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Full text of "On the classification and geographical distribution of the Mammalia, being a lecture on Sir Robert Reade's foundation, delivered before the University of Cambridge ... May 10, 1859. To which is added an appendix "On the gorilla," and "On the extinction and transmutation of species.""

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SSnftroftg of <ambrfoge, in tjjc &tiate*f^ouse, 

MAY 10, 1859. 














Gift of C. A. Kofoid 










I AVAIL 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, 



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 Kobert Keade'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 ' Hepl 
toTo/owi? 1 ,' animals generally, and by implication the 

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

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 modem 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 Pithecoida 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 
' Kodents' 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 Dischidce, or bisulcate quadrupeds, and 
Aschidce, 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 Ampkodonta, 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. 

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 1 ; 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. 761/09, 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. in. p. 289. 


To JOHN 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 Metliodica 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 1 . 

LINNAEUS first definitely and formally restored the great 
natural class I am now treating of to its Aristotelian inte- 
grity; 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, Linnseus and Buffon, had brought to light 
instances, e. 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 Ketocle. The 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 

1 'Kai avrl ovu< 





term proposed by Linnseus, 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 1 . 

Linnaeus, like Bay, 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 2 : 

[ Front teeth, none in either jaw . . . BRUTA. 
r TT 7 4. J Front teeth, cutters 2, laniaries o . . . GLIRES. 
3 r Ungmculate j Front ^ , ^ laniarieg x _ > PRIMATES. 

( Front teeth, piercers (6, 2, 10), laniaries i FER.E. 

S 1 TT 7 t J Front teeth, in both upper and lower jaw. BELLU^E. 

j t Front teeth, none in the upper jaw . . PECORA. 

^ t Mutkate -Teeth variable CETE. 

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 
Bay and Linnaeus 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 Linnaeus, however, it must be observed, 
that although he has followed Bay 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 l Le Regne Animal distribue d'apres 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 : 'TO,' (S 5&> /u.ti> 
Tobs) '5' &TOS, uffirep 5eA0is.' 

2 From the Systema Nalurw, ed. xn. Holmiae, Tom. I. p. 24. 

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 heterogeneous order which Linnaeus prepossessed 
in favour of the easily recognisable outward character by 
which he distinguished the class had characterised by the 
1 Mammce pectorales bince : dentes primores incisores : supe- 
riores i\ paralleli 1 ? 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 Linnsean genus Homo. The 
genus Simia of Linnaeus, with hands on the four extremities, 
became the order Quadrumana of Cuvier. The genus Ves- 
pertilio with the 'manus palmatse volitantes' formed the 
group Cheiroptera, answerable to the Dermaptera of Aristotle. 

KAY 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 (Echidna], and 
with the more strange duck-moles (Ornithorhynchus) of the 
same part of the world, to form the order Edentata of Cuvier, 
which answers to that called Bruta by Linnaeus, if the ele- 
phant and walrus be removed from it. The rest of Rays 
anomalous species exemplify the families Cheiroptera 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 Ferce of Linnaeus. 

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

1 Tom, cit. p. 24. 


unguiculate mammals of the rest of the world; 'some,' he 
writes, l corresponding with the Carnivora, some with the 
Rodentia, and others again with the Edentata, by their teeth 
and the nature of their food.' They formed a family of the 
Carnassiers in the first edition of the ' Rtyne Animal 1 , 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, c animaux a 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 Linnaean system, save that the manatee which Linna3us 
placed in the same group as the elephant is associated with 
the whale in the Regne Animal. 

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. 

2 . .' 




precedence of the colossal and sagacious elephant in the Cu- 
vierian scheme 1 . 

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 2 , and by having teats or mammae 
whence the name of the class. 

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

nutely cellular structure of the lungs, (fig. 1, ,) 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- 
nsean Society (Proceedings, 1857) and the Geological Society (Proceedings, Nov. 
1847, pp. 135140). 

