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Bonbon : 


[The Righl of Translation and Reproduction is Reserved.} 



Ami Bnngay, Suffolk. 


THE desire to acquire a knowledge of the structure of some 
portion at least of the Animal Kingdom, now becoming so 
general, is often checked by the difficulty of determining 
where to make a beginning amid the vast extent and variety 
of the materials at hard. 

I have selected for my first course of lectures on Com- 
parative Anatomy at the Royal College of Surgeons, the 
structure and modifications of the Skeleton, because, as 
the framework around which jthe rest of the body is built 
up, it gives, more than any other system, an outline of the 
general organization of the whole animal, and also because 
it is the most convenient for study, on account of the 
facility with which it can be preserved and examined. 

Moreover, Osteology has special importance in com 
parison with the study of any other system, inasmuch as 
large numbers of animals, all in fact of those not at presen 
existing on the earth, can be known to us by little else than 
the form of their bones. 

In endeavouring to gain anatomical knowledge, it sig- 
nifies little with which group of animals a commencement 
is made. 



The structure of Man has undoubtedly a more universal 
interest than that of any other organized being, and 
has, therefore, been more thoroughly worked out ; and as 
the majority of terms used in describing the parts com- 
posing the bodies of Vertebrate animals were originally 
bestowed on account of their form, relation, or real or 
fancied resemblance to some object, as they were met 
with in Man, there are advantages in commencing with 
members of the highest class, and mastering their essential 
characters before proceeding to acquire knowledge of the 
other groups. 

But as human anatomy may be taken as a point of de- 
parture from which to set out in the study of that of other 
Vertebrates, so, on the other hand, those whose special 
duty it is to become familiar with its details, will find 
themselves greatly assisted by some knowledge of the 
structure of lower forms. Thus the essential characters 
of the human skull will be much better understood if the 
student will also make himself acquainted with those of 
some simpler condition of Mammalian cranium, as that of 
the dog or sheep. 

Although the present work contains the substance of a 
course of lectures, the form has been changed, so as the 
better to adapt it as a handbook for students. Theoretical 
views have been almost entirely excluded ; and while it is 
impossible in a scientific treatise to avoid the employment 
of technical terms, it has been my endeavour to use no 
more than are absolutely necessary, and to exercise due 
care in selecting only those that seem most appropriate, or 
which have received the sanction of general adoption. 


With very few exceptions, all the illustrations have been 
drawn expressly for this work, with great care and fidelity, 
by Mr. R. W. Sherwin, from specimens in the Museum of 
the Royal College of Surgeons. 

September 2^1 ft, 1870. 


This edition has been revised throughout. Several new 
illustrations, a table shewing the number of the vertebra? 
of a large number of species, and many references to the 
more important recent memoirs on the subjects referred 
to, have been introduced. 

Odobtr ist, 1885. 



















THE RIBS 1 06 




























INDEX 375 






IT is not the object of the present work to enter in 
detail into the subject of the Classification of the Mam- 
malia ; but, as it will be necessary to refer frequently to the 
principal subdivisions in which the various animals treated 
of are arranged, a brief outline of the system adopted will be 

A perfect arrangement of any group of animals can only 
be attained simultaneously with a perfect knowledge of their 
structure and life-history. We are still so far from this 
that any classification now advanced must be regarded as 
provisional, and merely representing our present state of 
knowledge. Moreover, as naturalists will estimate differ- 
ently the importance to be attached to different structural 
modifications as indicative of affinity, it must be long before 
there will be any general agreement upon this subject 


The classification and the names of the subdivisions used 
throughout this work correspond, in the main, with those 
given by Professor Huxley in his " Introduction to the 
Classification of Animals," 1869. 

The modifications of this classification which appear to be 
necessitated by more recent acquisitions of knowledge will 
be found fully detailed in the author's article, Mammalia, 
in the ninth edition of the Encyclopaedia Britannica, vol. xv. 
(1883), from which the following introductory observations 
are extracted : 

One of the most certain and fundamental points in the 
classification of the Mammalia is that all the animals now 
composing the class can be grouped primarily in three- 
natural divisions, which, presenting very marked differen- 
tiating characters and having no existing or yet certainly 
demonstrated extinct intermediate or transitional forms, may 
be considered as sub-classes of equal value, taxonomically 
speaking, though very different in the numbers and import- 
ance of the animals at present composing them. 

These three groups are often called by the names 
originally proposed for them by De Blainville : (i) Or- 
nithodelphia, (2) Didclphia, (3) Monodelphia^ the first being 
equivalent to the order Monotremata, the second to the 
Marsupialia, and the third including all the remaining 
members of the class. Although actual palseontological 
proof is wanting, there is much reason to believe that each 
of these, as now existing, are survivors of distinct branches, 
to which the earliest forms of Mammals have successively 
given rise, and for which hypothetical branches or stages 
Huxley has proposed the names of Prototheria, Metatheria, 
and Eutheria}- 

I. The characters of the Ornithodelphia or Prototheria 
1 Proceedings of the Zoological Society, 1880, p. 649. 


can only be deduced from the two existing families, as 
hitherto no extinct animals which can be referred to other 
divisions of this remarkable and well-characterised group 
have been discovered. These two isolated forms, in many 
respects dissimilar, yet having numerous common characters, 
which unite them together and distinguish them from the 
rest of the Mammalia, are the Ornithorhynchida and the 
Echidnidie, both restricted in their geographical range to the 
Australian region of the globe. Taken altogether they 
represent the lowest known type of evolution of the Mam- 
malian class, and most of the characters in which they differ 
from the other two sub-classes, tend to connect them with 
the inferior vertebrates, the Sauropsida and Amphibia , 
for though the name Ornithodelphia owes its origin to the 
resemblance of the structure of the female reproductive 
organs to those of birds, there is nothing especially bird-like 
about them. 

Their principal distinctive characters are these : The 
brain has a very large anterior commissure, and a very small 
corpus callosum, agreeing exactly in this respect with the 
next group. The cerebral hemispheres, in Echidna at least, 
are well developed and convoluted on the surface. The 
auditory ossicles present a low grade of development, the 
malleus being very large, the incus small, and the stapes 
columelliform. They have no true teeth, though the jaws 
of Ornithorhynchns are provided with horny productions, 
which functionally supply their place. The coracoid bone 
is complete and articulates with the sternum, and there is a 
large " interclavicle " or episternum in front of the sternum 
and connecting it with the clavicles. There are also 
"epipubic" bones. The oviducts (not differentiated into 
uterine and Fallopian portions) are completely distinct, and 
open as in other oviparous vertebrates separately into a cloacal 


chamber, and there is no distinct vagina. The testes of the 
male are abdominal in position throughout life, and the vasa 
deferentia open into the cloaca, not into a distinct urethral 
passage. The penis, attached to the ventral wall of the 
cloaca, is perforated by a canal in the greater part of its 
length, but not at the base, which is open, as in reptiles and 
those birds which have such an organ, and brought only 
temporarily in contact with the termination of the vasa 
deferentia, so as to form a seminal urethra when required, 
but never transmits the urinary secretion. This condition 
is a distinct advance on that of the Sauropsida in the 
direction of the more complete development of these 
parts in most of the other Mammalia. The ureters do not 
open into the bladder, but behind it in the dorsal wall of 
the genito-urinary passage. The mammary glands have 
no distinct nipple, but pour out their secretion through 
numerous apertures in the skin ; the whole mammary area 
is restricted to a slightly cup-shaped invagination of the 
abdominal skin, forming a mammary marsupium, especially 
developed in the females during lactation, but hardly recog- 
nisable in the males. The Monotremata lay eggs and hatch 
them in the abdominal pouch. The primary egg undergoes 
meroblastic segmentation as in the Sauropsida ; the early 
stages of the development of the young are not yet fully 
known, but they appear never to be nourished by means of 
an allantoic placenta. 

II. The Didelphia, Marsupialia, or Metatheria are repre- 
sented at present by numerous species, presenting great 
diversities of general appearance, structure, and habits, al- 
though all united by many essential anatomical and physio- 
logical characters, which, taken altogether, give them an inter- 
mediate position between the Prototheria and Eutheria. In 
the structure of the brain and the presence of epipubic bones 


they agree with the former, while in the structure of the 
earbones and the shoulder-girdle, and the presence of 
true teeth, and of teats on the mammary glands, they 
resemble the latter; the reproductive organs belonging to 
neither one nor the other type, but presenting a special 
character representing an intermediate grade of development. 
The ureters open into the base of the bladder. The 
oviducts are differentiated into uterine and Fallopian por- 
tions, and open into a long and distinct vagina, quite- 
separate from the cystic urethra. The penis is large, but 
its crura are not directly attached to the ischia. The 
spongy body has a large bifurcated bulb. The young are 
born in an exceedingly rudimentary condition, and are 
never nourished by means of an allantoic placenta, but are 
transferred to the nipple of the mother, to which they remain 
firmly attached for a considerable time, nourished by the milk 
injected into the mouth by compression of the muscle cover- 
ing the mammary gland. They are, therefore, the most 
typically mammalian of the whole class. The nipples are 
nearly always concealed in a fold of the abdominal integu- 
ment or "pouch" (marsupium), which serves to support 
and protect the young in their early helpless condition. 
The existing species of this group are entirely confined to 
the Australian region and the American continent, though 
in former times they had a more extended geographical 
range. The earliest Mammals hitherto discovered appear 
(as far as the scanty evidence at present obtainable permits 
any such conclusion to be hazarded) to have belonged to 
this type, although it is reasonable to conclude that 
Prototheria (unless upon the improbable supposition that 
the existing forms have resulted from a process of degrada- 
tion), and perhaps Eutheria, were their contemporaries far 
back in the Mesozoic age. 


III. The Monodelphia, Eutheria, or "placental mammals " 
(so-called because the foetus is always nourished while 
within the uterus of the mother by means of an allantoic 
placenta), include at present by far the greater portion of 
the class. While the survivors of the other groups have 
probably been for a long time in a stationary condition, 
these have, as there is already good evidence to show 
throughout all the Tertiary geological age, and by inference 
for some time before, been multiplying in numbers and 
variations of form, and attaining higher stages of develop- 
ment and specialisation in various directions. They con- 
sequently exhibit far greater diversity of external or adaptive 
modification than is met in either of the other sub-classes 
some being fitted to live exclusively in the water as 
fishes, and others to emulate the aerial flight of birds. 

To facilitate the study of the different component members 
of this large group it is usual to separate them into certain 
divisions which are called "orders." In the main zoologists 
are now of accord as to the general number and limits of 
these divisions among the existing forms, but the affinities 
and relationships of the orders to one another are far from 
being understood, and there are very many extinct forms 
already discovered which do not fit at all satisfactorily into 
any of the orders as commonly defined. 

Commencing with the most easily distinguished, we may 
first separate a group called Edentata, composed of several 
very distinct forms the Sloths, Anteaters, and Armadillos 
which under great modifications of characters, of limbs, and 
digestive organs, as well as habits of life, have just enough 
in common to make it probable that they are the very 
specialised survivors of an ancient group, most of the 
members of which are extinct, but which the researches of 
palaeontology have not yet revealed to us. The characters 


of their cerebral, dental, and in many cases of their repro- 
ductive organs show an inferior grade of organisation to thai 
of the generality of the sub-class. The next order, about 
the limits of which there is no difficulty, is the Sirenia 
aquatic vegetable-eating animals, with complete absence of 
hind limbs, and low cerebral organisation represented in our 
present state of knowledge by but two existing genera, the 
Dugongs and Manatees, and by a few extinct forms which, 
though approaching a more generalised Mammalian type, 
show no special characters allying them to any of the other 

Another equally well marked and equally isolated, though 
far more numerously represented and diversified, order is 
that of the Cetacca, composed of the various forms of 
Whales, Dolphins, and Porpoises. In aquatic habits, 
external fish-like form and absence of hind limbs, they 
resemble the last, though in all other characters they are 
as widely removed from them as from any other order 
among the Monodelphia. The association by systematists 
of the Cetacea and Sirenia in our group can only be made 
either in ignorance of their true structure, or in an avowedly 
artificial system. 

All the remaining orders are more nearly allied together, 
the steps by which they have become modified from one 
general type being, in most cases, not difficult to realise. 
Their dentition especially, however diversified in detail* 
always responds to some modification of the well-known 
formula incisors %, canines |, premolars *, molars . Al- 
though the existing forms are broken up into groups, in 
most cases easy of definition, the discoveries already made 
in palaeontology have in great measure filled up the gaps 
between them. 

Very isolated among existing Monodelphia are the two 


species of elephants, constituting the group called Proboscidea, 
These, however, are now known to be the survivors of a large 
series of similar animals Mammoths, Mastodons, and Dino- 
theria which as we pass backwards in time gradually assume 
a more ordinary or generalised type ; and the interval which 
was lately supposed to exist between even these and the 
rest of the class is partially bridged over by the discovery 
in American Eocene and early Miocene formations of the 
gigantic Dinocerata, evidently offshoots of the great group 
of hoofed animals or Ungtilata, represented in the actual 
fauna by the Horses, Rhinoceroses, Tapirs, Swine, and 

Almost as isolated as the Proboscidea among existing 
Mammals are the few small species constituting the genus 
Hyrax, and in their case palaeontology affords no help at 
present, and therefore, pending further discoveries, it has 
been thought advisable in most recent systems to give them 
the honour of an order to themselves under the name of 
Hyracoidea. But the number of extinct forms already 
known allied to the Ungulata, but not coming under the 
definition of either of the two groups (Artiodactyla and 
Perissodactyla) under which all existing species range them- 
selves, is so great that either many new orders must be 
made for their reception or the definition of the old order 
Ungulata so far extended as to receive them all, in which 
case both Proboscidea and Hyracoidea might be included 
within it. 

Again, the Rodentia or gnawing animals Rabbits, Rats, 
Squirrels, Porcupines, Beavers, &c. are, if we look only at 
the present state of the class, most isolated. No one can 
doubt what is meant by a rodent animal, or have any 
difficulty about defining it clearly, at least by its dental 
characters ; yet our definitions break down before the 


extinct South American Mesotherium, half Rodent and half 
Ungulate, which leads by an easy transition to the still 
more truly Ungulate Toxodon, for the reception of which a 
distinct order ( Toxodontia) has been proposed. 

The Insectivora and the Carnivora again are at present 
quite distinct orders, but they merge into one another 
through fossil forms, and are especially connected by the 
large group of primitive Carnirora, so abundantly repre- 
sented in the Eocene deposits both of America and Europe, 
to which Cope has given the name of Crcodonta. The 
transition from the Insectivora to the Lemurs is not great, 
and, strange to say, however different they now appear, the 
early forms of Lemurs are not easily distinguished from the 
primitive Ungulata. The Hats or Chiroptera are allied to 
the Insectivora in all characters but the extraordinary 
modification of their anterior extremities into wings ; but 
this, like the want of the hind limbs in the Cetacea and 
Sirenia, makes such a clear distinction between them and 
all other Mammals, that, in the absence of any knowledge 
of intermediate or transitional forms, they can be perfectly 
separated, constituting as well-defined an order as any in 
the class. 

Lastly, we have the important and well-characterised 
group called Primates, including all the Monkeys and Man, 
and the question is not yet solved as to how, and through 
what forms, it is linked on to the other groups. It is com- 
monly assumed that the Lemurs are nothing more than 
inferior Primates, but the interval between them in the 
actual fauna of the world is very great, and our knowledge 
of numerous extinct species recently discovered in America, 
said to be intermediate in character, is not yet sufficiently 
perfect to enable us to form a definite opinion upon the 


The accompanying diagram (page n) is intended to 
exhibit the relationships which appear to exist between the 
different groups of recent Mammalia. If all known extinct 
forms were inserted, many of the intervals between the 
boundary lines of the groups would be filled up ; other- 
wise no great modification would be required in their relative 
position. But our knowledge of the systematic position 
and relations of the past forms of Mammalian life is in 
general so imperfect and fragmentary, that it seemed best 
to confine the diagram to a representation of the present 
condition of Mammalian life upon the earth. 




THE term Skeleton, in its widest sense, is used to denote 
a system of hard parts forming a framework which supports 
or protects the softer organs and tissues of the body, and 
which may be either entirely external or superficial as 
regards those organs and tissues, or may be more or less 
embedded in enveloping softer structures. In the former 
case it is called an Exoskdeton, in the latter an Endoskeleton. 

It is of the Endoskeleton alone that this work proposes 
to treat, as in the class Mammalia the external skeleton, 
when it exists, performs a relatively subordinate part in the 
economy. 1 

The branch of anatomy called Osteology is commonly 
restricted to a study of such parts of the endoskeleton as 
are composed of bony or osseous tissue, a tissue charac- 
terised by a peculiar histological structure and chemical 
composition, being formed mainly of a gelatinous basis, 
strongly impregnated with calcium phosphates and car- 
bonates, and disposed in a definite manner, containing 
numerous minute nucleated spaces or cavities called 
lacuna, connected together by delicate channels called 
canaliculi, which radiate in all directions from the sides 

1 The Armadillos and their extinct allies are the only known mam- 
mals which have an ossified exoskeleton. 


of the lacunae. This structure is readily recognised when a 
thin section of bone is examined under a moderately high 
magnifying power. 

Parts composed of bone are, of all the tissues of the body 
(with the exception of the teeth), the most imperishable, 
often retaining their exact form and intimate structure ages 
after every trace of all other portions of the organisation has 
completely disappeared ; and thus in the case of extinct 
animals affording the only means of attaining a knowledge 
of their characters and affinities. 

It must, however, be remembered that, at one period of 
life, the parts composing the skeleton exist in a fibrous or 
a cartilaginous form, that their transformation into bone is 
a subsequent and gradual process, and that even in the 
Mammalia, though in a less degree than in some of the other 
Vertebrata, the whole of the internal skeletal system is 
never entirely osseous, but that portions may remain per- 
manently in a cartilaginous or fibrous condition. 

The different bones composing the skeleton are con 
nected together either by sutures, or by movable joints 
or articulations. 

In the first, the edges of the bones are in close contact, 
often interlocking by means of projections of one bone 
fitting into corresponding depressions of the other, and are 
held together by the periosteum, or fibrous membrane in- 
vesting the bones, passing directly from one to the other, 
permitting no motion, beyond, perhaps, a slight yielding to 
external pressure. The bones of the cranium are connected 
together in this manner. In old animals there is a great 
tendency for such bones to become joined together by 
the extension of ossification from one to the other and 
consequent obliteration of the suture. This process is 
called synostosis. 


The various forms of joints may be arranged under two 
principal heads. In one, the contiguous surfaces of the 
bones are connected by interposed fibrous tissue, passing 
directly from one to the other, filling up the space between 
them, and allowing of only a limited amount of motion, as 
is the case with the bodies of the vertebrae. 

The other and more frequent and more perfect form of 
joint is that in which the contiguous extremities of the bones 
are covered by a thin layer of very smooth cartilage, and 
surrounded by a capsular ligament, attached only round the 
edges of the articular surfaces, and which is lined by a 
synorial membrane, so called from its secreting a viscid 
lubricating fluid termed synovia. The amount of motion 
permitted in these " synovial joints " varies according to the 
form of the opposed articular surfaces and the arrangement 
of the ligaments which hold them together. When the two 
surfaces are nearly flat, and the bones firmly bound by strong 
short ligaments, as in those which compose the carpus and 
tarsus, the motion is reduced to an extremely slight gliding 
of one on the other. Joints in the form of a hinge, as at 
the elbow, allow of a free motion in one plane only. Ball 
and socket joints, as at the shoulder and hip, allow of the 
greatest variety of movements. 

The Endoskeleton is divided into an axial portion, be- 
longing to the head and trunk, and an appcndicular portion, 
belonging to the limbs. There are also certain bones called 
splanchnic, being developed within the substance of some of 
the viscera. Such are the os cordis and os penis found in 
some Mammals. These, however, are more appropriately 
treated of with the anatomy of the organs of which they 
form a part. 

The Axial Skeleton consists of the vertebral column, the 
skull, the sternum, and the ribs. 



General Characters. The Vertebral Column consists of a 
series of distinct bones called Vertebra, arranged in close- 
connection with each other along the dorsal side of the neck 
and trunk, and in the median line. It is generally prolonged 
posteriorly beyond the trunk to form the axial support of the 
appendage called the tail. Anteriorly it is articulated with 
the occipital region of the skull. 1 

The number of distinct bones of which the vertebral 
column is composed varies greatly among the Mammalia, 
the main variation being due to the elongation or otherwise 
of the tail. Apart from this, in most Mammals, the number 
is not far from thirty, though it may fall as low as twenty- 
six (as in some Bats) or rise as high as forty (ffyrax and 

The different vertebrae, with some exceptions, remain 
through life quite distinct from each other, though closely 
connected by means of fibrous structures which allow of a 
certain, but limited, amount of motion between them. 

1 For the sake of uniformity, in all the following descriptions of the 
vertebral column, the long axis of the body is supposed to be in the 
horizontal position. 

8 These numbers are not exact, owing to the uncertainty in the mode 
of reckoning the sacral vertebrae. 


The exceptions are : Near the posterior part of the trunk, 
in nearly all Mammals which possess completely developed 
hinder limbs, two or more vertebrae become ankylosed 
together to form the " sacrum," the portion of the vertebral 
column to which the pelvic girdle is attached. As a rule, 
none of the other vertebrae are normally united by bone, 
but in some species there are constant ossific unions of 
certain vertebrae, more particularly in the region of the 
neck. These will be specially noticed presently. 

FIG. 2. Anterior surface of human thoracic vertebra^fourth), . c body or centrum ; 
r neural canal ; / pedicle and / lamina of the arch ; / transverse process ; az 
anterior zygapophysis. 

Although the vertebrae of different regions of the column 
of the same animal, or of different animals, present great 
diversities of form, there is a certain general resemblance 
among them, or a common plan on which they are con- 
structed, which is more or less modified by alteration of 
form or proportions, or by the superaddition or suppression 
of parts to fit them to fulfil their special purpose in the 

An ordinary vertebra (see Fig. 2) consists in the first place 
of a solid piece of bone, the body or centrum (c), of the 


form of a disk or short cylinder. The bodies of contiguous 
vertebras are connected together by a very dense, tough, and 
elastic fibrous material, called the interverttbral substance, of 
peculiar and complex arrangement. This substance forms 
the main, and in some cases the only, union between the 
vertebrae. Its elasticity provides for the vertebrae always 
returning to their normal relation to each other and to the 
column generally, when they have been disturbed therefrom 
by muscular action. 

A process (/) rises on each side from the dorsal surface 
of the body. These meeting in the middle line above form 
together an arch, surrounding a space or short canal (tic). 
As in this space lies the posterior prolongation of the great 
cerebro-spinal nervous axis, or spinal cord, it is called the 
neural canal, and the arch is called the neural arch, in con- 
tradistinction to another arch on the ventral surface of the 
body of the vertebra, called the hcetnal arch. 1 The last is, 
however, never formed in Mammals by any part of the 
vertebra itself, but only by certain bones, placed more or 
less in apposition with it, and which will not here be con- 
sidered as parts of the vertebral column, strictly speaking. 

The lower portions of each side of the arch (/>), usually 
thick and more or less vertical in direction, constitute its 
pedicles. The upper more compressed and more horizontal 
portions (/) are the lamina. The pedicles are usually 
notched in front and behind, but most deeply behind, to 
form the sides of the interrertebral foramina for the trans- 
mission of the nerves issuing from the spinal cord. Occa- 
sionally the foramina for these nerves perforate the pedicles, 
instead of being truly intervertebral. 

The laminae meet in the median line above, at a more or 

1 So called because it incloses the heart and the great central blood- 


less open angle. At the point of their junction there is 
usually a single median process projecting dorsally, called 
the spinous process or neural spine. 

In most cases upon the anterior and posterior edges of the 
laminae of the arch are flattened, slightly projecting, more or 
less oval, smooth surfaces or facets, which in the natural state 
are covered by a thin layer of cartilage, and come into 
contact and articulate (by synovial joints) with the corre- 
sponding surfaces of the immediately antecedent and suc- 
ceeding vertebra?. These have been called by Sir Richard 
Owen zygapophyses ; that placed on the front edge of the 
arch being the anterior zygafophysis, that on the hinder 
edge the posterior zygapophysis. As a general rule the latter 
have their faces directed downwards, overlying the upward 
directed anterior zygapophyses of the vertebra next behind. 
This is a useful rule to remember in ascertaining which is 
the front and which the posterior surface of a vertebra. 
Sometimes, especially in the lumbar region, the posterior 
zygapophyses have their faces directed outwards, in which 
case the corresponding anterior zygapophyses look inwards 
(Fig. 3, az). 

These articular surfaces on the arch constitute a second 
mode by which the vertebrae are united, and their size and 
conformation aid to regulate the amount of motion allowed 
between the component parts of the column. They are often 
entirely wanting when flexibility is more needed than 
strength, as in the greater part of the caudal region of 
long-tailed animals. 

In addition to the body and the arch, there are certain 
projecting parts called processes, more or less developed in 
different vertebrae. Many difficulties exist about the signi- 
fication, homologies, and terminology of these processes. 
Probably, when more is known of the development of the 


vertebrae in a large series of animals, some further light will 
be thrown on the subject ; but at present it does not appear 
that there is that uniformity in the plan of construction of 
all vertebne which has often been supposed, and definitions 
of the different parts applicable in every case have not yet 
been arrived at, and it may even be doubted whether this 
will ever be possible. 

FIG. 3. Anterior surface of the lumbar vertebra of Hare (Ltfus tintiJtis). s spinou 
process ; m meupophysis ; as anterior zygapophysis ; t transverse process ; 

The principal processes commonly met with are as 
follows : 

1. From the middle of the upper part of the arch, a process 
(Fig. 3, s) generally single, but sometimes bifid at the end, 
grows out vertically. This is the spinous process, or neural 
spine already mentioned ; about its homology in different 
vertebrae there never can be any question. It may, however, 
be completely absent when the arch is round or smooth 
above, as in the cervical region in some animals ; on the 
other hand, it may grow out into a very long conspicuous 
rod of bone, as in the anterior dorsal region of others. 

2. Occasionally a process grows in the median line from 
the under-surface of the body. This may be single and long 
and slender, as in the anterior lumbar vertebrae of the Hare 


(Fig. 3 , h\ or a sharp median ridge, as in the cervical verte- 
brae of many Ungulata and the cervical and caudal vertebrae 
of the Ornithorhynchus, or double, as in the atlas vertebra of 
the last-named animal and the caudal vertebros of many others. 
This is termed a hypapophysis. Most commonly there is not 
even a trace of any such process. 

3. From the sides of the lower part of the arch, or from 
the body, lateral processes project more or less directly out- 
wards. These are called transverse processes. There may be 
but one, or there may be two, superior and inferior, on each 
side of a vertebra. In the latter case the superior is some- 
times called a diapophysis^ and the inferior a parapophysis : 
though it is questionable whether the processes to which 
these terms have been applied can always be regarded as 
strictly homologous. 

4. Besides these principal laterally projecting processes, 
there are often others arising from the side of the arch, more 
especially developed in the lumbar region, though by no 
means constant even there. Of these there may be one or 
two on each side. They have often been called accessory 


processes ; but in the more precise system of nomenclature 
introduced by Sir Richard Owen, the one which is situated 
highest on the arch (see Fig. 4, m), projects more or less 
forwards as well as outwards, is usually thick and rounded, 
and is nearly always in relation with the anterior zygapo- 
physis, is termed metapophysis ; l the one placed rather lower 
(Fig. 4, a), and which projects more or less backwards, and 
is generally rather slender or styliform, is called anapophysis. 
These, with the zygapophyses before mentioned, sometimes 
called oblique processes, but which are rather articular surfaces 
than true processes, are all the processes commonly met with 
on any Mammalian vertebra. 

Development of the Vertebra. The first indication of the 
formation of a vertebral column in the embryo is the 
appearance of a longitudinal primitive dorsal groove in the 
germinal membrane, the edges of which (lamina dorsales) 
rise up and meet above, so as to convert the groove into a 
canal. From the tissue lining this canal (uppermost layer 
of the germinal membrane) the brain and spinal cord are 
developed, and in its walls are formed anteriorly the cranium, 
and posteriorly the vertebral column ; the canal itself 
becoming the cerebral cavity and the neural canal of the 

In the floor of this canal, formed by a horizontal lamina 
which separates it from another and larger, ventral or haemal 
canal (formed by the approximation in the middle line below 
of the lamina rentrales), a slender rod of peculiar structure 
is developed. This is the notodwrd or chorda dorsalis, 
around which the bodies of the future vertebrae are deve- 
loped. In the Mammalia it almost completely disappears at 

1 It is also called " mammillary process " in some works on Human 


a very early period, traces only remaining in the axis of the 
intervertebral substance, though in many of the inferior 
Vertebrata it is persistent as a continuous rod for a longer 
period, and sometimes permanently. 

The formation of the arches of the vertebrae in the 
lamina dorsalcs is preceded by the appearance of dark- 
looking cellular masses called proto-vertebrce or somatomes, 
corresponding in number, though not exactly in situation, to 
the future vertebrae, and which undergo a series of changes 
(for a description of which the student is referred to special 
treatises on embryology) out of which ultimately results a 
vertebra, similar in shape to that which it presents in adult 
life, but formed of a continuous piece of hyaline cartilage. 

The mode of ossification of this cartilaginous vertebra in 
the different groups of Mammals still offers an interesting 
field for investigation, but the following is a summary of the 
most important facts ascertained regarding it. 

Leaving out for the present the greatly modified two 
anterior vertebrae, the atlas and the axis, which must be 
specially considered afterwards, and also the comparatively 
rudimentary vertebrae of the caudal region, each vertebra 
consists at one period of three pieces of bone, as distinct 
from each other, and remaining so for as long a period, as 
many of the separate elements of the skull. 

One constitutes the greater part, but usually not the whole, 
of the body or centrum. Each of the others forms one side 
of the arch, and usually more or less of the upper lateral 
part of the body. These last ultimately unite to each other 
in the middle line above, and to the central piece on each 
side below. The line of union between them and the central 
piece is readily distinguishable in all vertebrae up to the time 
the animal is about half-grown, and is named by Professor 
Huxley the neuro-central suture. (See Fig. 8, p. 33, ncs.) 


As a general rule all the processes (except the hypapo- 
physes) arise from the part of the vertebra situated above 
the neuro-central suture, but there are notable exceptions. 

The body of the vertebra is nearly always completed by 
the addition of a thin disk-like epiphysis at each end, which 
for a considerable period after it is fully ossified remains 
adhering by a rough surface to the central or main part 
of the body, and is easily separated from it by maceration. 
Its coalescence with the remainder of the body, especially in 
the thoracic region, is one of the last acts in the completion 
of the bony skeleton, and does not take place until after all 
the epiphyses of the limb bones are firmly united. Hence 
it may be taken as a safe indication that the animal is 
thoroughly adult. 

It must be noted that the epiphysis covers the whole 
surface of the end of the body, whether ossified from the 
centrum or the arch, and is therefore quite independent of 
the position of the neuro-central suture. 

These terminal epiphyses to the bodies of the vertebrae 
are peculiar to the Mammalia, but not found universally 
throughout the class, as they are wanting, or, at all events, 
very imperfectly developed, in the Monotremata and the 
Sirenia. In man, the highest apes, and also in some of the 
Marsupialia, they have less solidity and importance than in 
other Mammals, being often mere thin osseous rings, repre- 
senting the circumferential portion only of the ordinary 

The various processes of the vertebrae have been divided 
into those that are autogenous, or formed from separate 
ossific centres, and exogenous, or outgrowths from either of 
the just-mentioned primary vertebral constituents. 

There can be no doubt but that an autogenous process 
of one vertebra of an animal may be serially represented 


by an exogenous process in another vertebra of the same 
animal ; l and likewise that the corresponding processes of 
the same vertebra may be developed exogenously in one 
animal and autogenously in another. 

In nearly all the more prominent processes, moreover, 
whether formed by exogenous or autogenous ossification, 
the extreme tip remains cartilaginous for a considerable 
time ; and at a comparatively late period in developmental 
life (near the approach of maturity) a small ossific centre 
forms in it. This spreads through the cartilage, and then 
constitutes an epiphysis, which ultimately unites to, and 
becomes indistinguishably incorporated with, the remainder 
of the process. 2 

The spinous process is either formed by the coalescence 
of outgrowths from the two pieces forming the neural arch, 
or the greater part of it may be (as in the long spines of the 
anterior thoracic vertebrae of Ungulates) formed by a very 
early autogenous ossification, which soon becomes united to 
the upper part of the arch. In either case it is usually com- 
pleted by an epiphysis of comparatively late ossification. 

There is one part connected with certain vertebrae which 
requires some particular consideration, on account of the 
great modifications it presents, being in some regions a 
largely developed independent bone, articulated with the 

1 Even the arches of some of the caudal vertebra; appear to be 
ossified directly from the body, and not independently, as is the rule 
in the thoracic and lumbar vertebras. 

2 These epiphyses are sources of considerable difficulty in tracing 
homologous parts, as it is questionable whether they should be treated 
as separate elements of the skeleton, and, if not, where to draw the 
line between an epiphysis and an element. They often appear mere 
conveniences of growth, as it were, being developed upon the end of 
a process when it is long, and being absent in a corresponding part of 
stunted dimensions. 


vertebra by synovial joints, and in other regions a small 
rudiment, early and firmly united to, and incorporated with, 
the vertebra itself. 

The ribs in the thoracic region, though primarily formed 
from a rod of cartilage continuous with that of the vertebra, 
always become distinct, independent, and movably arti- 
culated bones; after their original segmentation they can 
never be properly said to constitute part of the vertebra. 
But it frequently happens that in certain of the vertebras 
anterior to the thoracic region, and in certain of those 
posterior to it, there are bony elements formed at an early 
period, which, though very different from ribs in the ordinary 

FIG. . Third cervical vertebra of a nearly full-grown Echidna (E. aculeata), the 
different pieces of which it is composed being slightly separated from one another. 
ma neural arch ; c centrum ; / transverse process ; v vertebrarterial canal ; IKS 
neuro-central suture. 

sense of the word, occupy a somewhat similar position 
in relation to the vertebrae to that which the ribs do in the 
thoracic region. These have hence been considered as 
modified conditions of the same part, and have been called 
pleurapophyses by Professor Owen. 

Perhaps the clearest case of the presence of rib elements 
in the vertebrae in any Mammal is afforded by the cervical 
vertebrae of the Monotremata, where the greater part of each 
transverse process ossifies separately from the rest of the ver- 
tebra, and remains for a long time only suturally connected 
with it (Fig. 5). They thus closely correspond to the cervical 


ribs of reptiles, which are unquestionably homologous serially 
with the thoracic ribs. 

The anterior, or more properly inferior, bar of the trans- 
verse process of the seventh, and occasionally of some of the 
other cervical vertebrae in Man, is autogenously developed, 
and has some characters by which it may be placed in the 
category of rudimentary ribs. 

The transverse processes of the anterior lumbar vertebra? 
of certain Mammals, as the Pig, are originally autogenous 
elements, though coalescing very early with the rest of the 

In the sacral region, the separate lateral ossifications which 
connect the vertebral column with the ilium present many 
characters allying them to ribs. (See Fig. 6.) 

FIG. 6. Anterior surface of first sacral vertebra (human) showing mode of develop- 
ment, na neural arch ; t centrum ; p distinct (pleurapophysial) ossification fur 
attachment of ilium. 

Finally, the transverse processes of the caudal vertebra? of 
some animals (as the Manatee and Beaver) are separately 
developed, though it is doubtful whether this circumstance 
alone is sufficient to entitle them to be considered as costal 

Division of the Vertebral Column into Regions. For con- 
venience of description the whole vertebral column has 


been divided into five regions, the cervical, thoracic? lumbar, 
sacral, and caudal. 

This divison is useful, especially as it is not entirely 
arbitrary, and in most cases is capable of ready definition, 
at least in the Mammalia ; but at the contiguous extre- 
mities of the regions, the characters of the vertebra of one 
are apt to blend into those of another region, either nor- 
mally, or as peculiarities of individual skeletons. 

i. The Cervical region constitutes the most anterior por- 
tion of the column, or that which joins the cranium. 

The vertebrae which belong to it are either entirely 
destitute of movable ribs, or, if they have any, these art- 
small, and do not join the sternum. 


FlG. 7. Anterior surface of sixth cervical vertebra of Dog, \. t spinous process ; 
as anterior zygapophysis ; v vertcbrarterial canal; / transverse process; f it* 
inferior lamella. 

As a general rule they have a considerable perforation 
through the base of the transverse process (the rertc 
brarterial canal Owen), or, as it is sometimes described, 

1 Generally called dorsal, but it would be better to reserve this term 
in morphology as relating to the upper surface of the body and opposed 
to ventral. 


they have two transverse processes, superior and inferior, 
which, meeting at their extremities, enclose a canal. (See 
Fig. 7 ; Fig. 8, p. 33 ; and Fig?. 17 and 18, p. 45, 46.) This, 
however, rarely applies to the last vertebra of the re- 
gion, in which only the upper transverse process is usually 

The transverse process moreover very often sends down 
near its extremity a more or less compressed plate (inferior 
lamella, Fig. 7, /"), which being considered to be serially homo- 
logus with the ribs of the thoracic vertebrae (though not 
developed autogenously) is often called " costal " or " pleura- 
pophysial " plate. This is usually largest on the sixth, and 
altogether wanting on the seventh vertebra. 

The first and second cervical vertebrae, called respectively 
atlas and axis, are specially modified for the function of 
supporting, and permitting the free movements of the head. 
They are not united together by an " intervertebral sub- 
stance," but connected only by ordinary ligaments and 
synovial joints. 

The cervical region in Mammals presents the remarkable 
peculiarity that, whatever the length or flexibility of the 
neck, the number of vertebroe is the same, viz. seven, with 
very few exceptions, which will be particularised further on. 

2. The Thoracic or Dorsal region consists of the vertebrae 
which succeed those of the neck, having ribs movably 
articulated to them. These ribs arch round the thorax, the 
anterior one, and most usually some of the others, being 
attached below to the sternum. 

The characters of the ribs and their mode of articulation 
with the vertebras will be considered further on, but it may 
now be stated that in the anterior part of the thorax the 
vertebral extremity of each rib is divided into " head " and 
" tubercle " ; that the former is attached to the side of the 


body of the vertebra, the latter to its transverse process : 
and that the former (capitular) attachment corresponds to 
the interspace between the vertebras, the head of the rib 
commonly articulating partly with the hinder edge of the 
body of the vertebra antecedent to that which bears its 
tubercle. Hence the body of the last cervical vertebra 
usually supports part of the head of the first rib. In the 
posterior part of the series the capitular and tubercular 
attachments commonly coalesce, and the rib is attached 
solely to its corresponding vertebra. 

3. The Lumbar region consists of those vertebra of the 
trunk in front of the sacrum (to be afterwards defined) which 
bear no movable ribs. It may happen that as the ribs 
decrease in size posteriorly, the last being sometimes more 
or less rudimentary, the step from the thoracic to the 
lumbar region may be gradual and rather undetermined in 
a given species. But most commonly this is not the case, 
and the distinction is as well defined here as in any other 

As a general rule there is a certain relation between the 
number of the thoracic and the lumbar vertebrae, the whole 
number being tolerably constant in a given group of 
animals, and any increase of the one being at the expense of 
the other. Thus in almost ail Artiodactyle Ungulata there 
are 19 thoracico-lumbar vertebras; but these may consist of 
12 thoracic and 7 lumbar, or 13 thoracic and 6 lumbar, or 
14 thoracic and 5 lumbar. 

The smallest number of thoracico-lumbar vertebrae in 
Mammals occurs in some Armadillos, which have but 14. 
The number found in Man, the higher Apes, and most Bats, 
viz. 17, is exceptionally low ; 19 prevails in the Artiodactyles, 
nearly all Marsupials, and very many Rodents; 20 or 21 in 


Carnivora and most Insectivora, 23 in Perissodactyla. The 
highest and quite exceptional numbers are in the two-toed 
sloth (Chold'pus) 27, and Hyrax 30. 

The prevailing number of rib-bearing vertebrae is 12 
or 13, any variation being generally in excess of these 

4. The Sacral region offers more difficulties of definition, 
especially at its posterior portion. 

Taking the human " os sacrum " as a guide for comparison, 
it is generally defined as consisting of those vertebra?, between 
the lumbar and caudal regions, which are ankylosed together 
in the adult state to form a single bone. It happens, how- 
ever, that the number of such vertebrae varies in different 
individuals of the same or nearly allied species, especially as 
age advances, when a certain number of the tail vertebrae 
generally become incorporated with the true sacrum. 

A more certain criterion is derived from the fact that some 
of the anterior vertebrae of the sacral region have distinct 
additional (pleurapophysial) centres of ossification, between 
the body and the ilium (see Fig. 6, p. 26). To these perhaps 
the term sacral ought properly to be restricted, the remaining 
ankylosed vertebrae being called pseudo-sacral, as suggested 
by Gegenbaur. Our knowledge of the development of the 
sacrum in different animals is not sufficient at present to 
apply this test universally, but it appears probable that two 
is the most usual number of true sacral vertebrae, as thus 
defined in the Mammalia. 

5. The Caudal Vcrtebrcc are those placed behind the 
sacrum, and terminating the vertebral column. They van- 
in number greatly, being reduced to 5, 4, or even 3, in a 
most rudimentary condition, in Man, some Apes and Bats, 
and being numerous and powerfully developed with strong 


and complex processes in many Mammals, especially among 
the Edentata, Cetacea, and Marsupialia. The highes 
known number, 49, is possessed by the African Long-tailed 
Manis. (Af ant's mafrura.) A small Insectivora, Microgale 
longicaudata from Madagascar has sometimes as many as 
47, but in these cases there is considerable individual 



ORDER PRIMATES. The human cervical vertebrae (ex- 
cluding for the present the first and second) have short, 
wide, depressed bodies, hollowed in front from side to side, 
and behind from above downwards, 1 with wide neural canals, 
and short, broad, and usually bifid spines (considerably 
longer in the seventh vertebra than in the others), well 
marked, broad, flat, anterior and posterior zygapophyses, 
and short, sub-bifid, widely perforated transverse processes. 

These vertebrae are, as usual, developed mainly from three 
centres, one for each side of the arch, and one for the 
centrum (see Fig. 8), but it will be observed that the whole 
of the body is not formed from the latter, but that its lateral 
parts, with the transverse processes, are ossified from t In- 

Besides these main centres of ossification there are thin 
and imperfect disk-like epiphyses on the ends of the body, 

1 As before mentioned, the body is supposed to be placed horizontally, 
^o that the same terms of relative position may be used as when speak- 
ing of the vertebral column of the ordinary less modified animals of the 


very late in making their appearance, and not joined until 
long after the rest of the vertebra is completed. A small 
epiphysis is also formed on the end of the spinous 

Lastly, the inferior or ventral bar of the transverse process 
of the seventh vertebra is developed from a separate centre 
of ossification, and occasionally the same part of the sixth 
and fifth has its own separate nucleus. This bar of bone is 
connected internally with a projection from the side of the 
body, ossified from the arch ; externally with the end of the 
upper or true transverse process, which is an exogenous 

FIG. 8. Sixth cervical vertebra of a child, i. c centrum ; net neuro-central suture ; 
T' vcrtcbrarterial canal ; az anterior zygapophysis. 

growth from the arch, so that it is attached to the vertebra 
entirely above the neuro-central suture. Occasionally it 
acquires an abnormal development, and grows into a con- 
siderable rib-like bone, in which case it is usually united at 
its distal extremity with the first thoracic rib. 

The first vertebra or atlas (Fig. 9) is little more than an 
oval ring, thickened on each side into the so-called " lateral 
mass," which bears an articular surface before and behind. 
The anterior surfaces are very large, elongated from above 
downwards, and hollowed for the reception of the condyles of 
the occiput. The posterior articular surfaces are subcircular 



flattened, or slightly-concave. The transverse processes are 
short, stout, and perforated ; the arch presents scarcely a 
rudiment of a spinous process. On its anterior edge im- 
mediately above the articular surface is a deep notch or 
groove (g) of some importance, as it corresponds with the 
slight notch in front of the pedicle in other vertebras, which 
contributes with the deeper notch in the hinder border of 
the pedicle of the preceding vertebra to form the " inter- 
vertebral " foramen for the exit of a spinal nerve, and because, 
occasionally in man, and constantly in many animals, it is 
converted by a bridge of bone into a canal, through which 

FIG. 9. Human atlas, young, showing development. :]. in inferior arch ; as articula 

surface for occiput ; t transverse process ; g- groove for first spinal nerve am 

the first cervical (or subocdpitat) nerve passes out. The 
inferior arch of the atlas (ia) differs entirely from the bodies 
of the other vertebrae, being a simple, depressed, slightly 
curved bar of bone, with a smooth facet on its neural or 
upper surface, for articulation with the odontoid process of 
the axis. 

The second cervical vertebra, axis, epistropheus, or vertebra 
dentata (Fig. 10), has a body terminating anteriorly in a 
large subconical median projection, the odontoid process 
(o), which is received into, and articulates with, the con- 
cavity of the inferior arch of the atlas. It is retained in 




its place by means of a strong transverse ligament passing 
between the lateral masses of that bone, and separating its 
canal into an upper or neural portion for the passage of the 
spinal cord, and an inferior portion for the reception of the 
odontoid process. 

The axis has posterior zygapophyses placed on the arch, 
serially continuous with those of the rest of the vertebrae, but 
its anterior articular facets, like those of the atlas, do not 
belong to the arch proper, but partly to the body and partly 
to the arch, and are therefore not exactly serially homologous 

FIG. 10. Diagram showing mode of ossification of human axis (ha:mal or vential 
surface), o odontoid process, or centrum of atlas ; c proper centrum of axis ; na 
neural arch ; as anterior articular surface ; e t e epiphyses, completing the ends of 
the centra. 

with the zygapophyses of the other vertebrae. The trans- 
verse processes are short, single, and perforated. The arch 
is high, with a stout bifid spine. 

The development of the atlas and axis offers some im- 
portant points for consideration. 

The arch of each is ossified from two centres, one on each 
side, as in other vertebrae ; but if the axis is examined a year 
or two after birth (Fig. 10), its body appears to be composed 
of two parts, one placed in front of the other, the first includ- 
ing the odontoid process and the anterior part of the body, 

1) 2 


the second all the remainder of the body. The arch is 
united to both. On the other hand, the atlas at the time 
of birth has nothing corresponding to the centrum of other 
vertebras, its inferior arch being still cartilaginous. 

It is therefore a generally received opinion among ana- 
tomists that the anterior ossification of the axis is essentially 
the body of the atlas, which unites with both arch and 
centrum of the vertebra behind it. It must be observed, 
however, that in its mode of ossification, at least in Man, it 
differs from the contra of all the other vertebras, as at one 
period it consists of two distinct lateral pieces, which after 
a while coalesce in the middle line. The usual disk-like 
epiphyses of the vertebral bodies are represented by one 
at the posterior extremity of the body, by a small osseous 
nodule which completes the odontoid process in front, and 
by some irregular ossifications found between the two main 
portions of which the body is composed. 

The inferior arch of the atlas ossifies soon after birth 
from one or more centres, and the resulting piece of bone 
(Fig. 9, id] ultimately unites with the two pieces forming 
the neural arch about the same time as that at which they 
join together in the middle line above. This piece may 
probably be regarded as a detached " hypapophysial " seg- 
ment of the first vertebral centrum, the remainder of which 
forms the odontoid portion of the body of the axis. 

The cervical vertebras of the other PRIMATES resemble 
those of Man generally, the most noticeable deviations being 
the following : 

In the atlas the groove for the first cervical nerve is usually 
converted into a foramen ; and a median hypapophysial 
tubercle or spine often projects backwards from its inferior 
arch under the axis (especially in Mycetes and Lagothrix). 

The spinous processes, especially of the third, fourth, 

iv.] CARN1VORA. 37 

fifth, and sixth cervical vertebrae, are immensely elongated 
in the Gorilla, and considerably so in the Chimpanzee and 
Orang. These processes as a rule are not bifid, as in Man, 
but occasionally (as in Mycetes) they are trifid, having a pair 
of lateral backward-projecting processes developed near 
their extremity. 1 

The inferior lamellae of the transverse processes are 
generally larger proportionally than in Man, especially in the 
Lemurina. In the seventh vertebra, the transverse processes 
vary much as to their perforate or imperforate condition. 

