conjectwra Cabbalistica, fol. (1662)—p. 175.

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CAMBRIDGE: PRINTED BY C. J. CLAY, M.A. AT THE UNIVERSITY PRESS.

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