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


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, b) 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 


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 oesophagus 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- 

The trunk- vertebras 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 vertebras are seven in 
number, neither more nor less. The lumbar vertebras 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 

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 gristly or bony abdominal ribs or 


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 

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 Ibone is of a finer 
arid more delicate structure than in Keptiles, and forms a 


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

The Mammalia, like Eeptilia and Pisces, include a few 
genera and species that are devoid of teeth ; the true ant- 
eaters (Myrmecophaga), the scaly anteaters or pangolins 
.(Mam's), and the spiny monotrematous anteater (Echidna), 
are examples of strictly edentulous Mammals. The Orni- 
thorhynchus has horny teeth, and the whales (Balcena and 
Balcenoptera) 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 

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 incisors; 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 


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 Bumi- 
nantia and most Bruta (Sloths, Armadillos, Orycteropes). In 
most Mammals teeth are situated in all the bones above men- 

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- 

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

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


species the tooth-matrix does not develope the germ of a 
second tooth, destined to succeed 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. p) ; 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 1 or those that generate a 

s, once ; 0tfw, I generate ; oSovs, tooth. 


single set of teeth, and the Diphyodonts 1 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,' e canines,' and 
' molars,' were given to the teeth, in Man and certain Mam- 
mals, as in Keptiles 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 Garni vora, 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 

Those teeth which are implanted in the premaxillary bones, 
and in the corresponding part of the lower jaw, are called 
'incisors' (fig. 2, *'), 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 Sis, twice ; <piju and odovs. See Philosophical Transactions, 1850, p. 493. 



teeth, which are not displaced by vertical successors, are the 
'molars' properly so called (m. 14). 

Fig. i. 

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. I 3). This difference forms a character of an 

Lower Jaw of a young Pig (Sus.) 

ordinal group in the mammalian class l . 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, 

1 Outlines of a Classification of the Mammalia, Trans. Zool. Soc. Vol. u. 
P- 330 ( l8 39)- 


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^>. 4, together with m. 3, are more 
or less incomplete, and will be found concealed in their closed 
alveoli 1 . 

The last deciduous molar, J.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 : 

. 33 1 1 44 33 

*-^> c -r=T' p-4=i> -3^ =44 >' 

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 being 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 I 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 


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, l 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 
zoplogy, 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 


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 offspring 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 


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 1 .' 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 3 , I pointed out that the Mammalia might be divided 
into 'placental' and ' implacentalV 

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, Cyclopcedia of Anatomy and Physiology, Vo\. 111^.383. 

3 Art. MAESUPIALIA, torn. cit. p. 257. 

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


group never presenting the same degree of correspondence of 
organic structure as the smaller moiety, I continued to pursue 
investigations, with the view of gaming 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 

Nevertheless there were genera of Mammals, e. g. the 
sloths, anteaters, armadillos, roussettes, giraffes, rhinoceroses, 
&c. 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 Linnaean 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 1 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 t,7ie Structure of the Brain in Marsupial Animals, PkiJos. Tram. 1837. 


concerned in the nourishment of the progeny before birth, 
called t 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, LYENCEPHALA I , 
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. 

Fig- 5- 

Fig. 4 . 

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 

tfw, to loose ; 

, brain. 


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 LISSEN- 

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 GYRENCEPHALA, which 
I propose for the third subclass of Mammalia 2 . 

Fig. 7. 

Fig. 6. 

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 \i<7<rds, smooth ; tyiftyaXos, brain. 

2 "yvpbii), to wind about ; eyK^(f>a\os, brain. 


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. Fig . 8 

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- 
est form 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 
malia, for which I 
propose the name 
(tig. 9) . Ib. Side view, one-third nat. size. 

Brain of Negro, upper view. 

Fig. 9. 

, to overrule ; fy*l0aXos, brain. 


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 LYENCEPHALA 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 
1 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 Echidna 
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 LYENCEPHELA 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 


duplicature of the abdominal integument, which in the males 
is everted, forming a pendulous bag, arid 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 1 (fig. 2) : the angle of the 
lower jaw (ib. a) is more or less inverted 2 . 

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 whicli 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 

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. n. 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. n. p. 379 (1838). 


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 oesophagus. These correlated modifica- 
tions of maternal and foetal structures, designed with especial 

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

The LISSENCEPHALA, or smooth-brained placental Mam- 
malia, form a group which I consider as equivalent to 
the LYENCEPHALA or Implacentals ; and which includes the 
following orders, Rodentfa, Insectivora, Cheiroptera and 

The RODENTIA 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 fossa?. 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 


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 
mammas 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 Edentata 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 Yertebrata by the 
supernumerary cervical vertebras 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 (Cholcepus), recalls a 
lacertine structure. The same tendency to an inferior type 


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 1 . 