In the CARNIVORA, the atlas (Fig. n) has very deep 
anterior articular surfaces for the condyles of the skull. The 

FlG. ii. Inferior surface of atlas of I>og, J. in foramen for first spinal nerve ; 

' canal. 

first spinal nerve passes through a complete foramen. The 
transverse processes are large, wing-like, flattened from above 
downwards, and perforated by the vertebrarterial canal. 

The axis (Fig. 12) has a long conical odontoid process, 
and a large compressed neural spine, greatly extended from 
before backwards, and especially produced forwards. 

The remaining cervical vertebras have small, narrow, 
compressed, usually simple spines, gradually lengthening to 
the seventh, and large transverse processes, with greatly 

1 These are named hyperapophysu by Mivart, who has called par- 
ticular attention to them : "On the Axial Skeleton in the Primates : " 
Proc. Zool. Soc. 1865, p. 545. 


developed inferior lamellae (see Fig. 7, p. 27, /') especially 
large in the fifth and sixth. In the latter the lower edge of 
this lamella is frequently hollowed in the middle, and pro- 
duced at each extremity, so that the transverse process has 
a trifid appearance. This is especially marked in the Felida. 
The transverse process of the seventh vertebra has no inferior 
lamella, and its base is imperforate. 

Metapophyses are generally more or less developed on the 
cervical vertebrae of the Carnivora, and there are also in 
some genera small backward projecting tubercles (Jiyperapo- 
physes y Mivart) situated on the laminae of the arch, rather 

FIG. 12. Side view of axis of Dog, jj. s spinous process; o odontoid process ; 
pz posterior zygapophysis ; / transverse process ; v vertebrarterial canal. 

internal to the posterior zygapophyses, not usually found in 
other vertebra;. 

In the INSECTIVORA the cervical vertebras vary consider- 
ably in their characters. The atlas has usually short trans- 
verse processes. Generally the spinous process of the axis is 
large and prominent, and that of the other vertebrae very 
small, but in Centetes and Pofanwgale they are all more or less 
elongated. The neural arches in some (as Myogale and 
Sorex) are reduced to mere filaments. In the mole (Talpa} 
the transverse processes of the fourth, fifth, and sixth 
vertebra; are much expanded antero-posteriorly, and overlap 

iv.] RODENTIA. 39 

each other. Large single hypapophyses are developed from 
the inferior surface of most of the cervical vertebras in the 
Shrews (Sorex) and some of their allies, and in Galeopithecus 
each vertebra bears at its hinder end a pair of hypapophysial 

In the CHIROPTERA all the cervical vertebrae are broad, 
very short from before backwards, with slender neural arches 
from which (except in the axis) no distinct spinous processes 
are developed. In certain forms (as Vesptrugo) some of the 
vertebras have distinct double hypapophysial spines project- 
ing backwards. 

In the RODENTIA the atlas has usually broad, moderately 
long, wing-like transverse processes. The odontoid process 
is long and slender ; the spinous process of the axis is much 
developed, while as a rule that of the other cervical vertebras 
is exceedingly small. The transverse processes of the fifth 
and sixth have large inferior lamellas ; that of the seventh 
is sometimes perforated at the base (as in Ltptts), and 
sometimes imperforate (as in Hydrochcerus). 

In the Capybara (Hydrochcerus) and some others, the side 
of the arch of the atlas is perforated near its anterior border 
for the exit of the first spinal (sub-occipital) nerve, and also 
near its hinder border for the second cervical nerve. 

In the Jerboas (Difus) a very exceptional condition of 
the cervical vertebras occurs. The atlas is free, but all the 
others are ankylosed together by both bodies and arches, 
and the bodies are very wide and depressed, as in the 

Among the UXGULATA, the atlas (Fig. 13) in the Pecora is 
very long, with deep articular cavities for the occipital con- 
dyles. The transverse processes are not wide, but much 
extended from before backwards, and flattened from above 
downwards. Each is perforated by a foramen (sn) which gives 



exit to the inferior division of the first cervical nerve, but not 
by the vertebral artery, which usually enters the neural canal 
between the arches of the second and third vertebra;. The 
odontoid process of the axis (Fig. 14) is of peculiar shape, 

being like a spout, or hollow half-cylinder, with a prominent 
sharp semicircular rim. The canal for the second cervical 

spinal nerve pierces the lamina of the axis near its anterior 
border. The other vertebra? have more or less elongated 
bodies, which are opisthocalous, i.e. concave behind and 
convex in front. They are keeled below, the keel being often 

iv.] UNGULATA. 41 

developed into a hypapophysial spine posteriorly ; the neural 
spines are moderately long, and inclined forwards. The 
transverse processes of the fifth, and especially of the sixth 
have large inferior lamella;. That of the seventh is usually 
im perforate. 

In the Giraffe the bodies of the cervical vertebra; are very 
long. The transverse processes are short, but so extended 
from before backwards as to become divided into two, one 
at the anterior and one at the posterior end of the vertebra. 
That of the seventh is perforated. 

In the Tylopoda (Camels and Llamas) the vertebrarterial 
canal passes obliquely through the anterior part of the 
pedicle of the arch, being in its posterior half confluent with 
the neural canal. A similar condition occurs in Afacrau- 
chenix, an extinct South American Perissodactyle Ungulate. 

The Suina and Tragulina differ from the remaining exist- 
ing Artiodactyles in the form of the odontoid process, which 
is conical ; while on the other hand the Horse and Tapir 
among the Perissodactyles have this process wide, flat, and 
hollowed above, approaching the form it presents in the 
Ruminants. In the Pig, the broad pedicles of all the cervical 
vertebra; are perforated by canals for the passage of the 
upper division of the spinal nerves. 

The bodies of the cervical vertebra; in the Rhinoceros, 
Tapir, and Horse are markedly opisthocoelous. but in the 
Pig and Hippopotamus very slightly so. 

In the Horse the bodies of the cervical vertebra; are 
elongated, with a strong keel and hypapophysial spines. 
The neural lamina; are very broad, the spines almost obso- 
lete, except in the seventh, and the transverse processes not 
largely developed. The seventh is not perforated by the 
vertebrarterial canal. 

In the Rhinoceros, on the other hand, the bodies are 


comparatively short, and not keeled, the laminae narrow, the 
spines well marked, and the transverse processes greatly 
developed, especially those of the atlas. 

In the Elephant (order PROBOSCIDEA), the atlas much 
resembles the human atlas. The axis has a short conical odon- 
toid process and a very massive spine, broad above and bifid 
posteriorly. The bodies of the other vertebrae are very short, 
flattened, sub-circular disks, very slightly opisthoccelous. 
Excepting the seventh they all have short spinous processes, 
and short, broad, and largely perforated transverse processes. 
The seventh has a high spine, an imperforate transverse pro- 
cess, and on the hinder edge of its body a very distinct arti- 
cular cavity for the head of the first rib. In the young animal 
this is divided into two equal parts by the neuro-central 

In the order SIRENIA, the Dugong (Halicore) has seven 
cervical vertebrae, as in the Mammalia generally. The atlas 
has short imperforate conical transverse processes. The axis 
has a high arch and massive neural spine, a short rounded 
odontoid process, and very rudimentary transverse processes. 
The others have short and wide bodies, small spines, and 
irregularly developed transverse processes, often not com- 
pletely enclosing a vertebrarterial canal. 

The Rhytitta, a large animal of this order, which became 
extinct towards the close of last century, had also seven 
cervical vertebrae, and the Miocene Halitherium had the 
same number. 

The Manatees (genus Manatus), of which there are two 
well-marked forms, one inhabiting the west coast of 
Africa, and the other the east coast of Central and South 
America, never have more than six vertebras in the cervical 
region. These resemble generally those of the Dugong, 
having short and wide bodies, and very irregular transverse 

iv.] CETACEA. 43 

processes. In a specimen of Manatus senegalensis^ in the 
Museum of the College of Surgeons, the second and third 
are ankylosed by their bodies, and the neural arches of most 
of the others are widely open above. In the skeletons of M. 
americanuS) in the same museum, the vertebrae are all free, 
and the arches, though slender, are complete, and with very 
slightly developed spinous processes. In a specimen of J/. 
americanus in the Museum of Cambridge the axis and atlas 
have large neural spines ; the vertebrarterial canal is incom- 
plete in the 2nd, 3rd and 4th vertebra, wanting in the 5th, 
but complete in the 6th which carries a distinct movable 
cervical rib. 

In the Cetacea the seven cervical vertebrae usually found 
in the Mammalia are always present, though often so short 
and blended together, that it is not easy at first sight to re- 
cognise their existence. In some genera of both sub-orders 
all the vertebne are free, though never allowing of much 
motion between them ; but more commonly certain of them 
are firmly united together by bone. Even where the atlas 
and axis are separate the odontoid rarely forms a distinct 
process (it is most distinct in Platanista), but still it is devel- 
oped from an ossific centre of its own, as in other Mammals. 

Among the Mystacoceti, in the Right Whales (genus 
Balana) the whole of the seven cervical vertebra? are usually 
united into one mass by their bodies, though sometimes the 
seventh is free. The arches are also more or less united 
above, though generally not in a continuous mass. Small 
slit-like openings between the narrow pedicles of the arches 
permit the exit of the cervical spinal nerves, and in the adult 
condition afford the only indications by which the number of 
the united vertebrae can be ascertained. Already before 
birth most of the bodies have coalesced, and it is even 
doubtful whether they ever exist in a separate condition. 




The Fin Whales or Rorquals (genus Bal&noptcra) present 
a totally different condition of cervical vertebra?, as these are, 
as a rule, all distinct and free, though occasionally, as an 
individual peculiarity, an irregular ankylosis may take place 
between two or more of them. 1 

FIG. 15. Section through middle line of united cervical vertebra; of Greenland Right 
Whale (Balcena inysticctus), J. particular surface for occipital condyle ; e epi 
physis on posterior end of body of seventh cervical vertebra : sn foramen in arch 
of atlas for first spinal nerve ; i arch of atlas ; 2 3 4 5 6 conjoined arches of the 
a.\is and four following vertebra; ; 7 arch of seventh vertebra. 

In the common large Fin Whale of our coasts (//. 
musculns) the atlas (Fig. 16) has short, stout, conical, imper- 
forate transverse processes. The axis (Fig. 17) has a broad 
oval body, high massive arch, very short odontoid process, 
and very wide, oblong wing-like transverse processes directed 
somewhat backwards, and with an oval perforation near the 

1 Sec Professor Struthers ' 
Articulations in Fin Whales." 
November 1872.) 

On the Cervical Vertebra: and their 
(Journal of Anatomy and Physiology, 



base. The other cervical vertebrae (Fig. 18) have similar 
broad, very short bodies, small arches, without spines, and 
very long transverse processes, composed of slender upper 

Fie. 16 Anterior surface of at las of common Fin Whale (!>al<rnt>ftfra ntuscntits). ,'.. 
in foramen for first spinal nerve. 

and lower bars, widely separated at their bases, but united at 
their extremities so as to enclose a very large space between 
them. In the seventh the upper process only exists, and the 

Flc. 17. Anterior surface of axis of common Fin Whale (Balitncptera. tnufctilus), i . 
a odontoid process. 

lower one is occasionally imperfect in the sixth. 1 In very 
young animals these processes are formed only of cartilage 

1 Professor Turner has shown that, in a foetal BaLmoftera si 
the inferior transverse process of the seventh is present in a cartilagi- 
nous condition. (Journal of Anatomy and Physiology, May 1871.) 

4 6 



and as ossification takes place gradually from within out- 
wards, and does not reach the outer extremity until the 
animal approaches maturity, specimens are frequently met 
with in museums, which, instead of completely annular 
transverse processes, show only truncated upper and lower 
bars. In some species, however (as in Megaptera boops), 
most of the cervical vertebras remain permanently in this 

FIG. 18. Anterior surface of fourth cervical vertebra of the same animal, J. as ante- 
rior of zygapophysis ; t upper transverse process ; t' lower transverse process. 

Among the Odontoceti, all the cervical vertebree are free in 
the Gangetic Dolphin (Platanistd), and in the allied South 
American genera Inia and Pontoporia, also in the Nar- 
whal (Monodon) and the Beluga, or the White Whale. In 
most of these genera the atlas has a large hypapophysial 
process, projecting under and articulating with the body of 
the axis, which develops no distinct odontoid. In the 
Narwhal irregular ankyloses between the bodies of the cervical 
vertebrae are very frequent. In all the other Delphinidcc 
(including Delphinus, Orca, Psendorca, Glolncef>Jialns, Pho- 
ama, &c.), at least the first and second cervical vertebras 
are united by both body and spine, and most commonly 
some of the succeeding vertebrae are joined to them. If 
any are free, it is always those situated most posteriorly, 
and they have extremely thin, sub-circular disk-like bodies, 

iv.] EDENTATA. 47 

and irregular and comparatively rudimentary transverse 

In Hyptrcodoti) the whole of the cervical vertebrae are 
ankylosed together. In the other Ziphioids several of the 
posterior vertebne are free, and the allied Cachalot, or Sperm 
Whale (Physeter), presents a condition not met with in any 
other known Cetacean : the atlas is free, and all the other 
neck vertebrae are completely united. 

Among the various members of the order EDENTATA, the 
cervical vertebne present very different conditions. 

In the Armadillos (Dasypodidcc) the bodies are extremely 
short, broad, and depressed, and several are commonly anky- 
losed together ; the corresponding neural arches being also 
united, the neural, and to a certain extent the vertebrarterial, 
canals form continuous tubes. The orifices for the spinal 
nerves perforate the united pedicles. The atlas is always 
free. The vertebrae that are united are the second and third, 
or the second, third, and fourth ( Tatusia pcba), and in some 
species the fifth also. The spinous processes of the axis and 
those of the fused vertebrae in general are very large, but the 
neural arches of the hinder free vertebrae are extremely 
narrow, and the spinous processes rudimentary. The trans- 
verse process of the seventh has an inferior lamella, nearly 
as large as that of the sixth, and is not perforated. 

In Orycteropits, the Pangolins (Mam's), and the Anteaters 
(Myrmccophagd)) the neck vertebra? are more normal in form, 
and are not ankylosed. In the last-named genus, the verte- 
brarterial canal of several (2, 3, 4, in M. jubata) of the 
vertebras perforates the pedicle obliquely, and enters the 
neural canal posteriorly, much as in the Camels. 

Among the leaf-eating Edentates, or Sloths, the neck 
vertebrae present some remarkable peculiarities, especially as 
to number. 


All the known species of three-toed Sloths (genus Brady- 
pus] have nine cervical vertebra, i.e. nine vertebrae in front 
of the one which bears the first thoracic rib (or first rib con- 
nected with the sternum, and corresponding in its general 
relations with the first rib of other Mammals), but the ninth, 
and sometimes the eighth, bears a pair of short movable 
ribs. The eighth is perforated by the vertebrarterial canal, 
but not the ninth. 

The common species of two-toed Sloth (C/wlcepus didac- 
lylus] has seven cervical vertebra?, but a closely allied 
species (C. Iwffmannii) has but six. In both cases the 
vertebrarterial canal is continued to the end of the series, 
the sixth and seventh cervical vertebra: respectively. 

In the very heterogeneous order MARSUPIALIA (sub-class 
Didelphia) the cervical vertebrae vary much in their characters, 
though the number is always seven, as in the great majority 
of the Mammalia. 

One of the most important variations is in the mode of 
ossification of the atlas. In the Wombat (Pliascolomys}. 
Koala (Phascolarctos), P/ialangisfa, and Kangaroo (Macro- 
pus), there is no distinct osoific nucleus in the inferior arch 
of the bone, which remains either permanently open in the 
middle line below, or (as in some of the smaller Kangaroos) 
is completed by the union of prolongations of the arches 
inwards. This, however, is not the case with the carnivorous 
Marsupials. In the Thylacine (see Fig. 19) there is a distinct 
heart-shaped piece of bone in the centre of the inferior arch 
of the atlas, which appears never to become united to the 
remainder, as it is still attached by ligament in skeletons 
otherwise perfectly mature, and is commonly lost in mace- 
ration. In Perameles and Didelphys the atlas is completely 
ossified below by a wide intermediate piece, quite as in 
ordinary Mammals. 


As to the other vertebne, in the Kangaroos the transverse 
processes are long and slender, and (including the seventh) 
have a very small perforation close to the base. The inferior 
lamella arises near the base of the process, and is very 
large in the sixth, but generally absent in the seventh 

In the Wombat, the bodies are wide and depressed. The 
transverse processes are perforated in all ; the inferior 
lamella of the sixth is much developed antero-posteriorly. 
The spines of all are rather short. 

Fir,. 19. Inferior surface of atlas of Thylacine (Tkyl&cinMt ejmotefilialiii). \. A dis- 
tinct ossification in centre of inferior arch, with pointed hypapophysial pro- 

In Pframeles lagotis the greater jxirt of the transverse pro- 
cess of the axis is ossified separately from the rest of the 
vertebra, and remains sometime distinct, as in the Mono- 
tremata. In this genus, as in the other carnivorous Marsu- 
pials, the inferior lamellae of the transverse processes of the 
fourth, fifth, and sixth vertebrae, but especially of the latter, 
are particularly large. 

Some species of American Opossums (as Didelphvs 
Virginia na and its nearest allies) have the spinous processes 
of the second, third, fourth, and fifth cervical vertebrae very 
high, square, and massive, and being closely applied to each 
other by flattened surfaces, form a solid wall of bone along 
the top of the neck. 


In both genera of MONOTREMATA (sub-class Ornitho- 
delphia) the cervical vertebrae are seven in number, and in 
both the inferior arch of the atlas is completely ossified, 
apparently from a separate centre ; but in Ornithorhynchus 
a large bifurcated hypapophysis is developed, which is 
wanting in Echidna. 

In Ornithorhynchus also all the other cervical vertebras 
have a single median hypapophysial spine, equally wanting 
in Echidna. 

In both, the axis has a high compressed spine, and the 
odontoid portion remains long distinct from the true centrum 
of the bone. In both, the transverse processes are of auto- 
genous formation, and remain suturally connected with the 
remainder of the vertebra, until the animal is nearly full- 
grown (see Fig. 5, p. 25); that of the axis is still distinct in 
an adult Ornithorhynchus. Though in this respect they 
present an approximation to the Sauropsida (Reptiles and 
Birds), they differ from that group, inasmuch as there is not 
a gradual transition from these autogenous transverse pro- 
cesses of the neck (or cervical ribs, as they may be con- 
sidered) into the thoracic ribs, for in the seventh vertebra 
the costal element is much smaller than in the others, 
indicative of a very marked separation of neck from thorax, 
not seen in the Sauropsida. The vertebrarterial canal in 
Echidna bruijni is very wide, and passes through all the 
vertebrae from the second to the seventh. 



IT will be most convenient to consider the vertebrae of 
these two regions together. 

In Man, there are seventeen trunk vertebrae, twelve 
thoracic or rib-bearing, and five lumbar. 

The bodies increase in size from before backwards, and 
also change their form. The first is like a cervical vertebra, 
broad and depressed. They soon become more compressed, 
especially at the lower part, so as to be sub-triangular when 
seen from one end (Fig. 2, p. 16); after the middle of the 
thoracic region they become more circular in outline, and 
in the lumbar region they are wide transversely. The ends 
of the bodies are flat or slightly concave. 

The neural canal does not alter greatly in size throughout 
this region, though it does somewhat in form. In the first 
vertebra it is wider in proportion to its height than in any of 
the others. 

The arches have comparatively narrow pedicles arising 
from the anterior half of the body, deeply notched behind 
for the canal for the exit of the spinal nerves. The lamina 
are broad. The spines are moderately long, sub-equal 


throughout the series, rather slender, and sloping backwards 
in the thoracic region ; broader (in the antero-posterior 
direction) and more upright in the lumbar region, and pre- 
senting but scarcely any indication of that convergence 
towards a point in the posterior thoracic region so frequently 
seen in other Mammals. They are generally simple and 
slightly dilated at their ends ; but in the lumbar region, the 
posterior edge is often more or less bifid. 

The zygapophyses are well developed throughout. In 
the thoracic region they are oval, fiat facets, looking pretty 
nearly directly upwards (the anterior) and downwards (the 
posterior) : the anterior, developed on the top of the pedicle 
and projecting forwards, being supported by the " oblique 
process ; " the posterior is placed on the under-surface of the 
hinder part of the lamina. In the lumbar region, their form 
and position change, the anterior having their outer edges 
turned upwards, and supported by a short rounded meta- 
pophysis (inammillary process). The posterior ones have 
undergone a corresponding change, so that their faces, instead 
of looking downwards, are directed obliquely outwards ; they 
are also much curved. 

The transverse processes project throughout the series 
from the arch, near the junction of the pedicle with the 
lamina. In the greater part of the thoracic region they are 
tolerably long, project somewhat upwards and slightly for- 
wards, and are dilated and tuberous at the extremities, on 
the under-surface of which (except in the two last) they show 
a smooth concave facet for the attachment of the tubercle of 
the rib. In the posterior part of the thoracic region they 
are shorter, and begin to resolve themselves into three dis- 
tinct processes, generally conspicuous in the first lumbar. 
One of these projects outwards, and, elongating in the second 
and third lumbar, it forms the principal transverse process. 

v.] PRIMATES. 53 

One projects upwards and forwards, by the side of the 
anterior zygapophysis ; this is the metapophysis^ or mam- 
millary process. The other projects backwards, and 
represents in a rudimentary condition the process so largely 
developed in many animals called anapophysis. This 
gradually becomes smaller in the second and third lumbar 
vertebrae, and generally disappears in the fourth. 

The lumbar transverse processes are thus not serially 
homologous with the thoracic ribs, but with the part of the 
transverse process of the thoracic vertebrae to which the 
tubercle of the rib is attached, and are complementary to 
the ribs, becoming greatly augmented in size directly these 
cease. Neither are they normally developed autogenously. 1 

The sides of the bodies of the thoracic vertebrae bear 
facets for the articulations of the heads of the ribs. Except 
the last three or four, each vertebra supports a portion of the 
head of two ribs, having a large facet near its anterior edge 
(placed partly on the body and partly on the side of the 
pedicle) for the head of its own rib (i.e. the rib which arti- 
culates also with the transverse process), and on the hinder 
border of the upper angle of the body a small facet to 
receive the anterior edge of the succeeding rib. In the 
hinder part of the thoracic region the rib is connected only 
with its corresponding vertebra, and not with the one in 

Among the remaining Primates, 19 is the prevailing 
number of trunk vertebrae, of which usually 12 to 14 bear 
ribs. The Gorilla and Chimpanzee (genus Troglodytes), 
agree with Man in having 17. The Orang (Simia) has 

1 There are several specimens in the College Museum which show 
the co-existence, on the first lumbar vertebra, of a rudimentary (supple- 
mental) rib, with a transverse process serially homologous with the 
transverse processes of the other lumbar vertebrae. 


usually but 16. The Gibbons (Hylobates) and Spider 
Monkeys (Ateles) have mostly 18. Among the Lemurina, 
Loris and Nycticebus have as many as 23 or 24. 

Of thoracic vertebrae, the Gorilla and Chimpanzee have 
13, the Orang 12, the Gibbons usually 13 ; other Old World 
Monkeys mostly 12 ; the American Monkeys from 12 to 15 : 
the Lemurs from 12 to 16. 

As a general rule the vertebral column, taken as a whole, 
is straighter than it is in Man, showing- a much less marked 
sigmoid curve. 

Except in the anthropoid Apes, and a few others, the 
spinous processes of the anterior thoracic vertebrae lean 
backwards, and those of the lumbar and some of the 
posterior dorsal vertebras forwards, so that they converge to 
a point near the hinder part of the thoracic region, some- 
times called " the centre of motion " of the vertebral column. 1 
This may be between two vertebras, but more often there is 
one, which has an upright spine, towards which the others 
are directed ; this is the " anticlinal vertebra." It is at this 
point that the thoracic vertebrae begin to change their 
characters, and assume those of the lumbar vertebrae ; and 
the simple elongated transverse processes break up as it 
were into the metapophyses, anapophyses, and lumbar 
transverse processes, all of which are conspicuous in these 

The transverse processes of the lumbar vertebrae are 
usually placed lower on the sides of the vertebras than 
in Man. 

In Galago the hinder edges of the neural spines of 
the lumbar vertebrae bear a pair of backward-projecting 

1 This disposition of the spines of the trunk vertebrae is still more 
marked in many of the inferior mammals, especially the terrestrial 

v.] CARNIVORA. 55 

processes, which clasp the anterior edge of the succeeding 
spine. Similar processes are developed, but to a less extent, 
in the Howling Monkeys (Mycetes) and in Lagothrix. 

The foramina for the exit of the spinal nerves, instead of 
being " intervertebral," perforate the pedicles of the arches 
in the Potto (Perodicticus). In the same genus, two or three 
of the anterior thoracic vertebrae have very long slender 
spinous processes, which in the living animal project beyond 
the general level of the skin, forming distinct conical 
prominences, covered only by an exceedingly thin and 
naked integument. 

In the CARNIVORA, the trunk vertebrae are nearly always 
20 or 21 in number. The genera Felis, Canis, and Viverra 
have 13 thoracic and 7 lumbar, Hyana 15 and 5, Mustela, 
Nasua, Procyon, and Ursus 14 and 6, Meles 15 and 5 > 
Mephitis has the exceptionally high number of 16 and 6; 
Mellirora but 14 and 4. Among the Seals, Cystophora, Otaria 
and Phoca have 15 and 5, Trichecus and Stenorhynchus 
14 and 6. 

The spines of the anterior thoracic vertebrae are long and 
slender, and slope back to about the eleventh (the antidinai), 
after which they are shorter, thicker (from before backwards), 
and lean forwards. From this point also metapophyses and 
anapophyses become distinctly developed ; the latter are 
especially large in the Fdidce. The lumbar vertebra? have 
long transverse processes directed forwards and rather 
downwards, and short, stout, compressed spinous pro- 

In the Seals, the trunk vertebrae present much the same 
characters, but the anapophyses are usually but slightly 
developed, or may be altogether absent, and the spinous 
processes show no convergence to a " centre of motion." 

Among the INSECTIVORA, the number of the trunk vertebrae 


varies much in the different genera, from 1 8 (13 thoracic and 
5 lumbar) in Tupaia, 19 (13 and 6) in Talpa and most 
Soricidcz, 19 (14 and 5) in Gahopithecus, 21 (15 and 6) in 
Erinaceus, 22 (19 and 3) in Chrysochloris, to 24 (19 and 5) 
in Centetes. 

There are also great differences- in the development of the 
processes of the vertebrae, which appear to accord with the 
diversities in the habits and movements of the animal. The 
transverse processes of the lumber vertebras are very short 
in the comparatively slow-moving, running, or burrowing 
Hedgehogs (Erinaceus), Shrews (So rex) and Moles (Talpa), 
but they are very long, broad, and inclined downwards in the 
jumping Macroscelides and Rhynchocyon, where the lumbar 
muscles are greatly developed and the hinder extremities 
disproportionately large. 

In the Mole, there are distinct, small, oval, flat ossicles on 
the under-surfaces of the interspaces between the lumbar 
vertebras. Similar ossicles, but in a more rudimentary con- 
dition, are occasionally found in the same situation in some 
other Insectivora, as the Hedgehog, but not in any other 

The usual number of thoracic and lumbar vertebrae in the 
CHIROPTERA is 17 or 18, of which from n to 14 may bear 
movable ribs. The transverse processes of the lumbar 
vertebras are almost obsolete, as are also the spinous pro- 
cesses throughout the series. 

Among the RODENTIA, the most prevalent number is 19 ; 
but it rises as high as 23 (16 and 7) in Capromys, and even 
as 25 (17 and 8) in Loncheres. 

The characters of the vertebras vary much in the different 
genera, as among the Insectivora. In the Hares (genus 
Lepus) the anterior thoracic vertebrae have long slender 
spinous processes; the lumbar vertebras (see Fig. 3, p. 18) 


have very long and slender transverse processes directed 
downwards and forwards and widening at their extremities ; 
long metapophyses projecting upwards and forwards, small 
anapophyses, and remarkably long, single, compressed 
median hypapophyses. These latter are not found in the 
Rodentia generally. 

In the UNGULATA, the bodies of the trunk vertebrae are 
generally slightly opisthocoelous. The spinous processes in 
the anterior thoracic region are exceedingly High and com- 
pressed. The transverse processes of the lumbar vertebra; 
are long, flattened, and project horizontally outwards or 
slightly forwards from the arch. The metapophyses are 
moderately developed, and there are no anapophyses. The 
canals for the exit of the spinal nerves frequently pierce the 
pedicle of the neural arch (as in the Eland and Cape Buffalo). 

In the Artiodactyle sub-order the number of thoracic and 
lumbar vertebrae together is almost always 19, though the 
former may vary from 12 to 15. Among the Perissodactyles 
the number 23 is equally constant, the Horse and Tapir 
having 18, and the Rhinoceros 19 thoracic vertebra. 

Some species of Hyrax (ff. capensis) have as many as 
22 thoracic and 8 lumbar vertebra, making altogether 30, 
the highest number in any terrestrial Mammal ; whilst 
in Dendrohyrax the numbers are 21 and 7. 

The Elephants have 23 in all, 19 or 20 of which bear ribs. 

In the order SIRENIA, the thoracic vertebrae are numerous 
and the lumbar very few ; thus the Dugong (f/alicore) has 
19 thoracic and 4 lumbar, and the Manatee (Manatus) 17 
and 2. The bodies are rather triangular, being compressed 
and keeled below, and in the young state have no dis- 
tinctly ossified terminal epiphyses. The bodies of all the 
thoracic vertebrae bear articular facets for the heads of the 
ribs. The spinous processes are not very high, but the 


zygapophyses are well developed throughout the series. The 
metapophyses are rudimentary, and there are no distinct 

As there is no sacrum in the CETACEA the lumbar re- 
gion passes directly into the caudal, and they can only be 
distinguished by the presence of " chevron bones " in the 

The thoracic vertebras vary in number from 9 in Hyper- 
oodon, to 15 and occasionally 16 in some Fin-Whales 
(Balanopterd), and the lumbar vertebrae from 3 in Inia (the 
Amazonian fresh- water Dolphin) to 24 or even more in some 
of the true Dolphins (Delphinus). 

The bodies are short in the anterior part of the thoracic 
region, but posteriorly become more or less elongated and 
cylindrical. Their terminal epiphyses are strongly ossified 
disks, very distinct in young animals, but coalescing com- 
pletely with the rest of the body in adult age. The spinous 
processes are high and compressed. The zygapophyses are 
very little developed, and only found in the anterior thoracic 
region. The metapophyses are distinct (see Fig. 20, m], 
placed at first near the ends of the transverse processes, 
but gradually rising on the arch, are ultimately transferred 
to the sides of the anterior edge of the neural spine, from 
which they project forwards, clasping between them the 
hinder edge of the spine of the vertebra in front. 

In most Cetacea the transverse processes in the anterior 
thoracic region arise rather high on the side of the neural 
arch of the vertebra, but in the hinder part of the same 
region become gradually placed lower, until finally they are 
transferred to near the middle of the side of the body, 
which position they occupy in the lumbar region (see 
Fig. 20). The transverse processes of the lumbar vertebrae 
are thus evidently serially homologous with the transverse 




processes of the anterior dorsal vertebrae, which, in their turn, 
continue backwards the upper series of cervical transverse 

In the Physetfridie. (comprising P/iysekr, Hyptroodon, 
Ziphius, and the allied forms) a very different and peculiar 
arrangement occurs (Fig. 21). The transverse processes in 
the anterior thoracic region (/) are placed quite similarly to 

F<;. ao. Anterior surface of vertebra: of Dolphin (Globicef>kal*t me/at). ]. A fifth 
thoracic ; H seventh thoracic ; i eighth thoracic ; t> first lumbar ; r rib ; m meta- 
pophysis ; / transverse process. The dotted lines indicate the position of the 
Deuro-central suture. 

those of the ordinary Dolphins ; but passing backwards, 
instead of changing their position on the vertebrae, they 
gradually become smaller, and finally disappear: while, 
simultaneously with their diminution in size, other processes 
(/) rise from the body of the vertebra, in the situation of the 
capitular attachment of the rib, which, rapidly increasing in 
length, become continuous serially with the lumbar transverse 



processes. In two or three vertebrae the two co-exist (Fig. 
21, B and c), resembling the upper and lower transverse pro- 
cesses of the neck, and sometimes even meeting at their 
extremities so as to enclose a canal. The lumbar transverse 
processes in this case therefore are not serially homologous 
with the transverse processes of the anterior thoracic region, 

FIG. 21 .Anterior surface of vertebra; of Sperm Whale (I'/iyseter itmcrocef>hahts), ^ . 
A eighth thoracic ; u ninth tlioracic ; c tenth thoracic ; D fifth lumbar ; r rib ; 
in rnetapophysis ; / upper transverse process ; /' lower transverse process. 

and with the upper transverse processes of the neck, as in the 
former case, but rather with the lower transverse processes 
of that region ; and yet tried by every other test, the special 
homology of the transverse processes of the lumbar vertebrae 
of a Dolphin (Fig 20, D /) and a Sperm Whale (Fig. 21, D /) 
is perfectly evident. 

v.] EDENTATA. 61 

The mode of ossification of the thoracic and lumbar 
vertebrae of the Cetacea appears, so far as it has been 
ascertained, to differ from that of all other Mammals, inas- 
much as the neuro-central suture (see Fig. 20) is always 
placed a little above the junction of the arch and the body, 
the whole of the latter, with any process which may arise 
from it, being ossified from the central nucleus. Conse- 
quently, in the thoracic vertebne of the Dolphins, the trans- 
verse process is anteriorly an outgrowth from the arch, then 
partly from the arch and partly from the body, and finally 
from the body alone a condition quite unknown in other 

It would appear, from the conflicting statements on the 
subject, that the transverse processes of the lumbar region 
are sometimes ossified autogenously, and sometimes exo- 
gcnously from the centrum. 

The members of the order EDKNTATA present some great 
peculiarities in the condition of the trunk vertebrae, especially 
those of the lumbar region. 

As to numbers, the Three-toed Sloths (Bradypus) have 
19 altogether (either 16 and 3, or r$ and 4); and the Two- 
toed Sloths (C/iola'pus) have sometimes as many as 24 tho- 
racic and 3 lumbar, making altogether 27 trunk vertebrae. 
The great Anteater (Afyrwecophaga) has 18 (15 and 3. 
R. Coll. Surg. ; 1 6 and 2, Mus. Cam.) ; the little Two-toed 
Anteater (Cydothurus\ 17 (15 and 2). The Armadillos 
have 14 to 16, the Pangolins (Manis) commonly 18 to 21 
and the Cape Anteater (Oryctcropus) 21 (13 and 8). 

The vertebral column of the Sloths is remarkable for th e 
extremely broad, flat laminae and short neural spines, lyin 
backwards on the next succeeding vertebrae, throughout the 
whole column down to the sacrum. All the processes are 
very short, and the spines are bifid in the lumbar region. 



In Bradypits a small (anapophysial) process projects 
backwards from the hinder edge of the transverse process 
of each lumbar vertebra, having on its inner surface a facet, 
which articulates with a corresponding facet on the anterior 
edge of the arch of the succeeding vertebra, below the 
ordinary zygapophysis. 

In Megatherium, Myrmecophaga, Cycloturus, and Dasypus 
(in fact, all the remaining American Edentates), a disposition 
thus slightly indicated in the Sloths, is carried out to a great 

l-'lG. 22. - Side view of twelfth and thirteenth thoracic vertebra; of Great Anteater 
(Myrmtcophaga, jubata), 3. / metapophysis ; tc facet for articulation of tubercle 
of rib ; cc ditto for capitulum of lib ; az anterior zygapophysis ; a: 1 additional 
anterior articular facet ; pz posterior zygapophysis ; pz 1 and pz- additional pos- 
terior articular facets. 

extent, and results in a very complex and altogether peculiar 
method of articulation between the vertebrae. 

It will be most convenient to describe it from one species, 
the Great Anteater (My rmecophaga jubata), but it is the same 
in principle in all the above-named genera. 

The anterior thoracic vertebrae articulate in a perfectly 
normal manner by large anterior and posterior zygapophyses. 
These retain the horizontal position of their facets through- 
out. On the eleventh dorsal vertebra, the upper surface of 
the backward projecting process which bears the posterior 



zygapophysis (pz) below, develops an articular surface 
(/z 1 ) which looks upwards and articulates with a corre- 
sponding downward directed process (az l ) developed on the 
upper part of the arch of the following vertebra, rather 
below the metapophysis (///). Thus the vertebra has a 
process projecting backwards, with flattened articular facets 
on its upper and under surface, fitting into a deep recess 
on the anterior edge of the arch of the vertebra behind, 

FIG. 23. Posterior surface of second 
lumbar vertebra of Great Anteater, . 
/ transverse process ; pz posterior 
zygapophysis ; /s 1 , fz j . and /z 3 . ad- 
ditional posterior articular Cicets. 

FIG. 24. Anterior surface of third lum- 
bar vertebra of Great Anteater, j. 

articular facets. 

and the articulation is now by two zygapophysial surfaces 
on each side of the arch instead of one. 

In the thirteenth ' thoracic vertebra a third articular facet 
(PZ*) is developed on the hinder margin of the lamina of 
the arch, still higher than the last additional one (f>z l ), 
and separated from it by a deep notch. This looks mainly 
outwards, and articulates with a corresponding facet (az 2 , 
Fig 24) on the anterior edge of the arch of the fourteenth 

1 In the Cambridge specimen this description applies to the fourteenth 
and fifteenth vertebrae respectively. 


vertebra, placed to the inner side of the metapophysis, which 
is now situated on a process projecting forwards into the 
notch between the two upper articular facets of the antece- 
dent vertebra. So that there are now three distinct articu- 
lations connecting the arches of the vertebraj on each side, 
the processes of the vertebrae which bear them interlocking 
in a " tenon and mortise " fashion. 

This condition continues as far as the second lumbar, in 
which, in addition to these three facets, a fourth (ps 3 is) 
developed on the under-surface of the hinder edge of the 
transverse process near its outer extremity, which articulates 
with a similar facet (az 3) on the upper-surface of the 
transverse process of the third lumbar, so that there are now 
four pairs of articular facets, or zygapophyses, on each arch. 
The same occurs also between the third lumbar and the 
first sacral vertebra. 

In the Armadillos the lumbar metapophyses are very long, 
and project upwards, outwards, and forwards, supporting 
the bony carapace, while the broad transverse processes are 
exceedingly reduced. 

An allied extinct genus, Glyptodon, had the greater number 
of the trunk vertebrae completely ankylosed, a condition 
altogether unique in the Mammalia. 

In neither of the Old World Edentates, Mam's and 
Orycteropus, is there any development of the articular facets, 
other than the ordinary zygapophyses. In the former genus 
the metapophyses (contrary to the usual rule) project rather 
backwards than forwards. The anterior zygapophyses of the 
lumbar and posterior thoracic regions are largely developed) 
and very concave, completely embracing the semicylindrical 
surfaces of the posterior zygapophyses. There are no 
distinct anapophyses. In Orycteropus the lumbar vertebra? 
are numerous (8), with carinated bodies, long and slender 


spines inclined forwards, long, broad, and flat transverse 
processes pointing forwards and downwards, well developed 
metapophyses and rudimentary anapophyses. 

In the MARSUPIALIA, the number of thoracico-lumbar 
vertebras is invariably 19, although there are some apparent 
exceptions, in which the last lumbar assumes the form of a 
sacral vertebra. The rib-bearing vertebrae are always 13, 
except in the Koala (Phascolantos), which has but 1 1, and one 
species of Wombat (Phascolomys vombatus), which has 15. 
The Hair)-- nosed Wombat (P. latifrons) has the ordinary 

In the Kangaroos, the lumbar vertebras have largely 
developed metapophyses and anapophyses, and moderate- 
sized transverse processes much curved forwards. 

In the running and jumping Bandicoots (Perameles) the 
lumbar vertebrae have very slender, long, forward -directed 
spines, and long transverse processes. In the climbing 
Opossums (Didt'lphys), on the other hand, the spines are 
very short and broad from before backwards. 

The MONOTREMATA agree with the Marsupials in the total 
number of trunk vertebrae, but those that bear ribs are more 
numerous, viz. 16 in Echidna, and 17 in Ornithorhynehus. 

The spinous and transverse processes are very short, 
and the ribs have no articulation with the latter, but are 
attached to the bodies only, the greater part of the articular 
surface being below the neuro-central suture, the reverse of 
what occurs in the higher Mammals. In the thoracic ver- 
tebrae the canals for the exit of the spinal nerves perforate 
the neural arch. 



Sacral Vertebra. The difficulties in defining the sacral 
vertebras have been noticed at page 30. Their essential 
character is best illustrated by tracing it up from the simple 
condition it presents in the tailed Amphibians (as Menopoma). 
In these animals a series of similar small straight ribs are 
movably articulated to the ends of the transverse processes 
of all the trunk vertebra;, which are not distinctly divisible 
into separate regions. To the distal extremity of one of 
these the ilium is attached. This vertebra with its rib thus 
constitutes the "sacrum," and the ilium is clearly seen not 
to be a " pleurapophysis," as it is sometimes called, or any 
part of a vertebra, but a something distinct and superadded. 
In the Crocodiles there are two vertebras with strongly 
developed rib-like bones connecting them to the ilium, and 
remaining long only suturally united to their vertebras. 

The inferior ossification of the transverse processes of the 
true sacral vertebras in Mammals (see Fig. 6, p. 26) is clearly 
of the same nature, though more rudimentary in character, 
and coalescing at an earlier period with the remainder of the 
vertebras. It is not yet known that it exists in all Mammals, 


but this may be considered probable, as it is certainly found, 
at least in the first sacral vertebra, in such different forms as 
Man, the Chimpanzee, Orang, Cat, Sheep, Elephant, Sloth 
and Wombat. 

The ankylosis ot additional vertebrae in the Mammalia is 
probably related to the greater fixity and more complete 
attachment of the pelvis to the vertebral column in this class ; l 
for the innominate bone is not only articulated by its iliac 
portion to the true sacral vertebne, but it has also a posterior 
connection with the vertebral column by its ischial portion, 
by means either of very strong ligaments, or in some cases 
by bony union. 3 

In Man there are usually five ankylosed vertebrae, con- 
stituting the " os sacrum " of anthropotomy, but only two, or 
sometimes three, have distinct costal elements. The re- 
mainder may be called pseudo-sacral, and belong more pro- 
perly to the caudal series. The sacrum as a whole is broad, 
strongly curved in the longitudinal direction, with the con- 
cavity downwards, and its anterior extremity forms with the 
body of the last lumbar vertebra a more prominent " sacro- 
vertebral angle " than in other Mammals. 

In the Gorilla, Chimpanzee, and Orang, there are 
generally five ankylosed vertebrae, to which the last lumbar 
not unfrequently becomes united in old animals. The whole 
sacrum thus formed is long and narrow, gradually tapering 
posteriorly, and much less curved than in Man. In the 
other Monkeys, there are usually two or three, rarely four, 

1 This is carried to a still greater extent in birds. 

a Hence the following definition of the sacrum : " The posterior 
limit of the sacral region is characterized, not by the union of the 
different osseous pieces, which varies according to age, but by the place 
of insertion of the ischio-sacral ligaments." (A. MILNE EDWARDS, 
Famille Jes Chtvrotains, p. 52.) 

F 2 


ankylosed vertebrae ; the first two, or true sacrals, are broad . 
and behind these the sacrum suddenly contracts. 

In the Lemurina the number of united vertebrae varies 
from 2 to 5. 

In the CARNIVORA, there are generally few true sacral 
vertebrae, only 2 in the Puma, Proteles, and Hyena. 2 or 3 in 
the Tiger ; in the Bears and Seals as many as 4 or 5 may 
be united by bone in old animals. In the Dog there are 
usually 3 ankylosed vertebras. 

In most UXGULATA and RODF.NTIA the sacrum consists of 
one broad vertebra joining the ilia, and a series of narrow 
ones, varying in number with age, gradually diminishing in 
width, ankylosed to it behind. 

In the Beaver among Rodents, most Chiroptera, the Cape 
Anteater (Oryderopus) among Edentates, and the Wombat; 
among Marsupials, the sacrum consists of numerous anky- 
losed vertebrae, with widely-expanded transverse processes, 
which are longer in the hindermost vertebrae, and nearly 
meet the ischia. 

In most other EDENTATA, as the Sloths, Anteaters, 
Armadillos, and in Pteropus among Chiroptera, this modi- 
fication is carried further, and the transverse processes of the 
hinder pseudo-sacral vertebrae form a complete bony union 
with the ischia, converting into a foramen what is usually 
the sacro-sciatic notch. In some of the Armadillos as many 
as 10 vertebrae are thus firmly fused together, and with the 
innominate bones. 

In MARSUPIALIA usually but one vertebra supports the 
iliac bones, though another is commonly ankylosed with it. 

In the MONOTREMATA, the Ornithorhynchus has 2 
ankylosed sacral vertebrae, and the Echidna 3 or 4. 

The CETACEA having no iliac bones, have no part of the 
vertebral column specially modified into a sacrum : but in 




the SIRENIA, the rudimentary ilia are attached by ligament 
to the end of the transverse processes of one vertebra, which 
may hence be regarded as sacral. 

Fie. 35. Anterior surface of third caudal vertebra of Leopard (Felit tfofiantia), |. 
<u anterior zygapophysis ; /i posterior zygapophysis ; m metapophysis ; / trans- 
verse process 

Caudal Vertebra. The vertebrae of the tail vary greatly 
in number and in characters in different animals. When it 
is well developed, as, for example, in the long-tailed 
Carnivora, from one of which the accompanying figures are 

FIG. a6 Upper surface of the third caudal vertebra of I/eopard, |. a* anterior 
zygapophysis ; /* posterior zygapophysis ; m metapophysis ; / transverse process. 

taken, the anterior vertebrae (Figs. 25 and 26) are com- 
paratively short and broad, with complete neural arches, 
though without distinct spines, prominent metapophyses, 
and anterior and posterior zygapophyses (the latter especially 




being raised on pedicles), and well-developed single transverse 
processes. But a gradual change takes place in these 
characters (see Figs. 27 and 28), the body lengthens out and 
becomes more and more cylindrical ; the neural arch 
diminishes and finally disappears, leaving for a while a pair 
of processes at each extremity of the vertebra, the remains 
of the parts of the arch which bore the xygapophyses ; the 
transverse process is much reduced, and confined to the 

FIG. 27. Anterior surface of twelfth 
caudal vertebra of Leopard, jj. tit 
metapophysis ; p processes serially 
continuous with those which support 
the posterior zygapophyses in the an- 
terior vertebrae ; /; hypapophyses. The 
process on the side of the body be- 
tween MI and k is the anterior trans- 
verse process. 

FIG. 28. Upper surface of twelfth caudal 
vertebra of Leopard, 5. in met:ipo- 
physes ; p processes serially continuous 
with those which support the posterior 
zygapophyses in the anterior vertebra;. 
t transverse process ; t' anterior trans- 
verse process. 

posterior extremity of the body, a second one appearing at 
the anterior extremity. Even these rudiments of processes 
gradually cease to be perceptible, and nothing is left but a 
cylindrical rod of bone, representing the centrum alone of 
the vertebra. These diminish in size towards the apex of 
the tail, the last being usually a mere rounded nodule. 

Connected with the under-surface of the caudal vertebrae 
of many animals which have the tail well developed are 



certain bones, formed more or less in the form of an in- 
verted arch (Fig. 29), called chci'ron bones (French, Os en V ; 
German Untere Bogen ; hamapophyscs, Owen). These are 
always situated nearly opposite to an intervertebral space, 
and are generally articulated both to the vertebra in front and 
the vertebra behind ; but sometimes chiefly or entirely either 
to one or the other. They are usually articulated movably 
to prominences (hypapophyses) on the lower surface of the 
body of the vertebra, but occasionally become ankylosed to 

FIG. 29. Anterior surface of fourth caudal vertebra of Porpoise (Pkoffna com- 
munis), J. i spinous process ; m mctapophysis ; / transverse process ; ft chevron 

it. They ossify from two centres, one on each side, which 
usually coalesce in the median line below, though not un- 
frequently, especially at the beginning and end of the series, 
where they are less developed, the two lateral portions remain 
permanently separate. They serve to give a larger surface of 
attachment for the inferior muscles of the tail, and also to 
protect the caudal vessels, which nin within the canal formed 
by the series of these bony arches. They are always best 


developed near the anterior extremity of the tail, and are 
never found under the posterior rudimentary vertebrae. 

In Man the caudal vertebrae are quite rudimentary ; 
usually 4 in number, all ankylosed together, and constituting 
the coccyx, or oscoccygts, of anthropotomy. The first is some- 
times ankylosed to the sacrum. 

Among the Simiina there are but 4 to 5 caudal vertebrae 
in the Anthropoid Apes and in the Barbary and Black 
Macaques, no more than 10 in some Baboons, and as many 
as 32 in Semnopithecus, and 33 in some of the Spider Mon- 
keys (Ateles). 

In the latter the tail is prehensile, and the vertebra; are 
broader and altogether more strongly developed than in the 
weak, pendant, though almost equally long tails of the Old 
World monkeys. 

Chevron bones are found in all except those species that 
have the tails quite rudimentary. They are most fully de- 
veloped in Ateles, where the extremities are often bifurcated. 
They are attached to a pair of projections on the anterior 
end of the lower surface of the vertebra. 

In the Lemurina, the number of the caudal vertebrae 
varies from 7 to 29. 

Among the CARNIVORA, the Bears have very short tails, 
with from 8 to 10 vertebras, the Seals from 9 to 15 ; some 
of the Lynxes have but 13, but most of the animals of the 
order have tails of moderate or great length ; the greatest 
number of vertebrae being found in Paradoxurus, which 
may have as many as 36. 

Chevron bones are usually not much developed ; they 
are articulated (sometimes ankylosed) to the front ends of 
the vertebrae, as in the Primates. 

In the INSECTIVORA, the tail is very variable. It is short 
and simple in Erinaceus and Centetcs, long in Galeopithecus. 

vi.) RODENT1A. 73 

Solenodon, Gymnura, Potamogalc^ Aficrogale (43 vertebrae), 
Tupaia and Rhynchocyon ; in the last-named genus the 
chevron bones are well developed and bifid. 

In the CHIROPTF.RA, the tail is sometimes exceedingly 
rudimentary, as in Destnodus and Pteropns ; sometimes elon- 
gated, but composed of long, simple, slender, cylindrical 
vertebral bodies ; and generally enclosed in the interfemoral 
cutaneous expansion. 

Among the different members of the order RODENTIA, 
there are great differences in the condition of the caudal 

In the Hares, Guinea Pigs, Capybara, &c, the tail is 
almost rudimentary. In the Cape Jumping Hare (Ptdetes) 
it is nearly as long and powerful as in the Kangaroo, and 
has well-developed chevron bones. 

In the true Porcupines the tail is generally short ; but in 
some allied genera (Tree Porcupines) it is much elongated 
and prehensile. 

In the lleaver (Castor) there are 24 caudal vertebrae, all 
short, broad, and depressed, and with wide transverse pro- 
cesses, becoming double (anterior and posterior) about the 
middle of the tail, not by development of a new process, 
but by gradual division of the one existing in the anterior 

In the UNGULATA, the tail is variable in length, but of 
simple character and function ; it is never prehensile, 
nor has it ever chevron bones, although occasionally, as 
in the Ox, a pair of well-developed hypapophyses may be 
produced so as to meet in the median line, enclosing a 
small canal. 1 The vertebrae are most numerous (about 16 

1 The Eocene Anoplotherium appears to have had chevron bones 
beneath the vertebrae of its long tail. 


to 20), in the Oxen and in some Antelopes (Koodoo), and 
least so in some Deer, especially Moschus, in which animal 
the tail is quite rudimentary. 

The Elephant has a long tail, composed of 31 vertebras 
of simple character, without chevron bones. 

In the Hyrax the tail is short, consisting of about 4 to 9 

As the tail is the principal organ of locomotion in the 
CETACEA, it is always very well developed, and consists of 
numerous (from 18 to 30) vertebrae. 

Chevron bones are always present, and of simple character, 
though with long compressed median spines (see Fig. 29, 
p. 71). They are mainly attached to the posterior extremity 
of the vertebra immediately in front of them. 

The characters of the caudal vertebra in the various 
animals of the order are tolerably uniform, the tail having 
the same function in all. In the anterior part of the region 
the bodies are very massive and cylindrical ; the arches have 
high spines, with metapophyses on their anterior edges, and 
the transverse processes are tolerably long, and directed 
straight outwards. In passing backwards the arches and all 
the processes gradually disappear, and the bodies become 
much compressed, and elevated vertically. Suddenly a 
change takes place (at the spot where the end of the 
vertebral column becomes enclosed in the horizontal, 
laterally extended cutaneous expansions, constituting the 
" flukes " of the tail,) and the vertebrae altogether alter 
their characters, becoming much smaller, wide transversely 
and depressed. There is always one vertebra which is 
transitional in its character between these two forms. Most 
of the caudal vertebras are perforated by a vertical canal on 
each side, either passing through the base of the transverse 
process, or in the posterior vertebrae through the body of 

vi.] EDENTATA. 75 

the vertebra itself. This transmits an ascending branch of 
the caudal artery. 

The SIRENIA have numerous, much depressed caudal 
vertebra, with wide transverse processes, gradually dimin- 
ishing in length to about the 15th vertebra from the end. 
The processes then widen again, to decrease once more 
towards the end of the tail, thus corresponding more or less 
with the shape of the caudal fin. They are thus very different 
from those of the Cetacea. 

Among the EDENTATA, the Sloths have a quite rudi- 
mentary tail, consisting of from 6 to 10 depressed vertebrae 
without chevTon bones. 

FIG. 30. Anterior surface of third caudal vertebra of Great Armadillo (Priotion 
jfifai), }. t spinous jmxres-, ; m metapophyiis ; at anterior zygapophysis ; 
I transverse process ; h chevron with diverging processes. 

In the allied Megatherium the tail was greatly developed, 
with long processes and large chevron bones, as is the 
case with nearly all the Entomophagous Edentates, but 
mostly so in the Pangolins (Mam's), one species of which 
(M. macrura] has 46 to 49 tail vertebrae, the highest number 
known in any Mammal. Cydoturus has a prehensile tail 
of 40 vertebrae. The little Chlamydophorus has a rather 
short tail of 15 vertebras, remarkable for being expanded, 
depressed, and spatulate towards the end, the transverse 


processes increasing in size instead of gradually diminishing, 
as is almost universally the case. 

The chevron bones are usually much developed. They 
are Y-shaped, having long, simple, compressed spines in 
Orycteropus ; V-shaped in Manis and most Armadillos ; but 
in Priodon (Fig. 30) they have wide, diverging, lateral pro- 
cesses, instead of a median spine. They are attached 
rather to the vertebra in front than to that behind them. 

In the MARSUPIALIA, as might be supposed in so hetero- 
geneous a group, there is great diversity in the condition of 
the caudal vertebrae. 

In the Wombat (Phascolomys) and Koala (Phascolarctos) 
the tail is comparatively rudimentary. 

In the Kangaroo, on the other hand, it is very large, and 
serves as an organ of support when standing upright. It is 
composed of 2 1 to 25 vertebrae, the first few with short bodies 
and large processes ; afterwards the bodies lengthen out, be- 
coming cylinders contracted in the middle. The zygapophyses 
soon cease, but the metapophyses continue longer. The 
neural arch is not continued longer than about the middle 
of the tail ; the transverse processes are gradually placed 
further and further back on the vertebra, and then anew one 
arises near the anterior end, so that they become double. 

The chevron bones are placed quite between the verte- 
brae, so that it is difficult to say to which they most 
properly belong. In the proximal part of the tail their 
free edge is compressed and develops a process forwards 
and backwards, giving a hatchet shape when seen side- 
ways. Further back they also send out broad processes 
laterally, so as to be cruciform, with a flat inferior surface. 

In some Marsupials the tail is prehensile, as in the 
Opossums (Didelphys), with from 19 to 35 vertebrae, and the 
Phalangers (Phalangista) with from 21 1031. 


Chevron bones are generally present in the tails of all the 
Marsupials, except the Wombat and Koala. In the Thylacine 
they are few, and comparatively rudimentary. 

The tails of the two forms composing the order 
MONOTREMATA differ considerably. 

The Echidna has 12 caudal vertebrae. These have no 
hypapophyses, but there are two single median chevron 
bones near the middle of the tail, more like the lumbar 
subvertebral ossicles of the Mole than ordinary' chevron 
bones. The Omithorhynchus has 20 or 21 caudal vertebra, 
with single median hypapophyses, no chevron bones, and 
wide transverse processes which increase in length towards 
the middle of the tail, being shortest near the sacrum and 
end of the tail. 

The following Table, showing the number of vertebrae in 
a very large series of Mammals, is mainly taken from the Cata- 
logue of the Museum of the Royal College of Surgeons of 
England. 1 Those distinguished by an asterisk (*) are from 
the collection of the University of Cambridge, and have been 
added to make the list still more complete. The + after the 
number of the caudal vertebrae indicates that the termina- 
tion of this region (as too often happens with mounted 
skeletons) was defective. The others are presumed to be 
complete or very nearly so. 

In compiling this list the following rules have been 
observed : The cervical vertebrae are those in front of the 
first which bears a rib directly articulating with the sternum. 
The thoracic or dorsal vertebrae are those which bear 

1 Catalogue of the Specimens illustrating the Osteology and Dentition 
of Vertebrated Animals recent and extinct, contained in the Museum of 
the Royal College of Surgeons of England, part ii. Class Mammalia, other 
than Man, by \V. H. Flower, assisted by J. G. Garson. London, 1884. 



movably articulated ribs. The lumbar vertebrae are those 
that intervene between the thoracic and sacral regions. The 
artificial and conventional definition of the sacral vertebrae 
as those that are ankylosed together in the adult state to 
form a single bone has been generally adopted in default of 
a more scientific one of universal application. In the 
Cetacea, the first caudal is that which bears upon its hinder 
edge the first pair of chevron bones. 

It must never be forgotten that although the division of 
the vertebral column into distinct regions is convenient for 
descriptive purposes, at the contiguous extremities of the 
regions the characters of the vertebrae of one region are 
apt to blend into those of the next, either normally or as 
peculiarities of individual skeletons. 







Homo sapiens 





J A 

Troglodytes ni^er 


I \ 




Gorilla savagii 


I ^ 




Simia satyrus . . 





Hylobates syndactylus 
* lar 

- 7 







Colobus vellen>us. . 






Semnopithecus entellus 





schistaceus . 
Nasalis larvatus 


- 7 




17 + 


Cercopithecus patas 






sabaeus ... 





albigularis ... 
petauri.-ta ... 
Macacus cynomolgus 






19 + 




2O + 

pilentus ... 


















nemestrinus . 










j + 






Cervic Thormc 




Theropithecus gelada 
































I 4 + 









19 + 


18 + 



Cynoccphalus babouin 

anubis . . 


Chrysothrix sciurea 

Cebus capucinus 

Ateles geoflfroyi 

Lagothrix humboldtii 

Nyctipithccus vociferans 
Pithccia monachus .... 

Mycetes seniculus . . 

Midas oedipus 

Hapale jacchus 

Indris brevicaudatus 

Avahis laniger 

Hapalemur griseus .. . 

Lemur varius 


Chirogaleus furcifer 


Galago crassicaudatus . . 


Loris gracilis 



Perodicticus potto 

Tarsius spectrum 

Chiromys madagascaricnsis . . . 

FcHs leo 







manx variety 











Felis caracal 


J 3 



II + 







Cynselums jubatus 















in + 

Genetta tigrina 


1 1 




Hemigalea hard wick i 






Paradoxurus musanga 
Arctictis binturong 














Eupleres goudoti 





Galidia elegans 






10+ ' 

Herpestes griseus 






Crossarchus obscurus 






Suricata tetradactyla 











Proteles cristatus 






* ,, .... 




26 + 

Hysena bruncea .... 









Lycaon pictus 






Icticyon venaticus . 






Canis javanicus 








I 7 




familiaris (Dingo) 







(mastiff) . . 





(German boarhound) ... 
(French bloodhound) ... 







(greyhound) . 





(retriever)... . 






(Skye terrier) . 












*Canis lupus 





vulpes . .. 


















r e 4- 

Otocyon megalotis 





Mustela martes . . 











16 + 






I c 












Mustela siberica 












1 : 












I r 


I ; 



Clulo luscus 


1 ; 



I c 

(ialictis vittala 






Mellivora indica 



I r 

Taxidea arnericana 


1 ; 



16 + 

Meles taxus . 









Mydaus mcliccps 





Q + 

Arctonyx collaris 




Mephitis mcphitica 






Lutra vulgaris 















* lalandi 




22 + 

Enhydra lutris 








10 + 

Procyon loior 






Nasua rufa 




2 \ 

10 + 

Bassaris astuta 


I ! 




Cercoleptes candivolvulus ... 
Ailurus fulgens 










<; 6 


10 + 














Q + 



left side 




7 > 


. S. 





















1 1 


Otaria jubata 









9 + 






. Thorac 

. Lunib. 



Otaria stelleri 






* ,, 



left sidi 



( s 





















1 1 

Trichechus rosmarus 




8 + 







Halichocru^ rypus 


I C 


Phoca vitulina 










I c 




I T. 

grcrnlandica . 




i e 

Stenorhynchus leptonyx 
Cystophora cristata 








Macrorhinus leoninus 




I i 

Bos taurus ... 






*Bos primigenius 





"Bubalus depressicornis 





* caffer 






Bison americanus . 





174- : 






i r 







*Ovibos moschatus. . . 


I : 



Ovis aries 





7 4- 



I 7 




1 1 


tragelaphus ... 




Capra ibex 








Boselaphus tragocamelus 
Gazella dorcas 





1 J 

Saiga tartarica 




Tetraceros quadricornis 
Connochaetes tmu... 






8 + 

Antilocapra americana 





VI. J 



Corvic. Thorac. 

GiraflFa camelopardalis 7 14 

7 4 

Moschus moschifcrus ... 7 14 

7 3 

Cervulus muntjac \ 7 13 

Ccrvus davidianus ! 7 13 

canadcnsis 7 13 

clcphus 7 13 

damn ] 7 13 

hibcrnicus 7 '3 

Rangifer tarandus 7 14 

.. 7 '4 

Alces machlis J 7 13 

Capreolus caprea 7 13 

Hydropotes inermis 7 12 

Cariacus mexicanus 7 13 

Pudua humilis 7 13 

Tragulus javanicus 7 13 

Camel us bactrianus 7 12 

dromedarius 7 12 

Auchenia glama 7 12 

7 '2 

pacus 7 I2 

vicugna 7 12 

Dicotyles tajacu . 7 14 

.. 7 H 

7 '4 

Sus cristatus 7 13 

scrofa 7 '4 

7 M 

Babirussa alfurus j 7 13 

Phacochrerus africanus 7 13 

Hippopotamus amphibius 7 15 

,, 7 15 

Tapirus indicus 7 18 

amcricanus 7 18 

Equus caballus 7 18 

7 18 

.. 7 ' '9 

7 19 

asinus 7 ' 

zebra ' 7 > 18 

Rhinoceros unicornis 7 '9 

sumatrensis 7 19 


SacraL Caudal. 























14 + 




10 + 



'5 + 
'3 + 
'3 + 

9 + 







u + 



'5 + 


21 + 








Sacral. Caudal. 

Rhinoceros sumatrensis 
Myrax capensis 





7 4 

Dendrohyrax arboreus 




6 7 + 





6 8 

Elephas indicus 




4. 3O + 








4. 24. 

t africanus . 




C 24 + 

Halicorc dugong 








20 + 

' ' 




Manatus anicricanus 





senegalensis ... . 




Baltena mysticetus ... 










Bala;noptera rostrata 


1 1 



Physeter macrocephalus 
Hyperoodon rostratus 








Mesoplodon grayi 





Berardius arnuxi 





Platanista gangetica... 





Monodon monoceros 


1 1 







Delphinapterus leucas 


1 1 



Phocrcna communis 








Orca gladiator 


1 1 


23 + 

Pseudorca crassidens . 









Globicephalus melas 





Tursiops tursio 








Sotalia sinensis 











Sacra] . Caudal. 

Lagenorhynchus acutus 























20 ! 35 + 

V - 













6 7 



16 + 





32 ; 

20 + 

20+ ' 
25 + 
20 + 

10 + 

5 + 

Delphinus delphis. 

Sciuropterus volucclla 

Sciurus vulgaris 


Tamias asiaticus 

Spermophilus mongolicus ... 
Cynomys ludovicianus 

Arctomys marmotta 

Castor canadensis 

Muscardinus avellanarius 
Hydromys chrysogaster 
Cricctus frumcntarius 

Mus rattus. ... .... 





Hapalotis albijxrs 

Fiber zibcthicus 

Avicola amphibius 


Myodes lemmus 

Ellobitis talpinus 

Spalax typhlus 

Khizomys pruinosu 1 * 

Bathycrgus maritimus 

Gcorychus capensis 

Geomys bursarius . 

Xapus hudsonicus . 

Dipus sagitta 

Alactoga jaculus 

Pedetes caffer 






i Thorac. 








2 5 





19 + 





2 5 

Capromys pilorides 





17 + 

Aulacodus swindernianus 






6 + 

Hystrix cristata 


















10 + 



21 + 

Lagostomus trichodactylus ... 




10 + 

Coelogenys paca 




7 + 

Cavia porccllus 





Dolichotis patachonica 




9 + 

Hydrochoerus capybara 
Lepus timidus 










variabilis ... 












Chrysochloris aurea 





Centetes ecaudatus 





* ,, 




8 + 

Hemicentetes semispinosus... 
Ericulus setosus 






Microgale longicaudata 
Solenodon cubanus 


I c 







Talpa europira 


I ^ 




Myogale mo.schata . . 


I ^ 




pyrenaica . . . 


I 3 




Crocidura cterulescens . . 





17 + 

Sorex vulsjaris . 





Erinaceus europasus 





I [ 


I c 

I I 




Gymnura raffles! 





Galeopithecus volans 










Thorac I.umb. 



Galeopithecus volans 













































17 + 












2.3 + 


24 + 





Epomophorus franqueti 
Pteropus ruhricollis 


Cynopterus marginatus 
Phyllorhina tridens 

diaclema . ... 


Megadcnna frons 

Vesperugo scrotinus 

noctula .. , 


Miniopterus schreibersi 
Molossus obscurus 

Desmodus rufus 

Bradypus tridactylus 

Cholcvpus didactylus 

* > 


Megatherium americanum .. 
Mylodon robustus 

Myrmecophaga jubata . ... 


Tamandua tetradactyla 




Cervic. \ Thorac. ' Lumb. 


Cycloturus didactylus 










left side 

































Tatusia novemcincta 


Dasypus sexcinctus . 

villosus . 


Xenurus unicinctus 

Priodon giganteus 

Tolypeutes conurus 

Chlamydophorus truncatus ... 
Manis pentadactyla 

aurita . . 




Orycteropus capensis 

Phascolomys wombat 


Phascolarctos cinereus 

Phalangista vulpina 


Petaurus australis 


Macropus giganteus 



Dendrolagus ursinus 

Hypsiprymnus murinus 
Bettongia gaimardi . . 


16 + 

16 + 


12 + 

12 + 




13 + 




8 + 



17 + 
15 + 
10 + 




Marsupial ia. 

Cervic Thorac. Lumb. 


Pcrameles ol>esula 7 13 6 

Myrtnecohius fasciatus 7 12 7 

Phascogale flavij>es 7 '3 ^ 

Dasyurus maculatus 7 13 6 

viverrinus 7 13 6 

ursinus 7 13 6 

Thylacinus c>nocephalus 7 13 6 

Didelphys virginiana 7 13 6 

a?ane 7 . 13 6 

Chironcctcs varicgatu-s 7 13 6 


Kchidna aculcata 7 16 

liruijni 7 16 4 

Ornithorhynchus anatinus ... 7 17 2 



10 + 

19 + 
13 + 
16 + 






THE Sternum of Mammals is a bone, or generally a series 
of bones, placed longitudinally in the middle line, on the 
inferior or ventral aspect of the thorax, and connected on 
each side with the vertebral column by a series of more or 
less ossified bars called ribs. 1 

It is present in all Mammals, but varies much in character 
in the different groups. 

When in its usual and most complete form (see Fig. 31), 
it may be divided into three parts, called respectively 

1. Presternum, or " manubrium sterni " of human ana- 

2. Mesosternum, body of the sternum or gladiolus. 

3. Xiphisternum^ xiphoid or ensiform process of the ster- 

The mesosternum is usually composed of several distinct 
segments, which may become ankylosed together, but more 
often permanently retain their individuality, being con 
nected either by fibrous tissue or by synovial joints. 

1 For much valuable information upon the structure and development 
of the sternum, see W. K. Parker's " Monograph on the Shoulder-girdle 
and Sternum of the Vertebrata," published by the Ray Society, 1868. 




The ribs are attached to the sides of the sternum : the 
first pair to the presternum ; the second to the presternum 
and the mesosternum at their point of junction ; the 
remainder to the mesosternum, opposite the interspaces 
between each segment, though two or more pairs are often 
clustered round the last segment. The xiphisternum bears 
no ribs. 

Development. The osseous sternum is preceded by a 
continuous or non-segmented piece of cartilage ; and as the 


FIG. 31 Human sternum and sternal ribs, J. /i presternum ; mt mesosternum ; 
jet xiphisternum ; c point of attachment of clavicle ; i to to the cartilaginous 
-ternal rids. 

portion of the body in which this is developed is formed by 
the union in the middle line of the two " ventral laminae " of 
the embyro, traces of this original median division are generally 
seen in very young sterna, and are often persistent through 


life in the form of fissures or fenestrse in the middle line 
of the sternum. Each segment ossifies from a single nucleus, 
or from two nuclei placed one on each side of the middle 
line, and which usually become blended together in the 
course of growth. Sometimes epiphyses are added to the 
ends of the segments. The terminal portion of the 
xiphisternum generally remains cartilaginous through life. 

Special Characters of the Sternum in the various Orders. 

Order PRIMATES. In Man (see Fig. 31,) the presternum is 
broad and flat, hollowed in the middle line in front, and 
expanded laterally to give large surfaces for the attachment 
of the clavicles and the first pair of ribs. The mesosternum 
is elongated, but is also comparatively broad and flattened. 
It consists of four distinct segments. The xiphisternum is a 
more or less elongated posterior appendage, varying some- 
what in form and size in different individuals. 

The ossification of the human sternum is endosteal, or 
commencing within the substance of the primitive hyaline 
cartilage. The presternum ossifies from one, or sometimes 
two, centres, which may be placed side by side, or one in 
front of the other. Each of the segments of the mesosternum 
has a distinct centre, though these may be double in their 
earliest condition, and sometimes remain so for a long 

The segments of the mesosternum usually unite together 
so as to form one continuous bony piece, to which the pre- 
sternum often remains throughout life connected only by 
fibrous tissue, although it is not unfrequently ankylosed in 
old age. 

The xiphisternum ossifies irregularly and imperfectly. 

The Gorilla, Chimpanzee, Orang, and Gibbons, resemble 
Man, and differ from the other monkeys in the breadth 




and flatness of the sternum. It is broadest in proportion 
to its length in the Siamang (Hylobates syndactylus). In 
the Orang (Fig 32), each segment of the mesosternum 
is developed from a pair of lateral ossifications, which 
commonly remain separate until the animal is about half 
grown. 1 

In the lower Monkeys, the presternum is somewhat broad 
but the bones constituting the mesosternum are elongated 
and compressed, and are not ankylosed together as in Man 
and the highest Apes. Their number varies from three to 
five. In the Howling Monkey 
(Mycetcs) the original two halves of 
the presternum are separated by a 
broad fissure, and form a pair of 
diverging, horn-like, bifurcated pro- 
cessess (pro-ostta, Parker), which 
ossify separately, and support the 
clavicles and first pair of ribs. Ac- 
cording to Albrecht - this condition 
of the manubrium is in relation 
with the enormously developed thy- 
roid cartilage in the male Mycttes, and 
is therefore a secondarily acquired 

In the CARNIVORA, the sternum 
(Fig. 33) is generally composed of eight or nine pieces 
altogether, including the presternum and the xiphisternum. 

The presternum, or manubrium, is long and narrow, 
somewhat expanded near the front for the attachment of the 

1 The sternum of a young Gorilla in the Museum of the College of 
Surgeons presents the same condition. 

- Su> la Elbnens Alorpholo&iquts du Manubrium Sterni. (Brussels, 


FIG. 33. Sternum of young 
Orang (Sitnia latyrtts). fs 
presternum ; mt meso*ter- 
nuiii ; jcs xiphisternum. 




Parker, ectosteally. 

first pair of sternal ribs, and terminating anteriorly in a 
conical rounded projection. 

The segments of the mesosternum are elongated, and 
more or less four-sided, contracted at the middle, and 
widening at each extremity. They ossify, according to 
or from without inwards, the bony 
deposit commencing in the inner layer 
of the perichondrium, as in the shafts 
of long bones ; and they remain per- 
manently distinct from each other. 

The xiphisternum is long, narrow, 
and flat, and generally ends in an ex- 
panded flattened cartilage. 

In the Pinnipedia, the presternum 
is produced considerably in front of 
the attachment of the first pair of 

In the INSECTIVORA, the sternum 
is variable in form, but always more 
or less elongated and segmented. 
The presternum is always more or 
less expanded laterally in this clavi- 
culated group of aninials. It is bilo- 
FIG. 33. sternum and sternal bate in front in the Hedgehog 

ribs of Dog (Ca.nisfa.tni- , . \ ^ -i i ., ^i 01 

Han's), if. /.? presternum ; (fcrinaceits), trilobate in the bhrews 

jus mesosternum ; .rs xiphi- /r , \ T /-> , , v i j 

sternum. (Sorex). In Rliynchocyon it is broad 

in front, narrow posteriorly, strongly 

keeled below, and with two horn-like processes projecting 
outwards and forwards between the attachment of the 
clavicles and the first pair of ribs. 

The mesosternum is usually narrow, as in the Carnivora, 
but in the Hedgehog, where it consists of three segments, 
it is broad and flat posteriorly, and to the last segment three 




ribs are attached. In this genus the xiphisternum is rudi- 
mentary, whereas in the Shrews it is long and ends in a flat 
expanded cartilage. 

The Mole (Talpd) and its nearest allies have a remark- 
ably developed presternum, which is as long as the whole 
of the mesosternum (Fig. 34). It is strongly keeled below, 
except at the front part, which is much thickened. On its 


KIG. 34. Sternum and siernal rib< of the Common Mole (Talfa eurofaa)v\lh the 
clavicles (cl) and humeri (//). Nat. size. 

superior or inner surface it is grooved in the middle line. 
Laterally it gives off a pair of wing-like processes, behind 
which the first ribs are attached. It is distinctly separated 
from the mesosternum, which consists of five segments of 
nearly equal width. The xiphisternum has a broad oval 
cartilaginous expansion posteriorly. 


As the clavicle is supported at the anterior extremity of 
the elongated presternum, it is widely separated from the 
first rib, and the anterior extremities are brought into such 
close juxtaposition with the head that the animal appears to 
have no neck. 

In the CHIROPTERA, the sternum presents a considerable 
general resemblance to that of Man. The presternum is 
large, trilobate in front, and strongly keeled. The meso- 
sternum is likewise frequently strongly keeled (Pferop2ts, 
Cynonycteris), but this keel is separated from that of the 
presternum by a deep notch. In many of the Insectivorous 
Bats the segments of the mesosternum are (at least in adults) 
firmly ankylosed together, but in some of the frugivorous 
Bats (Pteropus, &c.) they continue separate. 

In the RODKNTIA, the sternum is long and narrow, consist- 
ing of a presternum (which is generally broad in the forms 
which have the clavicle well developed, as the Rats, Beavers, 
&c.), a mesosternum of three, or more usually four, seg- 
ments, and a long xiphisternum, with a broad cartilaginous 
terminal expansion. The segments of the mesosternum 
often have epiphyses at each end. 

The presternum is compressed and produced forwards in 
those species in which the clavicle is absent or rudimentary, 
as the Aguti, the Hares, and the Capybara. In the latter it 
much resembles that of the Horse or Tapir. 

Order UNGULATA. In the Ruminantia there are usually 
seven segments altogether in the sternum (Fig. 35). The 
presternum is narrow, rounded in front, and bearing the first 
pair of sternal ribs close to its apex. The succeeding pieces 
gradually widen, the posterior segments of the mesosternum 
being square, flat, and rather massive (especially in the 
Giraffe) ; they are hollowed at the middle of their lateral 
borders. The xiphisternum is thin and flat. 




In the Pig (Fig. 36) and Hippopotamus the presternum is 
( ompressed and keeled ; the articular facets for the first 
pair of ribs are close together on its upper surface ; but the 
mesosternum is broad and flat, the first segment being 
transitional, compressed in front, and broad posteriorly. 

FIG. 35. Sternum and sternal ribs of 
Red Deer (Cftvn t elaf-hut), \. fs 


presternum ; mi 



KH;. 36 Sternum of the Pig (S*i 
scro/a), \. fit presternum ; tin meso- 
sternum ; .rt xii>histcmum. 

The xiphisternum is narrow and pointed. The sternum of 
the Pig very often retains indications of the primordial 
median fissure through life. 

The Horse and the Tapir have a very peculiar sternum. 



The presternum is extremely compressed and projects 
forward like the prow of a boat. In the Tapir, its anterior 
portion is originally, and commonly remains, a distinct 
ossification (pro-ostcon, Parker). The segments which 
follow gradually widen, and the hinder part of the sternum 
is broad and flat. The last mesosternal segment in the 
Tapir is generally divided in the middle line. 

The sternum of the Rhinoceros is very narrow through- 
out, with a long, rather spatulate xiphisternum. 

Order CKTACF.A. Each of the two primary divisions of 
this order has a distinct form of sternum. 

Among the Odontcceii, the typical Dolphins have a very- 
broad presternum of peculiar form, emarginate in the middle 
line in front, and with a pair of lateral processes behind the 
attachment of the first pair of ribs. This is followed by two 
or three mesosternal segments but no xiphisternum. An 
indication of the primordial median fissure can generally be 
traced, except in very old animals, either as a hole in the 
presternum, or as a division of the posterior mesosternal 

In the Porpoise (P/iocfcna) the sternum is shorter and 
broader than in most Dolphins, and its various elements 
early coalesce into a single bone. 

In the Cachalot (Physeter macrocephalus) the sternum 
ossifies from three distinct pairs of nuclei, and a large 
median fontanelle remains between the first and second 
pair. 1 In the specimen in the Museum of the Royal College 
of Surgeons (Fig. 37), which is very nearly adult, each half 
of the presternum (ps) has coalesced with the corresponding 
half of the first segment of the mesosternum (;//-$'), but the 
resulting pieces are not united by bone across the middle 
line, while the second or last pair of mesosternal segments 
1 I have observed this in animals evidently of great age. 




(ms-) are ankylosed together mesially, but not with the 
portion of the sternum in front of them. 

FIG. 37. Sternum of Cachalot or Sperm Whale (fkyseter tnacrorefkalux), fa. 

In the Whalebone Whales (Afystacoceti} the sternum is 
comparatively rudimentary, consisting only of a broad, 

FlC. 38. Sternum of Greenland Right Whale (Baftma myslicetut), /,. 

flattened presternum, produced posteriorly into a xiphoid 
process in some species. There are never any mesosternal 

H 2 



segments, and consequently no ribs, other than the first 
pair, are attached to it. 

The presternum is ossified from one, or perhaps a pair of 
symmetrical nuclei. In the Right Whales (Balana, Fig. 38) 
it is heart-shaped, or longitudinally oval; in the Fin Whales 
(Batenoptera, Figs. 39, 40, and 41) it is transversely oval 
or trilobate, with a backward projecting xiphoid process. 1 
In young animals ossification of the cartilaginous sternum 
advances forwards on each side of the middle line, so that 

FIG. 39. Sternum of Common Rorqual or Fin FIG. 40. Sternum of Pike Win 
Whale (Bal&noptera musculus), ,V (Balifiwptera rostrata), ,*,. 

the ossified portion at one period appears deeply notched in 
front ; as the bone meets across the middle line anteriorly, 
this notch usually becomes converted into a hole (see 
Fig. 39), which finally closes with complete maturity. 

1 In the cartilaginous sternum of a young F!al<cnoptcra sibbaLiii 
Professor Turner found the xiphisternum to be quite distinct from the 
presternum, and connected with it by fibrous tissue. (Journal of 
Anatomy, May 1870.) In most Whales the sternum shows no such 
evidence of segmentation. 




In the Lesser Fin Whale (/?. rostratd) the sternum is 
cross-shaped (Fig. 40), the first ribs being attached behind 
the Literal arms of the cross. 

FlG. 41 Sternum of Rudolphi's Whale (RaJtrnafltra A.'rra/is). showing its relation 
l'i the inferior extremities uf the first pair of ribs. From " Proceedings Zoological 
Society. 1883." 

In the SIKKNIA the sternum is a simple, flattened, some- 
what elongated bone, which in the adult shows no trace of 

FIG. 42. Sternum of a young Dugong (Halicore indicia), \. From a specimen in 
the Leydcn Museum. 

In a young Dugong (Ilalicore), Fig. 42, there are two 
distinct ossifications, a presternum (/j), to which the first 


pair of ribs are attached, and a xiphisternum (xs). The 
second, third, and fourth pairs of sternal ribs are attached 
to the intermediate unossified portion, representing a rudi- 
mentary mesosternum. 

In the Manatee (Manatus) the sternum is of somewhat 
similar form, and has three pairs of ribs attached to its 
lateral margins near the middle. 

Among the EDENTATA there is considerable variation in 
the characters of the sternum. 

In the Cape Anteater (Orycteropus) the sternum is of 
quite a normal form. The presternum is trefoil-shaped, 
expanding laterally near the front to meet the largely 
developed clavicles, then contracting to the width of the 
mesosternal segments, which are four in number, simple, 
flattened, oblong, with lateral margins nearly parallel, 
rather broader above than below, united together by fibrous 
tissue, and succeeded posteriorly by a moderately developed 

In Manis aurita the sternum is flat, consisting of seven 
segments, several of which are sometimes divided in the 
middle line by synovial cavities. The xiphisternum is 
very long, partially cleft in the middle line, and ending in 
a large flattened cartilaginous expansion. In the Long- 
tailed Pangolin (Manis macrura] the xiphisternum is of a 
remarkable form, being prolonged into a pair of cartila- 
ginous processes, each about nine inches long, and con- 
nected posteriorly with some rudimentary abdominal ribs. ' 

In the Anteaters (Myrmecophaga) the presternum is broad, 
flat, and oval. The segments of the mesosternum (Fig. 43) 
are eight in number, short, deep, broad above, and sending 
a club-shaped process downwards ; each is ossified from a 
principal endosteal centre and eight epiphyses, is connected 

1 Parker, op. fit. p. 208. 

vii.] EDENTATA. 103 

by synovial articulations with the segment before and behind, 
and has at either end an upper and lower hollowed surface, 
which, with the corresponding surfaces on the contiguous 
segment, form articulating facets for the double-headed 
sternal ribs. This mode of articulation curiously resembles 
that at the vertebral end of the rib. The xiphisternum is 
rather long and simple. 

In the small Tree Anteater (Cycloturus didactylus) the 
presternum is very broad and trilobate, sending out lateral 
expansions behind the attachment of the clavicles to meet 


FIG. 43 Side view of three mesosternal segment* of a young Anteater (Myrnte- 
cofHaga t<iman<iua\ showing the mode of articulation of the sternal rib (jr). 
copied from Parker's figure, nisi thi upper or inner surface of the mesostcrnal 
segment ; ty the synovial articulation between the segments. 

the first pair of ribs. The hinder, narrow part of the manu- 
brium is segmented off from the larger anterior part, and 
resembles a mesosternal segment ; but it is in front of the 
attachment of the second pair of ribs. The true mesosternal 
segments are six in number, of nearly equal width, high, 
rounded above, and con-. pressed below, with a synovial 
cavity between each. The sternal ribs are articulated by a 
single oval condyle. The xiphisternum is long, stout, and 


In the Armadillos (Dasypodidce) the presternum is broad, 
and in Priodon gigas (Fig. 45, p. 114) strongly keeled. The 
mesosternal segments, four to six in number, are broad above, 
but very narrow below. They are connected by synovial 
joints to each other, and to the strongly ossified sternal ribs, 
which have broad, sub-bifid heads. The xiphisternum 
expands posteriorly into a wide cartilaginous flap. 

In the Sloths the sternum is long and narrow. The Three- 
toed species {Bradypus} have a rather broad presternum, but 
with no prolongation in front of the attachment of the first 
rib. This is followed by eight small mesosternal segments, 
and a very small rounded xiphisternum. In the Two-toed 
Sloths (Cholazpus) the presternum is narrow, slightly keeled, 
and forms a considerable projection in front of the attach- 
ment of the first rib. The mesosternum has twelve segments, 
and the xiphisternum is rudimentary or absent. 

In the MARSUPIALIA the sternum presents no especial 
aberrant characteristics. The presternum is rather broad at 
the point of attachment of the first pair of ribs. Its anterior 
extremity often does not ossify. There are usually four 
quite distinct, elongated segments to the mesosternum, con- 
nected to one another by fibrous tissue, and sometimes 
completed at each end by epiphyses. The xiphisternum 
has an elongated, narrow, ossified portion, and terminates 
in a laterally expanded cartilage, which may contain one 
or two endosteal bony patches. 

In the MONOTREMATA the Ornithorhynchus (Fig. 44) has 
a broad presternum (ps\ with a small partially-ossified 
pro-osteon (po) in front of it ; three keeled mesosternal 
segments (;;), which commence to ossify in pairs, and 
no xiphisternum. The Echidna agrees in all important 
respects, but it has an ossified xiphisternum, which in E. 
Bruijni consists of three metameric portions. 



The T-shaped bone, interclavicle or episternum. (if) in front 
of the presternum, which connects it with the clavicle 
and is often completely fused with it, appears to have 

FIG. 44. Sternum and adjacent parts of the skeleton of a young Ornithorhynchus 
(l). anatini4i) ( clavicle ; if interclavicle ; fx> pro osteon (a part of the true 
kternum) ; fs prcstcrnum ; nis mcsostcrnum ; sr sternal ribs ; ir intermediate ribs ; 
vr vertebral nbs. 

no homologue among the other Mammalia, and belongs 
more properly to the shoulder-girdle than to the sternal 



THE ribs form a series of long, narrow, and more or less 
flattened bones, extending laterally from the sides of the 
vertebral column, curving downwards towards the median 
line of the body below, and mostly joining the sides of 
the sternum. 

Free ribs are normally only attached to the thoracic ver- 
tebras, although, as before shown, certain parts, which may 
be serially homologous with ribs, are found in other regions 
of the vertebral column ; but in such cases they become, 
almost always, ankylosed with their corresponding vertebras. 
In the thoracic region, the ribs are never normally ankylosed 
with the vertebrae, but are articulated to them by synovial 
joints, which permit a certain, though limited, amount of 

As a general rule, the first thoracic rib joins the pre- 
sternum or manubrium ; sometimes, as in the Whalebone 
Whales, this is the only rib united to the sternum ; but 
usually a larger number are so connected, while the more 
posterior are either attached by their extremities to the 
edges of the ribs in front of them, and thus indirectly join 
the sternum, or else they are quite free below, meeting no 


part of the skeleton. These differences have given rise to 
the division into true ribs and false ribs (by no means good 
expressions), signifying those that join the sternum directly 
and those that do not ; and of the latter, those that are 
free below are called floating ribs. 

Each primary piece of cartilage, out of which one of the 
half hoops or ribs is developed, is, moreover, divided trans- 
versely into two portions, which assume different characters, 
as they usually undergo a different mode of ossification, and 
remain more or less distinguishable from each other during 
life. The portion nearest the vertebral column is called 
the vertebral rib. This is the larger segment, and becomes 
firmly ossified at an early period by ectostosis ; l it is the 
bone commonly spoken of as a "rib." 

The portion towards the sternal extremity or sternal rib 
is usually imperfectly ossified, and always at first (except 
in Monotremes) by cndostosis? Sometimes it remains per- 
manently in a cartilaginous state ; but, on the other hand, in 
some cases (as in Armadillos and Sloths) it becomes as firmly 
ossified as the vertebral ribs. 

The vertebral ribs are variously connected with the sternal : 
by continuous cartilage, by intercalation of fibrous tracts, 
or by synovial joints. 

Occasionally, in the Mammalia, an " intermediate " 
portion of the rib is segmented off, as in Reptiles ; this is 
best developed in the Monotremata (Fig. 44, />), where, 
however, it is only partially ossified by cndostosis. In all 
other instances in which it occurs (e.g., Odontoccti} it is 
quite rudimentary. 

1 The bony deposit commencing at the surface, and advancing in- 
wards, as in ordinary long bones. 

' J The bony deposit being irregularly scattered throughout the carti- 
lage, often beginning near its central part. 


The vertebral ribs, when in their most typical condition, 
have two points of attachment to the vertebrae : the tubercle 
(tubcrcului)i) and the head (capituluiri). The former is 
superior or dorsal and posterior, and attached to the trans- 
verse process of the vertebra ; the latter, inferior or ventral 
and anterior, and attached to the body of the vertebra, or 
the inferior part of the arch near the body, and always very 
near the neurocentral suture. Commonly, in fact, the 
articular surface is cut by this suture. Sometimes, as in Man, 
the greater part of the articulation is above the suture; 
or, on the other hand, it may be, as in the Monotremes, 
below the suture. The distinction between the two points 
of attachment is most marked in the anterior ribs ; in 
passing backwards they approach nearer to each other, 
sometimes becoming blended, or sometimes either one or 
the other (generally the tubercular) attachment is lost in the 
hindermost ribs. 

The tubercle articulates, by a nearly flat or slightly con- 
vex surface, to a facet on the under-surface of the extremity 
of the transverse process of the corresponding vertebra, but 
the more rounded capitulum (at least in the anterior ribs) is 
placed opposite to the intervertebral space in front of this 
vertebra, and portions of two vertebra commonly contribute 
to form the articular cavity for its reception. Thus the first 
rib is articulated by its tubercle to the transverse process of 
the first thoracic vertebra, and by its head to the hinder 
part of the seventh cervical and front part of the first 
thoracic vertebra, and so on. The posterior ribs, as a rule, 
are connected solely with their own corresponding vertebra 

The amount of motion permitted by these articulations 
is sufficient to allow the thorax to expand and contract in 
respiration. In inspiration the ribs are drawn forwards, and 
approach nearer to a right angle with the vertebral column, 


while in expiration they fall back, and occupy a more 
oblique position to the axis of the column. 

The sternal ribs are connected with the sternum either by 
interposed fibrous tissue or by distinct synovial joints. The 
first is attached to the side of the presternum, the second 
opposite to the junction of the presternum and the first 
mesosternal segment, and the succeeding ones opposite to 
the interspaces between the other mesosternal segments ; 
though two or more may be attached to the hinder end of 
the last of these segments. The inferior ends of the so- 
called " false ribs " are attached by fibrous tissue, or by 
synovial joints, to the hinder borders of the sternal ribs in 
front ; though, as before said, the most posterior are free or 
" floating." 

The ribs of existing Mammals never have "uncinate 
processes," like those found in Birds and Reptiles. 

The most prevalent number is thirteen pairs ; the lowest 
is nine (in Hyptroodoii), the highest twenty-four (in the Two- 
toed Sloth, Cholocpus). 

Special Characters of the Ribs in the various Groups rf 

Order PRIMATES. In man there are normally twelve 
pairs of ribs, of which the first seven are reckoned as true 
ribs and the last two as floating ribs. The last pair may 
be rudimentary or absent ; or, on the other hand, the seventh 
cervical or the first lumbar vertebra may have an additional 
movable rib articulated with it. 

The first vertebral rib is much shorter, broader, flatter, 
and more curved than the others. These gradually increase 
in length until the seventh, after which they again diminish 
to the twelfth. In breadth they gradually decrease from the 
first to the last. 


The portion of the rib between the head and the tubercle 
is called the neck ; it is wanting in the last two ribs, in which 
the two attachments are blended. The greatest point of curva- 
ture on the external surface of the rib is called the angle. 

Each vertebral rib has a main centre of ossification and 
two epiphyses, one for the head and (except in the last 
two) one for the tubercle. 

The sternal ribs generally remain cartilaginous throughout 
life, being only partially ossified by endostosis in old age or 
under abnormal conditions. They are not distinctly sepa- 
rated from the vertebral ribs except by their difference of 
structure ; but synovial joints are (except in the first) inter- 
posed between their inferior extremities and the sternum. 

Among the higher Simiina the ribs do not differ very 
notably from those of Man, except in number; but in the 
lower forms, and especially in the Lemitrina, they more 
resemble those of the Carnivora. Among the Old World 
Monkeys the number varies from n to 13 pairs. The 
Gorilla, Chimpanzee (Troglodytes} and Gibbon (Ifylobates) 
have 13, and the Orang (Simia] 12. In the American 
Monkeys there are from 12 to 15 pairs; in the Lemurs 
from 12 to 17 pairs. 

In the most typical forms of CARNIVORA the vertebral 
ribs are comparatively slender, subcylindrical, and little 
curved. The most anterior espjcially are short and straight, 
the thorax being thus more compressed in front than it is in 
Man and the higher Primates. The sternal ribs (see Fig. 
33, p. 94) are long, slender, have a feeble granular ossifica- 
tion, and are not otherwise segmented off from the vertebral 
ribs. In all the Fdidcc and Canidce- there are 13 pairs, in 
the Viverridcc 13 or 14, in the Hyienidtc 14 or 15, in the 
Mustdidic 14 to 1 6, in the Procyonid<z 14, in the Ursidtc 14 
or 15, in the Pinnipcdia 14 or 15, 


In the UNGULATA the ribs are generally more or less 
flattened and broad, notably so in the Ox and Camel, and 
least so in the Perissodactyla. The anterior ribs have 
scarcely any curve, the thorax being very narrow in this 
region. The sternal ribs (see Fig. 35, p. 97), especially 
those near the front of the series, are short, stout, rather 
flattened or prismatic, tolerably well ossified, and articulated 
with the vertebral ribs by a cup and-ball synovial joint. 
The Artiodactyles have from 12 to 15 pairs of ribs, the 
Horse and Tapir 18, the Rhinoceros 19, the Elephant 19 
or 20, and the Hyrax 20 to 22. 

In the SIRKNIA the total number of ribs is very great, 
though but tew are attached to the sternum. In the Manatee 
they acquire an extraordinary thickness and SDlidity of 
texture. This animal has 17 pairs, of which but three are 
attached by flexible cartilages to the sternum. 

Order CETACEA. In the Whalebone Whales the ribs differ 
greatly from those of the rest of the Mammalia in their ex- 
tremely loose connection, both with the vertebral column 
above and with the sternum below, probably to allow of greater 
alteration in the capacity of the thorax in respiration neces- 
sitated by the prolonged immersion beneath the surface of 
the water which these animals undergo. 

At their vertebral extremities they are attached only by 
their tubercle to near the end of the transverse process, but 
apparently not by synovial articulation. The heads of only 
a few of the anterior ribs are developed, and are rarely 
sufficiently long to reach the bodies of the vertebrae, their 
place being supplied by a ligamentous band. The first rib 
is the only one connected with the sternum, either directly 
or indirectly, the whole of the remainder being free or 
floating ribs. The sternal ribs are mere cartilaginous rudi- 
ments, connected by an intermediate layer of fibrous tissue 


to the inferior extremity of the vertebral rib ; at least such 
is their condition in the foetus of Balcena mysticetus, as 
described by Eschricht and Reinhardt. 