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, I may remark that the LYENCEPHALA cannot 
be regarded as equivalent merely to one of the orders, say 
Rodentia, of the LISSENCEPHALA, without undervaluing the 
anatomical characters which are so remarkable and distinct 
in the marsupial and monotrematous animals. The anato- 
mical peculiarities of the edentulous LYENCEPHALA 2 appear to 
me to be, at least, of ordinal importance. In these deduc- 
tions I hold the mean between those who, with Geoffrey 
St Hilaire, would make a distinct class of the Monotremata, 
and those who, with Cuvier, would make the Monotremes 
a mere family of the Edentata. In like manner, whilst I 
regard the LYENCEPHALA 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 LYENCEPHALA 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 mo turn 
suum validissime retinebat, postquam exemptum erat e corpore per semiho- 
rium : exempto corde cseteris visceribus multo post se movebat et pedes lente 
contrahebat sicut dormituriens solet.' Buffon, who quotes the above from the 
Historia Naturalis Brasilia, 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. xm. p. 45. 

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


primary divisions or subclasses of the Mammalia; its true 
equivalency is with the LISSENCEPHALA, 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 : 

Rhizophaga ' Burrowing Rodentia. 

Poephaga l Dipodidce and Leporidce. 

Petaurus Pteromys. 

Phalangistidce Sciuridce and prehensile-tailed 

arboreal Rodents. 

Phascolarctos Bradypus. 

Perameles and Myrmecobius. Erinaceidce. 

Chceropus Macroscelis. 

Diddphys and PJiascogale . Soricidce. 

Dasyuridce Centetes, Gymnura. 

Echidna Manis. 

Besides the more general characters by which the LISSEN- 
CEPHALA, in common with the LYENCEPHALA, 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 vertebras 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 ( Clirysochlora aurea) ; 
the proventriculus of the Dormouse and Beaver; the pre- 

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

P- 3i5> l8 39- 



valence of disproportionate development of the hind limbs in 
the Rodentia, coupled, in the Jerboa, with confluence of the 
three chief metatarsals into oiie bone, as in birds ; the keeled 
sternum and wings of the Bats ; the aptitude of the Cheirop- 
tera, Insectivora, and certain Rodentia to fall, like Keptiles, 
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 LYEN- 
CEPHALA to the Oviparous air-breathing Yertebrata, have 
mainly prevailed with me against an acquiescence in the 
elevation of different groups of the LISSENCEPHALA 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 1 , and of the shorter-clawed Shrews, Moles and' 
Hedgehogs, as well as the Bats, with the Carnivora 2 ; of the 
Sloths with the Quadrumana 3 ; of the Bats with the same 
high order 4 ; and of the Insectivora and Rodentia in immediate 
sequence after the Linnean 'Primates,' as in the latest pub- 
lished < System of Mammalogy,' from a distinguished French 
author 5 . 

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 GYRENCEPHALA WQ 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, i2mo, 1843, p. xii. 

2 Cuvier, R^gne Animal, 1829, p. 'no. 

3 De Blainville, Osteographie, 4to, Fasc. i. p. 47 (1839). 

4 Linnaeus, Systema Natures, Ed. 12, Tom. I. p. 26. 

5 Prof. Gervais, Zoologie et PaUontologie Franpais, 4to, 1852, p. 194. 


y 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 GYRENCEPHALA 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 
LYENCEPHALA and LISSENCEPHALA with the unguiculate GYREN- 
CEPHALA into one great primary group, as in the Mammalian 
systems of Ray, Linnasus 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 GYRENCEPHALA are of 
higher value than the ordinal divisions of the LISSENCEPHALA ; 
just as those orders are of higher value than the representative 
families of the LYENCEPHALA. 

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 c testiconda,' and 



have no 'vesiculse seminales.' The mammae 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 (Balcena mysticetus) is represented by a 
distinct species (Balo&na 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 SIEENIA, 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 mamma3 are pectoral ; 
they are ' testiconda,' but have 'vesiculae seminales.' The 
Sirenia exist near coasts or ascend large rivers ; browsing on 
fuci, water plants, or the 1 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 Eed 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, 


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

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 (Elephas) 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 PEOBOSCIDIA given to the order. The feet 
are pentadactyle, but the toes are indicated only by divisions of 
the hoof; the placenta is annular ; the mammse 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. Humboldtii) 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 texianus) 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. 


abundant remains of this Elephas primigenius (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 

Both the proboscidian and toxodontal orders of UNGULATA 
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 Sirenia in its re- 
markable modifications. 

The typical Ungulate quadrupeds are divided, according 
to the odd or even number of the toes, into PERTSSODACTYLA 
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 UNGULATA, the dorso- 
lumbar vertebrae 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 ectocunei- 
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 7repi<T(roddKTV\os, qui digitos habet impares numero ; and dprios, 
par, SciKTuXoj, digitus. Quarterly Journal of the Geological Society, No. 14, 
May, 1848. 