Balanoptera rostrata, the smallest of the Whalebone 
Whales, has but n pairs of ribs, Megaptera longimana 14 
pairs, the Greenland Right Whale (Balcena mysticetus) 
usually 13, and the larger Fin Whales (Balcenoptera musculns 
and sibbaldt) 15, and occasionally 16, the highest number 
known in any Cetacean. In these last it not unfrequently 
happens that the hindermost rib, having only the middle or 
lower portion developed, is separated by a wide interval 
from the vertebral column a very rare condition, as in most 
other cases where the hinder ribs are rudimentary the part 
in immediate connection with the vertebra remains. 

The first rib presents a very anomalous condition in some 
Whalebone Whales, being apparently double, probably owing 
to the coalescence of a supplemental cervical rib with the 
ordinary first thoracic rib. In some species (as Bahcnoptera 
borealis] this appears to be of almost constant occurrence ; in 
others it is occasional. 1 

In the Odontoceti or Toothed-whales, as the Common Dol 
phiri and Porpoise, the ribs (usually 12 or 13 pairs) are long 
and slender. The first four or five have tubercles, by which 
they articulate with the transverse process of the thoracic 
vertebrae, and long necks and heads, reaching to the side of 
the antecedent vertebra, rear the junction of the body and 
the arch (see Fig. 20, p. 59). The posterior ribs, however, 
lose the neck, and are solely articulated by the tubercle to 
the transverse process. There are usually 7 pairs of rather 
short, straight, but strongly ossified sternal ribs nnd often 
small intermediate ribs, sometimes distinctly ossified. 

1 See Professor Turner " On the so-called Two-headed Ribs in Whales 
and in Man." (Journal of Anatomy and Physiology, May 1871.) 

v in. ] CE TA CEA. EDENTA TA. 1 1 ; 

In the aberrant Physfteiidte y including tlie Sperm Whale, 
Hyperoodon, and various forms of Ziphioids, the ribs are 
connected to the vertebrae throughout the greater part of the 
series by both head and tubercle; but a few of the most 
posterior have only a single point of attachment, in con- 
sequence of the changes which take place in the condition 
ot the transverse processes of the vertebra.', described at 
p. 59. In this family the sternal ribs are either j>ermanently 
cartilaginous, or very imperfectly ossified. The Hyperoodon 
has but 9 pairs of vertebral ribs, the smallest number 
known in any Mammal; the Sperm Whale (Physder) u, 
of which the last is quite rudimentary; Ziphins, 10, and 
A'o;/V7, 14. 

Among the EDENTATA the Sloths have very numerous 
ribs (from 15 to 24 pairs, the latter number in Choltcfnis 
didactvhis, of which 12 are true ribs). In the anterior part of 
the thorax the sternal ribs are firmly ossified, and indistin- 
guishable from the vertebral ribs (at least in adult age), 
but posteriorly they are separated from the latter by a less 
perfectly ossified intermediate rib. 

In the Armadillos (Fig. 45) the ribs are comparatively few 
(10 to 12 pairs), and are broad and flat, the first extremely so. 
The first sternal rib is very short and incorporated with the 
vertebral rib, but the others are very strongly ossified, and 
articulated by synovial joints with the sternum, with each 
other, and with the vertebral ribs. 

The peculiar double articulation of the sternal ribs with 
the sternum in the Anteater (Afyrmteophaga) has been already 
described (see p. 102). The ribs of the small climbing Two- 
toed Anteater (Cydoturus dideutylus) are remarkable for a 
thin lamelliform expansion of their hinder border, over- 
lapping the succeeding rib. This animal has 15 pairs, the 



great Anteater 16, the Cape Anteater (Orycteropus) 13, of 
which the last is sometimes very short. 

The MARSUPIALIA have nearly always 13 pairs of ribs : 
the Koala (Phascolarctos) with but n, and the common 
Wombat (Phascolomys vombatus] with 15, being the only 
known exceptions. The sternal ribs are articulated by 
synovial joints with the sternum, but are not distinctly 
segmented from the vertebral ribs, and are but feebly 
ossified bv endostosis. There are no intermediate ribs. 

Fir,. 45. Sternum nnd ribs nf the Great Armadillo (Priodon f>igas), \. 
ps presternum ; xs xiphisternum. 

In the MONOTREMATA the intermediate ribs are well 
marked (see Fig. 44, p. 105), and only partly ossified by 
endostosis, while the sternal ribs (except the first) are, 
according to Parker, strongly ossified ectosteally, as in Birds. 

viii.] MONOTREMATA. 115 

The hinder sternal ribs are very broad and flat. The 
Echidna has 16, and the Ornithorhynchus 17 pairs of 
vertebral ribs ; they do not divide above into head and 
tubercle, but are attached only to the sides of the bodies 
of the vertebra;. The last 5 sternal ribs of Echidna, especi- 
ally E. bruijni, are exceedingly broad and overlap each 

I 2 



THE skull is the term commonly applied to the expanded 
anterior portion of the axial skeleton situated within the 

It consists mainly of the cranium, a strong bony case or 
frame, enclosing the brain, and affording support and pro- 
tection to the organs of smell, sight, hearing, and taste, and 
formed by the close union, either by sutures or by synostosis< 
of numerous bones. 

To the inferior surface of the cranium are suspended (i) 
the Mandible, or lower jaw, movably articulated by a syno- 
vial joint; and (2) a group of skeletal structures called the 
hyoidean apparatus. 

The diagram at p. 126 is intended to show at a single view 
the names applied to the various bones of which the skull is 
composed, and to give some idea of their relative position. 

It will be well to commence the study of the skull by 
describing that of a Dog, as a good average specimen of the 
class, and one which is easily procurable at various ages : 
and the student is strongly advised to follow the descrip- 
tion with a skull in his hand, or better still, two, in one of 
which a longitudinal median section has been made. In 
the other, the various bones should be separated from each 




other. For this purpose a young animal, still retaining the 
milk teeth, will be best. 1 

The skull has a longitudinal central axis (the cranio-facial 
axis, Huxley), around which all its parts are arranged, and 



Km. 46 longitudinal and vertical section of the skull of a Do({ (Canit/amiliarii), 
with mandible ami hyoid arch, J. an anterior narial aperture; MT maxillo- bone; KT ethmo-turbinal ; A'<* nasal; ME ossified portion of the 
mesethmoid ; CK cribriform plate of the clhmo-turbinal ; /> frontal; Pa parietal ; 
//' interparietal : SO supraoccipital ; / jcO exoccipital ; BO basioccipital : I'tr 
periotic ; ItS basisphenoid ; /'/ ptcrygoid ; AS alisphenoid ; OS orbitosphenoid ; 
PS presphenoid : /'/ palatine ; I'o vomer ; MX maxilla ; PMx premaxilla : 
sk stylohyal ; tk epihyal ; ch ceratohyal ; bk basihyal ; tk thyrohyal ; s symphysis 
of mandible ; c/ coronoid process ; <-</condyle ; a angle ; id infenor dental canal ; 
the mandible is displaced downwards to show its entire form ; the * indicates the 
part of the cranium to which the condyle is articulated. 

its structure will be best understood by commencing with 
the description of the bones forming this axis. 

1 When the zoologist wishes to throw into the strongest relief the 
distinctive characters of different species, he selects for comparison fully 
adult examples ; when the anatomist wishes to trace their community of 
structure and their resemblances, younger specimens are better adapted 
for his purpose. 


When the skull remains in connection with the vertebral 
column, it will be seen that its axis is a continuation for- 
wards of the axis of that column, consisting of the bodies of 
the vertebrae ; and that its hinder termination is placed in 
the same line with the odontoid process of the second cer- 
vical vertebra, the anterior termination of the axis of the 
spinal column. 

The large cavity above the axis of the skull (cerebral 
cavity) is in direct continuity with the spinal canal above 
the axis of the vertebral column. 

Beginning at the posterior end of the axis, the section 
will be seen to have passed through a flat, elongated bone 
(Fig. 46, BO\ terminating freely behind at the inferior 
margin of the great opening (foramen magnum} at the hinder 
end of the cerebral cavity, by which this cavity is continued 
into the vertebral canal, and through which the backward 
prolongation of the brain (the medulla spinalis) passes. This 
bone is the basiocdpital. 

Immediately in front of this is a bone (the basisphenoid, 
BS] not quite so elongated from before backwards, but of 
greater vertical depth ; the interior being more or less 
cellular in structure. The under surface is flat, but the 
upper surface is hollowed in the middle, and raised at each 
extremity. This hollow corresponds to the part called 
" sella turtica " in the human skull, and lodges the pituitary 
body of the brain. 

Further forwards, and likewise separated by a vertical 
fissure, is a bone (PS) of about the same length as the last, 
but still more elevated, and very cellular within. Its in- 
ferior contour is perfectly straight, but above it is somewhat 
irregular. This is the presphenoid. 

So far the cranio-facial axis consists of bones placed in 
a continuous line, more or less depressed, and broad from 

ix.] OF THE DOG. 119 

side to side, and forming the floor of the cranial cavity ; 
but the continuation of the axis forward is of a different 
character. The anterior end of the presphenoid narrows 
considerably, and the segment in front of it, in very young 
skulls, is a much compressed vertical plate of cartilage, of 
very considerable size, both from before backwards and 
from above downwards. Ossification of this cartilage com- 
mences in the posterior end and upper part, and spreads 
forwards and downwards, but it never or very rarely reaches 
its anterior extremity ; and in the animal now described a 
narrow inferior margin remains jxjrmanently cartilaginous. 
The ossified portion of this cartilage {ME} constitutes the 
lamina ferpendicularis of the ethmoid bone, the anterior 
unossified portion the septal cartilage of the nose, which is 
the anterior termination of the cranio-facial axis. The 
term mesethnwid may be applied to the whole of this element 
of the skull, whether ossified or not. 

Above all the posterior, or basicranial, part of this axis, 
constituted by the three first-mentioned bones, is the cere 
bral cavity, the walls of which constitute the " brain-case." 

These walls are formed by several more or less expanded 
and curved bones, which rise up from the sides of the axis 
or floor of the cavity below, and, meeting in the middle line, 
roof in the cavity above. These bones are arranged in 
three sets from behind forwards, each corresponding with 
one of the axial bones, and with the latter constituting one 
of the three segments or bony rings into which the brain- 
case may be divided. 

The hindermost (or occipital) segment consists of the 
basioccipital below ; next on each side the exocdpitah (EO), 
and a large, median, flat bone above, with its upper ex- 
tremity prolonged forwards in the middle line between the 
bones of the next segment, called the supraocripital (SO). 


These four bones surround the foramen magnum behind, and 
all take share in its circumference, though the exoccipitals, 
which bound it laterally, contribute most. On each side of 
the foramen, and rather below than above, are the occipital 
condyles, by which the skull articulates with the first cervical 
vertebra; and externally to these, separated by a deep de- 
pression, is a prominent process for muscular attachments, 
called the paroccipital (or paramastoid] process. The con- 
dyles in the Dog are formed by the exoccipitals alone. The 
part (IP] which appears to be an anterior prolongation of 
the upper extremity of the supraoccipital, wedged in be- 
tween the parietals, is ossified from a separate centre, and 
in some animals remains permanently as a distinct bone. It 
is then called intcrparietal. 

The middle (or parietal) segment is formed by the 
basisphenoid below. From the sides of this a pair of 
wing-like bones (AS) extend outwards and upwards, 
called alisphenoids ; and above these are large square- 
shaped bones (/h), meeting in the middle line above, the 

The occipital and the parietal segments are in contact 
below and above, but there would be a considerable open 
space between them laterally were it not for the inter- 
position of a group of bones, which do not form part of the 
segmented wall of the brain-case proper, but are more or 
less connected with the organ of hearing, and will therefore 
be described hereafter. These are the bones which, being 
all united into one in Man, constitute the so-called temporal 
bone of human anatomy. 

The anterior (or frontal) segment is formed by the pre- 
sphenoid below ; then by the wing-like (OS} projecting 
from its sides, smaller than those of the second segment, 
called orbito sphenoids ; and finally by two greatly expanded 

ix.] OF THE DOG. 121 

bones (Ft), curving inwards above and in front, so as to close 
in the cerebral cavity in these directions, by meeting in the 
middle line. These are the frontal bones. 

Between the middle bones of the parietal and frontal 
segments (alisphenoid and orbitosphenoicl) is an irregular 
vacuity, called the foramen lace rum a uteri us, or sphenoidal 
fissure, through which several nerves pass to the orbit. This 
is the second vacuity in the side wall of the skull, the first 
being the one between the occipital and parietal segment, 
partially filled by the periotic bone. 

As the occipital segment is not closed behind, so in the 
same way the frontal segment is open in front, the aperture 
being bounded by all the bones which enter into its com- 
position presphenoid, orbitosphenoids, and frontals. The 
hinder edge of the mesethmoid rising up to meet the 
frontals makes a median partition (the crista galli of human 
anatomy) to this aperture, which is further closed by a 
special ossification (CE) connected with the of smell, 
the cribriform />/<//<?. 

Thus the brain case may be described as a tube, dilated 
in the middle, composed of three bony rings or segments, 
with an aperture at each end, and a fissure or space at the 
sides between each of the segments. 

The cranial cavity thus formed is of a general oval form, 
but broader behind than in front. The floor is compara- 
tively straight ; the upper surface arched. It is imperfectly 
divided by bony ridges into three compartments. The 
most posterior of these, marked off in front by a sharp 
ridge along the periotic bone (Per), extending from near the 
junction of the basisphenoid and basioccipital, upwards, 
outwards, and backwards, along the line of junction of 
the parietal and supraoccipital, and strongly marked by an 
inward shelf-like projection from the former (the ossified 


tcntorium cerebelli], is called the cercbellar fossa, as it 
lodges the division of the brain named "cerebellum." The 
most anterior and smallest compartment is marked off by a 
vertical ridge on the orbitosphenoid and the frontal. Its 
walls are chiefly formed by the cribriform plate. This is the 
olfactory fossa (rliinencephalic fossa, Owen), for the lodgment 
of the olfactory lobe. Between these two is the great cere- 
bral fossa, in which the hemisphere of the cerebrum lies. 
This is very imperfectly divided below into two compart- 
ments, by a slight ridge at the hinder edge of the orbito- 
sphenoid and continued thence outwards at the junction of 
the frontal and alisphenoid. This ridge corresponds with 
the Sylvian fissure of the brain ; the part of the cerebral 
fossa in front of it lodges the frontal lobe of the cerebrum, 
that behind it the temporal lobe. 

Through the lateral parts of the floor of the cranial cavity 
are various perforations, or foramina, either holes passing 
directly through the bones, or vacuities occasioned by want 
of contact, for a limited space, of contiguous bones. These 
are mainly for the purpose of allowing of the exit of the 
various nerves which take origin from the brain ; and as 
they are extremely constant in their position, and offer useful 
landmarks for determining the homologies of the bones 
throughout the vertebrate series, it is important that they 
should be well known. (See diagram at p. 126.) 

1. The most anterior is the space, before spoken of, in 
front of the anterior segment, occupied by the hinder part 
of the ethmo-turbinal, commonly called the " cribriform 
plate." The numerous perforations in this plate transmit 
the olfactory nerves arising from the olfactory lobes. 

2. Near the hinder border of the orbitosphenoid is a con- 
spicuous, nearly round hole, through which the optic nerve 
passes, and hence called optic foramen. 

ix.] OF THE DOG. 123 

3. At a very short distance behind this is a more irregular 
oval opening, between the orbitosphenoid and the ali- 
sphenoid. This is the sphenoidal or orbital fissure, or 
foramen lacerum anterius. It leads into the orbit, and 
allows the exit of the motor nerves of the eyeball, or third, 
fourth, and sixth cranial nerves, and also the first division 
of the trigeminal or fifth nerve. 

4 and 5. The alisphenoid near its base is perforated by 
two foramina : the anterior small and somewhat round, the 
posterior larger and oval ; these are the foramen rotntnhtm 
and \\\z foramen male, and transmit respectively the second 
and third divisions of the fifth nerve. 

6. Between the alisphenoid anil the exoccipital is a large- 
space, almost entirely filled by the bony capsule of the organ 
of hearing, the periotic bone. In front of the inner end of 
this bone is an opening (foramen lacerum medium faisis 
cranii), through which the internal carotid artery sometimes 
enters the craninl cavity. 

7. Near the middle of the inner surface of the periolic 
is the meatus anditorius infernus, into which the seventh 
and eighth nerves enter : the former (the facial nerve) 
passes through the bone and emerges on the other side 
(by the stylo-tnastoid foramen] ; the latter, the auditory, is 
distributed to the internal organ of hearing within the 
periotic bone. 

A deep depression seen above the internal auditory 
meatus, and of nearly the same size, is not a foramen but a 
fossa, lying within the concavity of the superior semicircular 
canal. It lodges the flocculus, a small process of the cere- 

8. Between the periotic and the exoccipital an irregular 
space is left (the foramen lacerum poster tus), through which 
the glossopharyngeal, pneumogastric, and spinal accessory 


nerves (the ninth, tenth, and eleventh) pass out of the 

9. The exoccipital is perforated, a little in front of the 
condyle, by the condylar foramen, which gives exit to the 
twelfth, or hypoglossal, nerve. 

10. Lastly, the large median opening, behind the bones 
of the posterior segment of the skull, is the fora men magnum. 
through which the spinal cord passes out. 

It will be seen from the foregoing description that the 
three organs of special sense, situated in the walls of the 
cranium, have definite relations with the three osseous 
segments. The first, or organ of smell, is situated in front 
of the frontal segment ; the second, or organ of sight, 
receives its nerves by apertures situated between the frontal 
and parietal segments or perforating the former; the third, 
or organ of hearing, is intercalated between the parietal 
and occipital segment. 

The portion of the skull anterior to the junction of the 
presphenoid and the mesethmoid constitutes the face. This 
differs entirely from the cranial cavity in having a complete 
median partition, and consists mainly of two tubular cavities 
placed one on each side of this partition. These are the 
nasal cavities. They are open at each end, the orifices 
being termed respectively anterior and posterior nares. 

Each of these elongated cavities is deepest vertically in 
its posterior part, where it is partially divided into an upper 
and lower chamber : the upper one, the olfactory chamber, 
being closed behind by the cribriform plate ; the lower, the 
narial passage, terminating in the posterior nares. 

Each nasal cavity may be described as having an inner 
wall, an outer wall, a floor, and a roof. 

The inner wall is formed mainly by the partially ossified 

ix.] OF THE DOG. 12.5 

mcsethmoid cartilage (ME), but the lower part of the pos- 
terior two-thirds also by the vomer ( Vo). The greater part 
of this bone has the form of a trough, hollow above, em- 
bracing the inferior, thickened border of the mesethmoid 
cartilage, and extending a little way behind this so as also 
to underlie the anterior portion of the presphenoid ; but it 
also develops from its under surface in the middle line a 
thin plate, which passes vertically down to the centre of 
the floor of the nasal passages, and completes the septum 

Above, rather behind the middle of the bone, the lateral 
plates of the vomer, which embrace the mesethmoid carti- 
lage, send out sideways a pair of wing-like processes, which 
join the side walls of the nasal cavity, and form the partial 
horizontal partition, dividing the narial passage from the 
olfactory chamber. 

The outer wall of the nasal cavity is formed mainly by 
four bones : (i) a somewhat quadrate, thin, nearly vertical 
plate of bone (/V), the pterygoid, attached above to the under 
surface of the basisphenoid and presphenoid, supported 
externally by a strong descending process of the alisphen- 
oid, the- external pterygoid plate, ending posteriorly and 
inferiorly by a free border, and articulating in front with (2) 
the palatine. This bone (7Y) is of much greater extent ; for, 
besides its vertical portion, forming the outer wall of the 
nasal canal in front of the pterygoid, it sends from its upper 
edge a lamina inwards to meet the horizontal plate of the 
vomer, and aid in forming the roof of the hinder part of the 
narial passage. It also sends a strong horizontal lamina 
inwards from its lower edge, which, meeting its fellow in the 
middle line, forms the posterior part of the floor of the narial 
passage. In addition to these it sends a broad plate upwards 
and forwards in the inner wall of the orbit. 

[Continued on p. 128. 

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3. In front of this is the maxilla (Mx], a still more con- 
siderable bone. It not only forms the chief part of the outer 
wall of the nasal cavity, but it also sends inwards a hori- 
zontal plate, forming the middle part of its floor. At the 
junction of its vertical and horizontal portions is the alveolar 
border, in which the canine, premolar, and molar teeth are 

4. The most anterior bone of this series is the premaxilla 
(PMx), which also has an ascending or vertical plate, 
forming the outer wall of the nasal cavity, and a horizontal 
plate forming the anterior part of its floor ; at their junc- 
tion its alveolar border lodges the incisor teeth. The pre- 
maxilla forms the outer and lower boundary of the anterior 

Besides these four, there is a small bone which enters into 
the outer wall of the upper part of the nasal cavity, between 
the ascending process of the palatine, the maxilla and the 
frontal. This is perforated by the duct, which conveys the 
tears from the orbit into the nasal cavity, and is hence 
called la cry in al. 

Above this a process from the frontal completes the 
upper and posterior part of the outer wall of the olfactory 

The floor of the nasal cavity is formed, as above said, by 
the horizontal plates of the palatine, maxilla, and premaxilla 
meeting the corresponding bones of the opposite side in the 
middle line. The inferior surface of this same horizontal 
layer of bone is the roof of the mouth, or bony palate. 

The roof of the nasal cavity is formed, posteriorly by 
the continuation of the frontal forwards beyond the cere- 
bral cavity, the " nasal process of the frontal" but mainly 
by a long narrow bone, the nasal bone (^Vtf). The hinder 
extremity of this lies upon the nasal process of the frontal ; 

ix.] OF THE DOG. 129 

its anterior end is free, and forms the upper boundary of 
the anterior nares ; its outer side is in contact with the 
frontal, maxilla, and premaxilla ; and its straight inner edge 
lies against that of the corresponding bone of the opposite 

Within each nasal cavity are two very singular bones, 
each being composed of a number of delicate lamella; 
folded and arranged in an exceedingly complex manner, 
forming a mass with so many passages and perforations that 
the term " spongy bones " has been applied to them. 

The most posterior is the larger, and placed rather higher 
than the other, its anterior extremity (ET) overlapping it. 
It completely fills the proper olfactory chamber ; its hinder 
extremity occupying the gap left in the cranial wall in front 
of the anterior segment of the brain-case. The various 
laminae are all connected together and to the hinder end of 
the mesethmoid, by a plate of bone (CE) so full of perfora- 
tions of varied form and si/e that it is called the cribriform 
plate, and from it the name of ethmoid (or sieve-like) is 
commonly .applied to all the bony structures with which 
it is united. On their outer side these laminae are con- 
nected to a thin flat plate of bone (the so-called os planum} 
which lies against the inner wall of the maxillary, but does 
not ordinarily contract any union with it. 

This bone results from the ossification of the complexly 
folded cartilage, over the surface of which the olfactory 
nerves are spread, the division into laminae permitting a 
great increase of sensitive surface. As, although originally 
distinct, it subsequently unites with the mesethmoid, by 
means of the cribriform plate, it is considered in human 
anatomy as part of the same bone, and described under 
the name of "lateral mass of the ethmoid," being further 
divided into the superior and middle " turbinated bones " ; 



but the name ethmoturbinal, applied to it by Sir Richard 
Owen, is perhaps more appropriate. 

The uppermost of the lamellae of the ethmoturbinal, lying 
immediately under the nasal bones, is rather distinct from 
the others, and extends much further forwards ; and as in 
certain Mammals it becomes united by bone with the 
nasal, it is sometimes distinguished under the name of 

In front, and on a rather lower level, a similar, but smaller 
and less complex bone (AfT), consisting chiefly of horizontal 
lamellae, is placed. This, though originally developed from 
the same cartilage lining the outer wall of the nasal chamber 
as the last, ossifies quite distinctly from it, and contracts a 
bony union by a horizontal lamella on its outer side with 
the maxilla. This is the maxiUoturbinal, and corresponds 
with the " inferior turbinated bone " of human anatomy. 

It will be observed, that while the ethmoturbinal is placed 
high in the nasal cavity, and above the direct channel by 
which the air passes to the posterior nares, the maxiUo- 
turbinal, situated nearer the front of the chamber, before it 
has divided into an upper true olfactory chamber and a 
lower narial passage, nearly blocks up the whole cavity, so 
that air passing through in inspiration is filtered between its 
meshes. The moist membrane which covers its bony plates 
in life is supplied with nerves chiefly from the fifth pair, 
and not from the olfactory ; so that it does not perform the 
function of an organ of smell like the ethmoturbinal, but 
rather serves to guard the entrance of the respiratory 
passages from foreign substances, and perhaps to warm the 
inspired air. 

In describing the walls of the cranium, a large space was 
mentioned on each side, between the posterior and middle 

ix.] OF THE DOC. 131 

cranial segment, in which were inserted certain bones not 
yet noticed. These bones form a definite group by them- 
selves, at all events locally connected, though very different 
in function and structure. 

In a mass of cartilage, in the position just indicated, 
ossification takes place from several centres (three, called 
respectively pro-otic, ebiotic, and opisthotic, according to 
Professor Huxley and others, in the human skull; 1 but the 
process has not been accurately traced in other Mammals). 
These very rapidly unite to form a single bone, which com- 
pletely encloses the labyrinth or essential organ of hearing, 
consisting of the vestibule, semicircular canals, and cochlea. 
This bone is the periotic (Pfr). It is divided into two por- 
tions : an antero-internal, which forms a somewhat angular 
projection within the cranial cavity, and is of remarkable 
density the petrous portion ; and a postero external, a sort 
of process from the former, smaller, less dense, and forming 
a small portion of the wall of the cranium, appearing 
externally just in front of the exoccipital the mastoid 

The petrous is of course the more important, and has 
constant characters throughout the class, while the mastoid 
is very variable, and sometimes can scarcely be said to exist. 
It is in no case a separate bone ; and, although a portion of 
it may develop originally from a separate centre, it is always 
before birth firmly united with the petrous, so that they 
will be spoken of here as one bone, under the name of 

The essential characters of the petrous portion of the 
periotic are, that it contains within it the internal ear ; 
that it has on its inner or cranial side a foramen, through 
which the facial and the auditory nerves leave the cranial 

1 " Elements of Comparative Anatomy," (1864), p. 148. 
K 2 


cavity, the former to pass through the bone, escaping by 
the stylomastoid foramen on the outer and under surface, 
the latter to be distributed on the sensitive portions of the 
organ of hearing ; and that it has on its outer side two 
holes, one placed above the other, the fenestra ovalis and 
the fenestra rotunda, through which the internal ear com- 
municates with the cavity of the tympanum, or middle-ear, 
which is situated on the outer side of the petrous part of 
the periotic bone. 

Externally to the periotic bone are placed two bones 
separately developed in fibrous tissue, which often acquire a 
very close connection with the periotic, occasionally, as in 
Man, becoming firmly ankylosed with it. 

The upper one of these is the squamosal (Figs. 48 and 49, 
Sy), which has a broad, scale-like, vertical portion spreading 
out over the side of the cranial wall, uniting with the supra- 
occipital behind, overlapping the lower edge of the parietal 
and the hinder part of the alisphenoid, and also appearing 
for a very small space in the inner side of the cranial wall. 
From near its lower border it sends a strong process out- 
wards, which soon curves forwards, called the zygomatic 
process. This articulates with another bone ( Ma), the malar 
or jngal, which connects it with the maxilla, and so forms 
the strong, lateral, nearly horizontal, or slightly arched 
osseous bridge, which passes from the face to the hinder 
part of the cranium, called the zygomatic arch. On the 
under surface of the base of the zygomatic process of the 
squamosal is a laterally extended, oblong surface, concave 
from before backwards, for the articulation of the condyle 
of the lower jaw, called the glenoid fossa (gf\ the hinder 
edge of which is projected into \\\Q postglenoid process (gp). 

The lower bone, on the outer side of the periotic, is 
the tympanic (TV). At birth this .is a mere osseous ring, 

ix.] OF THE DOG. 133 

incomplete above, surrounding the inferior three-fourths of 
the membrana tympani, but it undergoes a considerable 
development in the course of the first few months. The 
external edge of the ring is produced horizontally outwards 
to form the short, bony, external auditory meatus ; while 
the under and inner surface is greatly expanded, to form 
the conspicuous rounded prominence, hollow within, called 
the auditory bulla, which abuts against the outer edge of 
the basioccipital below. 1 

The space that is left among this group of bones 
bounded by the periotic (the part in which the before- 
mentioned fenestne are situated) within, the periotic and 
squamosal above, the tympanic and its bullate expansion 
below, behind, and in front, and by the meatus auditorius 
externus closed in the natural state by the membrana 
tympani, to the outer side is called the tympanic carity. 

It contains within it the ossicula auditus, three small bones 
called malleus, incus, and stapes, which articulated together, 
stretch across the cavity from the membrana tympani to the 
fenestra tn-alis.- The cavity has an opening at its antero- 

1 The whole of the bulla is generally considered as belonging to the 
tympanic bone, but its inner part in many Mammals is developed in a 
distinct cartilaginous lamella, interposed between the lower edge of the 
tympanic ring and the base of the skull. This may ossify from a sepa- 
rate nucleus, or by extension of bony deposition inwards from the true 
tympanic The development of this region of the skull in the Mam- 
malia still offers an interesting field for investigation, 

* As these bones are, in the Mammalia, completely sul>servient to 
the organ of hearing, their modifications will not be described in the 
present work. A very detailed and fully illustrated account of them 
has been given by Alban Doran in the Transactions of the Linnean 
S<>ciety t Second Series, vol. i. " Zoology," 1879. 

Much has been written on the still unsettled question of the develop- 
ment and the homologies of the auditory ossicles by ReUhert, Peters, 
Huxley, Parker, Koelliker and Gegenbaur ; see also IV. Saltmky, 


internal angle, through which the Eustachian tube connecting 
the tympanum with the pharynx passes. 

The inner side of the bulla is perforated lengthwise by a 
canal, which commences posteriorly within the margin of 
the foramen lacerum posterius (between the auditory bulla 
and the exoccipital), and transmits the internal carotid artery. 
This vessel appears again on the surface, at the anterior 
extremity of the bulla, close to the Kustachian orifice ; then 
runs upwards and inwards and enters the cranium through 
the foramen lacerum medium. 

This completes the enumeration of the bones of the 
cranium. Before proceeding further, it will be desirable to 
take a general survey of this part as a whole, pointing out 
the most prominent features of its various surfaces. 

The posterior surface is, in a general sense, a vertical wall 
somewhat triangular in form, broad below and pointed above. 
In the middle line, at its lowest border, is the nearly round 
foramen magnum, bounded by the supraoccipital above, the 
exoccipitals on each side and the basioccipital below. On 
the sides of the foramen magnum, and approaching each 
other in the middle line below, but diverging above, are 
smooth eminences, the occipital condyles. Further outwards, 
and separated from these by a deep valley, are the par- 
occipital processes, projecting backwards and downwards. 
Outside of the upper part or origin of these processes, the 

"Beitrage zur Entwicklung. der knorpeligen gehorknochelchen. " 
Morphol. Jahrb. VI. 1880. Fraser, "On the Development of the 
Ossicula Auditus of the higher Mammalia," Philosoph. Transact, vol. 
173, 1883, p. 901. /'. Albrecht, " Sur la valeur morphologique ck- 
1'articulation mandibulaire, &c.," Bruxelles, 1883. M. L. Dallo, "On 
the malleus of the I.acertilia, and the malar and quadrate bones of 
Mammalia," Quart. Journ. Science, vol. xxiii. N. 8. p. 579, 
1883. The last contains a complete list of the literature on the subject 
and a tabular arrangement of the views held by previous authors. 

ix.] OF THE DOG. 135 

mastoid portion of the periotic appears on the hinder wall of 
the skull The remainder of the region is formed by the 
supraoccipital, and it is distinctly marked off laterally by 
ridges, which, commencing in the median line above, runs 
downwards and outwards, at the junction of the parietals and 
supraoccipital, and are continued on the squamosal in front 
of the mastoid to the upper edge of the external auditory 
meatus. The ridges of each side taken together form the 
lambdoid or occipital crest. They are far more conspicuous 
in old than in young animals. 

The superior surface of the skull (Fig. 48) may be divided 
into a cranial and facial portion. The former is of a some- 
what oval form. On its upper surface posteriorly, in full- 
grown dogs, is a median ridge joining behind with the 
superior angle of the occipital crest, and dividing anteriorly 
into two less elevated ridges which curve outwards to the 
superior posterior angle of the orbit. This ridge, as long as 
it is single and median, is called the sagittal crest. It bounds 
superiorly a large surface on the side of the skull, limited 
behind by the occipital crest, and below by the zygoma, 
called the '' temporal fossa," from which the temporal muscle 
takes its origin. In young dogs the upper boundary of the 
surface for the origin of this muscle is of less extent, not 
reaching so high as the middle line of the cranium, and is but 
obscurely indicated on the comparatively smooth surface of 
the skull. As the muscle increases in development its surface 
of origin gradually ascends until it reaches the middle line, 
and with advancing age a still larger space is afforded for it 
by the gradual growth of the sagittal crest. 

These changes in the upper part of the skull during growth 
have been particularly noticed, because they take place in very 
many animals, and, without altering in the least the actual 
form of the brain-case give rise to a very different external 



appearance of the skull, either in members of the same 
species, or in different but allied species. 

The upper part of the skull, in front of the diverging 
boundary lines of the temporal fossa, is expanded and some- 

Fr;. 48. Upper surface of cranium of a Dog, J. SO supra-occipital; IP inter- 
parietal ; Pa parietal ; Sq squamosal ; Fr frontal ; Ma malar ; L lacrymal ; 
.l/.r maxilla ; A'a nasal ; PMx premaxilla : tip anterior palatine foramen ; to infra- 
orbital foramen ; pof postorbital process of frontal bone. 

what flattened, and has on each side a triangular process 
(pof\ which curves somewhat downwards, and indicates the 
division of the temporal fossa behind from the orbit in front. 

ix.] OF THE DOC. 137 

This is the postorbital process of the frontal bone. It is con- 
nected by a ligamentous band, in the living animal, with a 
corresponding process arising from the zygoma ; but when 
this is removed the orbit and temporal fossa are widely 
continuous, their respective limits being only indicated by 
the above-mentioned processes. 

The face is produced considerably in front of the orbits, 
and not only becomes more depressed, but also more com- 
pressed laterally, and is obliquely truncated anteriorly, 
terminating in the rounded incisor border of the premaxilla 
(PMx); above which is placed the subcircular orifice of the 
anterior nares. 

The upper, and a considerable portion of the lateral surface 
of the cranium behind is formed by the parietal bones (Pa), 
having the narrow interparietal (//*) ankylosed with the supra- 
occipital (SO), extending between them for about half their 
length. In front of this the parietals are commonly united 
together by bone in old dogs. 

Anteriorly to the parietals, the upper part of the temporal 
fossa, the frontal plateau between the orbits, and the upper 
half of the inner wall of the orbit, are formed by the frontal 
bone. The remaining or lower portion of the temporal fossa 
is formed by the squamosal behind and by the alisphenoid 
in front On the inner or cranial surface of the confluent 
orbital and temporal fossae, a wide groove runs obliquely 
downwards and backwards, which may be considered as the 
boundary lines of these two regions. In the lower part of 
this groove are several large foramina placed in linear series. 
The highest is the optic foramen, the next (of larger sue) the 
sphenoidal fissure, and the third the foramen rotundum and 
anterior opening of the alisphenoid canal. The orbit has no 
floor except for a very short space in front ; the lower border 
of its inner wall passing directly into the outer surface of the 


vertical ridge formed by the pterygoid, the pterygoid process 
of the alisphenoid and the palatine bones, and continuing the 
outer wall of the narial passage backwards beyond the bony 
palate. The palatine bone forms a considerable part of 
the inner wall of the orbit, joining the frontal anteriorly 
though separated from it for a considerable space posteriorly 
by the orbitosphenoid. The lacrymal (L) appears in the 
anterior boundary of the orbit, the malar (Ma) joins its outer 
boundary, and the upper surface of the hinder end of the 
alveolar border of the maxilla projects backwards, so as to 
form a partial floor to its anterior extremity. In front of 
the orbits the face is formed above by the long narrow nasals 
(Na) pointed behind, and widening and obliquely truncated 
in front to form the upper border of the narial aperture ; on 
each side by the maxillae (Jlfx) having near the middle oi 
their surface the large infraorbital foramen (/<?), through 
which the terminal branches of the second division of the 
fifth or sensory nerve of the face pass to be distributed to the 
upper lips and whiskers ; and quite anteriorly by the pre- 
maxillae (PMx\ which complete the boundaries of the nares, 
and send up narrow processes between the nasals and 
maxillae towards, though not meeting with, similar processes 
which run from the frontals. 

The inferior surface of the skull (Fig. 47) is formed an- 
teriorly by the nearly flat, elongated surface of the palate, 
narrower in front than behind, composed anteriorly of the 
premaxillae (PMx) ; then of the maxillae (Mx), which diverge 
posteriorly and allow the palatines (PI) which form the 
hinder border to reach in the middle line almost to the 
centre of the palatal surface. In front and at the sides this 
surface is bounded by the alveolar borders of the premaxillae 
and maxillae in which the teeth are set. Anteriorly are two 
considerable oval foramina (#//), placed longitudinally very 




near the middle line, formed mainly in the premaxillae, 
though their boundary is completed posteriorly by the 

FIG. 49. Under surface of the cranium of a Dog. J. SO siipraoccipital : ExO 
exoccipital ; BO basioccipital ; Per mastoid portion of periotic ; Ty tympanic 
bulla : BS basisphenoid : Sq ic process of squamosal ; Ma malar ; - 1-V 
alisphenoid: ft pterygoid : PS presphenoid ; Ar frontal; Vo vomer \Pl palatine 
MX maxilla ; PMx premaxilla ; fin foramen magnum ; oc occipital condyle 
// paroccipital process ; cf condylar foramen ; flp foramen lacerum posterius ; nn 
stylo-mastoid foramen ; earn external auditory meat us ; pgf postglenoid foramen 
gp postglenoid process ; gf glenpid fossa;///* foramen lacerum medium \ fo 
foramen ovale ; its posterior opening of alisphenoid canal ; fr foramen rotundum 
and anterior opening of alisphenoid canal ; sf sphenoidal fissure or foramen 
lacerum an terms ; op optic foramen ; ///" posterior palatine foramen ; apf anterior 
palatine foramen. 

maxillae ; these are the anterior palatine or incisive foramina. 


The naso-palatine nerve descends through them to spread 
over the anterior surface of the soft palate. Not far from 
the hinder border of the palate, and more distant from the 
middle line, near the suture between the maxilla and palatine, 
are several much smaller foramina (posterior palatine), also 
for the transmission of branches of the fifth nerve and blood 

The truncated median part of the hinder edge of the 
palate forms the lower margin of the posterior narial aperture. 
Laterally, the palate bones are continued backwards as 
vertical plates, thick and rounded below at first, but gradually 
becoming more compressed. These are continued still 
further backwards by the compressed pterygoid bones (Pt), 
ending in the backward projecting hamular processes, and 
supported externally by the descending (pterygoid) plate of 
the alisphenoid. The groove between these descending 
lamellae of bone continues the narial passage backwards. It 
has for its roof the vomer ( Vo] in front, then the presphenoid 
(PS), and posteriorly a portion of the basisphenoid (BS) ', 
but the palatines and pterygoids arch over so much towards 
the middle line that they only leave a small strip of these 
bones exposed. Inferiorly, this groove is not closed by 
bone, but in the living animal the soft palate is stretched 
across it. 

The base of the skull, behind this " mesopterygoid " fossa 
presents in the middle a nearly flat elongated surface, con- 
sisting of the basisphenoid (-BS) and basioccipital (-BO] ; the 
latter, roughened for the attachment of muscles and ter- 
minating posteriorly at the inferior border of the foramen 
magnum (////), flanked on each side by the occipital 
condyles (oc). The nearly straight lateral edges of the 
anterior half of the basioccipital rise up to abut against the 
prominent smooth rounded auditory bullce (Tj), which form 

IX.] OF THE DOG. 141 

so conspicuous a feature in this region of the skull, and 
which are produced outwards into the lower wall of the 
external auditory meatus (tarn}. In the antero-internal angle 
of the bulla is seen the irregular orifice of the Eustachian 
canal. Close to the inner side of this is an oval aperture 
which is at the same time the anterior extremity of the carotid 
canal and the entrance to the foramen lacerum medium (/////) 
through which the internal carotid artery enters the cranial 
cavity. In front, and rather to the outer side of this, is the 
foramen ovale (fo) piercing the alisphenoid, and immediately 
before this is a round aperture (as) leading to a short canal 
running horizontally forwards through the same bone at the 
root of its pterygoid process, and opening anteriorly into the 
foramen rotundum (fr). Through this the external carotid 
artery runs lor part of its course, and it has been called the 
alisphenoid canal. * 

In front of the outer side of the auditory bulla' is the 
glenoid fossa for the articulation of the mandible, bordered 
behind by the conspicuous curved postglenoid process (.<?/). 
Immediately behind this the root of the zygoma is pierced 
by a large hole, postglcnoid foramen ( f>gf), through which a 
vein passes out from the lateral sinus within. 

Behind the auditory bulla, to the inner side, is the large 
foramen lacerum paste rius (///), and situated deeply within its 
recesses, the posterior opening of the internal carotid canal. 
On a ridge of the exoccipital, between this large foramen 
and the depression immediately in front of the condyle, is 
the small, nearly circular condy la r foramen (cf\ and at the 
outer termination of the same ridge rises the conical paroc- 
cipital process (//>), abutting at its base against the hinder 

1 See H. N. Turner's " Observations relating to some of the Foramina 
in the Base of the Skull in Mammalia," &c., Proc. Zool. Soc. 1848, 
p. 63. 


end of the auditory bulla. Immediately behind the bulla, 
and to the outer side of the abutment of the paroccipital 
process against it, is an oval hole (sm\ partially divided by 
a constriction into an inner and an outer division. In 
the inner division the end of a small cylindrical plug of 
bone, the tympanohyal, can generally be seen. The outer 
division is the stylomastoid foramen, through which the 
seventh nerve, or portio dura, makes its exit. The bone 
forming its outer boundary is the mastoid portion of the 

Connected with the posterior lateral parts of the cranium 
are two appended bony parts : the lower jaw or mandible, 
and the hyoidean apparatus. The former forms the frame- 
work for the floor of the mouth, and supports the lower 
series of teeth ; the latter gives a firm yet movable point 
of attachment to the root of the tongue and to the larynx, 
or organ of voice. 

The mandible consists of two symmetrical elongated rami 
(see Fig. 46, p. 117), diverging behind, and coming in contact 
in front at the middle line, by a roughened surface called the 
symphysis (s) ; here they are firmly held together by inter- 
posed fibrous tissue, or in old animals they may become 

Each ramus is compressed from side to side, has a 
thickened rounded lower border, slightly curved in the 
longitudinal direction, and a nearly straight upper alreolar 
border, in which the teeth are implanted. The inferior 
border inclines upwards in front to meet the alveolar border 
at the front of the symphysis. Near the posterior extremity 
is the condyle (cd), a transversely-extended projection, with its 
upper surface rounded in the antero-posterior direction, and 
which, fitting into the glenoid cavity of the squamosal bone, 
forms the hinge-like synovial articulation by which the lo\ver 

ix.] OF THE DOG. 143 

jaw moves on the skull. The upper border, between the 
condyle and the hindermost tooth, rises into a high, com- 
pressed, recurved process (the coronoid process, cp), to which 
the temporal muscle is attached. The outer surface of this 
process gradually subsides into a considerable hollow in the 
side of the ramus, with prominent anterior, inferior, and 
posterior edges, to which the masseter, another powerful 
muscle for closing the jaw, is attached. 

The point at which the vertical hinder edge of the ramus, 
descending from the condyle, meets the horizontal inferior 
border, is called the angle, which in the Dog is prolonged 
into a conspicuous compressed process with an upturned and 
slightly inverted pointed extremity, the angular process (a). 
On the inner side of the ramus, a little way in front of and 
below the condyle, is the inferior dental foramen (id), for the 
admission of the inferior dental nerve (from the fifth pair) 
and artery. On the outer side of the ramus, near its anterior 
extremity, is the mental foramen, through which a branch of 
the same nerve passes out to the lower lip and surrounding 

The hyoidean apparatus (Fig. 50) consists of a median 
portion below, the basihyal (bK), from which two pairs 
of half arches, or " cornua," extend upwards and outwards. 
The anterior (ch to s/i) is the largest, and connects it with 
the cranium. The posterior (//;) is united externally or 
superiorly with the thyroid cartilage of the larynx. In the 
Dog there are four distinct ossifications in the anterior arch. 
The first is a small cylindrical piece of bone lying in a 
canal between the tympanic and periotic bones, immediately 
to the inner and anterior side of the stylomastoid foramen, 
and by its upper end firmly ankylosed with the surround- 
ing bones. It can be seen much more distinctly in 
some dogs' skulls than others, and is more conspicuously 


developed in some other Mammals. This I have called tym- 
panohyal, as it is always in relation with the hinder edge 
of the tympanic bone, generally more or less surrounded by 
it, and it extends upwards, embedded in, and afterwards 
ankylosed with, the periotic, to the hinder wall of the 
tympanic caviiy. Its lower end is truncated and con- 
tinued into a band of cartilage, which connects it with 
the proximal end of the bone which has been generally 
recognised as the uppermost of the series forming the 


FIG. 50. Extracranial portion of hyoidean apparatus of Dog, front view, sh stylo- 
hyal ; eh epihyal ; ch ceratohyal (these three constitute the "anterior cornu"); 
b'k basihyal or " body " of hyoid ; th thyrohyal, or "posterior cornu." 

anterior hyoidean arch, the stylohyal (s/i). The two suc- 
ceeding bones (ep and ch} are named by Sir Richard Owen 
respectively epihyal and ceratohyal. All three are elon- 
gated, compressed, slightly curved or twisted on them- 
selves, tipped at each end with cartilage, and connected 
with each other by synovial joints. The stylohyal and 
epihyal are nearly equal in length, the ceratohyal shorter 
and stouter. 

The basihyal (bit] is a transversely-extended, flattened bar, 


with its extremities rather upturned and thickened. The 
posterior cornu (th) consists of a single, nearly straight, 
compressed bone, the thyrohyal, articulated inferiorly with 
the outer end of the basihyal, just below the attachment 
of the ceratohyal, and truncated at its superior extremity, 
to which the thyroid cartilage of the larynx is suspended. 

Development of the Skull. For a detailed and beautifully 
illustrated account of the early development of the Mam 
malian skull, we must refer to Professor Parker's monograph 
"On the Structure and Development of the Skull in the 
Pig" {Philosophical Transactions, 1874). As however the 
prevailing views regarding the morphology of the skull have 
been considerably modified even since the publication of 
that memoir, we take the following abstract from Balfour's 
Treatise on Comparative Embryology, 1881. 

The primitive cartilaginous cranium is formed by a 
differentiation within the membranous cranium, and is 
composed of the following parts : 

(i.) A pair of cartilaginous plates on each side of the 
anterior extremity of the notochord, known as the para- 
chordals (Fig. 51, pa. ch.}. The continuous plate, formed 
by them and the notochord, is known as the basilar or 
basicranial plate. 

(2.) A pair of bars forming the floor for the fore-brain, 
known as the trabeculae (tr) ; these are continued forward 
from the parachordals. They meet behind and embrace the 
front end of the notochord ; and after separating for some 
distance bend in again in such a way as to inclose the 
pituitary body (py). In front of this they coalesce, and 
together with the olfactory capsules (ol) give rise to the pre- 
sphenoidal and ethmoidal regions of the cranium (Fig. 47, 1.). 

(3.) The cartilaginous capsules of the sense organs. Of 
these the auditory and olfactory capsules unite more or less 




intimately with the cranial walls. The basicranial plate 
grows up as an arch over the occipital region of the skull, 
and coalescing with the auditory capsules gives rise to the 
primordial skeleton of the occipital, periotic, and basi- 
sphenoidal regions of the skull : the parietal and frontal 
regions being afterwards completed by ossification in mem- 
brane surrounding the cranial cavity. 