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 Tapir, 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 csecum, 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 UNGVLATA. 
Many extinct genera, e. g. Coryphodon, Pliolophus,Lophiodon, 
Tapir other ium, Palceotherium, Ancitherium, Hipparion, Acero- 
therium, Elasmotherium, &c., have been discovered, which 
once linked together the now broken series of Perissodactyles, 
represented by the existing genera Rhinoceros, 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 Hyrax, 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 vertebras 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 vertebrae, which those ribs charac- 
terize, being fifteen in the Hippopotamus and twelve in the 


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 vertebras, and their non-recognition as 
parts of a vertebra equivalent to the neurapophyses and other 
autogenous elements. 

The vertebral formulae of the Artiodactyle skeletons shew 
that the difference in the number of the so-called dorsal and 
lumbar vertebras 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, I. 
5 = 19; the Hippopotamus has d. 15, I. 4 = 19 ; the Gnu and 
Aurochs have d. 14, I. 5 = 19 ; the Ox and most of the true 
Ruminants have d. 13, I. 6 = 19; the aberrant Ruminants have 
d. 12, I. 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 


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 Kuminants ; the comparatively small and simple caecum 
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 UNGULATA. The placenta is diffused in the Camel-tribe 
and non-ruminants ; is cotyledonal in the true Ruminants. 
Many extinct genera, e. g. Chceropotamus, Anthracotheriumj 
Hyopotamus, JEntelodon, Dichodon, Merycopotamus, Xiphodon, 
Dichobune, Anoplotkerium, 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, Mosckus, Camelo- 
pardalis, Cervus, Antilope, Owis, 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,' i. 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 
grinding surface marked by two double crescents, the con- 


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 1 adduced evidence to shew that the right progression 
of the affinities of the UNGULATA 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 

It is interesting, in relation to the needs of mankind, to find 
that, whilst some groups of UNGULATA, 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- J 35 


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 or a 
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 
Camelidce 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. 

Palaeontology 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 Giraffe 
or Hippopotamus were ever other than Old- World forms of 

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


tyles] in tlie 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 in either 
division of the New World, nor has any Dicotyles been found 
fossil in the Old World of the geographer. Phacochoerus 
(Wart-hogs) is a genus of the Hog-tribe at present peculiar 
to Africa. 

The Khinoceros 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 1 , 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 Equidce, and Elephantidce, 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 Equus, like species of Mastodon, coexisted with 
the Megatherium and Megalonyx in both South and North 
America, and perished with them, apparently before the 
human period. 

The third division of the GYRENCEPHALA enjoy a higher 
degree of the sense of touch than the Ungulates through the 
greater 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. 


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 UNGUICULATA, 
applied to this group, which, however, is here more restricted 
and natural than the group to which Linnaeus extended the 
term. All the species are ' diphyodont,' and the teeth have 
a simple investment of enamel. 

The first order, CAKNIVORA, includes the 'beasts of prey, 
properly so called. With the exception of a few Seals the 

33 . , ,, 11 . 

incisors are - - in number ; the canines - - , always longer 

o o 1 1 

than the other teeth, and usually exhibiting a full and perfect 
development as lethal weapons; the molars graduate from a 
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 
vesiculse seminales ; the teats are abdominal ; the placenta is 

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, 


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) , Nasua (Coati-mondis) and Cercolep- 
tes (Kinkajous) are peculiarly American : other plantigrade 
genera, e. g. Mydaus, Ailurus, and Arctictis, are peculiarly 

The genus Hysena 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 ( Tigris) 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 Hycenodons 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 CTYRENCEPHALA the paw is 
perfected as an instrument for retaining and lacerating a strug- 
gling prey by the superadded elastic structures for retracting 
the claws and maintaining them sharp. We next find in 


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- 
2 2 o o 

culate molars ; perfect clavicles ; pectoral mammae ; 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 tenni- 

Q Q 

nal nostrils, with much modified incisors, commonly - - ; 

o o 

o q n n 

premolars - - or in number, and molars with sharp 

o o 2 2 

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 

The Platyrhines are those with the nostrils subterminal 

q p 

and wide apart ; premolars - - in number, the molars with 

O' O 

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- 

2 2 

molars are - 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. 