FIG 51. Klements of skull (seen from below) of an embryo Pig two-thirds of ;in 
inch long, parachordal cartilage; tic notochord ; au auditory capsule ; py 
pituitary body; tr trabeculai ; trabecular cornu ; / prenasal cartilage; en 
external nasal opening ; ol nasal capsule ; j>.f>g palatopterygoid tract inclosed in 
the maxillopalatine process; mn mandibular arch; ky hyoid arch; th.h first 
branchial arch ; ya facial nerve ; 8a glossopharyngeal; ?>b vagus ; 9 hypoglossal. 

Besides the parts composing the cranium proper, the 
Mammalian skull is completed by the labial cartilages, the 
meaning of which is still very obscure, and a series of three 
paired descending cartilaginous bars or arches, developed in 
the visceral laminae, constituting the sides of the face and 

The first visceral (mandibular) arch divides itself into 
a short proximal or dorsal, and a much longer distal or 
ventral end (Meckefs cartilage}. The first sends out 
forwards towards the roof of the mouth-cavity, a process 

ix.] DEVELOPMENT. 147 

(pterygo-palatine} which gives rise to the pterygoid, palatine, 
maxillary and malar regions (Fig. 47, II.). 

The latter, Meckefs cartilage (Fig. 52, mk ; Fig. 47, III.), a 
slender rod, is proximally in relation with the periotic region 
of the skull. In the Mammalia, the upper extremity of this 
rod becomes converted into the malleus, one of the small 
bones in the tympanic cavity, while in connection with the 
lower or distal part, the ramus of the mandible or lower jaw 
is developed, partly by conversion of the cartilage itself, but 
principally by the ossification of fibrous or cartilaginous 
tissue deposited around it. The upper end of the ramus 


Ku;. 52. Side view of mandibular and hyoid arches of an embryo Pig, an inch and 
a third long. The main arch i seen as displaced backwards after segmentation 
from the incus, tg tongue ; mk Meckelian cartilage ; ml body of malleus : ntl> 
mnnubrium or handle of the malleus; t.ty legmen tympani ; /'incus: tt stapes : 
if>y interhyal ligament ; st.h stylohyal cartilage ; It.h hypohyal ; bh basi-branchial ; 
th.k rudiment of first branchial arch ; ^a. facial nerve. 1 

afterwards acquires a secondary articular connection with 
the squamosal bone, its primitive connection with the 
malleus entirely disappearing. 

The rod of cartilage forming the second visceral arch 
(Fig. 47, IV.) becomes the anterior hyoid arch, its proximal 
extremity being modified into the incus. 

From the third arch, which corresponds to the first 
branchial of branchiate vertebrates, is formed the posterior 
hyoid arch, or thyrohyal ( (Fig. 47, V.). 

1 Figs. 51 and 52, reduced from the original figures of Parker, are 
taken with permission from Balfour's Comparative Embryology. 

L 2 

1 4 8 THE SKULL. [CHAP. 

The "cleft" between the palato-pterygoid (II.) and the 
mandibular (III.) bar is the mouth. The first true cleft, 
however, is the one between the mandibular and the anterior 
hyoid arch (IV.). It becomes contracted into the Eustachian 
tube, tympanic cavity, and meatus auditorius externus, which 
would form a. canal of communication between the pharynx 
and the external surface but for the interposition of the 
delicate membrana tympani. 

Some of the changes which take place in the cranium 
while advancing from youth to maturity have already been 
noticed ; but it will be well, before proceeding to describe 
the modifications of the Mammalian skull, to mention certain 
others which take place, to a greater or less degree, in all 

These depend mainly on the fact that the brain, and con- 
sequently the cavity which contains it, and also the sense 
capsules, increase in size in a much smaller ratio than the 
external parts of the head, especially the jaws and pro- 
minences for the attachment of muscles. The dispropor- 
tionate growth and alteration of form of these parts, 
concomitant with little or no change in the brain-case, is 
effected partly by increase in thickness of the bones, but 
mainly by the expansion of their walls and the develop- 
ment of cells within, which greatly extend the outer surface 
without adding to the weight of the bone. 

In the Dog these cells are developed chiefly in the fore- 
part of the frontal bones, constituting the frontal sinuses, 
and in the presphenoid, constituting the sphenoidal sinuses. 
Air passes freely into them from the nasal passages. In 
many animals they attain a much larger extent than in the 
Dog, reaching their maximum in the Elephant (see Fig. 64, 
p. 205), where the alteration of the external form of skull 
during growth, without material change in the shape or size 

ix ] DEVELOPMENT. 149 

of the cerebral cavity, is strikingly shown. At the same 
time the alveolar borders of the jaws gradually enlarge to 
adapt themselves to the increased size of the permanent 
teeth which they have to support, and the various ridges and 
tuberosities for the attachment of muscles become more 

During these changes a gradual consolidation takes place 
in the structure of the skull generally, by the partial or com- 
plete union of certain of the bones by synostosis. The 
union of the different bones generally proceeds in a certain 
definite order, which, however, varies much in different 
species. Sometimes it extends so far as to lead to complete 
obliteration of all the cranial sutures. 



Order PRIMATES. Man. On comparing a longitudinal 
and vertical section of a young human skull in which most 
of the sutures are still distinctly seen (Fig. 53), with that of 
the Dog, it will be seen to be composed of the same bones, 
having very nearly the same connections, and yet the whole 
form is greatly modified. This modification is mainly due 
to the immense expansion of the upper part of the middle 
or cerebral fossa of the brain cavity, which not only carries 
the roof of the cavity a great distance from the basicranial 
axis, but also forces, as it were, the anterior and posterior 
walls from the vertical nearly to the horizontal position, so 
that they are, roughly speaking, in the same line with the 
short basicranial axis, instead of being perpendicular to it. 
In addition to this great difference, the facial portion of the 
skull is deeper from above downwards, and very much 
shorter from before backwards. 

Taking a survey of the human skull in the same order 
as was done with that of the Dog, we find the craniofacial 
axis, composed of the basioccipital bone (BO\ terminating 
at the anterior border of the foramen magnum (/;;/) behind, 

CHAP. X.] 



FIG. 53. Vertical, longitudinal, median section of a young human skull, with the first 
dentition. J. As in the other sections of a skull figured, the mandible is displaced 
downwards, so as to show its entire form. PMx premaxilla ; MT maxillo- 
turbinal ; KT ethmo-turbinal ; .I/A" ossified portion of the mesethmoid ; Na nasal ; 
eg crista galli of the mesethmoid; OS prbttosphenoid. or lesser wing of the 
sphenoid ; AS alisphenoid. or greater wing of the sphenoid ; /> frontal ; /'a 
parietal ; SO supraoccipital ; M inastoid portion of the periotic ; Sy squamosal; 
Per petrous portion of the periotic : the large foramen below the end of the line 
is the internal auditory meatus. the small depression above it is the nearly- 
obliterated floccular fossa. E.\O exoccipital, the line points to the condylar fora- 
men \ _fm foramen magnum ; HO basioccipital ; BS basisphenoid ; it sella turcica ; 
/'.V presphenoid. aiikylosed with the basisphenoid, forming the " body of the 
sphenoid ;" /'/ pterygoid ; PI palatine ; Vo vomer ; MX maxilla ; i symphysis of 
mandible ; cp coronoid process ; cd articular condyle ; a angle ; ih stylohyal, or 
' styloid process of temporal;" ch ceratohyal, or lesser cornu of hyoid ; bk 
basihyal, or body of hyoid ; tk thyrohyal, or greater cornu of hyoid. 


and in this young skull still separated from the basisphenoid 
in front by a vertical fissure. The basisphenoid (BS} is short 
and deep, and has a strongly marked pituitary fossa or " sella 
turcica" (sf) above. It has completely united with the pre- 
sphenoid (PS), though at birth the line of separation (below 
the spot called the olivary process or tubercuhim se/lce) is still 
visible. In adult age large air-cells fill the interior of this con- 
joined bone, which is the "body" of the so-called "sphenoid " 
of human anatomy. Anteriorly the presphenoid narrows to 
a sharp vertical edge, which is in contact with the mesethmoid 
(ME) above and the vomer ( Vo) below. The whole of the 
upper part of the mesethmoid is ossified in the specimen 
described, constituting the "lamina perpendicularis," but the 
anterior and lower part forms the septal cartilage of the nose. 
Its upper border forms a strong compressed triangular pro- 
jection into the cranial cavity, called the " crista galli " (eg). 
The posterior segment of the brain-case is completed, as 
in the Dog, by the pair of exoccipitals (ExO), and a large 
supraoccipital (SO). 1 The triangular upper part of the latter 
may be considered to represent the interparietal, though it 
very soon becomes incorporated with the rest of the supra- 
occipital. The middle segment is completed by large ali- 
sphenoids (AS), the "greater wings of the sphenoid bone," 
and enormously extended, somewhat square- shaped parietals 
(Pa) ; the frontal segment by narrow triangular orbito- 
sphenoids (OS), the " lesser wings of the sphenoid bone," 2 
and by large arched frontals (Fr). 

1 The "occipital bone" of human anatomy is formed by the coales- 
cence of the basioccipital, exoccipitals, and supraoccipital. 

2 The "sphenoid bone" of human anatomy is formed by the union 
of the basisphenoid, presphenoid, alisphenoids, orbitosphenoids, and 
the pterygoids. The basal portion ultimately ankyloses with the 

x.] MAN. 153 

Of the fossae into which the cranial cavity is divided, the 
olfactory fossa is very small, rather narrow, elongated, and 
shallow. The cribriform plate which closes it in front, in- 
stead of being vertical as in the Dog, is horizontal, and 
almost in the same line with the basicranial axis. It is 
bounded in the median line, and separated from the cor- 
responding fossa of the other side by the prominent crista 
galli of the mesethmoid. The middle fossa is, as before 
said, of comparatively enormous extent ; it is bounded pos- 
teriorly by the tentorial ridge, having the same relations to 
bones as in the Dog, but lying more horizontally and being 
far less prominent, having no osseous shelf like inward exten- 
sion. This fossa is distinctly divided into an anterior and 
posterior portion, by the strongly projecting hinder ridge of 
the orbitosphenoid The floor of the anterior portion is 
arched in consequence of the inward projection of the roof 
of the orbit, while the floor of the posterior or " temporal 
fossa " is deeply concave. The cerebellar fossa is of mode- 
rate si/e, and lies entirely underneath the hinder part of 
the cerebral fossa. 

The "sclla turcica," or depression in the basisphenoid for 
the lodgment of the pituitary body of the brain, is bounded 
posteriorly by an elevated transverse ridge, the corners of 
which are called the " posterior clinoid processes." Cor- 
responding processes projecting backwards from the orbito- 
sphenoids are called " anterior clinoid processes." 

The foramina in the base of the skull scarcely differ from 
those of the Dog. i. The olfactory has been already 
described. 2. The optic is a large round hole close to the 
inner and posterior part of the orbitosphenoid. 3. The 
sphenoidal fissure is larger than in the Dog. and produced 
externally into a long narrow slit. 4 and 5. The foramen 
rotundum and the foramen ovale pierce the alisphenoid, one 


near its anterior, the other near its posterior border. Close 
behind the last named is a small hole (" foramen spinosum "), 
through which a branch of the external carotid artery 
(middle meningeal) enters the brain-cavity. 6 and 8. The 
foramen lacerum medium basis cranii and the foramen 
lacerum posterius have the same functions and relation as 
in the Dog. 7. Between these two the periotic has the 
conspicuous meatus auditorius internus on its inner side- 
The depression above this, for the lodgment of the floc- 
culus, is distinctly seen in fcetal human skulls up to the 
time of birth, but it afterwards becomes gradually oblite- 
rated. 9. The condylar foramen perforates the exoccipital, 
as in the Dog; and lastly (10), the foramen magnum has 
the same general subcircular form, and is bounded by the 
same bones, but differs greatly in direction, its plane looking 
mainly downwards instead of backwards. 

The nasal cavities differ chiefly from those of the Dog in 
their shortness and greater vertical height. In their inner 
wall, the descending median plate of the vomer ( Vo) is 
much more developed. The pterygoids (Pi) are extended 
vertically, are narrow from before backwards, end below in 
a marked " hamular " process, and soon ankylose with the 
pterygoid plates of the alisphenoid anteriorly, but posteriorly 
are separated from them by a well-marked " pterygoid 
fossa." l The palatines (PI) and maxillae (Mx) are short 
from before backwards. The premaxillae (PMx) are small 
and early ankylosed with the maxillre. 2 The nasals (Net) 

1 The pterygoid, not being recognised as a distinct bone, is commonly 
described in works on human anatomy as " the internal pteryrroid plate 
of the sphenoid ; " the pterygoid process of the alisphenoid being the 
"external pterygoid plate." 

2 The premaxilla is a distinct bone in the human foetus, but is covered 
on its external or facial aspect by a process of the maxilla, which extends 
over it towards the middle line, and becomes completely fused with it 

x.] MAN. 155 

are short and nearly vertical, broad below and narrow 
above. The anterior nares are also nearly vertical. The 
turbinal bones are comparatively little developed, and of 
simple structure, especially the lower or maxillo-turbinal 
(MT). The flat bony plate on the outer side of the ethmo- 
turbinal or " os planum," instead of lying against the inner 
side of the maxilla, forms part of the outer wall of the nasal 
cavity and inner wall of the orbit, uniting with the frontal 
above, the lacrymal in front, the maxilla below, and the 
palatine behind. 

The group of bones placed around the organ of hearing, 
periotic, squamosal, and tympanic, though originally dis- 
tinct, become united together soon after birth to form the 
so-called "temporal bone." They differ from the corre- 
sponding bones in the Dog in the following particulars. The 
periotic has a very much larger mastoid portion (J/), which 
forms a considerable part of the wall of the cerebellar fossa. 
In the new-born infant its outer surface is smooth and flat, 
but as life advances, air-cells become developed within it, 
communicating with the tympanic cavity, and a strongly- 
marked descending projection, the " mastoid process," 
appears on the lower and anterior part of its outer surface. 
The squamosal (Sq) is a large flat vertical plate, forming a 
considerable part of the wall of the posterior cerebral fossa, 
behind the alisphenoid. Its zygomatic process is com- 
paratively slender and straight The tympanic forms a 
long tubular external auditory meatus, but its inner part 

before birth, so that no trace of the maxillo-premaxillary suture is ever 
seen on the outer side of the face. On the inner and palatal aspect of 
the bones the suture is always evident at birth, and can often be traced 
even in adult skulls. See G. IV. Callender " On the Formation and 
Early Growth of the Bones of the Human Face." (Phil. Trans. 1869, 
p. 163.) Also : P. Albrecht, " Sur la fente maxillaire et les os inter- 
maxillaires," etc. 1883. 


joins the periotic, forming the floor of the tympanic cavity 
without being inflated into an auditory bulla. Its under 
surface is produced into a rough ridge, to the inner side of 
which the large carotid canal perforates the base of the 
periotic, being directed obliquely forwards and inwards. 
In adult skulls the stylohyal often becomes ankylosed with 
the tympanic and periotic, constituting the " styloid process 
of the temporal bone." 

In examining the external aspect of the skull, the large 
smooth subglobular or oval brain-case, constituting by far the 
larger part of the whole cranium, is strikingly different from 
that of the Dog. The occipital surface, instead of being 
vertical, is nearly horizontal. The condyles, instead of 
being at the hindermost part of the skull, are not far from 
the middle of the base. The paroccipital processes of the 
exoccipitals are represented by mere rudiments, the so-called 
"jugular eminences"; on the other hand as before men- 
tioned, the mastoid processes, almost obsolete in the Dog, 
are very greatly developed. The occipital crest is repre- 
sented by a slightly raised and roughened line, the " superior 
curved line," and the sagittal crest is absent. 

The sutures connecting the bones of the upper surface 
of the cranium are remarkable for their wavy or indented 
character, processes from one bone interlocking with those 
from the other in a most complex manner, at least on the 
external surface, for seen from within they appear com- 
paratively straight and simple. There are very often 
irregular ossifications, separated from the contiguous bones, 
lying among the indentations of the occipito-parietal suture, 
called "Wormian bones." 1 The temporal fossa? are but 

1 In works on human anatomy, the occipito-parietal suture is com- 
monly called " lambdoid ; " the interparietal, "sagittal;" and the 
front j- parietal, " coronal." 

x.] MAN. 157 

indistinctly marked out by a curved line above, and are 
separated from each other by a wide expanse formed by 
the smooth rounded upper part of the parietal and frontal 
bones. The orbit is completely encircled by bone, the 
outer margin being formed by a process from the malar 
ascending to join the post orbital process of the frontal ; and 
it is, moreover, in great part sepaiated from the temporal 
fossa by an extension inwards of the ascending process of 
the malar meeting the alisphenoid, although a communica- 
tion is left between the two cavities below in the " spheno- 
maxillary fissure." The axis of the orbital cavity is directed 
more forwards than in the Dog. The face is altogether very 
much shorter, broader, and flatter. 

In the inferior surface of the skull the palate is seen 
to be much shorter and wider than that of the Dog, es- 
pecially anteriorly, where its outline forms an almost semi- 
circular curve. The maxillo-palatine suture is nearly 
straight transversely, and so is the hinder border of the 
palate, though produced backwards into an obtuse spine at 
the middle line. The distance between the hinder border 
of the palate and the foramen magnum is much shorter 
relatively, the space between the pterygoids being particularly 
short and wide. The true pterygoids and pterygoid plates 
of the alisphenoid are widely separated posteriorly, leaving 
a considerable fossa between them ; and the latter are larger 
and project further backwards than the former. The under 
surface of the tympano-periotic region is rough and irregular, 
instead of being smooth and bullate, and the perforation 
for the internal carotid artery is very conspicuous. There 
is no alisphenoid canal, scarcely any postglenoid process, no 
distinct glenoid venous foramen, a very small paroccipital, 
and a very large mastoid process. By the inclination of the 
occipital surface downwards, instead of backwards, an inferior 


view of the skull includes nearly all this surface, with the 
large foramen magnum and the condyles. 

In accordance with the general form of the face the 
mandible is short. The two rami of which it is originally 
formed unite together at the symphysis within a year after 
birth. They are widely divergent behind, and approach in 
front at a much more obtuse angle than in the Dog. The 
horizontal portion of each ramus is deep and compressed, 
the lower margin straight or slightly concave, and produced 
anteriorly rather in front of the alveolar margin so as to 
occasion the mental prominence characteristic of the human 
lower jaw. The anterior symphysial margin (Fig. 53, s), 
therefore, instead of sloping upwards from behind forwards, 
is vertical, or rather inclined in the other direction. Pos- 
teriorly, the condyle (cd) is more elevated than in the Dog 
and is less transversely extended. The coronoid process 
(cp) is smaller and less recurved. The posterior border, 
between the condyle and the angle (a\ is nearly straight 
and vertical, and the angle is rounded, compressed, slightly 
everted, and not produced into any hook-like process, as in 
the Dog. The depression for the masseter muscle is very 
faintly marked. 

The hyoidean apparatus differs in several particulars from 
that of the Dog. The tympanohyal can generally be 
recognised in the skull of an infant at birth and for a few 
years after, as a cylindrical piece of bone, with a truncated 
lower extremity, about one-twentieth of an inch in diameter, 
seated in a depression in the hinder border of the tympanic 
immediately to the anterior and inner side of the stylo- 
mastoid foramen. Its upper end becomes soon ankylosed 
with the periotic. The tympanic is produced around it 
anteriorly, constituting the " vaginal process." The stylohyal 
(s/i), at first a long styliform piece of cartilage, continuous 

x.] MAN. 159 

with the tympanohyal, commences to ossify by a separate 
centre before birth, and, at a very variable period after- 
wards, is often (but by no means constantly) ankylosed 
with the tympanohyal and surrounding cranial bones, con- 
stituting the so-called "styloid process." This is a con- 
dition not met with in any other Mammal. Below the 
stylohyal the greater part of the anterior hyoid arch is 
represented by a slender ligament (the " stylohyoid " liga- 
ment), there being no ossification corresponding to the 
Dog's epihyal ; but the ceratohyal (ch) to which the liga- 
ment is attached below, is a small bony nodule, the 
" lesser cornu of the hyoid " of human anatomy, which 
is articulated synovially to the upper corner of the 
outer extremity of the basihyal, though sometimes in old 
age becoming ankylosed. The basihyal (bh), or " body 
of the hyoid," is transversely oblong, hollowed posteriorly, 
and deeper from above downwards than in the Dog. The 
thyrohyals (///) or "greater cornua of the hyoid," are elon- 
gated, nearly straight and somewhat compressed. They 
usually become ankylosed before middle life with the outer 
extremities of the basihyal. 

The remainder of the true Primates (Simii/ia, or Monkeys) 
have the skull formed generally on the same plan as that 
of Man, with certain modifications in detail. 

The facial portion is enlarged and elongated as compared 
with the cerebral potion, though to a very variable extent 
in different members of the group. 

In nearly all, the brain-cavity maintains the same general 
form as in Man, though it is usually of less comparative 
vertical extent. With few exceptions, the middle compart- 
ment for the lodgment of the cerebrum retains its 
relative situation and superiority in size to the cerebellar 


and the olfactory fossae, completely overlying them both ; 
and consequently the occipital region of the skull with the 
foramen magnum behind, and the cribriform plate of the 
ethmoid in front, are in the same general horizontal line 
with the basicranial axis as in Man, and not perpendicular 
to them as in the Dog. 

It is remarkable that the deviations from this general rule, 
especially as regards the plane of the occipital surface, are 
not in relation to the general position of the animals in a 
descending series, from Man to the lowest Monkeys ; for the 
occipital surface is nearly vertical in the anthropoid Gibbons 
(Hylobatcs), especially // syndactylus (the Siamang), and 
completely so in the American Howling Monkeys (Mycetes), 
where the cerebral fossa does not project in the least degree 
behind the cerebellar fossa; while in the Baboons (Cyno- 
cephalus], among the Old World Monkeys, and still more 
in some of the smaller forms of American Monkeys 
(as Saimiris], the posterior development of the cerebral 
fossa is so great as to throw the supraoccipital bone con- 
siderably more into the posteriorly prolonged base of the 
skull even than in man. 

The olfactory fossa is always small. It is not only very 
short, but, in consequence of the considerable projection 
inwards of the portion of the frontal forming the roof of the 
orbit on each side of it, is both narrow from side to side, 
and deep from above downwards. 

In most of the Siwiina, including the Gorilla and Chim- 
panzee, the frontals meet along the middle line over the 
presphenoid, between the mesethmoid in front and thr 
orbitosphenoids behind ; but the Orang agrees with Man in 
wanting this postethmoid union of the frontals, and so also 
do some of the Cebidce. 

The fossa on the inner surface of the periotic for the 

x.] PRIMATES. 161 

floccular process of the cetebellum is almost obliterated in 
the adult Gorilla, Chimpanzee, Orang, and Gibbon, but is 
persistent and often very large in all other Monkeys. 

A partial ossification of the tentorium from the inner 
edge of the periotic takes place in some of the American 
Monkeys, as Mycdes and Cebns. 

The suture between the basisphenoid and the presphenoid 
remains distinct in the Baboons and all the lower Monkeys, 
until the animal has nearly attained its full size and acquired 
its permanent teeth ; but it is completely obliterated, and 
the cancellous structure of the two bones is continuous, in 
the Gorilla, Chimpanzee, and Orang, while the animal still 
retains all its milk-teeth. 

The nasal cavities, with their surrounding bones, are 
generally longer and of less vertical extent than in Man, 
but, as in the case of the inclination of the occipital plane, 
not following any regular serial descent. Thus the propor- 
tions of these parts are more like those of Man in many 
of the smaller American Ccbidie than in the long-faced or 
" Dog-headed" Baboons (Cynocefhali} of the Old World. 

The vomer is generally longer, and of less vertical extent 
than in Man. The turbinals have much the same general 
characters, their relative situation of course varying with 
the elongation, or otherwise, of the nasal passages. The os 
planum of the ethmoturbinals always forms part of the 
inner wall of the orbit, having the same relations as in 

The pterygoid plate of the alisphenoid is usually largely 
developed, and generally projects considerably backwards 
beyond the pterygoid bone (which is narrow, and has a very 
distinct hamular process), and there is always a wide and 
deep fossa between them. 

The premaxilla is ahvays distinct on the facial surface, 


and the suture between it and the maxilla is only obliterated 
in aged specimens. It generally extends upwards on the 
side of the anterior nares, so far as to meet the nasal and 
completely exclude the maxilla from taking any part in the 
boundary of this opening. 

The lacrymal foramen is never situated externally to the 
orbit, although, in the lower forms, it may be close upon 
the margin. 

As in Man the postorbital process of the frontal meets 
the orbital process of the malar so as completely to encircle 
the outer side of the orbit ; and an extension backwards and 
inwards of these bones joining the alisphenoid divides the 
orbit from the temporal fossa. 

The nasal bones vary much in length and breadth, but 
they present the peculiarity throughout the order of a great 
tendency to ankylose together in the middle line, even at a 
comparatively early age. 

In all the smaller and middle-sized Monkeys the general 
surface of the calvaria is oval and smooth, but in the larger 
Baboons and Orangs there are well-marked supraorbital, 
sagittal, and occipital ridges. These attain their greatest 
development in the adult male Gorilla, where they com- 
pletely mask the original form of the cranium. Their size, 
in this sex, appears to increase with age ; while in the 
oldest females, on the other hand, they are but slightly 

The paroccipital process is always rudimentary, as in 

The squamosal in the higher forms is developed much as 
in Man ; but in the lower forms it is more reduced, and 
takes a smaller share in the formation of the side wall of 
the cranium. It generally comes in contact, at its upper 
anterior angle, with the frontal, but not in the Orang or in 

x.] PRIMATES. 163 

the Cebida, in which animals the union of the parietal and 
the alisphenoid separates the frontal from the squamosal, as 
is usually the case with Man. The glenoid surface is 
flatter than in Man, and there is a well-marked postglenoid 

In the PlatyrrhincK) with the individual exception of 
certain Afycetcs and Aides, the parietal and malar bones are 
in contact with each other, separating the frontal from the 
alisphenoid on the side wall of the skull. In the Catar- 
r/rituc the reverse takes place, the alisphenoid and frontal 
bones meeting each other and separating the malar from 
the parietal. Another point of difference between the Old 
and New World Monkeys is that in the latter is a small 
slit in the malar wing not far from the point where the 
malar, parietal and alisphenoid bones meet. This slit, 
together with the infraorbito-temporal foramen, is the last 
remnant of a wide orbito-temporal foramen. 

The zygoma is usually narrow, horizontal, and slightly 
arched outwards. 

The periotic generally resembles that of Man, and the 
mastoid portion is conspicious on the outer side of the 
skull between the squamosal and the exoccipital ; but its 
surface is smooth and rounded, without any distinct mas- 
toid process. 

In all the Old World species, the tympanic forms an elon- 
gated inferior wall to the external auditory meatus, which 
has consequently a considerable bony tube ; but in all the 
American Monkeys this bone retains more or less its primi- 
tive annular condition, and the cavity of the tympanum is 
close to the external wall of the cranium. This character 
alone will readily serve to determine to which of the two 
great divisions of Monkeys a skull may belong. 

No auditory bulla is developed in any of the Old World 

M 2 


Monkeys, but in all the Cebidx and Hapalidtz the inferior 
surface of the ankylosed periotic and tympanic is somewhat 

The carotid canal is always very conspicuous, entering 
the under surface of the periotic near its hinder border. 
There is often a glenoid foramen, but never an alisphenoid 

The foramen rotundum perforates the alisphenoid, but 
the foramen ovale is usually a notch on its posterior border, 
completed by the periotic behind. 

The mandible presents the same general characters as 
that of Man, but the horizontal portion of the ramus 
is usually more elongated, and the anterior border slopes 
upwards and forwards, there being a complete absence of 
mental protuberance. The condyle is extended trans- 
versely, the coronoid process well developed and recurved. 
The posterior or ascending portion of the ramus is broad 
and flat ; the angle well developed, square, or more or 
less rounded, but without any special pointed process as in 
the Dog. 

In the Howling Monkeys (Mycetes) the hinder or ascend- 
ing portion of the ramus is remarkable for its extent both 
vertically and an tero- posteriorly, corresponding to a certain 
extent with the extraordinary development of the vocal 
organs, which it partially covers and protects. 

The Sim Una are remarkable in never, or very rarely, 
having an ossified stylohyal ; but on looking closely at the 
base of the periotic, immediately to the anterior and inner 
side of the stylomastoid foramen, a very small depression, 
in which there is sometimes a minute ossified tympano-hyal, 
can generally be seen. To this the ligament representing 
the stylohyal is attached. 

In very few of the Old World Monkeys is there any 




ossification in the anterior hyoid arch (see Fig. 54) ; but in 
some Cercopitheci a short, bony, cerato-hyal is found. This 
occurs also in the American Monkeys (Fig. 55), with occa- 
sionally the addition of a second piece (epihyal). 

The thyrohyals are always well-developed, long, narrow, 
nearly straight, and somewhat flattened. 

The basihyal varies much in form. In the anthropoid Apes 
it is broad transversely ; but in nearly all the other Monkeys 
its antero-posterior extent exceeds its breadth, owing to a 
great development from the posterior border. It is generally 
convex below, and concave above and behind, forming a 

FlG. 54. Inferior surface of hyoid boots 
of Baboon (CyniKepkalm pamirim,), 
bh basihyal ; th thyrohyal. 

FIG. 55. Inferior surface of hyoid bones 
of an American Monkey (Lago'hruc 
humbolJtii). tk thyrohyal ; c h cerato- 
hyal ; ck epihyal. 

considerable cavity in which the median laryngeal air-sac 
is lodged. This condition is enormously exaggerated in 
the American Howling Monkey (Mycetes), where the basi- 
hyal is transformed into an immense subglobular, thin- 
walled, bony capsule, with a large orifice posteriorly, by 
which the laryngeal air-sac enters, and having the straight 
narrow thyrohyals attached on each side. In this genus 
there are no ossifications in the anterior arch. 

While in nearly all the characters in which the skull of 
Man differs from that of the Dog, the Simiina agree with 


the former ; the Lemnrina, on the other hand, more resemble 
the lower type. 

In the Common Lemur the general proportions of face to 
cerebral cavity, and the inclination of the occipital and 
olfactory planes of the cranium, are quite dog-like. The 
orbits, although completely surrounded behind by the 
junction of the postorbital processes of the frontal and the 
malar, are yet perfectly continuous with the temporal fossa 
beneath this bony bar ; that extension inwards of the 
frontal and malar to meet the alisphenoid, and thus form a 
posterior external wall of the orbit, so characteristic ol 
Man and all Monkeys, being absent. The lacrymal fora- 
men, situated on the facial part of the bone, is altogether 
external to the margin of the orbit. The os planum of 
the ethmo-turbinal does not enter into the inner wall of the 
orbit, but is shut out from it by the maxilla, as in most 
inferior Mammals. The inferior surface of the tympanic is 
developed into a large rounded bulla. The hyoid apparatus 
much resembles that of the Dog, having the stylohyal, 
epihyal, and ceratohyal all distinctly ossified in the anterior 
arch, and the basihyal in the form of a narrow transverse 

Some of the Lemurina have much shorter faces than the 
common species, though still possessing all the essential 
characters of the group. Among these, Tarsius is remark- 
able for the extraordinary size of the orbits, which are so 
expanded that their margins form prominent, thin, bony 
rings, and the interorbital part of the skull is reduced to an 
exceedingly delicate septum. The orbit is also partially 
separated from the temporal fossa as in the Simiina. 

The general characteristics of the skull of the CARNIVORA 
have been described, as seen in the Dog. The more 


obvious modifications from this type relate to the compara- 
tive length and compression or width of the facial portion, 
the strength and curve of the xygomatic arch, and the extent 
to which the various ridges and processes for the attachment 
of muscles are developed. Thus the Cats have short and 
round skulls, with wide zygomatic arches ; and in the Bears 
(especially the Polar Bear, Ursus maritimus) the whole 
skull is elongated, and the nasal cavities are greatly enlarged 
as compared with the brain-case, and the maxillo-turbinal 
bones are correspondingly developed. 

But there are certain other modifications of the cranial 
bones, which, being less obviously adaptive to functional 
purposes, and being constantly associated with structural 
modifications in other parts of the body, are of considerable 
value in classifying the members of the group. Of these 
the most important are related to the form and structure of 
the auditory bulla, and the surrounding parts of the base of 
the cranium. 

In the Bears, the auditory bulla is comparatively little 
inflated. It consists of a single bone (tympanic), readily 
detached from the cranium in skulls of young animals. Its 
form is more or less triangular, being broad and nearly 
straight at the inner edge, and produced outwards into a 
considerably elongated floor of the external auditory meatus. 
Its greatest prominence is along the inner border; from 
this it gradually slopes away towards the meatus. The 
entrance of the carolid canal is a considerable circular 
foramen, near the hinder part of the inner edge of the bulla. 
In old animals it is partly concealed by the prominent lip of 
the basioccipital, which abuts against the inner edge of the 
bulla ; and by the growth of this, and the paroccipital pro- 
cess, it becomes almost included in the deep fossa leading 
to the foramen lacerum posterius. When a section is made 

1 68 



through the auditory bulla (see Fig. 56) it is seen to be a 
simple thin-walled bony capsule, imperfect above, where it 
fits on to the petrosal and squamosal bones, and prolonged 
externally into the much thickened spout-like floor of the 
meatus externus. At the inner extremity of this floor is a 
freely projecting oval lip (/), which gives attachment to the 
membrana tympani, and which is the original and first ossified 
ring-like portion of the tympanic bone. In the front of the 


FIG. 56. Section of the left auditory bulla and surrounding bones of a Bear (Ursns 
fcrox). Si/ squamosal; T umpanic; BO basioccipital ; am external auditory 
meatus ; t tympanic ring ; c Eustachian canal ; Car carotid canal. (From Proc. 
Zool. Soc. 1869.) 

floor of the bulla is the groove for the Eustachian canal (e) : 
between this and the inferior part of the tympanic ring, a 
low and thin ridge of bone with a concave free margin rises 
from the floor of the cavity. This is the only indication of 
any septum or division of the cavity of the bulla. 

Behind the bulla, the prominent and tuberous paroccipital 
process projects downwards, outwards and backwards, stand- 

x.] CARNIVORA. 169 

ing quite ofl from the bulla, and only connected with it by a 
low laterally compressed ridge. Between the paroccipital 
process and the occipital condyle is a smooth concave surface 
the front of which is excavated into a deep notch, the 
posterior boundary of the foramen lacerum posterius, 
between which and the condyle is situated the condylar 
foramen, which transmits the hypoglossal nerve. At the 
outside of the bulla, just behind the external auditory 
mcatus, the mastoid process is distinct and prominent, and 
widely separated from the paroccipital. There is a very 
conspicuous glenoid foramen situated just behind the post 
glenoid process of the squamosal. 

All the Ursidtf, Procyonida, and Mustelida agree with 
the true Bears in the general characters of this region of the 
skull ; for even when (as in some of the smaller species) the 
auditory bulla is considerably dilated, it always has its 
greatest prominence near the middle of the inner border, and 
gradually slopes away from this point to a prolonged floor of 
the auditory meatus ; and though there are often trabecula; 
or partial septa passing mostly transversely across the lower 
part, 1 there is no distinct and definite septum dividing it 
into a separate outer and inner chamber. In all cases, on 
looking into the external auditory meatus (in the dried 
skull when the membrana tympani is removed) the oppo- 
site wall of the bulla can be seen ; or if a probe is passed 
into the meatus, no obstacle will prevent its touching the 
inner wall. 

In the Tiger, which may be taken as a type of the Felidce, 
the auditory bulla is very prominent, rounded and smooth 
on the surface, rather longer from before backwards than 
transversely, its greatest prominence being rather to the 

1 Especially developed in the Weasels (Mustela], in which also the 
entire parieties of the bulla are thickened and cancellous. 




inner side of the centre. The lower lip of the external 
auditory meatus is extremely short ; the meatus , in fact, 
looks like a large hole opening directly into the side of the 
bulla. On looking into this hole, at a very short distance 
(just beyond the tympanic ring), a wall of bone is seen quite 
impeding the view, or the passage of any instrument, into 
the greater part of the bulla. In the section (Fig. 57) it will 
be seen that this wall is a septum (s), which rises from the 

f the left auditory bulla of the Tiger (t'rlis tieris). 

mosal ; t't penotic ; />'(' basioccipital ; am external auditory meatus ; 

chamber ; ic the inner chamber : s th 

plum ; * the aperture of communication 

between the chambers, (from J J roc. Zool. Soc. 1869.) 

floor of the bulla along its outer side, and divides it almost 
completely into two distinct chambers ; one (cv), outer and 
anterior, is the true tympanic chamber, and contains the 
tympanic membrane and ossicula, and has at its anterior 
extremity the opening of the Eustachian tube (<>) ; while the 
other (/>), internal and posterior, is a simple but much 
larger cavity, having no aperture except a long but very 
narrow fissure (*) left between the hinder part of the top of 

x.] CARNIVORA. 171 

the septum and the promontory of the periotic, which 
fissure expands posteriorly, or rather at its outer end, into a 
triangular space, placed just over the fenestra rotunda, so 
that the opening of this fenestra is partly in the outer and 
partly in the inner chamber of the bulla. This chamber is 
formed by a simple capsule of very thin but dense bone.- 
deficient only at a small oval space in the roof, where the 
periotic projects into and fills up the gap, except such 
portion of it as is left to form the aperture of communica- 
tion with the outer chamber. 

Not only are these two chambers thus distinct, but they 
are originally developed in a totally different manner. At 
birth the only ossification in the whole structure is the in- 
complete ring of bone supporting the membrana tympani, 
and developed originally in fibrous tissue. Ossification 
extends from this, so as to complete the outer chamber, 
and the very limited lip of the meatus auditorius externus. 
The inner chamber is formed from a distinct piece of hya- 
line cartilage, which at birth is a narrow slip, pointed at 
each end, lying between the tympanic ring and the basi- 
occipital, applied closely to the surface of the already 
ossified periotic, and forming no distinct prominence on the 
under surface of the skull. Soon after birth this increases 
in size, and gradually assumes the bullate form of the wall 
of the inner chamber. In young animals, even some time 
after the ossification of the bulla is complete, the distinc- 
tion between the two parts is clearly seen externally ; not 
only are they marked off by a groove, but the tympanic 
portion has a more opaque appearance than the other. The 
septum is formed by an inversion of the walls of both ap- 
plied together, and ultimately perfectly fused in Felts, although 
remaining permanently distinct in some of the Virerrida. 

The carotid foramen in the TigQr is only represented by 


a minute groove deep in the recess of the foramen lacerum 
posterius. In the smaller Cats, this groove is more super- 
ficial, but always very minute, and apparently never converted 
into an actual foramen, except by the contiguous wall of the 

The paroccipital process is flattened over the back of the 
bulla, being applied closely to the whole of its prominent 
rounded hinder end, and projecting, as a rough tubercle, 
slightly beyond it. From the inner side of this process a 
sharp ridge runs towards the occipital condyle. This forms 
the posterior boundary of a deep fossa, at the bottom of 
which is the foramen lacerum posterius, and in the hinder 
part of which, under cover of the aforesaid ridge, the 
condylar foramen "opens. The mastoid process is a 
moderately conspicuous rough projection, not very widely 
separated from the paroccipital. There is no distinct 
glenoid foramen. 

The Viver rider, agree with the Felidce in having the auditory 
bulla divided into two cavities by a bony partition, in having 
the paroccipital spread over the hinder surface of the bulla, 
and in having no prolonged external auditory meatus ; but 
the bulla is more elongated and compressed, and the inner 
chamber is placed altogether behind the outer or true 
tympanic chamber. 

In the Hy?ena this region of the skull much resembles the 
same part in the Cats, but the bulla is simple and undivided. 
In the Dogs there is a partial septum, and otherwise the 
characters are intermediate between the two extremes of the 
Bears on one side, and the Cats on the other. 1 

1 For the various modifications of the structure of this part of the 
skull in the different genera of the order, see "On the Value of the 
Characters of the Base of the Cranium in the Classification of the Order 
Carnivora." (Proc. Zool. Soc. 1869, p. 5.) 

x.] CARNIVORA. 173 

In nearly all the Carnivora the hyoidean apparatus is con- 
structed on the same plan as described in the Dog, having a 
narrow, transversely elongated, curved basihyal, either round 
or compressed from above downwards, a nearly straight 
compressed thyrohyal, not ankylosed with the basihyal, and 
a well-ossified anterior cornu, composed of three distinct 
pieces of subequal length. In the Lion, Tiger, and Leopard, 
however, the anterior arch is imperfectly ossified, the dif- 
ferent bones being small, and connected together by long 
ligamentous intervals ; but in the Puma, Cheetah, Lynx, and 
Cat, the bones are large, and in close relation with each 

In the Seals the brain-cavity is very broad, round, and 
rather depressed. The orbits are large, and the interorbital 
portion of the skull greatly compressed. The olfactory 
chambers of the nasal cavities are consequently very narrow, 
and the ethmo-turbinals little developed ; but in front of the 
orbits the cavities widen, and the maxillo-turbinals are very 
large and complex. The mesethmoid is of considerable 
vertical extent, and its ossified portion, which is extensive, 
terminates anteriorly in a straight vertical line. In some 
forms, as Cystflp/tora, this ossification extends in front of the 
nasals. The hyoid resembles that of the typical Carnivora, 
all the elements being well ossified and distinct. The 
Otariidir, or Eared Seals, also called Sea Bears or Sea 
Lions, are intermediate in most of their cranial characters 
between the true Seals and the Bears. 

The skulls of the animals composing the Order INSEC- 
TIVORA present great variations. 

In some of the higher forms, as Galeopithccus, Tupaia, 
Macroscelides, and Rhynchocyon, the cranium much resembles 
that of the Ltmurina, having a considerable and vaulted 


cerebral cavity, large orbits, nearly vertical occipital plane, 
large olfactory fossre, a well-developed zygoma tic arch 
sending up a postorbital process to meet a corresponding 
one from the frontal so as either partially or completely to 
encircle the orbit behind, and tympanics ankylosed with 
the other cranial bones, dilated into a bulla, and pro- 
duced externally into a tubular auditory meatus. The 
face is generally elongated, and narrow anteriorly, but in 
Galcopithccus it is broad and depressed. 

In the Macroscelidce, or Elephant Shrews, the auditory 
meatus is very large, the tympanic bulla much inflated, and 
there are sometimes also very large mastoid bulla:. 

In Tupaia, the malar has a large, oval, longitudinal per- 
foration. There are also in this genus, as in some of the 
other Insectivora, vacuities in the palate, arising from defects 
of ossification, like those found in many Marsupials. 

In the remaining Insectivora the cranial cavity is of small 
relative size. The orbit and temporal fossa are completely 
continuous, and there is often not even a trace of a post- 
orbital process to the frontal or malar. 

In the Erinaceidce (Hedgehogs) and the Ccntetida' the 
tympanic is a mere ring unankylosed to the surrounding 
bones, but a kind of bulla is formed by a lamella pro- 
jecting from the basisphenoid to join its inner and inferior 

In Erinaccns and Gymmtra the zygomatic arch is com- 
plete, but slender, and formed chiefly by the processes of the 
maxilla and squamosal, "which meet each other, the malar 
being a small splint-like bone attached to the outer and 
under side of the middle of the arch. In the Ccntetidtc the 
malar is entirely absent, and, as the zygomatic processes of 
both maxilla and squamosal are very short, there is no bony 
arch. Centetes (the Tenrec) has a remarkably elongated 


and narrow skull (both cranium and face partaking of the 
same character), with very prominent occipital and sagittal 
crests. Both in this genus and in Erinaceus (the Hedge- 
hog) the mesoptervgoid fossa at the base of the skull is 
very deep, and ends posteriorly in a hemispherical depression 
in the basisphenoid, between the wing-like processes which 
abut against the inner wall of the tympanic. Both parocci- 
pital and mastoid processes are also well developed. 

The Moles (Tatyida:) have an elongated and depressed 
cranium, broad posteriorly, and gradually narrowing to the 
muzzle. The occiput slopes upwards and forwards, the 
supraoccipital being greatly developed. The zygomatic 
arches are complete, but very slender, and show no dis- 
tinct malar bones. There are no postorbital or paroccipital 
processes. A lamelliform expansion of the upper edge of 
the periotic (pterotic, Parker) forms part of the lateral wall of 
the cranium, as in the Echidna. The tympanic is united 
with the other bones of the cranium to form a flattened 
bulla, produced into a short meatus with a small external 
opening. There is a small " prenasal " ossicle at the anterior 
extremity of the mescthmoid cartilage, as in the Pig. 
Myogah agrees with the Shrews in having no zygoma nor 
postorbital process. 

In the Cape Golden Mole (Chrysochloris) the cranium is 
conical, very broad and rounded behind, and pointed in 
front. There are no postorbital processes. The zygoma is 
complete, and tolerably strong. The tympanics ankylose 
with the skull, and form a completely ossified bulla. 

In the Shrews (Soriddie) the cranium is broad behind and 
tapering forwards. The facial portion is long and narrow. 
The occiput slopes much forwards. There is no zygoma 
and no postorbital process. The postglenoid process of the 
squamosal is remarkably large. The tympanic is ring-like, 


and there is a large unossified space on each side of the 
base of the skull. 

The mandible in the Insectivora has generally an elongated 
and rather narrow horizontal portion, above which the 
transversely extended condyle is but slightly elevated, and 
there are well-developed coronoid and angular processes ; 
the latter is remarkably long and slender in the Shrews. 

The hyoid is formed generally like that of the Carnivora, 
with three complete extracranial ossifications in the anterior 
arch, a transversely extended basihyal, and tolerably long, 
stout, flattened thyrohyals, sometimes ankylosed with the 
basihyal. 1 

Order CHIROPTERA. In the large Frugivorous Bats 
(Pteropus), the cranium is generally elongated, the cerebral 
cavity large, oval, arched above, and contracted in front : 
its walls formed mainly by the greatly expanded parietals, 
both supraoccipital and frontals being small. In old indi- 
viduals of some species there are well-marked sagittal and 
occipital crests. The base of the cranium is elongated, flat, 
and thin. The facial part is long and rather compressed. 
The postorbital processes of the frontals are long and 
pointed, and partially define the orbits behind ; but there 
is usually no corresponding ascending process from the 
zygomatic arch, which is long and slender, and mainly 
formed by the processes of the maxillary and squamosal, 
the malar being a splint-like bone attached to their under 
and outer surface. The lacrymal foramen is situated outside 
the margin of the orbit. The nasals are long and narrow, 
and often ankylose together in the middle line. The pre- 
maxillse are small. The palate is elongated backwards : 

1 For details of the osteology of the Insectivora, see G. E. Dobson's 
valuable monograph of the group, now in course of publication. 

x.] CHIROPTERA. 177 

the horizontal plates of the palate bones being large and 
early united in the middle line, without defects of ossifica- 
tion. The pterygoids are small. There is no alisphenoid 
canal. The glenoid fossa is broad and shallow ; the post- 
glenoid process very little developed, and with a venous 
foramen behind it. The tympanics are very slightly con- 
nected with the neighbouring bones, and are consequently 
nearly always lost in macerated skulls. A wedge-shaped 
portion of the mastoid appears on the outside of the skull 
between the squamosal and the exoccipital. The par- 
occipital process is long and rather slender and directed 
downwards and backwards. The periotic has a large and 
deep fossa for the flocculus on its inner side. 

FIG. 58. Hyoid bones of Frugivorous Rat (Ptervfus) from below. M bosihyal ; 
ty thyrohyal ; sh stylohyaL 

The mandible has a high, broad, recurved coronoid pro- 
cess, a transversely extended condyle, and a flattened rounded 
angle, without a distinct process. 

The hyoid (Fig. 58) has a narrow, transversely extended 
basihyal, with which the elongated, laterally compressed and 
curved thyrohyals are commonly ankylosed. The anterior 
cornu contains three slender ossifications of nearly equal 

The Insectivorous Bats generally have the skull shorter 
and broader than the Pteropi. The cranial cavity in many 
species is almost globular, with thin smooth walls, though 
sometimes sagittal and occipital crests are developed. The 
occipital foramen is very large. The zygoma is slender. 
Postorbital processes are sometimes well developed, but 


more often small and rudimentary. The face is usually 
short and broad, in some (as Mojinops] bent upwards on the 
cranium in a remarkable manner, so that the plane of the 
palate is nearly perpendicular to the basicranial axis. The 
premaxilke are generally small, sometimes not meeting in 
the middle line, and sometimes (as in Megaderma) altogether 
wanting. The tympanics are annular, not ankylosed to the 
surrounding bones, nor prolonged into a bony canal ex- 
ternally, though often developing a partial bulla on their 
inner side. 

The mandible has a distinct angular process. 

Order RODENTIA. In the Rodentia the cerebral cavity is 
generally elongated, depressed, somewhat broad behind and 
narrow anteriorly. The occipital plane is more or less 
vertical : the cerebellar fossa altogether behind the cerebral, 
and the tentorial plane, or division between these fossae 
approaching the vertical. The anterior part of the cerebral 
fossa is contracted ; the olfactory fossa is of moderate size, 
and situated directly in front of the cerebral. 

The nasal cavities are very large, and both sets of 
turbinals well developed, including an upper or nasoturbinal 
lamella. The olfactory chambers attain their maximum of 
development in some of the Porcupines (JfystrL-c], where 
nearly all the bones of the upper part of the cranium are 
expanded by great air sinuses formed within their walls. In 
the Hare, and some others, the two optic foramina in the 
orbitosphenoids are confluent ; and in consequence, in the 
dried skull, there is a direct aperture of communication 
between the orbits above the craniofacial axis. 

The supraoccipital is more or less vertical, and does 
not extend far on to the upper surface of the cranium. 
There is often a distinct interparietal. There are generally 

x.] RODENTIA. 179 

moderately developed paroccipital processes, which in the 
Capybara (Hydrocharus) are of great length, curving for- 
wards and compressed laterally. They are also very large 
in the Coypu (Myopotamus). 

The parietals are moderate or small. The frontals, except 
in the Squirrels, Marmots, and Hares, have little more than 
a rudiment of a postorbital process, and there is never any 
marked corresponding process arising from the zygoma, so 
that the orbit is perfectly continuous with the temporal fossa. 
The latter is always very small. 

In a very remarkable East African genus, Lop/iiowys, a 
broad bony lamella extends from the upper part of the 
parietal outwards and downwards to join a similar ascending 
plate from the malar, and thus forms an arched covering 
to the temporal fossa, an arrangement unknown in any 
other Mammal, but recalling that met with in the Tortoises. 
The whole of the superior surface of the cranial bones of 
this animal are covered with miliary granulations, disposed 
with perfect regularity and symmetry. 1 

The nasals in Rodents are both long and wide, and 
generally extend so far forwards as to make the anterior 
nares quite terminal and vertical, or even with a downward 
inclination. In Hystrix their development is enormous, but 
this is chiefly owing to their breadth and backward extension 
over the great nasal chambers and air sinuses. They are 
narrowest in Jiathyergus and its allies. 

The premaxillai are large, and lodge the great curved 
incisor teeth,'- and always send a narrow prolongation back- 
wards by the side of the rasals to join the frontals. 

1 See A. Milne-Edwards, Xour. Archiv. du Museum, iii. 1867. 

2 These teeth, though first developed in the gum covering the pre- 
maxilla, have their roots, when fully developed, in the maxilla. This 
does not invalidate their determination as incisors. 

N 2 




In the larger number of Rodents there is a great vacuity 
in the anterior or maxillary root of the zygoma of varying size 
and form, apparently an enormous dilatation of the infra- 
orbital foramen, and through which a portion of the masseter 
muscle passes (see Fig. 59). It is sometimes as large as the 
orbit itself, with which it communicates freely posteriorly 
underneath a vertical bar, formed by the maxilla in front, and 
by the lacrymal and malar behind. Its inferior boundary 




A a nasal; P\Ix premaxilla ; 3Ix maxilla ; Ma, malar; 7V tympanic ; ExO ex- 
occipital ; Per points to the large supratympanic or mastoid bulla. 

is a slender zygoma-like horizontal bar, formed by the maxilla 
alone. In the Rats it is a vertical fissure dilated superiorly. 
In the Viscacha (Lagostomus), the true infraorbital foramen 
is separated from the large antorbital vacuity by a thin as- 
cending bony lamella. In Castor, Leptis, Bathyergus, Sa'urus, 
Arctomys and some others, the infraorbital foramen is of the 
usual size. 

In the Hares, the facial surface of the maxilla is curiously 

x.] RODENTIA. 181 

The zygoma is present in all, of various degrees of thick- 
ness, but always either straight or more or less curved 
downwards, and usually not much arched outwards. Its 
anterior and posterior roots are formed by the maxilla and 
squamosal respectively, the malar intervening. In many 
cases the last-named bone extends backwards, applied to the 
under surface of the zygomatic process of the squamosal to 
form the outer side of the glenoid articular surface. In 
the spotted Cavy (Caelogcnys), the zygoma has an enormous 
vertical expansion, with a rugose or pitted outer surface, 
and a large fossa in the inner side of the maxillary portion 
with which the cavity of the mouth communicates in a 
recent state. 

The lacrymal bone usually presents both orbital and 
facial surfaces, but the orifice of the canal (lacrymal foramen) 
is always well within the margin of the orbit. In the Beaver, 
and many others, the facial portion is reduced to a mere 
tubercle, and in the Hare the lacrymal is entirely within 
the orbit. 

The j>alate of the Rodents is usually narrow. In the long 
space intervening between the incisor and molar teeth, it 
has no definite lateral margins, but rounds off insensibly on 
to the sides of the face. In this region the anterior palatine 
foramina form very conspicuous longitudinal slits, of specially 
large size in the Hares. The portion of the palate situated 
between the molar teeth is often very narrow anteriorly 
and ends posteriorly in a thickened excavated border. In 
the Hare it is reduced to a short transverse bridge, ex- 
tending across the middle line between the premolar teeth. 
In the Capybara and Guinea Pig the alveolar border of the 
maxilla is very long, and presents the remarkable peculiarity 
of extending backwards beneath the orbit to unite with the 
squamosal at a level with the anterior border of the glenoid 


fossa. It thus forms the outer border of a large conical 
cavity, opening posteriorly, bounded on the inner side by 
the pterygoids, above by the alisphenoids, and below by the 

The pterygoids are always simple subquadratc lamella;, 
early ankyloscd with the basisphenoids, often sending a 
well-marked hamular process backwards, which unites with 
the auditory bulla in Hystrix, Lagostowtts, Bathycrgits, &c. 
There are usually well-marked ptcrygoid fossas, and the sides 
of the alisphenoids are often perforated by an alisphenoid 
canal. In Hystrix the hamular process is slightly bullate. 
The squamosal forms a considerable part of the outside wall 
of the cranium, but in consequence of the large size of the 
united tympano-periotic, the root of the zygomatic process is 
thrown very forward on the side of the skull, and the pos- 
terior part of the body of the squamosal which unites with 
the occipital is reduced to a long, rather narrow strip, 
interposed between the parietal and pcriotic. When the 
mastoid bulla; are greatly developed, as in Pcdetcs (see Fig. 
59), this does not reach as far backwards as the occipital, 
and is a slender curved process, which clasps the outside of 
the bulla, and appears to hold it in its place. 

The glenoid fossa is situated on the under side of the 
posterior root of the zygoma. In its most typical form (as 
in the Capybara, Viscacha, Aguti, Paca, etc.) it is narrow and 
concave transversely, with prominent inner and outer edges, 
the latter being often formed, as before mentioned, partly by 
the hinder end of the malar. In the Beaver the glenoid fossa 
has considerable breadth. In the Porcupines, Marmots, 
Squirrels, Rats, etc., no raised inner margin is developed, 
and the fossa passes insensibly into the side of the skull wall. 
In the Hare it is a transversely oval hollow, with a prominent 
rounded anterior margin. 

x.] RODENTIA. 183 

The tympanic is ankylosed to the periotic, but not to the 
squamosal ; it generally develops a tubular meatus, which in 
the Hare is directed upwards and backwards, in the Beaver, 
outwards and forwards. In the Porcupines, as well as in 
most of the smaller Rodents, the meatus is short. In the 
Capybara it is fissured below. 

There is always a considerable tympanic bulla, which is 
often supplemented by a bulla developed above the tympanic 
cavity apparently in the periotic. In some genera (Pedetes, 
Difus, Chinchilla'] this attains an enormous size (see Fig. 
59, Per), and forms a rounded prominence on the posterior 
external angle of the skull, interposed between the squamosal, 
parietal, and occipital. Usually, the mastoid portion of the 
periotic only appears on the surface for a small space in front 
of the exoccipital. In the Beaver, it forms a conspicuous 
angular process. 

The periotic is never ankylosed with any of the bones of 
the cranium, other than the tympanic. On its inner surface 
the floccular fossa is nearly always wide and deep, but it is 
absent, or nearly so, in the Capybara, Paca, and Porcupine. 
The place of attachment of the hyoid arch is an inconspicuous 
depression in the usual situation, and the tympanohyal is 
never distinct. This is in relation with the rudimentary 
condition of the anterior cornu of the hyoid. 

In the mandible, the symphysial portion is narrow, curving 
upwards, and rounded laterally, with a single large alveolus 
on each side. The coronoid process is never large, and is 
often rudimentary or absent, in relation to the small develop- 
ment of the temporal fossa and muscle ; while the portion 
adjacent to the angle is greatly developed, showing marked 
masseteric and pterygoid fossae, and often with its lower 
edge expanded laterally, or slightly incurved. The angle is 
rounded in the Hares, but it is more often produced into a 

1 84 THE SKULL. [CHAP. x. 

long backward process, more or less pointed and upturned. 
The condyle is considerably elevated, with a rounded 
articular surface, usually longer from before backwards than 
from side to side. 

The hyoid has a transversely extended basihyal and a 
straight compressed rod-like thyrohyal, often ankylosed with 
the basihyal. The anterior arch is long, but mostly liga- 
mentous, the ossification being usually confined to the lower 
part (cerato- and epi-hyals). In the Hares, the basihyal is 
deep from above downwards, and compressed, keeled, or 
pointed in front. 



Order UNGULATA ; Sub-order Perissodactyla. In the 
Horse the whole skull is greatly elongated, chiefly in con- 
sequence of the immense size of the face as compared with 
the hinder or true cranial portion. The basal line of the 
skull from the lower border of the foramen magnum to the 
incisor border of the palate is very nearly straight. The 
occipital and ethmoid planes are nearly perpendicular to this 
line, the latter inclining slightly forwards. The tentorial 
plane, strongly marked by inward projecting ridges of bone, 
slopes obliquely backwards at an angle of 45. The cerebral 
fossa is a smooth and regular oval, broad and rounded in 
front, and with no distinct division into anterior and pos- 
terior portions. The olfactory fossa is short, but deep from 
above downwards. The pituitary fossa is very shallow, and 
there are no distinct clinoid processes. The alisphenoid is 
very obliquely perforated by the foramen rotundum, but the 
foramen ovale is confluent with the large foramen lacerum 
medium behind. There are considerable frontal and sphen- 
oidal air sinuses, but the former do not extend any great 
distance over the brain-cavity. 


In front of the cerebral cavity, the great tubular nasal 
cavities are provided with well-developed turbinal bones, and 
are roofed over by very large nasals, broad behind, and ending 
in front in a narrow decurved point. The opening of the 
anterior nares is prolonged backwards on each side of the 
face between the nasals and the elongated slender premaxillo3. 
The latter expand in front, and are curved downwards to form 
the semicircular alveolar border which supports the large 
incisor teeth. 

The orbit is rather small in proportion to the size of the 
whole skull, but very distinctly marked, being completely 
surrounded by a strong ring of bone with prominent edges. 
The lacrymal occupies a considerable space on the flat 
surface of the cheek in front of the orbit, and below it the 
malar does the same. The latter sends a hori/ontal or 
slightly ascending process backwards below the orbit, to 
join the under surface of the /.ygomatic process of the 
squamosal, which is remarkably large, and instead of ending 
as usual, behind the orbit, runs forwards to join the greatly 
developed postorbital process of the frontal, and even forms 
part of the posterior and inferior boundary of the orbit a 
very exceptional arrangement (see Fig. 60). 

The palate is very narrow in the interval between the 
incisor and molar teeth, in which are situated the large 
anterior palatine foramina. .Between the molar teeth it is 
broader, but it does not extend further back than the pen- 
ultimate molar and ends in a rounded excavated border. It 
is mainly formed by the maxillai, as the palatines are very 
narrow. The pterygoids are delicate slender slips of bone 
attached to the hinder border of the palatines, and supported 
externally by, and generally ankylosed to, the rough pterygoid 
plates of the alisphenoid, with no pterygoid fossa between. 
They slope very obliquely forwards, and end in curved, 




compressed, hamular processes. There is a distinct alis- 
phenoid canal for the passage of the internal maxillary artery. 
The base of the cranium is long and narrow. The glenoid 
surface for the articulation of the mandible is greatly extended 
transversely, concave from side to side, convex from before 
backwards in front and hollow behind, and is bounded 

Fir,. 60. Side view of the posterior part of the skull of a Horse, j. Fr frontal (the 
line points to the jxisturbita! process); Sq squamosal ; 1'a parietal: SO supra- 
occipital; ExO cxoccipital : oc occipital condyle : // paroccipital process; /Vr 
nia-.ti.iil portion of periotic ; // post-tympanic process of squamosal : th tympano- 
hyal ; Ty tympanic : f>g postglenoid process of squamosal ; As alisphenoid (inc- 
line points to the plate of the bone which bridges over the alisphenoid canal); Ma 

posteriorly, at its inner part, by a prominent postglenoid 
process (Fig. 60, j>g). 

The squamosal (S</) enters considerably into the forma- 
tion of the temporal fossa, and besides sending the zygo. 
made process forwards, it sends down behind the meatus 
auditorius a post-tympanic process (//), which aids to 
hold in place the otherwise loose tympano-periotic bone. 

1 88 



Behind this the exoccipital gives off a very long paroccipital 
process (//). 

The periotic and tympanic are ankylosed together, but not 
with the squamosal. The former has a wide but shallow 
floccular fossa on its inner side, and sends backwards a 
considerable " pars mastoidea " which appears on the outer 
surface of the skull (Per) between the 
post-tympanic process of the squamosal 
and the exoccipital. The tympanic (Ty} 
forms a tubular meatus, directed out- 
wards and slightly backwards. It is 
not dilated into a distinct bulla, but 
ends in front in a pointed styliform 
process which forms part of the outer 
wall of the Eustachian canal. It com- 
pletely embraces the truncated cylin- 
drical tympanohyal (///), which is of 
great size, corresponding to the large 
development of the whole anterior arch 
of the hyoid. 

The stylohyal (Fig. 61, s/i) is of great 
size, compressed, and expanded at the 
upper end, where it sends off a tri- 
angular posterior process. Below the 
FIG. 61. Superior surface stylohyal, and usually becoming anky- 

of hyoid bones of horse, , , . , . ,, , , , 

i. sA stylohyal ; ch cera- losed with it, is a small nodular bone 

tohyal ; bh basihyal ; tk / > i\ 11 .1 , 

thyrohyal. ( 6 P ln y a V> ar >d tnen t' 16 ar h IS COm- 

pleted by a short cylindriform cera- 
tohyal (c/i). The basihyal (M) is rather flattened from 
above downwards, arched with the concavity behind, and 
sends forwards a long, median, pointed, compressed 
" glossohyal " process. The thyrohyals (///) are compressed 
bars projecting backwards from, and in adult animals 


completely ankylosed to, the lateral extremities of the 

Each ramus of themandible has a long, straight, compressed 
horizontal portion, gradually narrowing towards the symphysis, 
where it expands laterally to form with the ankylosed opposite 
ramus the wide semicircular, shallow, alveolar border for 
the incisor teeth. The region of the angle is expanded and 
compressed, with a thickened rounded border without any 
process. The condyle is greatly elevated above the alveolar 
border ; its articular surface is very wide transversely, and 
narrow and convex from before backwards. The coronoid 
process is slender, straight, and inclined backwards. 

The skull of the Rhinoceros resembles that of the Horse 
in many essential features, but the occipital region is of 
much greater extent vertically, the form of the cranial cavity 
being concealed externally by large occipito-parietal air- 
cells. There is no postorbital process to the frontal, so 
that the orbit is not divided from the temporal fossa. 
There is a conspicuous, rough, antorbital projection on the 
lacrymal bone just in front of the lacrymal foramen 
The nasals are very large and strong, early ankylosed to 
gether, arched from before backwards, and pointed anteriorly. 
The most elevated part of their upper surface is roughened, 
and supports the great median horn which characterises 
the genus. In some species a posterior rough, "but less 
elevated, surface indicates the attachment of a second 
horn. In some of the extinct species the mesethmoid 
cartilage was ossified nearly as far forwards as the extremity 
of the nasals, which is not the case with any existing 
species. The premaxilloe are very small, and do not extend 
anteriorly beyond the level of the front end of the nasals. 
The hinder border of the palate is deeply excavated, the 


horizontal plates of the palatines being very narrow. The 
pterygoids are very slender, as in the Horse, but placed more 
vertically. There is a distinct alisphenoid canal. The 
squamosal sends down a very long, conical, postglenoid 
process parallel with, and equalling, or sometimes exceeding, 
in length, the paroccipital process. In all existing two- 
horned species (7?. sumatrensis, bicornis and simus) the 
meatus auditorius lies in a deep groove between the post- 
glenoid and the post-tympanic processes of the squamosal. 
In the one-horned species (R. unicornis and sondaicus) these 
processes unite below so as to completely surround the 
meatus. The post-tympanic process articulates with the 
exoccipital, completely excluding the mastoid from the 
external surface of the skull. 1 

The tympanic and periotic are ankylosed together, but 
not with the squamosal. They are both very small. The 
under surface of the tympanic is rough, forms no distinct 
bulla, and is much encroached upon posteriorly by the very 
large tympanohyal, which presents a circular, slightly concave, 
rough, inferior surface, half an inch in diameter (in an adult 
Sumatran Rhinoceros). Externally, the tympanic is pro- 
duced into a rough, irregular, inferior wall to the auditory 
meatus. The periotic internally shows the internal auditory 
meatus near its lower part, but no distinct depression for 
the flocculus ; it is prolonged upwards and outwards into 
a small mastoid portion, which, as before said, does not 
appear on the outer surface of the skull. 

The mandible has a very wide condylar articular surface, 
and slender recurved coronoid process, a rounded, somewhat 
incurved angle, a compressed, rather narrow, horizontal por- 
tion, and a shallow depressed symphysis. 

1 See W. PI. Flower, " On seme Cranial and Dental characters of the 
existing species of Rhinoceros." (Proc. Zool. Soc. 1876.) 


The hyoid is much like that of the Horse, and has a 
glossohyal process from the middle of the basihyal. 

The Tapirs present some singular modifications of the 
same type of skull. 

As in the Rhinoceros, there is no separation between the 
orbit and temporal fossa, but the anterior nares are of 
immense size, and extend backward above the orbits, being 
separated from them only by a thin plate of bone instead 
of a broad flat surface, as in the Horse and Rhinoceros. 
The nasal bones are short, broad behind, pointed in front, 
much elevated, and supported by a tolerably well ossified 
mesethmoid, which spreads out laterally at its upper end. 1 
The inferior and lateral margins of the great narial apertures 
are entirely formed by the maxillae, which extend up to meet 
the nasals, the premaxillx not taking any share in them. The 
ethmo-turbinals are small, while the maxillo-turbinals, on the 
other hand, are very extensive, though their plications are 
comparatively simple. A conspicuous feature in the upper 
part of the face is a groove, which extends backwards on 
the side of the dilated hinder end of the nasal bone, and 
curves inwards to form a rounded depression over the naso- 
frontal suture. The form and size of this depression vary 
in different species. It lodges an air sinus, with cartilaginous 
walls extending upwards from the nasal chamber. In front 
of the nares the rostrum formed by the maxillsi with the 
premaxillna in front is produced, compressed anteriorly, and 
curved downwards. 

The base of the cranium resembles generally that of 
the other Perissodactyla. There is an alisphenoid canal, and 

1 In one species ( T. Dairdii) the ossification of the mesethmoid extends 
far in advance of the nasal bones, and is clasped and suppDrted below 
by ascending plates from the maxillae. 


large post-glenoid and post-tympanic processes ; the latter 
joins the paroccipital process of the exoccipital, but above 
their point of union a narrow slip of the mastoid appears 
on the surface of the skull. The periotic is not ankylosed 
to the squamosal or to the tympanic, which is exceedingly 
rudimentary, forming a small irregular floor to the tympanic 
cavity, with an oval lip for the attachment of the membrana 
tympani, and always becomes detached in macerated skulls. 

The mandible is chiefly noticeable for the great rounded 
incurved posterior projection of the angle. 

The hyoid has a simple, elongated stylohyal without pos- 
terior process at the upper end, but one other ossification in 
the anterior cornu, and no glossal process to the basihyal. 

Sub-Order Artiodadyla. The skull of the Sheep, as one 
of the best known and easily procurable examples of this 
group, may be first described, though in some respects it 
is rather peculiarly modified. 

On comparing the section of the cranium (Fig. 62) with 
that of the Dog, it will be seen that there is a great difference 
in the relation of the principal elements to each other, inas- 
much as the face is bent downwards on the basicranial axis, 
so that when the latter is horizontal, the upper surface of 
the face looks forwards and the palate backwards. The 
occipital foramen is terminal posteriorly, the tentorial plane 
nearly vertical, so that the cerebellar fossa is altogether 
behind the cerebral, but the plane of the cribriform plate 
is horizontal, and the olfactory fossa altogether beneath the 
anterior portion of the cerebral fossa. 

The occipital region is small and sloping forwards. 
There are long paroccipital processes (//). In very young 
skulls a distinct interparietal bone is present, but in the 
specimen figured this has coalesced with the supraocci- 
pital (SO). The two parietals (Pa) unite very early at the 




sagittal suture. The frontals (Fr) are large, and usually 
(except in some domestic races) develop from their outer 
surface conical, curved, bony processes, cancellous within, 


FIG. 62 Longitudinal and vertical section of skull of a Sheep (Otn's artts), i. 
I'Mt premaxilla ; MT roaxillo-turbinal ; Na nasal; E'J' ethmolurbinal ; ME 
mcsethinoid ; hr frontal ; OS orbitosphenoid : AS alisphenoid ; t'a parietal : 
SO supraoccipital ; Per periotic ; ExO exoccipital ; BO basioccipital ; fp par- 
occipital process ; Ty styhform process of tympanic; US basisphenoid ; PS pre- 
sphenoid ; Pt pter>'goid ; /'/ palatine ; Vo vomer ; MX maxilla ; cp coronoid 
process ; rd condyle ; s symphysis of mandible ; sA stylohyal ; eh epihyal ck 
ccratohyal ; M basihyal; th thyrohyal. 

which are called the " horn cores," as they form the internal 
support of the true horns. The nasals (Na) are long and 
pointed in front. The premaxillse (PMx) are slender, with 


a shallow alveolar border, bearing no teeth and forming the 
anterior and lateral boundaries of large anterior palatine 
foramina. The lacrymals are large, and form a considerable 
portion of the side of the face in front of the orbit, but the 
foramen is entirely within the margin. 

The olfactory chamber is larg *. The turbinals are greatly 
developed ; the upper lamina of the ethmoturbinal or " naso- 
turbinal " is distinct, and extends over the scroll-like maxillo- 
turbinal (MT\ but does not ankylose with the nasal. 

The orbit is large, nearly circular, with a complete, 
prominent margin, formed below by the large malar, which 
extends considerably on the side of the face, and posteriorly 
sends a process upwards to meet the postorbital process of 
the frontal, and is continued backwards to join the zygomatic 
process of the squamosal. 

The palate bones (PI) are of moderate, extent ; their 
horizontal plate is deeply notched posteriorly. The pterygoids 
(Pt) are broad above, but end below in a narrow lamella, 
with a hamular process projecting backwards. The basi- 
occipital (13O), seen from below, is square, with eminences 
for muscular attachments at each of its four angles. The 
basisphenoid (S) is much contracted laterally. The pos- 
terior clinoid processes are large, and the pituitary fossa 

The squamosal is small, and scarcely appears in the interior 
of the skull. The glenoid facet is rather extensive, and 
slightly convex, and there is a postglenoid process and 
foramen. The tympanic is not ankylosed to the periotic ; it 
forms a complete tubular external auditory meatus, and a 
considerable, but simple, bulla, narrowing to a sharp-pointed 
process anteriorly (Ty). The periotic (Per) is rather small, 
without any fossa for the flocculus ; its mastoid portion 
forms a distinct, narrow, rough surface on the outer side of 


the skull, between the hinder border of the squamosal and 
the exoccipitaL The tympanohyal is very large, cylindrical, 
curved, and almost completely embedded in the tympanic, 
between the inferior wall of the meatus and the outer wall 
of the bulla. 

The extracranial portion of the hyoid consists of large 
compressed stylohyals (sh), with a prominent posterior 
process near the upper end, short but well-ossified epi- 
hyals (eh] and ceratohyals (ch), and a basihyal represented 
by a small rounded nodule of bone, to which the straight 
thyrohyals (//*) are not ankylosed. 

The mandible has a broad flat condyle (cd], a long slender 
coronoid process (r/), a rounded angle, a rather slender 
horizontal portion, contracted and with a sharp upper edge 
in front of the molar teeth, and expanded anteriorly for the 
lodgment of the incisors. 

The Ox agrees generally with the Sheep in its cranial 
characters. The face is bent down on the basicranial axis 
almost in the same manner. The occipital surface is flat, 
and terminates above in a broad transverse ridge, which 
extends between the horn cores. The parietals are ex- 
tremely narrow above, and placed almost entirely behind 
this ridge. They unite very early with the interparietal and 
supraoccipital. The intercornual ridge of the frontals is 
excavated by large air-cells, communicating with those of 
the horn cores, and is especially developed when the horns 
are large. Unlike the parietals, the frontals are of very 
great extent, and have a broad and flattened upper surface. 

The tympanics are compressed and scarcely at all bullate. 
They end anteriorly in long compressed styliform processes, 
and become firmly ankylosed with the periotic and squa- 
mosal. The under surface of the meatus auditorius has a 

o 2 

i 9 6 THE SKULL. [CHAP. 

compressed ridge. The large tympanohyals are entirely em- 
bedded in the tympanic, only the rough lower surface for 
articulation with the stylohyal being exposed. 

The mandible and hyoid are like those of the Sheep, but 
the basihyal is rather more developed, and has a rounded, 
anterior, median projection. 

Many Ruminants (especially among the Cervidcc} have a 
vacuity of varying extent on the side wall of the face, 
between the frontal, lacrymal, maxillary and nasal bones, 
leading in the macerated skull into the nasal chamber, but 
closed in the living animal by membrane. Most of the 
Deer and Antelopes have also a large depression on the 
facial surface of the lacrymal bone, called the suborbital or 
lacrymal fossa, though it has nothing to do with the tears, 
but lodges a glandular fold of the integument, which secretes 
a peculiar unctuous and odorous substance. In most Deer 
the orifice of the lacrymal canal is double, and situated on 
the margin of the orbit, whereas in most of the hollow- 
horned ruminants it is single and placed well within the 
margin. There are however exceptions in both cases. 

In the Deer the axis of the face is nearly in the same 
line with that of the cranium, so that when the basicranial 
axis is horizontal the nose is directed forwards instead of 
downwards, as in the Sheep and Ox. The animals of this 
family have no permanent horn cores continuous with the 
cranium and ensheathed by true horns, but have short pro- 
cesses on the frontal bones (pedicles], from which branching 
antlers of true osseous structure are annually developed and 
shed. These, as a rule, are only present in the males, 
while the horns of the Bovida and Antilopida are usually 
common to both sexes. 
_. Among the Antelopes, the Saiga (Saiga tartaricd) is very 


remarkable for the conformation of the upper part of the 
face. The anterior nares extend backwards almost to a 
level with the front edge of the orbits, and have an unusual 
vertical expansion. The nasal bones are aborted or coa- 
lesced with the frontals. The turbinals are very short. 
The lacrymals enter largely into the side walls of the 
anterior nares. The ascending processes of the premaxillse 
are small, and very widely separated from the nasals. In 
the living animal the edges of these greatly expanded narial 
apertures are continued forwards into a truncated, almost 
proboscidifonn muzzle without any bony support, giving the 
contour of the face a totally different appearance from that 
presented by the skull. 

In the Elk (Alces), and a small Abyssinian Antelope 
(Neotragus saltiand), the nasal bones are very much shorter 
than they are in ordinary ruminants. 

The Tylopoda (Camels and Llamas) and the Tragulina 
differ from most of the Pecora^ and resemble the non-rumi- 
nating Artiodactyles in having the tympanic bulla filled with 
cancellated bony tissue. 

The skull of the Pig shows in section that the axis of the 
face is bent down upon the basicranial axis almost as much 
as in the Sheep, a disposition which increases with age. 
Though the form of the cranial cavity is not very different 
from that of the Sheep, the external appearance of the 
hinder-part of the skull is greatly changed by the elevated 
and backward sloping occipital crest, formed by the union 
of the supraoccipital (concave from side to side posteriorly) 
and the parietals. The latter have their outer and inner 
surfaces widely separated in the adult Pig by large air-cells. 

The frontal is broad and flat between the orbits, and 
sends out a small postorbital process, which does not join the 


zygoma. The face is greatly elongated, tapering forwards, 
and compressed laterally. The nasals are long and narro\v, 
and the apertures of the nares small and nearly terminal. 
The premaxillae send up long processes on each side of 
the nasals, which, however, do not meet the frontals. The 
lacrymal has a considerable facial portion ; and, as in 
other Ungulata, the malar encroaches considerably on the 
face, uniting with the lacrymal. 

At the anterior extremity of the mesethmoid a peculiar 
ossicle (prenasat) is developed, which strengthens the cartil- 
aginous snout. 

The palate is long and narrow, and extends posteriorly 
beyond the last molar tooth. The pterygoid fossae are well 
marked, being chiefly formed by the well-developed ptery- 
goid plates of the alisphenoid ; the true pterygoids are very 
slender. There are very long, slender, compressed par- 
occipital processes, curved forwards. 

The squamosal and tympanic are ankylosed together ; the 
floor of the long, narrow, upward-directed auditory meatus 
is formed by the tympanic, wedged in a cleft of the squa- 
mosal, between the hinder edge of the glenoid fossa (there 
being no postglenoid process) and a long descending post' 
tympanic process which articulates with the exoccipital. 

Inferiorly the tympanic is dilated into a very prominent 
bulla, peculiarly elongated vertically, and rather compressed 
from side to side. The interior of this bulla is filled with 
cancellous bony tissue. 

The periotic is small and not ankylosed to the tympanic 
or squamosal. The mastoid portion is quite rudimentary, 
being merely a short scale-like prolongation upwards and 
backwards, lying on the inner surface of the squamosal, and 
making no appearance on the external surface of the skull. 
The tympanohyals are very inconspicuous, being small, and 

xi.] ART10DACTYLA. 199 

situated at the bottom of a deep fossa on the outer and 
posterior side of the tympanic bulla. 

The mandible has a high ascending portion behind, a 
transverse condyle, a very small coronoid process, and a flat 
expanded angle, rounded posteriorly. 

The hyoid of the Pig is very different from that of most 
other Ungulata. The basihyal is very small. The thyrohyals 
are large, broad and flat, and ankylosed to the basihyal, but 
with their extremities cartilaginous even in old animals. 
There is a well-ossified ceratohyal, not ankylosed with the 
basihyal, but the greater part of the anterior arch is a long 
cartilaginous band, with one, or sometimes two, slender 
ossifications near the middle part, representing the stylohyal. 

The skull of the Hippopotamus resembles that of the 
Pig in many essential features, although its external form is 
greatly modified. The brain cavity is very small, and the 
face immensely developed. The orbits project outwards in 
an almost tubular manner, and their margins are nearly, and 
in some cases quite, complete posteriorly. The face is con- 
tracted laterally in front of the orbits, and then expands 
widely into a massive truncated muzzle, which supports the 
great canine and upper incisor teeth. 

The anterior narial orifice is nearly circular ; it is bounded 
by the extremities of the narrow but greatly elongated 
nasals above, and laterally by the prominent, rounded, 
rugged and massive premaxillae. At the anterior and lower 
part of the orbit, the lacrymal is dilated into a great thin- 
walled bony capsule, of such delicacy that it is nearly always 
destroyed in the skeletons preserved in museums. This 
opens into the nasal air-passages and has no connection 
with the lacrymal apparatus. A similar but smaller dilatation 
exists in many of the Pecora. 


The palate is long and narrow, and extends posteriorly a 
short distance behind the last molar teeth. The internal 
pterygoids end in well-marked stout hamular processes. The 
glenoid surface of the squamosal is very much extended, 
but not bounded externally by a projection from the malar 
as in the Pig, and the inner half of its posterior margin is 
produced into a tolerably well marked postglenoid process. 
The paroccipital process is long and conical, but far less 
conspicuous than in the Pig. The tympanic bulla is pro- 
portionately smaller, and of a trihedral form, ending in an 
antero-inferior pointed process. Its interior is filled with 
cancelli, as in the Pig. As in that animal, there is a long 
narrow meatus auditorius, directed upwards and backwards 
in a fissure between the postglenoid and post-tympanic 
processes of the squamosal, the floor being formed by 
a compressed, ridged prolongation of the tympanic, 
which is at a very early age completely fused with the 
squamosal. The periotic is very small, remains longer 
distinct, though ultimately ankylosing with the conjoined 
squamoso-tympanic, and has only a rudiment of a mas-, 
toid portion, which is quite confined to the interior of the 

The tympanohyal is slender, ankylosed to the back of the 
tympanic, and in the adult skull sunk in a deep fossa, 
between that bone and the exoccipital, which also gives exit 
to the facial nerve. 

The mandible is of immense size and weight. The 
condyles rise very little above the level of the molar teeth. 
The coronoid process is small and much recurved. The 
angle is greatly expanded and everted, rounded behind, and 
terminating below in a distinct process, projecting down- 
wards and forwards. The horizontal rami are compressed 
in their middle portion, but widen anteriorly into a very 

xi.] HYRAC01DEA. 201 

broad and massive truncated symphysial portion, which 
supports the huge incisor and canine teeth. 

In the hyoid apparatus, the basi- and thyro-hyals anky- 
lose, and are something like those of the Pig. The an- 
terior arch consists of three well-ossified pieces of subequal 

HYRACOIDEA. The skull of the Hyrax presents many 
affinities with that of the Perissodactyla, others with the 
Rodentia, and some characters peculiar to itself. 

The cranium is high and truncated behind, the occiput 
nearly vertical, the tentorial and olfactory planes oblique, 
the olfactory fossa rather small. 

There is a small distinct interparietal. The frontal region is 
broad and flat. The zygoma is tolerably strong, and consti- 
tuted mainly by the malar, which extends so far backwards as 
to form the outer wall of the glenoid fossa, but it is supported 
anteriorly by a strong process from the maxilla. The orbit 
is bounded posteriorly by well-marked postorbital processes 
which sometimes meet, the lower one from the malar, and 
the upper one from the parietal (a very unusual condition). 
The lacrymal is small, and scarcely extends at all on to 
the face, but sends outwards a strong antorbital process (as 
in the Rhinoceros and Elephant). The face is short, and 
compressed laterally. The nasal bones are wide posteriorly, 
and anteriorly are either truncated, or more produced at 
their outer than their inner margins. The premaxillae do 
not send up processes to meet the frontals, as in all 

The palate is not produced posteriorly beyond the middle 
of the last molar tooth. The palate bones are large. The 
ptervgoids very slender. There are well-marked pterygoid 
fossse, and alisphenoid canals. The paroccipital processes 


are long and slender. The glen old fossa is wide transversely, 
and with a considerable postglenoid process. The periotic 
and tympanic are ankylosed together, but usually remain 
distinct from the squamosal. The tympanic forms a 
moderate-sized bulla, and a spout-like floor to the external 
auditory meatus, between the glenoid and post-tympanic 
processes of the squamosal. The periotic has a very slight 
floccular depression, and sends backwards no distinct mas- 
toid process. The pituitary fossa is very shallow, without 
clinoid processes. The foramen rotundum and foramen 
ovale are distinct perforations through the alisphenoid. The 
optic foramen pierces the large orbitosphenoid near its 
hinder margin. 

The mandible has a high and exceedingly broad ascending 
portion, its hinder margin being produced far behind the 
condyle, but the angle is rounded, and without any distinct 
process. The condyle is much extended transversely, and 
narrow from before backwards, especially in its inner half, 
for externally it is somewhat rounded. The coronoid pro- 
cess is small and recurved. 

The hyoid apparatus of the Hyrax s unlike that of any 
other known Mammal. The basihyal is oval, transversely 
extended and flat, with a small median eminence on its 
anterior border, and an emarginate posterior border, only 
ossified in the centre, and prolonged laterally, without any 
definite segmentation, into broad, flattened, slightly curved 
cartilaginous thyrohyals. Articulated to the anterior and 
external angles of the basihyal are two large, triangular, 
flattened bones (ceratohyals), each with a long process 
projecting forwards and meeting in the middle line, so 
as to enclose (with the anterior margin of the basihyal) a 
triangular space. There is no other cartilage or bone in the 
anterior arch, unless a very minute pyramidal bone, described 


by Brandt as articulating with the mastoid process of the 
skull, represents the stylohyal. 1 

PROBOSCIDEA. The skull of the only existing animals of 
this group, the Elephants, presents many very remarkable 
features. As the brain-case increases but little in size 
during growth, and as the exterior wall of the skull is required 
to be of great superficial extent to support the trunk and the 
huge and ponderous incisor teeth or rusks, and to afford 
space for the attachment of muscles of sufficient size and 
strength to wield the skull thus heavily weighted, an extra- 
ordinary development of air-cells takes place in the can- 
cellous tissue between the outer and inner surface walls or 
" tables " of nearly all the bones of the cranium, separating 
them in some cases as much as twelve inches apart (see 
Fig. 64). These cells are not only formed in the walls of the 
cranium proper, but are also largely developed in the nasal 
bone and upper part of the premaxilla and maxilla, the bones 
forming the palate and the basicranial axis, and even 
extend into the interior of the ossified mesethmoid and 
the vomer. Where two originally distinct bones come in 
contact the cells pass freely from one to the other, and 
almost all the sutures become completely obliterated in old 

The intercellular lamellae in the great mass which sur- 
rounds the brain-cavity superiorly and laterally mostly 
radiate from the inner to the outer table, but in the other 
bones their direction is more irregular. Like the similar but 
less developed air-cells in the skulls of many other Mammals, 
they are entirely secondary to the original growth of the 
bones. In the young African Elephant's skull figured (from 

1 "Mem. de 1'Acad. Imp. de St.-Petersbourg," Vile Serie, tomexiv. 
No. 2, p. 68 (1869). 


an animal about six months old, see Fig. 63), their formation 
has scarcely commenced, and as the sutures are still quite 
distinct, and the bones not distorted by these cellular 
dilatations, they are in a much better state for studying 
their connections and characteristics. 

When the basicranial axis is placed in a horizontal position, 
it will be seen that the foramen magnum is quite posterior, 
and its plane nearly vertical. The cranial cavity is elongated 
and depressed (more so in the African than the Indian 
Elephant), the tentorial plane nearly vertical, so that the 
cerebellar fossa is altogether behind the cerebral fossa. The 
latter is broad behind and contracted laterally in front. The 
olfactory fossa is large, and placed altogether below the 
anterior part of the cerebral fossa, the cribriform plate being 
nearly horizontal. The ridge which separates the anterior 
from the posterior division of the cerebral fossa is very well 
marked. The pituitary fossa is very shallow, and there are 
no distinct clinoid processes. The supraoccipital (SO) is 
high, and inclines greatly forwards ; so that the occipital sur- 
face looks upwards as much as backwards. In the adult skull 
(Fig. 64) the lateral parts of the occipital region (rounded 
smoothly off in the young state) are vastly expanded and 
leave between them a deep median depression, with a rugged 
floor, and a partial bony septum at the bottom, into which 
the ligamentum nuchae is inserted. The median portion of 
the supraoccipital never becomes expanded by air-cells. 
The parietals (Pa) are very large, and form the greater part 
of the lateral walls of the cranium. The frontals (Fr) are 
narrow from before backwards, and produced laterally into 
elongate supraorbital processes, which send out small post- 
orbital processes, not, however, completely separating the 
small orbit from the large and high temporal fossa. 

Jb Sir 





FIG. 63. A section of the cranium of a very yvung African Elephant (ElrfJuu 
a/rjfanns). taken s tj the right of the middle line, so that the (me^cih- 
inoid and vomcr are removed, 4- The letters as in the other figures. 

FIG. 64. A section of the cranium of a fiill-gr^wn African Elephant, taken to the left 
of the middle line, and including the vomer (Yo), and the mesethmoid (ME), 
an anterior, and tit posterior narial aperture. i'j. 


The most remarkable feature in the face is the form and 
position of the anterior narial orifice (an). It is wide trans- 
versely, very short from above downwards, placed very high, 
and is directed upwards and forwards, almost as much as in 
the Whalebone Whales. The nasal bones (No) which bound 
it above are very thick, short, broad behind, and conical 
in front, and contain large air-cavities. The inferior and 
lateral margins of the orifice are formed entirely by the pre- 
maxillas (PMx), which send processes up to join the nasals 
and frontals. In front of the nares the face is prolonged 
into a somewhat quadrate, depressed, alveolar process, trun- 
cated in front, concave above, rounded laterally, formed by 
the premaxillse above and at the sides, and by the maxilla; 
below. This contains the roots of the great incisor teeth 
or tusks. 

The iacrymal is small, placed almost entirely within 
the margin of the orbit, and ends anteriorly in a projecting 
antorbital process. The zygomatic arch is slender and 
straight, the malar being small, and forming only the middle 
part of the arch, the anterior portion of which is (unlike 
that of all Ungulates) formed by the maxilla. 

The elongated, tubular nasal cavity forms a sigmoid 
curve, being directed (from below) at first forwards, then 
upwards, then forwards. The olfactory chamber is a 
comparatively small fossa in the middle third of its posterior 
wall, filled by the complex ethmoturbinals. The maxillo- 
turbinals are but rudimentary, the narial passage being quite 
free. 1 The floor of the palate is completed posteriorly by 
well-developed palatines (PI). The pterygoid (Pt) is slender 
and very early ankylosed with the pterygoid process of the 

1 The elongated proboscis probably supplies their place functionally 
in warming and cleaning the inspired air. 


alisphenoid, which is greatly expanded, and hollowed in 
front, being spread round the dilated posterior margin of 
the alveolar portion of the maxilla, and aiding to close the 
great alveolar cavity of the hindermost molar tooth. 

The squamosal (Sy) forms a considerable part of the 
cranial wall, extending outside the small alisphenoid to meet 
the frontal, and externally sends off a broad post-tympanic 
process, which meeting (though not uniting with) the hinder 
border of the glenoid fossa in front, bounds the bony ex- 
ternal auditory meatus, to which the tympanic contributes 
very little. The latter bone is, in the young specimen, com- 
pletely united with the periotic, but not with the squamosal. 
Inferiorly it forms a large, rounded, but not very prominent 
auditory bulla, deeply notched on its inner side by the 
canal for the internal carotid artery (a). The periotic 
(/Vr) presents a large surface within the cranium without 
any floccular fossa. The mastoid portion is very small, and 
does not appear on the surface of the cranium. There are 
no paroccipital or postglenoid processes. At the bottom 
of a deep fossa between the squamosal, exoccipital, and 
tympanic, the tympanohyal is distinctly seen, with the 
stylomastoid foramen to its outer side. The exoccipi- 
tals are not perforated by a condylar foramen, neither 
is the alisphenoid perforated, but it is grooved in front 
for the foramen rotundum, and behind for the foramen 

The mandible is of a very peculiar shape. The ascending 
portion of the ramus is high, and terminates in a rather 
small rounded condyle, wider from side to side than from 
before backwards. The posterior border is thick, but 
rounded off gradually into the inferior edge, without any 
projection at the angle. The coronoid process is compressed, 

208 THE SKULL. [CHAP. xi. 

and but very little elevated. The horizontal portion is very 
massive and rounded to support the great molar teeth ; it 
unites with its fellow in front in a narrow, prolonged, spout- 
like symphysis. 

The stylohyals are forked at their upper extremity, the 
posterior process being greatly developed. They taper 
below to a point which is connected by a long ligament with 
the basihyal. The thyrohyals are long, compressed, and 
ankylosed to the basihyal. 1 

3 See A. H. Garrod, Proc. Zool. Soc. 1875, P- 3^5- 



Order CETACEA. The animals of this order exhibit some 
remarkable modifications in the characters of the skull. 

I will first select for description that of a young example 
of one of the Odontoceti or Toothed Whales, the common 
round-headed Dolphin or Globictphalus of our coasts. Fig. 
65 represents a vertical median section of this skull. It 
will be seen that the cerebral cavity is of a very unusual 
shape, being short and broad, but extremely high and con- 
tracted above in fact, somewhat in the form of a truncated 
cone, with rounded edges. The bones of the basicranial 
axis are curved upwards at each extremity. They consist 
of the ankylosed basioccipital (BO) and basisphenoid (BS) 
separated by a vertical fissure from the presphenoid (PS) 
and mesethmoid (M), which are also ankylosed, though 
their original line of separation can still be traced. The 
pituitary fossa scarcely forms a distinct concavity, and the 
clinoid processes are almost obsolete. The mesethmoid is 
very large, and consists of (i) a high and broad vertical 
plate, which closes in the vacuity between the frontals in 
the anterior part of the cerebral cavity, and corresponds to 


the cribriform plate of the ordinary Mammal, though with 
but few and small perforations ; (2) an anterior rod-like, 
somewhat compressed, pointed prolongation from the lower 
part of this plate, which extends forwards in the groove of 
the vomer ( Vo) almost to the extremity of the rostrum, and 
which in great part remains permanently cartilaginous. 
This corresponds with the septal cartilage of the nose of 
other Mammals, although owing to the altered position of 
the nares it has here little relation with these passages. 

The cranial cavity is formed chiefly of the cerebral fossa, 
the cerebellar fossa being relatively small, and the olfactory 
fossa entirely wanting. 

The optic nerve passes out through a deep notch, some- 
times a foramen, in the hinder border of the orbitosphenoid. 
The alisphenoid is not perforated, the foramen rotundum 
being confluent with the large sphenoidal fissure, and the 
foramen ovale with a large infundibuliform opening between 
the alisphenoid, parietal, exoccipital, basioccipital, and basi- 
sphenoid, in the bottom of which is seen the inner surface 
of the periotic (Per), which in the Cetacea makes no projec- 
tion into the cerebral cavity. The anterior part of this 
opening corresponds to the foramen lacerum medium with 
the foramen ovale, the hinder part to the foramen lacerum 
posterius. 1 The squamosal (,5V;) appears in the outer 
boundary for a very small space, between the parietal and 
the exoccipital. The condylar foramen pierces the ex- 
occipital, near its anterior edge. The large or nearly 
circular carotid canal has a peculiar position, passing 
through the basisphenoid, near its middle, in a direction 
from below upwards, forwards, and inwards. 

1 In the adult of the same species, the foramen ovale is separated 
from the large opening common to the seventh and eighth pair of nerves 
by a strong bony partition formed by the ossified tentorium cerebelli. 