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 Semiiopitheques, Gibbons (Hylobates) , 
and Orangs (Pithecus} are peculiarly Asiatic. Palaeontology 
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 (Inuus 
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 (Mycetes}, 
but larger than any now known to exist, has left its remains 
in Brazil. 

Whilst adverting to the geographical distribution of 
Quadrumana, I 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 ARCHENCEPHALA, 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 BIMANA, on account of the 
opposable thumb being restricted to the upper pair of limbs. 
The mammse 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 



the first two premolars of the typical series on each side of 

"both jaws, the dental formula "being : 

. 2 2 11 2 2 3 3 _ 01 

'2=2' C T^T' ^2^2' m -3^ = 32 ' 

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 scapulae 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 terminated 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. 


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 Temple 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. 





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 Extinct Mammals of Australia, published in the 
Volume of Heports 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 palseotheres 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 aifecting a due 
supply of sustenance to animals in a state of nature which must 
have accompanied the slow alternations of land and sea brought 


about in the seons 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 


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 j 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 iSfalforiaj 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 Elephas 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 dinomis, 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 Mend 1 , 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 Keykianes, the S. 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 Alca impennis must form 
an important element in its decline towards extinction. The 
numbers of the bones of Alca impennis on the shores of Iceland, 
Greenland, and Denmark, attest the abundance of the bird in 

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


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 palaeozoic 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 seons, 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. 


In all cetacea the skeleton is well ossified, and the vertebrae 
are very numerous : the smallest cetaceans would be deemed large 
amongst land-mammals j 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 modem 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 Cetacea in newer 
secondary beds 1 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 Ichthyosaurus, Plio- 
saurus, or Plesiosawrus, been represented by species in the same 
ocean that was tempested by the Balsenodons 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. 

In 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. 


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 JSnaliosauria, 
Cetiosauria, and Crocodilia, played the same part and fulfilled 
similar offices in the seas from which the lias and oolites were 
precipitated, as the Delphinidce and Balcenidce 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 Enaliosauria. 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 'Echini are most common in the superior 
strata, Ammonites in those beneath, and Producti with numerous 
Encrini in the lowest' 1 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. 


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, Product^ 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 Palaeontology 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 of a higher and blessed race of intelligences. 

If the guide-post of Palaeontology 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. 


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 I have 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. 


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 palseontological 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 . 

1 Sedgwick, Address to the Geological Society, 1831. 




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 1 . He called it the Homo Sylvestris, or pigmy. It 
is noted by Linnseus, in some editions of his Systema Naturce, 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 Simla Troglodytes; 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 Tro- 
glodytes of Africa, by being covered with hair of a reddish-brown 
colour, and by having the anterior, or upper limbs, much longer 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 'Rdgne 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/ 4 to, 1699. 


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 


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 
vertebrae 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 vertebrae 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 vertebrae, 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 vertebrae 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 (humeru's) 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 


subject, and a retention of the character met with in the lower 
quadrnmana ; 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 
fronvtheir 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 Simla satyrus, and 



55 to 60 in the Troglodytes 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. First, 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 cristce 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 j one of a smaller size 
and somewhat more anthropoid ; and the larger species presenting 
the baboon-like cranium. 

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 1 847 I received 
a letter from Dr Savage, a church-missionary ab 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 Troglodytes 
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 


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 (Troglodytes), 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 e*nter 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 
siamangs, gibbons or long-armed apes (Hylobates). 

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 scapulae, 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. 


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 denned 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 1 . 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 ilii.' 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. Yery 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. Yol. n. (1855) p. 26, and in 
the Transactions of the Zoological Society, 1848. 


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 5^ 
inches; in man, at the same part, it averages 2J 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 progression 
on the lower limbs than any other ape. 


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- 


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 'alse:' 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 (Trogl. niger): this is due to an admixture of a few' 


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 Troglodytes 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 


long-armed gibbons, more nearly related to the genus Homo ? O 
the broad-breast-boned quadramana, are the knuckle-walkers or 
the brachiators, i.e. the long-armed gibbons, most nearly and essen- 
tially related to the human subject 1 ? 

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 palaeontologist, M. Lartet, and accepted 
by a high geological authority at home l . 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 
Cyclopaedia, cited by Lartet 2 , Professor Yrolik briefly treats of the 
osteology of the Quadrumana according to their natural families. 
In 'a first genus, Simla 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, Yrolik says, 
* its skeleton approaches most to that of man,' which may be true 

1 Sir C. Lyell, Supplement to tlie 5th Edition of a Manual of Elementary 
Geology, 1859, p. 15. 

2 Comptes Rendus de V Academic des Sciences, Juillet 28, 1856. 


in comparison with other gibbons, but certainly is not so as respects 
the higher Slmlce. 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 Hylobates lar, was probably 
the obvious character in Yrolik'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 Yrolik 1 , 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 Simla satyrus and Simla troglodytes, as 
deduced from the facial angle and dentition, are proportionally 
exaggerated in the Regne Animal 2 . 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, Cyclopaedia of Anatomy, Vol. iv. p. 195. 