The bones forming the walls of the cranial cavity are dis- 
posed in a very remarkable manner. The occipital surface 
is of great size, and slopes upwards and forwards. The 
foramen magnum is large, and looks directly backwards ; its 





FIG. 65. A section of the skull of a young Dolphin (Globictfikalui nielas), |. PMx 
premaxilla ; MX m.ixill.i ; ME ossified portion of the mesethmoid ; an anterior 
nares ; t<ln nasal ; //' interparietal ; /> frontal ; Pa parietal ; SO supraoccipital ; 
EjcO exoccipital ; KO basioccipital : Sq squamosal ; Per periotic ; AS alisphenoid; 
PS presphenoid ; Pt pterygoid ; />n posterior nares ; PI palatine ; Vo vomer : 
* symphysis of mandible ; id inferior dental canal ; ef cjronjid process ; erf con - 
dyle ; a angle ; sh stylohyal ; bk basihyal ; th thyrohyaL 

lower lateral margins are bounded by large oval condyles, 
which meet in the middle line below, and are formed by 
the exoccipitals, with a small portion of the basioccipital. 
Above the foramen, the immense supraoccipital (SO), with 
which an interparietal (IP) is ankylosed, extends forwards 


beyond the vertex, to be wedged in between the frontals, 
completely excluding the parietals from the upper region of 
the cranium. These latter (Pa) form the greater part of the 
sides of the narrow high temporal fossae, and are ankylosed 
with the supraoccipital above, although the different elements 
of the occipital are still distinct. The frontals (Fr) are broad 
from side to side, being prolonged outwards into the arched 
supraorbital plates, but are almost entirely covered by lamel- 
liform extensions of the maxillae, which leave but a thin 
strip of the frontals visible on the external surface of the 
cranium. The temporal fossa is bounded below and in 
front by a stout postorbital process of the frontal, very nearly 
meeting the broad zygomatic process of the squamosal. The 
orbit is elongated from before backwards ; at its anterior 
extremity is a rounded antorbital prominence, formed by 
the junction of the maxillae, frontal and malar ; below, it is 
bounded by a long and very slender styliform zygomatic 
process of the malar, which arises from near the anterior and 
inner angle of the body of the bone, and passes backwards, 
slightly curved downwards, to articulate with the extremity 
of the zygomatic process of the squamosal. There is no 
distinct lacrymal bone, or canal. 

The special modification of the bones of the face has 
relation chiefly to the peculiar position of the nasal passages, 
which instead of passing forwards above the roof of the 
mouth to the anterior extremity of the face, are directed 
upwards and somewhat backwards towards the vertex of the 
cranium ; the external narial orifices being situated quite on 
the top of the head, the part which first appears above the 
surface of the water when the animal rises for the purpose of 
respiration. The whole nasal cavities are small, and they are 
(as far as concerns their bony walls) simple canals, entirely 
destitute of turbinals. Though their direction is in the main 

xii.] CETACEA. 213 

vertical, they are not straight, but curve round the anterior 
end of the brain cavity, both upper and lower orifices (an and 
/) being directed somewhat backwards. They are com- 
pressed from before backwards above, but wider and more 
round below. The nasal bones (No), instead of being 
lamelliform and roofing over the nasal passages, are reduced 
to nodular masses, lying in depressions in the frontals, but 
forming as usual the hinder boundary of the anterior narial 

In front of these openings, the face stretches out into 
an elongated, depressed, pointed beak, or rostrum, formed 
by the premaxillae and maxillae surrounding the vomer and 
mesethmoid cartilage. The premaxillae send prolongations 
upwards to form the lateral boundaries of the narial orifice, 
and it is remarkable that these are not quite symmetrical, 
that of the left side being the shortest. The orifice itself, 
moreover, is rather inclined towards the left. Between the 
antorbital process of the maxilla and its rostral prolongation 
is a deep notch, the "antorbital notch." The upper surface 
of the face, near this notch, has several very large foramina 
for the transmission of branches of the fifth nerve. 

The elongated, pointed, and convex palate is formed 
chiefly by the maxillae (Mx), the premaxillae (PMx) only 
appearing for a short space near the apex. Behind the 
maxillae, the palatines (PI) are somewhat wide laterally, 
but towards the middle line form an exceedingly narrow 
strip, inserted between the maxillae and the pterygoids (Pt) ; 
the latter are greatly developed ; besides forming the outer 
wall of the posterior nares, each sends a lamella inwards, 
which nearly (in most Dolphins, completely) meets its fellow 
in the middle line, and so prolongs the bony palate back- 
wards. This process, moreover, is reflected outwards again 
from its inner or lower edge, and, joining with a projecting 




plate from the palatine, incloses a large cavity, open only 
behind, which contains the post-palatine air sinus. The 

FIG. 66. Under surface of the cranium of a young Dolphin (Globicepkalus welas). \. 
BO basioccipital ; ExO exoccipital ; Per posterior (mastoid) process of periotic ; 
Ty tympanic ; Sq squamosal ; AS alisphenoid ; OS orbitosphenoid ; ZM zygo- 
matic process of malar ; Fr supraorbital process of frontal ; Ma body of malar ; 
Pt pterygoid ; PI palatine ; MX maxilla ; PMx premaxilla ; Vo vomer ; gf 
glenoid fossa of squamosal ; tg deep groove on squamosal for meatus auditonus 
externus, leading to tympanic cavity ; cf condylar foramen. 

vomer ( Vo) is of great si?e, extends forwards nearly to the 
apex of the rostrum, embracing the mesethmoid cartilage, 

xii.] CETACEA. 215 

and posteriorly reaches for a considerable distance beneath 
the basisphenoid. It forms as usual the inner wall of the 
posterior narial apertures. Behind these apertures the base 
of the skull is flat in the middle line, but with prominent 
lateral elevations formed by the basioccipital, continuing 
the pterygoid ridge backwards. The glenoid fossa (gf) is 
a shallow, oval facet, on the inner and under surface of the 
zygomatic process of the squamosal. 

The periotic region of the skull differs much from that 
of most Mammals. On the side of the base of the cranium 
is a large recess, bounded below by the prominent edge of 
the basioccipital on the inner side, by a projecting edge of 
the exoccipital (paroccipital process) behind, by the base 
of the zygomatic process of the squamosal externally, and 
by a long curved process from the same bone in front, and 
communicating with the cranial cavity above by an irregular 
opening between the exoccipital and the alisphenoid. In 
this recess lies a bone of singular shape which, having only 
a ligamentous connection with the surrounding bones, is 
easily separated from the rest of the cranium in maceration, 
and is hence often wanted in specimens in museums. This 
is the united tympanic and periotic, ankylosed in the adult, 
but in young specimens still separable into its two com- 
ponent parts. 

The tympanic (Ty) is a hollow, bullate bone, broad, 
rounded and bilobate behind, and pointed in front. It 
is open above, the hinder part being, however, in relation 
with the periotic. Through the anterior spout-like end the 
Eustachian canal passes. At the upper border of the outer 
side, rather behind the middle, is an irregular or somewhat 
crescentic opening, bounded in front by a prominent lip, 
this is the meatus auditorius externus, closed in the living 
animal by the membrana tympani. 


The periotic is a rounded, very dense bone, characterised 
as usual by having on its inner or cerebral side the large 
meatus auditorius internus, and on the surface turned 
towards the tympanic cavity, the two fenestrae (ovalis and 
rotunda). The tympanic and periotic bones are separated 
along their inner margin by a narrow fissure, the "tympano- 
periotic fissure " ; but they are united externally in front of 
the external meatus auditorius, and more firmly posteriorly, 
where a tongue-shaped process (Per) projects backwards 
and outwards, fitting into a groove formed by the junction 
of the squamosal and exoccipital, and which is the principal 
point of attachment of the tympano-periotic bone to the 
rest of the skull. This process resembles in its relations the 
mastoid of ordinary Mammals, but in young Cetaceans it 
may be seen to be composed of two nearly equal parts, in 
close apposition with each, the inferior being derived from 
the tympanic, and the superior from the periotic, so that 
the latter alone can represent the "pars mastoidea " of 
other Mammals. 

The mandible (Fig. 65) consists of a pair of nearly straight 
compressed rami, wide behind and gradually narrowing to 
the symphysis (s), where they usually become ankylosed in 
adult animals. The condylar articular surface (cd) is small 
and looks almost directly backwards, being placed on the 
hinder edge of the ramus. The coronoid process (cp) is 
quite rudimentary. The angle is square and flat. The 
entrance to the dental foramen on the inner side is ex- 
tremely wide and infundibuliform. 

The ossified portion of the hyoid in the adult Dolphin 
consists of a large subcylindrical, slightly curved stylohyal (s/t) 
on each side, and a flattened crescentic median bone, com- 
posed of the ankylosed basihyal (b/i) and thyrohyals (t/i). The 
stylohyal is connected above by a slender cartilaginous rod 

xii.] CETACEA. 217 

to a small ossified tympanohyal, which becomes ankylosed to 
the periotic in the usual situation, close to the stylomastoid 
foramen ; it has also a strong ligamentous attachment to 
the prominent rough paroccipital process of the exoccipital. 
Between the stylohyal and the basihyal are one or two dis- 
tinct short cartilages articulated together by synovial joints, 
one of which occasionally becomes ossified. 

In many of the Delphinidic the rostral portion of the skull 
is proportionately more elongated and compressed than in 
the species just described ; notably so in Pontoporia, a South 
American genus. In this animal the mandible has a very 
long symphysial portion, the two rami being parallel and 
ankylosed for more than half their length, and diverging 
only in the posterior portion. 

The Sousou, or Platanista, a Dolphin inhabiting the 
rivers of South Asia, has also a remarkably elongated and 
compressed rostrum and mandible, and the cranial portion 
of the skull presents several structural peculiarities. The 
orbit is extremely small, the temporal fossa large, and 
the zygomatic processes of the squamosal are greatly 
developed. From the outer edge of the ascending plates 
of the maxilla;, which lie over the frontals, great crests of 
bone, smooth externally, but reticulated and laminated on 
their inner surface, rise upwards, and curving inwards, nearly 
meet in the middle line, above the upper part of the face. 

The PhyseteridiC, including the genera Ziphius, Hy- 
peroodon, Physeter, and their allies, present several special 
modifications of the skull. The bones of the face and 
cranium, meeting at the vertex, are raised so as to form a 
more or less elevated transverse prominence or crest behind 
the anterior nares, generally curved forwards at its upper 
edge. The bone which corresponds to the malar in other 


Dolphins is usually divided into two, one of which may 
represent the lacrymal. The pterygoid bones are thick, 
produced backwards, meeting in the middle line for a 
considerable space, concave on their outer side, but not 
involuted to form an outer wall to the post-palatine air sinus. 
In some of the Ziphiincz the rostrum is very dense, the 
anterior prolongation of the mesethmoid being either 
partially or completely ossified, and ankylosed with the 
surrounding bones. 

In Hypcroodon, the crest at the vertex is high and mas- 
sive, being formed by the nasals, the ascending plates of the 
premaxillas and maxillse, the frontals and supraoccipital. 
Separated from this crest by a depression, there is on each 
maxilla, at the commencement of the rostral portion of the 
skull, a very thick and high longitudinal ridge, which attains 
an extraordinary development in the old males of the com- 
mon Northern species. 

An easy transition from this cranium leads to that of 
the great Sperm Whale or Cachalot (Physefer macrocephalus), 
which of all Mammals is perhaps the most modified from 
the ordinary type. The transverse vertical crest and the 
longitudinal maxillary crests are united to form the walls 
of a great semicircular basin, surmounting the whole of 
the back part of the cranium, open only above and in 
front. The bones composing this wall are the same as in 
Hypcroodon, but excessively expanded and flattened. The 
rostrum is broad at the base, gradually narrowing to the 
front, and immensely elongated. The great supracranial 
cavity lodges the oily substance which, when refined, is 
known as spermaceti. 

The skull of the Cachalot is remarkable for its want of 
symmetry, especially in the region of the anterior narial 
apertures, of which the left is very much larger than the 

xii.] CETACEA. 219 

right. In consequence of the small increase in the size of 
the brain-cavity, compared with that of the external parts of 
the head in these enormous animals, the foramina through 
which the nerves pass out of the lateral parts of the base 
of the skull, are long channels excavated through immense 
bony masses. The petro-tympanic bone, which is scarcely 
larger than that of some of the small Dolphins, is situated 
at the bottom of such a channel, at the distance of fourteen 
inches from the inner wall of the brain cavity. In the 
general principle of their conformation, these bones do not 
differ from those of the ordinary Dolphins, but the tongue- 
shaped backward projection before described is greatly 
elongated and laminated, being composed of a large num- 
ber of distinct thin plates, only held together by their com- 
mon attachment to the tympanic. These fit into grooves 
between the squamosal and exoccipital, their extremities 
appearing on the outer surface of the skull, and they serve 
to attach the petro-tympanic more firmly to the cranium 
than is the case in the other Toothed Whales. 

The hyoid in Phyxeter and in the allied genus Kogia is 
remarkable for the great breadth and flatness of the basi- 
and the thyro-hyals, which, moreover, do not usually become 
ankylosed, as in most Dolphins. 

The cranium of the Whalebone Whales (sub-order Mysta- 
coceti) never shows that deviation from bilateral symmetry 
so frequent in the Toothed Whales. The cranial cavity- 
has much the same general form, and the bones around are 
disposed in a somewhat similar manner, but the parietals 
meet at the top of the skull, although completely overlaid 
and concealed externally by the great supraoccipitaL 

The anterior nares are not directed upwards and back- 
wards as in the Dolphins, but approach more in position to 
those of the ordinary Mammalia, being arched over by the 


frontals, which are of considerable antero-posterior thick 
ness at this part (see Fig. 67, Fr), and also by moderately- 
developed nasals (No), meeting by a flattened surface in 
the middle line. The nares are still near the most elevated 
part of the head, and the premaxillae and maxillae, with the 
vomer and mesethmoid cartilage, are produced in front of 
them into a long tapering rostrum, narrow, compressed, and 
much arched in the Right Whales (JBalcetia) ; broader, de- 
pressed, and nearly straight in the Rorquals (Balcenoptera). 

An essential difference between the Whales and the 
Dolphins is the presence in the former of an olfactory organ 
of the same type as in other Mammals, though in a compara- 
tively rudimentary condition. In the skull of an adult Green- 
land Whale (Balana mysticetus), the olfactory fossa of the 
cerebral cavity is eight and a-half inches in length, scarcely 
more than half an inch high, and one and a-half inch 
wide. It runs forward from the anterior part of the floor of 
the cerebral fossa, through the great mass of bone formed by 
the union of the frontal, mesethmoid and presphenoid. In 
front it divides into two compartments, each somewhat oval 
and dilated, with a concave floor about an inch in extent 
in either direction, perforated with foramina. This floor is 
the cribriform plate. In the hinder wall of the great narial 
passage is a narrow vertical slit, twelve inches in length, 
very near the middle line ; this is the opening of the 
olfactory chamber of the nasal cavity, which is bounded 
by the under surface of the cribriform plate above, by the 
flat mesethmoid on the inner side, and has its outer wall 
raised into several longitudinal elevations of very simple 
character, representing the ethmoturbinal bones. 

In the Rorquals (Balcenoptera) the olfactory fossa is less 
elongated, the foramina of the cribriform plate larger and 
more numerous, and the ethmoturbinals better developed. 




The supraorbital processes of the frontals are very long 
and narrow in the Right Whales, but broader in the Rorquals ; 
they are not covered by plates from the maxillae, as in the 
Dolphins. The orbit is small and completed below by a 
small curved malar, quite different from that of the 
Dolphins. There is a small wedge-shaped lacrymal inter- 
posed between the antorbital processes of the frontal and 
maxilla and the malar. 

FIG. 67 Section of the skull if a foclal Southern Right Whale (Baltrna austratis). 
i\. /M/r prcmaxilla ; \a nasal : /rfronlal ; fa parietal : SO cupmoccipital : OS 
orbitosphcnoid ; . I-V alisphenoid ; Kx<) exoccipital ; Per penotic ; BO basi- 
occipital : Ty tympanic ; US basisphenoid ; Sy glenoid articular surface of suua- 
mosal ; XV ptcryguid ; /V palatine; f'S presphenoid ; I'a vomer; /V.r maxilla: 
</ coronoid process uf mandible; cd condyle ; dg dental groove ; mg remains of 
groove which lodged Mcckel's cartilage. No part of the mcscthmoid is ossified. 

The palate is long and narrow, with a strong keel or 
ridge in the middle line. The surface on each side, sloping 
upwards and outwards, is perforated by many large foramina 
to permit the passage of blood-vessels and nerves to the 
matrix of the baleen, or " whalebone," which covers it in 
the living animal. It is chiefly formed by the maxilla, 


behind which are large palatines and very small and widely 
separated pterygoids. 

The zygomatic process of the squamosal is an immense 
trihedral pillar in Balcena, having the large shallow glenoid 
fossa on its under surface removed considerably from the 
middle of the cranium, so as to give sufficient width to the 
hinder part of the capacious mouth. The periotic and 
tympanic are formed much on the same principle as in the 
other sub-order, and in adult animals are completely excluded 
by a considerable distance from the cranial cavity, owing to 
the thickness of its walls. Instead of the small flattened 
tongue-shaped process projecting backwards from these bones, 
there is a long pyramidal tenon-like process, which fits into 
a groove in the squamosal and appears on the external sur- 
face of the skull like, though more solid than, that of the 
Cachalot. In addition to this another process projects out- 
wards and backwards, and the two together hold the bones 
much more firmly in their place than in the Toothed Whales. 
The tympanohyal is a large conical bony mass, with a 
truncated base, with which the stylohyal is connected, and 
firmly ankylosed by its apex to the periotic. 

The mandible differs much from that of the Toothed 
Whales. The two rami of which it is composed are not com- 
pressed and straight, but rounded and arched outwards, and 
never have extensive, flat, opposed symphysial surfaces, but, 
curving towards each other, meet at an angle in front, where 
they are held together by strong bands of fibrous tissue. 

The hyoid arch is formed essentially on the same plan as 
in the other Cetacea. The basihyal has a pair of processes 
placed side by side on its front edge, to which the anterior 
cornua are attached ; the hinder edge is excavated. In 
Balocna the thyrohyals are cylindrical, and thicker towards 
their free extremities. In Balanoptera musculus they are 

xii.] SIKENIA. 223 

cylindrical and tapering, in B. rostrata, flat and pointed 
externally. They always ankylose with the basihyaL 

Order SIRENIA. The animals belonging to this order, 
restricted at the present time to only two genera, which were 
formerly, but quite erroneously, included among the Cetacea, 
have skulls constructed on a very peculiar type, though with 
some affinities both to the Ungulata and the Proboscidea. 
Many of the special modifications are adaptations to their 
aquatic mode of life, and it is in these alone that they 
present any resemblances to the Cetacea. 

The skull of the African Manatee (Manatus senegalensis, 
Fig. 68), which may be taken as a type of the order, is remark- 
able for the massiveness and density of structure of the bones 
of which it is formed. There are no air sinuses in any part, 
and most of the bones when cut through appear as hard and 
solid as ivory. This character is not peculiar to the skull, 
but shared with it by the ribs, and other bones, and must 
add much to the general specific gravity of this slow-moving 
animal, and aid in keeping it to the bottom of the shallow 
water in which it dwells, while feeding on fuci and other 
aquatic vegetables. 

The cerebral cavity is very different from that of the 
Cetacea, being small as compared with the size of the 
animal, rather elongated and laterally compressed, truncated 
at each end, and with the upper surface flattened. The 
cerebellar fossa is large, and altogether behind the cerebral ; 
the olfactory fossa is distinct, but small and narrow, bounded 
on the inner side by a strongly-developed " crista galli " from 
the mesethmoid. The foramen magnum is of great size ; its 
plane looks backwards and downwards. The supraoccipital 
(SO) is inclined forwards, but does not extend beyond 
the ridge bounding the occipital region ; the roof of the 




cerebral fossa of the cranial cavity being formed by the pari- 
etals (Pa). The upper surface of the skull is very narrow, 
and flat or slightly arched in the longitudinal direction ; its 

FIG. 68. Section of the skull of an African Manatee (Manatiis senegalensis). \. 
PMx premaxilla; Vo vomer ; MX maxilla; Fr frontal; ET ethnioturbinal : 
M E mesethmoid : Fr frontal ; Pa parietal ; Sq squamosal ; SO supraoccipital ; 
E.rO exoccipital ; Per periotic ; BO basioccipital ; 'J y tympanic ; AS alisphenoid ; 
US basisphenoid ; /'.9 prcsphenoid ; Pt pterygoid ; PI palatine ; Vl/.r maxilla ; 
cp coronoid process of mandible ; cd condyle : a angle ; ^ symphysis ; sh stylo- 
hyal ; bh basihyal ; tk thyrohyal. 

sides, which are parallel for a considerable distance, join 
at a right angle the vertical inner wall of the great temporal 
fossa. The squamosal has an extremely massive and long 

xii.] SIXENIA. 22$ 

zygomatic process, flattened on the outer surface, and 
posteriorly it sends down a strong triangular post-tympanic 
process, articulating with a rough projecting edge of the 
exoccipital. Above this, between the squamosal, supra- 
occipital, and exoccipital, is a considerable vacuity in the 
cranial wall, partly filled by the periotic (Per). The lower 
vacuity, between the exoccipital and alisphenoid, common 
to all skulls, is of immense extent. 

The frontals (Fr) are narrow, and run backwards between 
the parietals to the upper part of the cerebral fossa of the 
brain cavity, and forwards a short distance over the nasal 
cavities ; each is produced anteriorly into a long narrow 
process, inclining outwards and downwards, between the 
temporal fossa behind, and the great anterior narial openings 
in front, forming the roof of the orbit. This cavity has a 
very prominent margin, especially below and in front, where 
it is formed by the very largely developed malar. This bone 
sends upwards a conspicuous postorbital process, which 
nearly (in some cases completely) meets that of the frontal, 
and then is continued below the zygomatic process of the 
squamosal as far as the wide shallow glenoid fossa, and 
sends down from its middle a broad flattened process with a 
thickened and rough inferior border. There is a very small 
scalelike and imperforate lacrymal in the usual situation at 
the anterior and inner angle of the orbit. The antorbital 
foramen of the maxilla is very large. 

One of the most peculiar features of the upper surface of 
the face is derived from the position of the anterior nares, 
which is a further modification of that met with in the Tapirs 
among the Ungulata, and presents some approach to that 
so characteristic of the Cetacea. Taken together they form 
a large lozenge-shaped aperture, which extends backwards 
considerably behind the orbits. Their sides are formed by 



the ascending processes of the premaxillse below, and by 
the supraorbital processes of the frontals above, no trace of 
nasals being found in most skulls, though these bones are 
occasionally present in a most rudimentary condition attached 
to the edge of the frontals, far away from the middle line, 
a condition quite unique among the Mammalia, or only 
approached in some of the Dolphins. In the floor of the 
great narial opening is seen the vomer ( Vo], of very delicate 
structure, and posteriorly the ossified portion of the mes- 
ethmoid (ME) of considerable vertical extent. The olfac- 
tory chamber of the nasal cavity is greatly compressed from 
side to side, and contains a series of simple, longitudinally 
placed ethmoturbinals, of which the upper one is very much 
the largest. There are no maxilloturbinals in any skulls 
which I have examined. 

In front of the narial opening the face is prolonged into 
a narrow rostrum, formed by the premaxillce, supported 
below and at the sides by the maxillae. The under surface 
of this is very rugose, and in life supports a horny plate. 
There is a large, oval, single, median anterior palatine 
foramen. The palate is long and narrow between the two 
parallel rows of numerous molar teeth. It does not extend 
beyond the last of these, and is formed almost entirely by 
the maxillae, the horizontal plates of the palate bones 
being very narrow. Behind each row of teeth is a massive 
descending rough process, formed by the union of the 
palatine, pterygoid plate of the alisphenoid, and true 
pterygoid. Posteriorly this has a longitudinal groove cor- 
responding to the pterygoid fossa. Behind this the base 
of the skull contracts in width, leaving a large opening on 
each side of the basioccipital. between the alisphenoid in 
front and the exoccipital behind, and only partially filled by 
the tympanic and periotic. 

xil.] SIREN/A. 227 

The two last-named bones are ankylosed together, but not 
to any of the other bones of the skull, and though freely 
movable in the dried skull, they are retained in their place 
by the overhanging process of the squamosal. 

The tympanic (Ty) consists of a large and very solid half- 
ring, with its lower margin considerably thickened and 
produced downwards ; but not forming any bulla, or any 
tubular meatus. It is only attached to the periotic by its 
extremities, and close to the inner side of its posterior 
attachment is a well-marked tympanohyal ankylosed to the 

The periotic (Per) is large and rounded externally. It 
forms a very considerable part of the inner wall of the 
cranium. Besides the portion containing the organ of hear- 
ing, it has a large solid upper part of somewhat kidney 
shape, lying in a groove in the squamosal (Sq). This solid 
part has a large anterior prominence, to which the anterior 
limb of the tympanic ring is ankylosed, and a smaller rounded 
posterior projection, corresponding with the mastoid of other 
Mammals, and mentioned before as appearing on the ex- 
ternal surface of the skull, in the vacuity between the supra- 
occipital, exoccipital, and squamosal. 

The foramina at the base of the skull are very few and 
simple, as nearly all the nerves appear to pass out only by 
the rather small sphenoidal fissure, and by the great confluent 
median and posterior foramina lacera. There is a small 
optic foramen passing through the middle of the orbito- 
sphenoid, but the alisphenoid is imperforate, and even the 
condylar foramen in the exoccipital for the hypoglossal 
nerve, so constant in all Mammals (except the Elephant), is 
represented by a groove (in some instances with a narrow 
bridge across it) on the anterior edge of the bone. There 
is no distinct carotid canal. 

The mandible is exceedingly different from that of the 
Q 2 

228 THE SKULL. [CHAP. xn. 

Cetacea, and is formed of the same dense heavy bone as the 
rest of the skull. The rami are firmly united by a symphysis 
(s) of moderate extent in front, and diverge widely behind. 
The posterior border is of considerable vertical depth, the 
condyle (cd], with its obliquely placed, oval, convex arti- 
cular surface, being raised high above the horizontal alveolar 
border. The coronoid process (cp) is large and directed for- 
wards. The angle (a) is well marked, thickened, and some- 
what inflexed, but does not form a distinct process. The 
lower border of the ramus is very thick, rounded from 
side to side, and concave from before backwards. The 
symphysial portion is compressed laterally, but its upper 
surface forms a somewhat expanded, rugose surface, concave 
in the middle line, to which a horny plate is attached in the 
living animal. 

The anterior arch of the hyoid has a single, slender, 
slightly-curved bone (stylohyal), three inches long, cylindrical 
at its upper end, and laterally compressed below, attached 
above by a broad, short ligament, chiefly to the exoccipital, 
but also to the squamosal and tympanic. The basihyal 
is a broad, flat, reniform plate, and the thyrohyals are not 

In the other existing Sirenian, the Dugong (Halicore}, from 
the Indian Seas, the skull resembles that of the Manatee in 
its essential characters, especially the form of the brain-case, 
the condition of the tympano-periotic bones, and the form 
and situation of the anterior nares ; but it differs mainly in 
the great development of the premaxillary bones, which curve 
downwards in front, and lodge large descending tusks The 
deep, compressed symphysial portion of the mandible is 
bent down in a corresponding manner. The zygoma is less 
massive, the orbit is not closed behind, and the lacrymal 
bone is more developed. The nasals are absent or quite 



Order EDENTATA. The different families of this hetero- 
geneous group present some remarkable variations in their 
cranial characters. 

One of the most extremely modified forms is the Great 
Anteater (Myrntecc'phaga jubata, Fig. 69). The whole skull 
is very greatly elongated and narrow, and its upper surface 
smooth and cylindriform. The occipital plane slopes upwards 
and forwards. The parietals are narrow, but the frontals much 
elongated. The olfactory fossa of the cerebral cavity is very 
large ; the cribriform plate greatly expanded, and the ethmo- 
turbinals much developed, and consisting of very numerous, 
delicate lamelke. Anteriorly, the face is produced into a 
very long, tubular rostrum, rounded above and flattened 
below, and with terminal nares. This rostrum is composed 
of the mesethmoid, ossified for more than half its length, 
the vomer, the maxillae, and the long and narrow nasal 
bones, the premaxillae (PAfx) being extremely short and 
confined to the margin of the anterior nares. There are no 
teeth in either jaw. The zygomatic arch is incomplete, the 
styliform malar (Afa) only articulating with the maxilla in 



FIG. i 



69. Under-surface of the cranium 
the Great Anteater (Myrmecophaga 

front, and not reaching the 
very short zygomatic process 
of the squamosal (Sq]. The 
lacrymals (Z) are distinct, and 
have a large perforation in 
front of the margin of the 
orbit. There are no postor- 
bital processes to the frontals, 
or any other demarcation 
between the orbits and the 
temporal fossae. 

The palate is extremely 
elongated, and produced back- 
wards as far as the level of 
the external auditory meatus 
by the meeting in the middle 
line of the largely-developed 
pterygoids (Pf). The glenoid 
fossa is a shallow oval facet, 
with its long diameter from 
before backwards. 

The periotic, tympanic, and 
squamosal are ankylosed to- 
gether. A small mastoid 
portion appears on the outer 
side of the skull, forming a 
roughened surface between the 
squamosal and the exoccipital. 
The tympanic (Ty] is some- 
what triangular in form, 
slightly bullate, and not pro- 
longed into an auditory meatu*. 
of the tympanic: 

xili.] EDENTATA. 231 

cavity, and freely communicating with it, is a considerable 
air sinus, formed between the pterygoid (Pf) and the ali- 
sphenoid (AS), and causing an oval prominence in the side 
of the palate. 

In another species of the same family, Tamandua tetra- 
dactyla, there is a second similar but smaller sinus anterior 
to this, in the side of the palatine bone. 

The mandible is very long and slender, with an exceed- 
ingly short symphysis, no distinct coronoid process, and 
a slightly elevated, elongated, flattened, condylar articular 

The anterior cornu of the hyoid is very long and slender. 
Its proximal end is ligamentous, and its distal portion con- 
tains three distinct ossifications. The thyrohyals become 
united by bone to the very narrow basihyal. 

The skull of the little Tree Anteater (Cycloturns didac 
tylus) besides being shorter, and much arched in the longi- 
tudinal direction, differs mainly from that of Myrmecophaga 
in not having the long canal of the posterior nares closed 
by bone below, as neither the pterygoids nor the greater 
part of the palatines meet in the middle line. The 
tympanic is more bullate. The mandible has a pro- 
minent, narrow, recurved coronoid, and a well-developed 
angular process ; it is strongly curved downwards in front. 

In the Armadillos of the restricted genus Dasyfits, in- 
cluding D. sexcinctus, rillosus, and minutus, the cranial 
portion of the skull is broad and depressed ; the facial 
portion triangular, pointed in front, and much depressed. 
The anterior narial orifice is small, terminal, and directed 
forwards and downwards. There is a completely ossified 
tympanic bulla, ankylosed with the rest of the skull, per- 
forated on the inner side by the carotid canal, and con- 
tinued externally into an elongated bony meatus auditorius, 


with its aperture directed upwards and backwards. The 
zygoma is complete. The pterygoids are small, and send 
no horizontal plates inwards to complete the bony palate, as 
in the Anteater. The mandible has a well-marked ascending 
posterior portion, supporting a transversely extended con- 
dyle, and a high, slender coronoid process. 

In all the other genera of Armadillos, Chlamydophorus 
excepted, the tympanic is a mere half-ring, loosely connected 
with the surrounding bones. 

The hyoid arch is strongly ossified. The anterior cornu 
consists of three bones. The thyrohyals ankylose with the 

In the Scaly Anteaters or Pangolins (genus Manis), the 
skull is somewhat in the form of an elongated cone, with 
the small end turned forwards, and very smooth and free 
from crests and ridges. The occipital plane slopes upwards 
and forwards. There is no distinction between the orbit and 
the temporal fossa, which together form a small oval depres- 
sion near the middle of the side of the skull. There are 
short zygomatic processes on the maxilla and the squa- 
mosal, but the arch is incomplete in most species, owing to 
the absence of the malar. There is likewise no distinct 
lacrymal bone. The plane of the anterior narial aperture 
looks forwards and upwards. The premaxilla is produced 
along the side of the nasals towards, but not reaching, the 
frontals. The palate is long and narrow. The pterygoids 
extend backwards as far as the tympanics, but do not meet 
in the middle line below. The tympanic is ankylosed to 
the surrounding bones, and more or less bullate, but not 
produced into a tubular auditory meatus. The hinder 
part of the squamosal is often dilated with air-cells, 
forming a rounded prominence at the outer posterior angle 
of the skull. 

xni.] EDENTATA. 233 

The rami of the mandible are edentulous, very slender 
and straight, without any angle or coronoid process. From 
near the anterior extremity of the upper edge a sharp conical 
tooth-like process projects upwards and outwards. The 
condyle is a slightly expanded flattened surface, not raised 
above the level of the rest of the ramus. 

In the Cape Anteater (Orycteropus) the skull is moderately 
elongated and dilated in front of the orbits. The facial por- 
tion is subcylindrical and slightly tapering. The lacrymal 
forms a considerable part of the side of the face. The 
zygoma is complete and slender. There is a small post- 
orbital process. The premaxillae are short and widely 
separated from the frontals. The palate ends posteriorly in 
the thickened transverse border of the palatines, and is not 
continued back by the pterygoids. The tympanic is annular 
and not ankylosed to the surrounding bones. 

The mandible is slender anteriorly, but rises high pos- 
teriorly, with a slender recurved coronoid, and an ascending 
pointed process on the hinder edge below the condyle, 
which is small, oval, and looks forwards as much as up- 

The hyoid arch is completely ossified. The basihyal 
is a thin bar, narrow in the middle. The thyrohyals are 
not ankylosed to it The ceratohyals are thick. There is 
a small (apparently epiphysial) ossification between the 
epihyal and the stylohynl. 

The Three-toed Sloths (genus Bradypus) have a high 
compressed skull, and an extremely short face. The cranial 
cavity is oblong, and rather high and compressed. There is 
no fossa on the periotic for the flocculus. The olfactory 
fossae are large. The plane of the occiput is vertical, or 
sloping slightly forwards and upwards. The frontal region 


is dilated with air sinuses. There is a small postorbital 
process. The lacrymal is very small, and the canal is 
external to the margin of the orbit. The malar is attached 
to the frontal, lacrymal and maxilla in front, curves down- 
wards and outwards, and then divides into a descending and 
a high ascending branch ; but neither of them join the 
straight zygomatic process of the squamosal. The nasals arc 
short and wide. The anterior nares are nearly vertical, or 
rather inclining downwards. The premaxiliee are exceedingly 
rudimentary, only the palatal portion being present, without 
any ascending process ; they unite with each other across 
the middle line, but not with the maxillae, hence they are 
generally lost in macerated skulls. The palate is narrow, 
especially posteriorly, and not produced behind the molar 
teeth. The pterygoids form large plates with prominent 
rounded borders, compressed in some, and inflated in 
other species. The glenoid fossa is narrow from side to side. 
The tympanic, squamosal, and periotic are ankylosed 
together. The former forms a considerable bulla, but no 
tubular meatus. There are large supratympanic air sinuses, 
and a well-marked ossified tympanohyal. 

The mandible has a comparatively high horizontal por- 
tion, rounded in front with a very small median triangular 
process at the upper border. The coronoid process is high 
and slender. The condyle is small ; its articular surface 
is convex from side to side, short and nearly straight 
from before backwards. The angle forms a broad com- 
pressed posterior projection, with a slightly incurved lower 

The stylohyals are large, compressed, and curved, with a 
prominent posterior process near their upper end. The 
basihyal is small, and ankylosed with the thyrohyals, so that 
they form together a V-shaped bone. 


The skull of the Two-toed Sloth (Cholcepus didactylus), 
though generally similar to the last, presents in some points 
marked deviations from it. Even in aged specimens, in 
which almost all the sutures are obliterated, the tympanic 
is a mere ring, incomplete at the upper margin, and but 
slightly connected with the other bones around. The 
premaxillae are more developed, and become ultimately 
ankylosed with the maxillae. The pterygoids are much 
smaller, but sometimes are bullate. 

The upper margin of the mandible is produced anteriorly 
into a spout-like process. The condyle is scarcely above 
the level of the molar teeth, and is wide from side to side. 
The angular projection is smaller and thicker. 

The hyoid (in an old specimen) has a strongly ossified 
anterior arch, consisting of two bones of nearly equal length, 
the proximal one with a stout-rounded process projecting 
backwards and outwards from near its upper end. The 
second is bent at a right angle near its lower end, and may 
result from the ossification of two elements. The basi- and 
thyro-hyals are ankylosed to form a wide U-shaped bone. 

Order MAKSUPIALIA. -The skull of the large carnivorous 
Marsupial, the Thylacine (Thylacinuscynocephalus), resembles 
so closely that of a Dog in its general aspect, that it will be 
well to commence an account of the peculiarities of the 
crania of Marsupials generally by comparing these two 

It will be seen by the section (Fig. 70) that the brain cavity 
of the Thylacine is very much smaller than that of the Dog 
(Fig. 46, p. 117) in relation to the size of the rest of the 
cranium, or to that of the whole animal, a sign of great in- 
feriority of organisation. This diminution affects chiefly the 
cerebral fossa ; the cerebellar fossa is nearly equal in size, 
but it is placed more directly behind the cerebral, and is 




not in the least overlapped by it, as in the Dog. The 
occipital plane is vertical, or even inclining forwards above. 
The tentorial plane is nearly horizontal. The olfactory 
fossa, though smaller in vertical extent than that of the Dog, 
is more produced anteriorly. Thus in form, as well as in 
relative size, the cerebral cavity is far more reptilian than 
that of the Dog. The basicranial axis is very straight, and 

FK;. 70, Section of the skull of the Thylacine (TJiylacinus cynocephalus), \. 
MT maxilloturbinal ; ET ethmoturbinal ; ME ossified portion of mesethmoid ; 
/> frontal ; Pa. parietal ; SO supraoccipital ; EjcO cxoccipital ; Per periotic ; 
BO basioccipital ; S<] squamosal ; AS alisphenoid ; TiS basisphenoid : OS orbito- 
sphenoid ; 1'S presphenoid : Pt pterygoid ; /V palatine; f'fvomer; M.\ maxilla ; 
PMx premaxilla; ai condyle of mandible ; a angular process. 

is continued forwards in the same line by the basifacial 
axis. The pituitary fossa forms no distinct impression, and 
there are no posterior clinoid processes. The ossified 
portion of the mesethmoid (AfE) is extensive, and termi- 
nates anteriorly in a nearly vertical line. The vomer ( Vo) 
is very shallow from above downwards. The turbinal 

xiii.] MARSUPIALIA, 237 

bones resemble generally in their extent and disposition 
those of the Dog. 

Externally the conformation of the zygomata, temporal 
fossae, orbits, maxillae, premaxillae, and nasals are strikingly 
similar to those of the Dog ; but the lacrymal bone of the 
Thylacine is larger both within and without the orbit, and 
it has one perforation within, and either one or two just 
external to the margin of the orbit. The palate has a pair 
of large oval vacuities between the molar teeth. The ptery- 
goid plates are very thin. The glenoid fossa is rather more 
expanded from before backwards than in the Dog, and the 
malar extends so far back beneath the zygomatic process of 
the squamosal as to form the outer edge of the glenoid cavity. 
The postglenoid and paroccipital processes are developed 
much as in the dog ; the former has rather greater lateral 
extent. A great difference is seen in the condition of the 
tympanic, which in the Thylacine is quite rudimentary, forms 
no bulla, and is not ankylosed to the other cranial bones, 
so that in the dried skull it is nearly always detached. On 
the other hand, the hinder part of the alisphenoid is dilated 
into an oval thin-walled capsule, which is connected with 
the front of the tympanic cavity. The periotic is not 
ankylosed to the squamosal, has a very large floccular fossa 
within, and a mastoid portion forming a long, narrow strip 
visible in the outer side of the occipital surface of the skull, 
between the squamosal and the exoccipital, almost exactly 
as in the Dog. The exoccipital is perforated by the con- 
dylar foramen ; but the carotid foramen, instead of passing 
through the inner edge of the tympanic bulla, perforates 
the basisphenoid, passing very obliquely forwards and in- 
wards. The large alisphenoid is pierced by the fora- 
men ovale near its posterior margin, and by the foramen 
rotundum near the front. The orbitosphenoid is very small, 


and not perforated, the optic nerve passing out through the 
sphenoidal fissure. 

The ascending ramus of the mandible is less elevated than 
that of the Dog, the condyle being almost on the same level 
as the molar teeth. The coronoid process has a more back- 
ward inclination. The masseteric fossa has a powerful, 
externally projecting lower border, and the angular process 
is flattened from above downwards, and inclined consider- 
ably inwards. 

The hyoid (Fig. 71) is constructed on a totally different 
type from that of the Dog. It consists of a small flat lozenge- 
shaped basihyal (/>//), surmounted by flattened, triangular, 

i it 

Fi<;. 71. Upper surface of hyoid of Thylacine (nat. size), bh basihyal ; ch ceratohyai 
(anterior cornu); th thyrohyal (posterior cornu). 

imperfectly ossified ceratohyals (ch}, partially ankylosed to 
the basihyal, and without any other ossifications in the 
anterior cornua. The thyrohyals (///) are tolerably long 
flattened bars, meeting in the middle line at their attachment 
to the basihyal, and with their free, or laryngeal, extremities 
expanded, but still cartilaginous in the perfectly adult animal 
from which the above figure was taken. 

All the other animals of the sub -class which contains the 
single order Marsupialia, however their skulls may differ in 

xiii.] MARSUPIAL1A. 239 

general appearance from that of the Thylacine, agree with it 
in the following important particulars : 

1. The brain cavity is small, with the cerebella fossa 
entirely behind, and the olfactory fossa entirely in front of 
the cerebral fossa. There are, however, degrees in this res- 
pect, the Kangaroos representing one extreme, with large, 
more vaulted cerebral fossa, and the Opossums and Dasyures 
the other. 

2. There is no distinct pituitary fossa, and the clinoid 
processes are obsolete. 

3. The ossification of the mesethmoid is extensive, and 
has an abrupt, nearly vertical, anterior termination. 

4. The nasal bones are large, and the anterior nares more 
or less terminal. 

5. The zygoma is complete, but the orbit has not a perfect 
posterior boundary. 

6. The malar is large, reaches the lacrymal anteriorly, 
and extends posteriorly beneath the zygomatic process of 
the squamosal, to form part of the outer wall of the glenoid 

7. The perforation in the lacrymal is usually upon, and 
frequently external to, the anterior boundary of the orbit. 

8. The ascending processes of the premaxillae never quite 
reach the frontals. 

9. The palate has often, but not always, large vacuities 
near its posterior margin. 

10. The pterygoids are always small and lamelliform. 

11. The alisphenoids are more or less dilated, and form 
the anterior wall of the tympanic cavity, which is often quite 
open below in the dried skull. It may be noted as a special 
peculiarity in the Kangaroos, that the alisphenoid extends 
backwards beneath the tympanic cavity to join the long 
paroccipital process of the exoccipital. In the larger mem- 




bers of the group it can scarcely be said to form a distinct 
bulla, but in some of the Hypsiprymni (Rat Kangaroos) it is 
immensely expanded. In the Koala (Phascolarctos)^ the 
alisphenoid bulla is very large, elongated vertically and 
compressed, having a very similar appearance in fact to the 
tympanic bulla of the Pig. 

12. The tympanic is small, simple, and annular in some ; 
in others it forms a short external auditory meatus, but 
it is never ankylosed to any of the other bones of the 

FIG. 72. Upper surface of hyoicl oi 
Wombat (fhascolomys latifrons). bh 
basihyal ; fh ceratohyal ; sh stylohyal ; 
tk thyrohyal. 

K;. 73. Hyoid of Kangaroo (Alacrn- 
J>us). bh basihyal ; ch ceratohyal ; th 

13. The periotic sends backwards a distinct mastoid, which 
appears as a narrow strip of bone of considerable vertical 
extent, between the squarnosal and exoccipital, on the side 
of the occipital region of the skull. 

14. There are almost always conspicuous paroccipita-1 

15. The internal carotid artery perforates the basi- 

1 6. The optic foramen is confluent with the sphenoidal 

17. The mandible has (Tarsipes excepted) an inverted 
border to the angle. 

xiii.] MONOTREMATA. 241 

1 8. The hyoid has a small, more or less lozenge-shaped 
basihyal, broad ceratohyals, the remainder of the anterior 
cornu usually unossified, and stout somewhat compressed 

Order MONOTREMATA. Both the animals of this group 
present very singular modifications of the cranium. 

The cerebral cavity, unlike that of the lower Marsupialia 
or the Reptiles, with which they have so many structural 
affinities, is large and hemispherical, flattened below and 
arched above, and about as broad as long. ' The broad cribri- 
form plate of the ethmoid is nearly horizontal. The walls 
are very thin, and smoothly rounded externally, and the 
sutures become completely obliterated in adult skulls so that 
it is very difficult to trace out the boundaries of the com- 
ponent bones. In both species, the broad occipital region 
slopes upwards and forwards, and the face is long and much 
depressed, though of very different shape in each. 

In the Echidna (Fig. 74) the squamosal is large and very 
compressed, the zygomatic process arising very far forward ; 
the slender hori/ontal zygoma being completed by a styliform 
malar, confluent with the maxilla. The face is produced 
into a long tapering rostrum, rounded above from side to 
side, and concave below in the same direction. The an- 
terior nares form an oval opening on the upper surface near 
the apex, bounded entirely by the premaxillaj, for the nasals 
do not appear to reach so far forwards. The alveolar borders 
are narrow and rounded, without trace of teeth. The palate 
is produced backwards, by very large palatine bones (/*/), 
considerably beyond the glenoid fossa. The narial canals 
have very little extent vertically, but the true olfactory 
chambers are large, and provided with complex turbinals, 
which, in accordance with the horizontal position of the 
cribriform plate, are mostly placed vertically. 




The pterygoids (Pt] are flattened, horizontal, oval plates, 
attached to the obliquely truncated postero -external extremi- 
ties of the palatines, and form part of the floor and the inner 
wall of the tympanic cavity, an arrangement not met with in 
any other Mammal. The tympanic (Ty) is a very slender 

ring, incomplete for a small space 
at the upper and outer end, and 
does not become ankylosed to 
the periotic. The latter (Per) is 
large, has no fossa for the floc- 
culus, sends out a large expansion 
(pterotic), forming a portion of 
the cranial wall between the 
squamosal, parietal, and occipital, 
the lower and hinder part at least 
of which corresponds with the 
mastoid portion. 

Each ramus of the mandible is 
a mere slender style, without any 
ascending portion, and with but 
rudiments of coronoid process 
and angle. The condyle is very 

FIG. 74,-Under surface of cranium Small, elongated from before 
of Echidna (Echidna aculcata), \. K. lr l,iv'ird<; nnd vr>rv mrrmv 

BO basioccipitai; ExO exocci- Dackwarob, ana vtry narrow . 

The hyoid (Fig. 75) has a well- 
ossified transversely extended, 
flattened and arched (with the 
concavity forwards) basihyal (bJi). The anterior cornu has 
only two ossifications (eh and ch\ apparently the epi- and 
cerato-hyal, as the upper one has a long ligamentous con- 
nection with the cranium in the situation of the stylohyal. 
There are broad, flattened, curved thyrohyals (tli) expanded 
at their laryngeal extremities. 

pital ; Per periotic ; m malleus 
Sq squamosal ; Ty tympanic ; Pt 
pterygoid ; /'/palatine ; MX max- 
illa ; PMx premaxilla. 



In the Ornithorhynchus the brain case is smaller than in 
the Echidna, and rather more depressed ; very broad behind 
and narrowing anteriorly. The olfactory fossa is compara- 
tively small. There are well-marked posterior clinoid 
processes. The falx cerebri is largely ossified, forming a 
strong median partition to the upper part of the cerebral 
cavity. The zygoma is compressed, and of considerable 
vertical depth, and sends up a well-marked postorbital 
process ; its hinder root arises very far back on the 

The glenoid fossa is wide and concave transversely. The 
zygomatic process of the maxilla is widened inferiorly into 

Fir.. 75. Lower surface of hyoid of Echidna (Echidna acttleata). th epihyal; 
ck ceratohyal ; //; basihyal ; th ihyrohyal. 

an oblong, concave, roughened surface for the attachment of 
the horny plate, which takes the place of the molar teeth. 