2 Ed. 1829, pp. 87, 89. 


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 

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 convolutions? in which is the cerebrum largest, as com- 
pared with the cerebellum 1 ? 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 caecum and the 
ischial integument : if he found in one subject of his comparisons 
(Troglodytes) a long 'appendix vermiformis cseci,' as in man, but 
no 'callosities,' and in another subject (Hylobates) the ischial 
callosities but only a .short rudiment of the csecal 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 csecum and nates were exhibited by the smaller- 
brained and longer-armed tailless gibbons. 

Pursuing the comparison through the complexities of the bony 


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 aiitibrachial 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 ilii,' 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 


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

The scapulae 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 in 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 (Bimana). 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 arid 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 


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 (Troglodytes) 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 vertebrae in man, and only so called, when they become 
long and free. The genera Homo, Troglodytes, and Pithecus, have 
precisely the same number of vertebrae : if Troglodytes, by the 
development and mobility of the pleurapophyses of the 20th ver- 
tebrae 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 vertebrae, 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 Simla, of 
Linnaeus. 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 


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 1 bulk by the 
gorilla, and of a more 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 (Troglo- 
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 



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 

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 Simice. 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 semnopitheques 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- 


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 
/Simice, 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 pvale, 
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. 



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 simise 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 simise 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 Journal 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 (Eopithecus) having its nearest affinities 
with 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 

/ 3 3 3 3\ 

molar dentition ( p - - , m - ^ ) to belong to the platyrrhine 
\ o o o o/ 

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 
Blainville, is so closely allied to the gibbons as to scarcely justify 
the generic separation which has been made for it under the name 

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


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 1 , 
seems to be 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 VAcademie des Sciences, Paris, Vol. XLIII. 


called DryopitJiecus, only by the figures published in the 43rd 
volume of the Comptes Eendus de TAcademie 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 
Dr+yopithecus 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 (Gorilla), 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, it came 
nearer to man than any quadrumanous species, living or fossil, 
before known to zoologists V 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. 


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 
fossil, 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 (Elais 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' (Parinarium excelsum), which bears a 
plum-like fruit. 


The papau tree (Carica papaya). 

The banana (Musa sapientium), and another species (Musa 

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 of a 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 
1 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 

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 


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 lire 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 


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 

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, 


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, l 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 
1 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 


were not able to take the men j 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 1 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 Bosnian, 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 Regne 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 

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 


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 


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 so long 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 joins a ' 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. 


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 oiitwards, 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 vertebrae, anchylosed to form the bone called *os 
coccygis.' The true vertebrae, 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 vertebrae 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 in a slight sigmoid flexure ; the 
humerus exceeds in length the bones of the fore-arm. The carpal 


bones are eight in number. The thumb is developed far beyond 
any degree exhibited by the highest quadrumaria, 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 cranii, 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 cristre. 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 vertebrae, 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 Patagoiiian 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 


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. We are 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, I 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 



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 cranii, 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 as 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 


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 cristse 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 cristse 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 


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 familiar is: 

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 Troglodytes 
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 


characteristic of the cranium of the genus Troglodytes 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 Troglodytes 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 t)f 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 Troglodytes gorilla began to be calcified 
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 


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 Tr. gorilla beyond the period at which they disappear 
in the Tr. 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 

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 Troglodytes 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 Troglodytes niger as com- 
pared with the Tr. 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 Troglodytes 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 Troglo- 
dytes gorilla parallels the Pithecus Wurmbii, so the Troglodytes 


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 

It is not without interest to observe, that as the generic forms 
of the Quadrumana approach the Bimanous order, they are repre- 
sented by fewer species. The gibbons (Hylobates) scarcely number 
more than half-a-dozen species ; the orangs (Pithecus) have but two 
species, or at most three; the chimpanzees (Troglodytes) 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 a real 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 mo*e nobly de- 
scended, better bred and born, and more skilfully taught by the 
purged faculties of his own minde 1 ." 

1 Henry More's Conjectura Cabbalistica, fol. (1662) p. 175. 





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