The face is broad and much flattened. It runs out 
anteriorly into two diverging processes, each formed by the 
premaxilla, supported by a pointed process of the nasal on 
the inner and of the maxilla on the outer side. These bend 
towards each other at their extremities, but do not meet in 
the middle line. They support the partly horny, partly 
membranous beak, which fills up the space between them, 
and extends considerably on each side and in front. There 
is a distinct median dumb-bell-shaped ossification in the 

R 2 

244 THE SKULL. [CHAP. xm. 

triangular interval between the diverging premaxillary bars, 
placed in front of the anterior extremity of the mesethmoid 
cartilage, on the palatal aspect of the jaw. This bone is 
not the homologue of the so-called prenasal of the Pig, but 
" it corresponds with that part of the intermaxilla, which lies 
between the incisive canal and the mesial palatal suture." : 
The infraorbital foramen is very large, corresponding to the 
large size of the nerves distributed to the sensitive sides of 
the beak. The periotic has a wide and deep floccular fossa. 
The mandible has, rather behind the middle of each ramus, 
an oblong expansion for a horny tooth, corresponding to that 
on the maxilla. Behind this it curves gradually upwards to 
the expanded and transversely extended articular surface. 
There is no distinct angle, the coronoid process is small and 
directed much inwards, and on the external surface there is 
a very deep masseteric fossa. Anteriorly the rami of the 
mandible have a very slight symphysial connection, in front 
of which their expanded, flattened terminations again diverge 
from each other. The apertures for the entrance and for 
the exit of the branches of the inferior dental nerve are 
remarkably large. 

1 W. Turner, " The dumb-bell-shaped Bone in the Palate of Ornitho- 
rhynchus compared with the prenasal Bone of the Pig" (Journ. Anat. 
Phys. xix. 1885, p. 214). P. Albrecht, " Sur la Fente Maxillaire et let. 
quartre Os Intermaxillaires de 1'Ornithorynque. " Bruxelles, 1883. 



HAVING finished the consideration of the trunk and 
skull we now turn to that of the two pairs of limbs, anterior 
and posterior. 

The anterior limb is present, and fully developed, in all 
Mammals, being composed of a shoulder girdle and three 
divisions belonging to the limb proper, viz. the upper arm 
or brachium, the fore-arm or antibrachium, and the hand or 

The SHOULDER GIRDLE in the large majority of Mammals 
is in a comparatively rudimentary, or rather modified, con- 
dition. Its true structure and its relations to the pelvic 
girdle can only be understood by a reference to its condition 
in the lower vertebrates. 1 

Each side of the girdle consists primitively of a curved 
rod of cartilage, placed vertically (in the horizontal position 
of the body), the upper or dorsal end being free, the inner 
side lying upon, though not united with, some of the anterior 
thoracic vertebrae or ribs, and the inferior or ventral end 
being attached to the side of the presternum. 

1 On this subject see W. K. Parker's valuable work before cited, and 
also Gegenbaur's " Untersuchungen zur vergleichenden Anatomic," 2 tes 
Heft, 1865. 


Near, or rather below, the middle of the outer surface of 
this rod is a more or less cup-shaped depression, the glenoid 
cavity, to which the head of the humerus, or bone of the 
upper arm, is articulated. The whole rod is separated at 
this spot into two divisions, which ossify from separate 
nuclei. The upper or dorsal division is the scapula, the 
lower or ventral division is the coracoid. 

In all Mammals above the Ornithodelphia the greater part 
of the coracoid is aborted, but a portion of its upper ex- 
tremity always remains attached to the scapula as a process, 
or sometimes as a most inconspicuous nodule, and occa- 
sionally rudiments of the lower end are found attached to 
the sternum. The scapula, on the other hand, is always 
greatly developed. 

A supplementary bone, developed in a different manner, 
being, at least in the greater part of its extent, ossified from 
membrane, 1 frequently forms an anterior bar, in front of the 
coracoid, passing between the scapula and the anterior end 
of the presternum : this constitutes the clavicle. It is 
often rudimentary, and very frequently entirely absent, in 

Though the scapula may be considered as essentially an 
elongated rod or bar of bone (a condition most nearly re_ 
tained in the Mole), it usually has three projecting plates 
or ridges arranged around its longitudinal axis, and three 
surfaces or fossae bounded by these, the variations in the 
extent and form of which give rise to the principal diversities 
in the form of this bone in different Mammals. 

In the most usual position of the scapula (see Fig. 76), 
one of these plates (of] projects forwards, this is the 

1 When it has a cartilaginous basis (as has been described by 
Ge^enbaur in Man) this is not a portion of the true primitive shoulder 




prascapula of Parker, and its edge constitutes the anterior 
border (cb) of the scapula ; another (//) projects backwards, 
constituting \hepostscapula, and its edge is the posterior border 
(gb) ; a third projects outwards, constituting the mesoscapula 
of Parker, called more commonly the spine (s). The first- 
named border (cb) terminates below by joining the coracoid, 
and hence, to avoid the inconvenience of a term which is 
only expressive when the bone is in a particular position, it 


FIG. 76. Right scapula of Dog (Cam's familiaris), \. pf postscapular fossa ; aj fossa ; gb glcnoid or posterior border ; cb coracoid or anterior border : 
s spine ; a acromion ; gc glcnoid cavity ; c coracoid ; ess indicates the position of 
the coraco-sc.ipular suture, obliterated in adult animals by the complete ankylosis 
of the two bones ; si suprascapular border. 

may be called coracoid border ; the second (gb) joins the 
prominent margin of the glenoid fossa, and may, for the 
same reason, be called glenoid border ; the third (s) has a 
free end, usually more or less prolonged into a curved, 
flattened process, called the acromion (a). 

The flat or concave surfaces or fossae between these pro- 
jecting lamellae are (i) the prescapular or anterior fossa 


(af), between the coracoid border and the spine, called, in 
works on human anatomy, "supraspinous fossa;" (2) the 
postscapular fossa (pf\ between the glenoid border and the 
spine, also called " infraspinous fossa ; " and (3) the sub- 
scapular fossa, between the coracoid and glenoid borders 
on the side of the scapula opposite to the spine. 

The greater part of the scapula is ossified by ectostosis (as 
the shaft of a long bone) from a single centre, which is 
placed not far from the middle of the bone ; but this ossifi- 
cation does not extend into a certain portion near the 
superior (or dorsal) extremity. This part (suprascapula 
of Gegenbaur) either remains cartilaginous or " is feebly 
ossified by one or more endosteal patches, or by the creeping 
upwards of such deposit from within the main bone " 
(Parker). When the spine runs out into a projecting 
acromial process, more or less of its terminal portion is 
ossified separately as an epiphysis. 

The coracoid always ossifies from one or more separate 
centres, and remains for some time suturally connected 
with the scapula, though firmly ankylosing with it by the 
time the animal has attained maturity. Sometimes (as in 
the Sloths) it forms a considerable part of the glenoid fossa ; 
sometimes (as in most Carnivora and Ungulata) it is a mere 
nodule, which becomes blended with the anterior margin of 
the fossa. 

In the Ornithodelphia (see Fig. 84), as in Birds and Rep- 
tiles, the coracoid is largely developed, and articulates with 
the presternum, and there is in addition a plate of bone in 
relation with its ventral edge called epicoracoid. A small 
plate of cartilage or bone, often found attached to the side 
of the presternum in certain Rodents and Insectivores, is 
considered by Parker as representing the epicoracoid of 
the Ornithodelphia, and by Gegenbaur as the sternal ex- 

xiv.] PRIMATES. 249 

tremity of the true coracoid, the middle part of which is 

The clavicular arch, when completely developed, extends 
from the free acromial extremity of the spine of the scapula 
to the anterior extremity of the presternum. 

It consists mainly of an elongated rod of bone, ossified 
usually in fibrous tissue, but at either extremity are certain 
patches of true cartilage, which may become converted into 
bone, or may sometimes degenerate into fibre-cartilage. 
These are thus described and named by Parker. At the 
scapular extremity of the clavicle, there is often a piece of 
cartilage considered to be segmented off from the end 
of the mesoscapula, and hence called mesoscapular segment 
(Fig. 77, mss). At the sternal extremity there may be two 
distinct pieces, the one (pc) nearest the clavicle being the 
supposed homologue of a displaced fragment of the pre- 
coracoid (Gegenbaur and Parker) of the lower vertebrates. 
The one (osf) nearest the sternum is called omosternum by 
Parker, and episttrnum by Gegenbaur, who considers it 
homologous with the so-called episternum (jntcrclaviele, 
Parker) of the Ornithodelphia and Lizards. 

Special Characters of the Shoulder Girdle in the Different 
Groups of the Mammalia. 

Order PRIMATES. Man. In the ordinary erect position 
of the human body, the suprascapular border is directed 
backwards and inwards, and is commonly called the " base " 
or " vertebral border " of the scapula ; the glenoid cavity 
looks forwards and outwards; the glenoid border, called 
" external " or " axillary," looks downwards and rather 
forwards; and the coracoid border is "superior." The 




postscapular fossa is much developed, and the suprascapular 
border is long, straight, and forms an acute angle with the 
glenoid border. At the junction of the coracoid border of 
the scapula with the coracoid bone, there is a more or less 
well-marked notch (coraco-scapular notcJi). The spine is 
well developed, and the acromion large and curved forwards 
near its extremity. 

IG. 77. The hitman sternum and right shoulder girdle at a very early period of 
development (from an embryo si inches long) after Parker, i\. The dotted parts 
are still cartilaginous ; the inner surface of the sternum and clavicle, and outer 
surface of the scapula are represented, ost omosternum, afterwards developed 
into the interarticular fibro-cartilaginous disk; pc precoracoid of Parker; cl shaft 
of the clavicle ; mss mesoscapular segment of Parker ; a acromion ; c coracoid ; 
gc glenoid cavity of scapula ; gb glenoid border ; cb coracoid border ; of anterior, 
or ''supraspinous," fossa; ^/"posterior, or '' infraspinous," fossa ; S.T suprascapular 

The coracoid forms a well-marked hook-like process ; it 
contributes a very small part to the glenoid fossa, and unites 
with the scapula about the time of puberty. 

The clavicle (Fig. 77, d] is a strongly-developed sigmoid 
bone, remarkable for the very early age at which it com- 
mences to ossify, in fact before any other bone of the body. 
The outer extremity is, in the young state, tipped with 

xi v.l INSECTIVORA. 251 

cartilage (the mesoscapular segment, Parker, tnss), which 
ossifies by extension of bone from the rest of the clavicle. 
It is connected with the acromion by a small oval, flat, 
synovial articulation. The inner end (pt) is also cartila- 
ginous for some time, but ossifies separately by endostosis, 
forming an epiphysis. This extremity is attached to the 
presternum by synovial articulation, but with a disk-like 
fibro- cartilage (ost) interposed, which, according to Parker, 
is a degeneration of the " omosternal " element. 

In the Gorilla the scapula is very like that of Man. In 
the Chimpanzee it is peculiarly elongated, the suprascapular 
margin being extremely oblique and long, at the expense 
of the greatly reduced coracoid border. The acromion and 
coracoid are largely developed. In the lower Monkeys the 
form of the scapula is quite different, the coracoid and 
glenoid borders being nearly equal, and the suprascapular 
border comparatively short and straight. 

The clavicle is well developed in all the Primates, 
and all its correlates are present ; the omosternum being 
generally converted during growth into a fibro-cartilaginous 
intra-articular disk. 

In the various members of the Order INSECTIVORA there 
is a great difference in the construction of the shoulder 

In the Mole (Talpa) and its immediate allies, Sea lops and 
Condylura, the scapula is extremely high and narrow, and 
appears to be ossified entirely from one centre. The spine 
and acromion are very little developed. The bone com- 
monly called clavicle, but which may be a combination of 
coracoid and clavicle, is of remarkable form, being nearly 
cuboid. It is formed primitively of a mass of cartilage, on 
the anterior aspect of which the true (membrane-developed) 
clavicle is engrafted. It articulates inferiorly with the pre- 



sternum, and superiorly with the humerus, and is connected 
with the scapula only by a fibrous band. The two articula- 
tions of the upper end of the humerus, the one with the 
scapula, and the other with the coraco-clavicle, are separated 
by a strong ligamentous partition. 

In the Cape Golden Mole (Chrysochloris) the condition of 
these parts is quite different. The scapula is long and 
narrow, but flattened. The spine sends a flat process back- 
wards near its middle, and a long slender " metacromial" 

FIG. 78. Shoulder girdle, with upper end of sternum (inner surface) of Shrew (Sorcx), 
after Parker, X 7. ps presternum ; sr l first sternal rib ; sr* second sternal rib; 
ec partially ossified " epicoracoid " of Parker, or rudiment of the sternal extremity 
of the coracoid ; ost omosternum ; pc rudiment of precoracoid (Parker) ; cl cla- 
vicle ; mss ossified " mesoscapular segment;" a acromion : ma metacromial 
process ; c coracoid. 

process from its extremity. The clavicle is very long, 
slender, and curved. The " mesoscapular segment " forms 
a distinct, though minute, bone between the clavicle and 

In the Shrews (Soricidee) the scapula (see Fig. 78) is 
also long and narrow, and the slender acromion ends in 
two long diverging processes, of which the anterior (a) 
supports the clavicle and the posterior (ma) is called 

xiv.] RODENTIA. 253 

"metacromion." The mesoscapular segment (mss) is a 
distinct bone. The clavicle (cf) is long and slender. It has 
a small piece of cartilage (pc] attached to its inner end. 
The omosternum (osf) is cartilaginous or partially ossified, 
and there is a considerable triangular flattened rudiment 
of the inner end of the coracoid (tc) attached to the pre 

In the other Insectivora the general form of the scapula 
is more normal. In Galeopithecus the coracoid is greatly- 
developed and bifurcated. 

All known members of the order have large clavicles, with 
the exception of Fotamogale, a rare aquatic form from 
West Africa. 

In the CHIROPTERA the scapula is large, of an oval form, 
and chiefly formed by the postscapular fossa, the anterior 
fossa being extremely small. The former is divided into 
two or three subfossse by ridges. The spine is short and 
moderately high, with a large and simple acromion. The 
coracoid is long and curved, often simple (as in Pteropus) 
sometimes forked (as in Pipistrellus). 

The clavicle is very long and curved. The " mesoscapular 
segment " on the outer end is soon lost, but the precoracoid 
ossifies separately. The omosternum is reduced to a 
cuneiform fibro-cartilage. A rudiment of the sternal end of 
the coracoid is often present as "a flat, reniform flap of 
cartilage, feebly ossified by endostosis, wedged in between 
the clavicle and the first rib " (Parker). 

The RODENTIA offer great diversities in the condition of 
the shoulder girdle. The scapula is generally high and 
narrow, and the acromion long. Sometimes the acromio- 
scapular notch is so deep, that the actual spine only occupies 
a short space near the suprascapular border, and it is com- 
pleted by a very long and slender acromion (as in the Coypu, 




Myopotamus]. There is often a long metacromion, as in the 
Hare ; but in others, as the Beaver, there is no such process. 
The coracoid is always a small blunt hook. 

In a few forms the clavicle is altogether absent, in some it 
is well developed, and various intermediate stages between 
these two extremes are met with. In some species, as the 
Guinea Pig and Rabbit, although no trace of this bone is 
found at birth, it becomes developed at a later period. In 
both of these it is very short, and is suspended by long 


FIG. 79. Shoulder girdle, with upper end of sternum (inner surface), of a young 
Rabbit (Lc/>ns citniculus), after Parker. 3. ps presrenuim ; sr l first sternal rib; 
ost om asternal cartilage ; fc precoracoid cartilage ; c/ossified clavicle ; HISS carti- 
laginous mesoscapular segment ; c coracoid ; a acromion ; ma metacromion : af 
anterior fossa ; pf posterior fossa. 

ligaments between the scapula and the sternum. (See Fig. 
79.) In many species, as in the Porcupines, in which the 
clavicular arch is more complete, the true clavicle is con- 
nected with the presternum by a long cartilaginous omo- 
sternum. In others, as the Beaver, this is replaced by a 
ligamentous band. Rudiments of the sternal end of the 
coracoid are often present, sometimes cartilaginous, some- 
times ossified. 

xiv.] CETACEA. 255 

In the CARNIVORA the anterior and posterior fossae of 
the scapula are nearly equal in area. (See Fig. 76, p. 247.) 
The spine and acromion are fairly developed, the latter often 
with a broad metacromial process. The coracoid is much 
reduced. According to Parker a portion of the scapula, 
near the coracoid border, ossifies from an independent 
centre. The clavicle is sometimes absent, and when present 
varies much in its development, but is always rudimentary 
and suspended in the muscles, never reaching either the 
acromion or sternum. In the Felida it is slender and 
curved, being longer than in any other members of the order. 
In the Canidce it is very short, and rather broad and flat. 
In most of the Ursida it is absent. 

In the Seals both acromion and coracoid are much re- 
duced, but the latter is a distinct bone in young animals, 
and forms a considerable part of the glenoid cavity. The 
whole scapula is much curved backwards, being almost 
sickle-shaped, and the suprascapular epiphysis is very large 
and slowly ossified. 

In the Eared Seals (Otaria) the scapula has a different 
form, the prescapufar fossa being very much larger than the 
posterior, and with a strong vertical ridge, parallel to the 

None of the Pinnipedia have clavicles. 

Order CETACEA. In the true Dolphins and nearly all the 
Odontoceti the scapula is usually very broad and flat, or fan- 
shaped. (See Fig. 80.) The prescapular fossa (af) is ex- 
tremely reduced ; the acromion (a) is a long flat process 
with a very narrow base of attachment, projecting for- 
wards ; the coracoid (c) is rather long, flattened, and parallel 
with the acromion. 

In the Cachalots (Phystter) the scapula is formed on the 
same general plan, but is comparatively high and narrow. 




The postcapular fossa is very concave, and the subscapular 
fossa convex. The Gangetic Fresh-water Dolphin (Platanista) 
has a flat flabelliform scapula, with the prescapular fossa 
entirely absent, and the acromion placed on the anterior 
edge, the spine and the coracoid border having coalesced. 

Among the Whalebone Whales, Balcenoptera has a broad 
fan-shaped scapula, like that of the true Dolphins, with long 
parallel acromion and coracoid processes, and a suprascapular 

IG. 80. Right scapula of Dolphin (Delphintis turtle), \. gc glenoid cavity ; 
a acruinion ; c coracoid ; pf postscapular fossa ; of prescapular fossa. 

border which remains permanently in a cartilaginous con- 
dition. In the Right Whales (Bal&nd) the scapula is more 
massive and not so broad, and the coracoid is much reduced. 
In Megaptera the scapula is triangular, and neither the 
coracoid nor the acromion forms a distinct process. 

None of the Cetacea possess clavicles. 

The scapula of the SIREXIA is formed on quite a different 
plan, being rather like that of the Seals in shape, narrow, 
and curved backwards. The anterior fossa is nearly as 
large as the posterior. The spine is moderately developed. 

xiv. UNGULATA. 257 

and the slender acromion points downwards. The coracoid 
forms a moderate-sized conical process. There are no 

In the UNGULATA the scapula is always high and rather 
narrow. The prescapular and postscapular fossa; are often 

Fl<;. 81. Right scapula of Red Deer (Crrrus eltf-has), \. st partially ossified 
_suprascapular border: pf postscapular fossa; af anterior or prescapular fossa; 
a acromion ; c coracoid : gc glcnoid cavity. 

subequal. The acromion and coracoid are never much 
developed. The clavicle is always absent. 

The Pecora (see Fig. 81) have all a very large and very 
slowly and imperfectly ossified suprascapular region (ss) ; 
when this is removed, as is almost always the case with 
macerated bones, the upper border of the scapula is very 
straight. The acromion usually forms a distinct process, 


but is quite absent in the Giraffe, which has the longest and 
narrowest scapula of the group. 

In the Horse the scapula is long and slender, the supra- 
scapular border is rounded, and slowly and imperfectly 
ossified. The spine is very slightly developed ; rather above 
the middle its edge is thickened and somewhat turned back- 
wards ; it gradually subsides at the lower extremity without 
forming any acromial process. The coracoid is a prominent 
rounded nodule. 

In the other Perissodactyles, and in the Pigs and Peccaries, 
there is a strongly-marked retroverted triangular process 
on the middle of the edge of the spine, and no true 
acromion ; but in the Hippopotamus there is a small 
acromion and no distinct mid-spinous process. In this 
animal the coracoid is rather long and upturned. 

In the Tapir the coraco-scapular notch is remarkably 

The Hyrax manifests, its affinity with the Ungulata in the 
form of the scapula, which is generally triangular, with a 
small spine, most prominent and with a retroverted edge 
near the middle, and gradually subsiding at each extremity) 
so that there is no trace of an acromial process. 

The Elephant has a largely-developed postscapular fossa 
and a narrow anterior fossa. The glenoid border is short, 
and forms a very prominent angle posteriorly with the 
unusually long suprascapular border. The spine is promi- 
nent, and has a very strongly-marked process projecting 
backwards from near the middle and a moderate-sized 
acromion. The coracoid is small and rounded. 

The EDENTATA present some very interesting conditions 
of the shoulder girdle. 

In the Cape Anteater (Orycteropus) the scapula is of 
the most normal form, with well-developed acromion and 

xiv.] EDENTATA. 259 

coracoid. The middle of the border of the spine is thick- 
ened and retroverted, and there is a well-marked meta- 
cromion. The clavicle is strong, curved, and dilated at its 
sternal end. 

In the Pangolins (Alan is) the scapula is broad, and rounded 
above, the anterior margin gently passing into the superior. 
The prescapular fossa is broader than the postscapular. The- 
suprascapular region remains cartilaginous. The acromion 
is very small. The coracoid is extremely rudimentary, but 
with a separate ossific nucleus. There are no clavicles. 

In the Anteaters (Afyrmecophagidai) the scapula is also 
broad and rounded, so that there is no distinct angle between 
the anterior and superior margin. The anterior margin is 
produced, to meet the large adze-shaped coracoid over the 
coraco-scapular notch, converting it into a foramen. The 
spine has a triangular process in the middle, and a long 
slender acromion, without distinct metacromion. The post- 
scapular fossa is nearly equally divided by a second spine. 

Rudimentary flat clavicles imbedded in muscles are 
present in both Tamandua tetradactyla and Myrmecophaga 

The small climbing Cydoturus didactylus has moderate 
gently curved clavicles. 

In the Armadillos (Dasypodidai) the scapula is rather 
varied in form. The acromion is always very long and 
curved ; in many cases it has a distinct articular facet on its 
inner surface for the upper end of the humerus. (See Fig. 
82, //.) There is a second spine on the postscapular fossa, 
and always a well-developed clavicle. 

In the Sloths (Bradypodida, Fig. 83) the prescapular 
region (of) is larger than the postscapular (pf\ The spine 
arises from little more than the middle third of the bone, 
vertically. In the young of both genera of this family the 




acromion is a long strip of cartilage connecting the spine 
with the end of the coracoid, while the coracoid border of 
the scapula and the coracoid bone join each other in front 
converting the coraco-scapular notch into a small oval fora- 
men (csf\ The acromion and coracoid remain united in the 
Two- toed Sloth (Cholxpus) and the extinct Megatherium; 
but in Bradypus the acromion gradually becomes reduced in 

FIG. 82. Right scapula of Great Armadillo (friodon gigas), \. // postscaptilar 
fossa ; rt/prescapular fossa : gc glenoid cavity ; csn coraco-scapular notch ; c cora- 
coid ; a acromion ; h articular surface for humerus. 

size, losing its connection with the coracoid, and finally 
remains a mere styliform, or slightly flattened process. 

The coracoid in both forms is unusually large, ossifies 
ectosteally according to Parker, and has an epiphysis on its 
free hook-like extremity. 

The clavicle (cl) of Cholcepus is well developed, attached 
external to the loop of bone on the scapula, formed by the 

xiv.] EDENTATA. 261 

united extremities of the acromion (a) and coracoid (^), and 
internally by the intervention of a long fibre-cartilaginous 
" omosternum " (degenerating into a mere ligament in the 
adult) to the presternum. In Bradypus the clavicle is very 
small, and separated by a long interval from the sternum. 
It originally articulates at its scapular end, as in Cholapus ; 
but in consequence of the atrophy of the acromion, it is 

FIG. 83. Right scapula and clavicle of Two-toed Sloth (Cholafius hoffntanm), i\. 
af prescapular fossa ; />/ postcapular fossa ; %c glenoid cavity ; a acromion 
c coracoid ; / coraco-scapular foramen ; cl clavicle. 

left attached to the end of the coracoid, in which unusual 
situation it remains through adult life. 

In the MARSUPIALIA the scapula is tolerably uniform in 
shape. The acromion is long, and the coracoid small, of a 
somewhat hooked form, and thick at the base. It ossifies 
by a separate endosteal nucleus. 

The clavicle is present in all known Marsupials except 


the Bandicoots (Peramelida). It has always a "mesosca- 
pular segment" as its outer end, and a " precoracoid seg- 
ment " at its sternal end ; these are, however, not ossified. 
Most generally it is attached to the acromion by a rather 
strong ligament, but in the Wombat by a synovial articu- 
lation. It is usually connected to the presternum by omo- 
sternal cartilages of varying length, best developed in the 

The shoulder girdle of the MOXOTREMATA (see Figs. 84 and 
85) differs widely, in many points, from that of any other 
Mammal, and far more resembles that of the Lizards. 

The scapula is rather long and narrow, and (especially in 
the Ornithorhynchus) curved backwards and pointed, sickle- 
like, at its upper end. Instead of three it presents but two 
distinct borders and two surfaces ; but the more convex 
border (s), which is turned forwards and outwards in its 
natural position, has a small projection (a) near its lower 
end, which affords attachment to the clavicle, and is evidently 
the acromion ; and the whole border may be considered to 
represent the spine. Following the indications afforded by 
the attachment of the muscles, it appears probable that the 
whole inner surface represents the prescapular fossa of the 
ordinary Mammalian scapula, and that the anterior portion 
of the outer surface (pf) is the postcapular fossa, and the 
posterior portion of the same surface (ssf) the subscapular 
fossa, these two being divided below by a slight ridge (gb), 
which runs to the edge of the glenoid cavity, and from which 
the long head of the triceps muscle takes origin. This 
ridge then answers to the posterior or glenoid border of the 
ordinary Mammal, and the hinder border of the Monotreme's 
scapula (cb) would correspond to the anterior or coracoid 
border. If this is really the case, the scapula of the Mono- 
treme and that of the Cetacean offer the widest contrast, 

xiv.] MONOTREMATA. 263 

the supposed primitive trihedral rod being flattened in 
opposite directions. In the Cetacean scapula there are two 
nearly parallel surfaces, the postscapular and the subsca- 
pular fossse ; while the third, the prescapular fossa, is reduced 
to the smallest possible width quite obsolete, in fact in 
Platanista. In the Monotreme the last-named fossa is so 
expanded that the other two, instead of being parallel to 

Fio. 84. Side view of right shoulder girdle of a young Echidna (Echidna hystrix), , 
ss suprascapular epipnysis ; uf subscapular fossa ; //" postscapular fossa ; cb 
coracoid border ; gf> Ctenoid border ; s spine ; a acromion ; ess coraco-scapular 
suture ; gc glcnoid cavity ; c coracoid ; ec epicoracoid ; cl clavicle ; ic interclavicle ; 
pi prcstcrnum. 

each other on opposite sides of the bone, are brought almost 
into one plane, which is parallel and opposite to the sub- 
scapular fossa. 

The coracoid (c) is a stout subcylindrical bone, expanded 
at its extremities, taking at its upper end a considerable 
share in the formation of the glenoid cavity, and becoming 
firmly ankylosed with the scapula. At its lower end it 


articulates to the side of the presternum, just in front of the 
first rib. 

Placed in front of the inner end of the coracoid is a 
broad, flat, shield-like plate of bone (epicoracoid, ec Fig. 84, 
Prc, Fig. 85), the rounded inner border of which passes 
beyond the median line, overlapping the corresponding bone 
of the opposite side. In the Echidna the left lies superfici- 
ally to the right, while in the Ornithorhynchus this disposition 
is reversed. 

Upon the front end of the presternum, lying below its 
anterior continuation (proosteon, see p. 104) and also below 

FIG. 85. Ventral view of shoulder-girdle and anterior portion of sternum and 
sternal ribs of Ornithorhynchus (O. ^anatinns) ; . sc scapula; cl calvicle ; 
c coracoid ; Prc epicjracoid ; Kp interclavicle or episternum. 

the epicoracoids, is a large azygous T-shaped bone (ic Figs. 
84 and 44 ; Ep Fig. 85), which has no homologue in any 
other Mammal, called interclavicle or episternum. Its lower 
end is broad, and rests on the expanded straight upper 
margin of the presternum ; it contracts somewhat above 
before dividing into a pair of nearly horizontal, slightly-curved 
arms, which extend outwards towards, though not quite 
reaching, the acromion. This bone differs from the pre- 
sternum, and the small proosteal plate behind its lower 

xiv.] MONOTREMATA. 265 

extremity, as well as the coracoids and epicoracoids, in being 
developed in membrane. 

The clavicles (d) are simple, elongated, slightly curved, 
thin, splint-like bones, resting upon the anterior surface of 
the arms of the interclavicle, pointed and not quite meeting 
internally, and dilated and articulating directly with the 
acromion at their outer end. 



IN the upper segment of the limb proper there is always 
one bone, the If it merits ; in the second segment, two bones 
placed side by side, the Radius and Ulna. 

The Humerus (except in some of its extreme modifica- 
tions) is more or less elongated and cylindrical. It is 
described as having a shaft, and two extremities. The upper 
or proximal extremity has a smooth, convex, generally more 
or less rounded head (Fig. 86, //), the axis of which is 
directed upwards and backwards. 1 This, in the living 
animal, is covered with a thin layer of cartilage, and 
articulates by a synovial joint with the glenoid cavity of the 
shoulder girdle. The head is marked off from the shaft 
very indistinctly by a constriction called the neck, imme- 
diately below which, upon the anterior surface of the bone, 
are two rough prominences (/ and /') for the attachment of 
muscles, called tnberosities, separated from each other by a 
groove (bg) called the bicipital groove, as the tendon of the 
biceps muscle runs in it after arising from the margin of the 
glenoid fossa. The tuberosities are generally distinguished 

1 The terms of relative position here used are those which the bone 
assumes in the ordinary attitude of a quadruped while standing or 


as great (/) and small (/') from their relative size in Man and 
most other Mammals ; the former is also called external, and 
the latter internal, from their relative situation in the most 
usual position of the bone. 

The lower or distal extremity of the humerus is some- 
what flattened, and usually has a broad, semi-cylindrical 
articular surface, which is received into a corresponding 

FIG. 86. Anterior surface of right humerus of Wombat {Phascolomys romtatus), J 
k head ; l>g bicipitnl groove ; / great or external tuberosity ; f small or internal 
tuberosity ; fir deltoid ridge ; sr supinator ridge ; cf supra-condylar foramen ; 
ec external condyle ; ic internal condyle ; ar articular surface for radius ; au arti- 
cular surface for ulna. 

concavity on the upper end of the bones of the fore-arm. 
This is called the trochlea (ar and au). On each side, and 
rather above this surface, is a prominence called the condyle, 
one of which is external, or radial (ec), the other internal, or 
ulnar (ic). The latter is usually the most prominent. 1 In 

1 "In anthropotomy, the term 'condyle' rightly applied to the 
prominent articular convexities of the occipital, mandibular and femoral 


the middle of the lower end, between the condyles, and 
above the thickened articular border, the bone is very thin, 
having a hollow both in front and behind. The latter, 
which is the deepest, is called the anconeal fossa, as it 
receives a projecting part of the anconeal process, or ole- 
cranon, of the ulna, when the fore -arm is fully extended. 
It often happens that these two fossas are so deep that they 
meet, and there is in consequence a vacuity in the bone 
called the siipratrochlear or intercondylar foramen. There is 
usually a prominent ridge running upwards for some distance 
on the shaft from the external condyle, called the ectocondylar 
or supinator ridge (sr), which affords a wide surface of origin 
for the supinator muscles of the fore-arm. When much 
developed this ridge terminates above at the groove for the 
passage of the musculo-spiral nerve. When the edge of the 
bone above the inner condyle is much developed, it is some- 
times grooved, but more often obliquely perforated from 
above, downwards and forwards, by a supracondylar foramen 
(cf\ through which the median nerve and brachial artery 
may pass. Lastly, somewhere towards the anterior sur- 
face of the middle of the shaft, on the outer side, there is 
usually a roughened, elevated, longitudinal ridge (dr), some- 
times developed into a tuberosity, for the insertion of the 
deltoid muscle, and hence called the deltoid ridge. The 
lower end of this ridge is separated from the upper end of 
the supinator ridge by a wide and shallow groove, winding 
in a spiral manner downwards and forwards round the outer 

bones, is transferred from the distal articular prominence of the humerus 
to the processes for attachment of muscles above the joint surfaces. 
I have found it convenient in comparative osteology to indicate the 
homologies of the ' external condyle ' and ' internal condyle ' of the 
human humerus by the terms ' ectepicondyle ' and ' entepicondyle ' 
(Owen)." By French anatomists they are called epicondyle and epi- 
trochlea respectively. 


side of the shaft of the bone, and indicating the course of 
the musculo-spiral nerve. 

The whole of the shaft of the humerus is developed 
ectosteally from a single centre of ossification, but at each 
extremity there is a large epiphysis : the upper one includes 
the head and both tuberosities, and is usually formed by the 
coalescence of two distinct endosteal nuclei ; the lower one 
includes the whole inferior articular surface with the condyles, 
and is formed of three or four originally distinct centres of 
ossification. The lower epiphysis unites to the shaft before 
the upper one. 

The skeleton of the second segment of the upper limb, 
the fore-arm or antibrachium, consists of two bones called 
radius and ulna, placed side by side, articulating with the 
humerus at their proximal, and with the carpus at their 
distal, extremity. 

In their primitive or unmodified condition, these bones* 
may be considered as placed one on each border of the 
iimb, the radius being preaxial, and the ulna postaxial. 1 
The radius articulates above with the preaxial (external) 
side of the humerus, the ulna with the postaxial (internal) 
side of the humerus. 

This position is best illustrated in the fore-limb of the 
Cetacea (see Fig. 104), where the two bones are fixed side by 
side and parallel to each other, the preaxial border being 
external, and the postaxial border internal, in their whole 

In the greater number of Mammals, the bones assume a 

1 So termed (by Prof. Huxley) in relation to the assumed central axis 
of the limb, when it is extended out from the body in a certain stage of 
its embryonic position, the extensor surface of the arm and back of the 
hand being upwards or dorsal, and the flexor surface of the arm and palm 
of the hand being downwards or ventral. 


very modified and adaptive position (as will be explained 
more fully in the chapter on the comparison of the fore and 
hind limbs), usually crossing each other in the fore-arm, the 
radius in front of the ulna, so that the preaxial bone 
(radius), though external (in the ordinary position of the 
limb) at the upper end, is internal at the lower end ; and 
the hand being mainly fixed to the radius, also has its pre- 
axial border internal. In the large majority of Mammals 
the bones are fixed in this position ; but in some few, as 
in Man, a free movement of crossing and uncrossing, or 
pronation and supinaiion, as it is termed, is allowed between 
them, so that they can be placed in their primitive parallel 
condition, when the hand (which moves with the radius) is 
said to be supine, or they may be crossed, when the hand is 
said to be prone. 

In most Mammals which walk on four limbs, and in which 
the hand is permanently prone, the ulna is much reduced in 
size, and the radius increased, especially at the upper end ; 
and the articular surface of the latter, instead of being con- 
fined to the external side of the trochlea of the humerus, 
extends all across its anterior surface, and the two bones, 
instead of being external and internal, are anterior and 
posterior (see Figs. 87, 88, and 89). 

The ulna is always characterised by a conspicuous, more 
or less compressed prolongation, extending upwards beyond 
the excavated humeral articular surface (sigmoid notch), and 
serving as the point of attachment to the extensor muscles 
of the fore-arm, called the olecranon or anconeal process. 

Each of the bones of the fore -arm has commonly a 
principal centre of ossification for the shaft, and an epiphysis 
at either end. 



Special Characters of the Bones of the Arm and 
Fore-ami in the different Groups. 

Order PRIMATES. In Man the humerus is long, slender, 
and straight, with a large globular head. Neither the tube- 
rosities, nor the deltoid and supinator ridges, are much 
developed. The internal condyle is prominent, but there is 
no supracondylar foramen as a normal condition. The 
anconeal fossa is wide and deep, and sometimes, though 
not usually, jxjrforated. The lower articular surface is divided 
by a groove into a pulley-like internal portion (trochlea} for 
the ulna, and a smaller rounded portion (capitelluni), confined 
to the front side of the bone, for the radius. 

The whole bone is somewhat twisted on its longitudina 
axis. Supposing it is so placed that a line drawn horizontally 
though the axis of the head passes directly backwards, 
another line drawn through the condyles would not cross 
this at a right angle, as in most of the inferior Mammals, but 
its outer end would be directed forwards. This appears to be 
caused by a process of resorbtion and new deposition of bony 
tissue during the growth of the embryo, producing an effect 
resembling torsion of the axis to an extent of about 35. a 

The radius has the head or proximal end expanded, disk- 
shaped, and cupped at its extremity, which is applied to the 
capitelluni of the humerus (see Fig. 87). Below this 
expanded head the bone is comparatively slender, but 
increases in size as it approaches the lower end, which is 
wide from side to side ; the surface next the ulna being 
hollowed to receive the lower end of that bone, while the 

1 See Gcgenbaur, " Jenaischc Zeitschrift," Bd. IV. p. 50, nlso his 
" I,ehrl)uch der Anatomic des Mcnschen." Leipzig, 1883. For an 
opposite view, Albrecht, " Bcitragzur Torsionstheorie, etc." Kiel, 1875 


opposite side is produced into the radial styloid process. 
The inferior surface is hollowed for articulation with the 
carpus. The whole bone is slightly curved. Not far below 
the head is a rough prominence, into which the tendon of 
the biceps flexor muscle is inserted. 

The ulna has a large sigmoid excavation above for articu- 
lation with the trochlea of the humerus ; the pointed elevated 
anterior edge of this is called the coronoid process. The 
olecranon is scarcely produced upwards beyond the hinder 
edge of the articular surface. Below this, on the radial side, 
is a smaller excavation, in which the edge of the disk-like 
head of the radius plays, being held in its place in the living 
state by a strong annular ligament, which encircles it. The 
ulna is straighter than the radius, and gradually diminishes 
in size to the lower end, where it terminates in a rounded 
surface, which articulates with the hollow in the lower end 
of the radius ; and also, though not very directly with the 
proximal surface of the carpus. The ulnar side is produced 
into a small conical process, the ulnar styloid process. 

The movement of these bones upon the humerus at the 
elbow-joint is simply that of a hinge, formed mainly by the 
articular surface of the ulna. In pronation and supination, 
which is more free and complete than in any other Mammal, 
the ulna is stationary, and the radius moves ; the upper end 
only on its own axis, being fixed to the side of the ulna by 
the annular ligament, but the lower end rotates round the 
lower end of the ulna, carrying the hand with it. 

The higher Apes have the axis of the humerus almost as 
much twisted on itself as in Man ; and they also, almost 
alone among Mammals, resemble Man in not having the 
olecranon process of the ulna prolonged upwards beyond 
the sigmoid notch. Even in the Baboons these special an- 
thropoid characters are wanting. The humerus has no supra- 




condylar perforation in any of the Old World Simiina, nor in 
Ateles, Mycetes or Hapale among the American Monkeys ; 
but in the remaining genera of Cebida, and in most of the 
Lemurs, such a perforation is found. In the Aye-Aye 
(Chiromys) the supinator ridge is remarkably developed. 
The radius and ulna are distinct in all ; in the higher forms 
(especially the Gorilla) greatly curved, leaving a large space 

FIG. 87. FIG. 88. FIG. 89. 

Anterior aspect of the bones forming the right elbow-joint of Man (Fig. 87) ; of the 
Dog (Fig. 88) ; of the Red Deer (Fig. 89) ; all J. h humerus ; r radius ; u ulna. 

between them in the middle of the fore-arm. The power of 
supination and pronation, which in the higher forms almost 
equals that enjoyed by Man, is much reduced in the inferior 
types of the order, although never entirely lost. 

In the CARNIVORA the head of the humerus has no longer 
that hemispherical form, so well marked in the higher 
Primates. The tuberosities are strong and rough, and 
project upwards beyond the level of the head. The shaft 


is much curved forwards. The deltoid ridge is strong, and 
extends far down on the bone, especially in the Bears. The 
inner condyle is prominent. The anconeal fossa is deep. A 
supracondylar foramen exists in the Felidce, and in most of 
the Viverridce, Mustelidtz, and Procyonidce, but not in the 
Canidce, Hycenida, or Ursidiz. 

The radius differs from that of Man, inasmuch as its 
upper end is broad, flattened, and extends further across the 
front of the humeral articular surface, forming part of the 
hinge (see Fig. 88) ; and although it is never ankylosed with 
the ulna, scarcely any appreciable amount of movement is 
allowed between them. The ulna has a large compressed 
olecranon, and a shaft gradually tapering to the lower 

In the Pinnipedia the bones of the anterior limb are very 
short and stout. The humerus has a remarkably prominent 
deltoid ridge, and usually no supracondylar foramen, though 
this is present in the Common Seal (P/wca ritulina}. The 
upper end of the ulna, and conversely the lower end of the 
radius, are much expanded. 

In most of the INSECTIVORA the bones of the arm gene- 
rally resemble those of the Carnivora. In the Hedgehog 
(Erinaceus) there is no supracondylar foramen in the 
humerus, but a large supratrochlear perforation. In Centetes 
Rhynehocyon, and nearly all the other genera, there is a 
supracondylar foramen. 

The radius and ulna are generally completely developed 
and distinct ; but in Galeopithecus, Macroscelides and Petro- 
dromus, they are fused together distally. 

The Mole (Talpa) and its allies have a humerus of ex- 
traordinary form (see Fig. 34, p. 95), being very short, and 
extremely broad and flattened at both extremities, though 
contracted in the middle. In addition to the narrow oval 


head for articulation with the glenoid cavity ot the scapula, 
there is a larger saddle-shaped surface, which articulates by 
a separate synovial joint with the outer end of the coraco- 
clavicle. The deltoid ridge is very prominent, joining the 
inner tuberosity above. From each condyle a slender bony 
process extends upwards. There is a supracondylar fora- 
men. The ulna has a greatly developed olecranon, with a 
narrow keel behind, and expanded laterally at the extremity. 

In the Cape Golden Mole (Chrysochloris) the humerus 
is much more slender generally than in the true Moles, but 
the inner condyle is extremely elongated. The olecranon 
is long, narrow, and incurved The fore-arm has a third 
bone, extending from the palmer surface of the carpus 
almost to the elbow, where it has a free termination. 
This appears to be an ossification in one of the flexor 

In the CHIROPTERA both arm and fore -arm are long and 
slender. The humerus has a slight sigmoid curve, its ulnar 
tuberosity is large, and there is no supracondylar perfora- 
tion. The ulna is extremely reduced, only the proximal 
third being present, and that ankylosed with the radius, which 
forms almost the whole of the lower articular surface of the 
elbow-joint. There is a detached sesamoid ossicle near the 
olecranon, in the tendon of the triceps muscle. 

In the RODENTIA the humerus varies much in its charac- 
ters. It is long, slender, and straight, with a very slight 
deltoid ridge, a narrow and laterally compressed inferior 
end, and without prominent condyles in the Hares and 
Agutis. But in the Beaver the deltoid and supinator ridges, 
and the inner condyle, are strongly developed. All inter- 
mediate conditions occur in different genera. As a general 
rule there is a large supratrochlear perforation, but no 
supracondylar foramen. In the Coypu (Myopotamus) the 

T 2 


deltoid ridge is an extremely salient, compressed, and 
everted tuberosity. 

In the fore-arm the two bones are nearly always dis- 
tinct, though closely applied to each other. The breadth 
of the upper end of the radius, and the amount of 
rotation permitted upon the ulna, vary much in different 

In the great order UNGULATA the hnmerus is stout and 
rather short. The outer tuberosity is very large, and gene- 
rally sends a strong curved process inwards, overhanging the 
bicipital groove (not, however, in the Horse and Camel). 
The deltoid ridge is usually not strongly marked, and placed 
rather high on the bone ; but in the Rhinoceros it is a very 
salient ridge. The lower end is always particularly straight 
and flat on the inner side (see Fig. 89, p. 273), the condyle 
forming no prominence, and there is never a supracondylar 
oramen. The outer condyle and the ridge above it are 
rather more developed. 

The radius is large at both ends, and superiorly extends 
across the whole of the humeral trochlear surface (see Fig. 
89). The ulna is a complete and distinct bone in the Pig, 
Hippopotamus, Tapir, and Rhinoceros. In the Ruminants 
it is more or less rudimentary and fixed behind the radius. 
In the Camel the two bones become completely coalesced. 
In the Horse the olecranon and proximal part of the shaft 
alone remain, firmly ankylosed to the radius. 

In the PROBOSCIDEA the humerus is remarkable for the 
great development of the supinator ridge. The ulna and 
radius are quite distinct, and permanently crossed. The 
upper end of the latter is small, while the ulna not only 
contributes the principal part of the articular surface for the 
humerus, but has its distal end actually larger than that of 
the radius, a Condition unique among Mammals. 

xv.] CETACEA. 277 

In Hyrax the humerus is straight, with a very prominent 
outer tuberosity, moderate deltoid ridge, rather compressed 
inferior extremity, large supratrochlear, but no supracon- 
dylar, perforation. The ulna and radius are complete and 
subequal, often ankylosing together in old animals. 

In the CETACEA, the bones of the arm and fore-arm are 
usually very short, broad, and simple in their characters 
(see Fig. 103). The humerus has a large globular head, 
which moves freely in the glenoid cavity of the scapula, 
the tuberosities are fused into one, the bicipital groove 
being absent ; the lower end is broad and flattened, and 
its inferior surface is divided into two nearly equal flat 
surfaces placed side by side (one external, the other internal), 
and meeting at a very obtuse angle. The equally flat 
upper surfaces of the radius and ulna are applied to these, 
and so united that scarcely any motion is permitted between 
them, and often in old animals ankylosis takes place at 
the joint. 

The ulna and radius are parallel to each other without 
any indication of crossing : the former has a tolerably well- 
developed olecranon process projecting directly outwards 
from the shaft of the bone ; the radius is extremely simple 
in form, wider distally than proximally. 

In the Rorquals (Balanoptera and Megaptera) these bones 
are considerably elongated. 

In the SIRENIA the bones of the fore-limb are formed on 
a different type, as there is a distinct, though small and 
simple, trochlear articulation at the elbow-joint. In the 
Dugong, the humerus is small in the middle of the shaft, 
and expanded at each end. The tuberosities are very pro- 
minent, especially the outer one, and the bicipital groove is 
distinct The internal condyle is prominent, the anconeal 
fossa small, and there is no supracondylar perforation. In 


the humerus of the Manatee the bicipital groove is obsolete, 
the two tuberosities coalescing, as in the Cetacea. In other 
respects it resembles that of the Dugong. 

The two bones of the fore-arm are, in both genera, about 
equally developed, and generally ankylose together at both 

Order EDENTATA. In the Sloths the humerus is long 
and straight, slender and cylindrical in the greater part, but 
flattened and laterally expanded at the lower end. The 
head is hemispherical, the tuberosities moderately developed, 
and subequal in size, the deltoid ridge very indistinct. In 
the Two-toed Sloths (Chol&pus) the humerus is shorter and 
broader than in Brady pus, and has a large supracondylar 
perforation, which is wanting in the latter genus. 

The radius and ulna somewhat recall those of the 
Primates in their form, and they are capable of a con- 
siderable amount of pronation and supination. The 
olecranon process scarcely projects beyond the sigmoicl 
articular surface. 

The humerus in all the remaining Edentates is stout and 
broad, and remarkable for the great development of the 
points of muscular attachment, as the tuberosities, deltoid, 
and supinator ridges, and internal condyle. These all 
reach their maximum of development in the Armadillos, 
animals which make great use of their fore-limbs in 
scratching and burrowing. The supracondylar foramen is 
present in all. 

The radius and ulna are also well developed and 
distinct in all, but with no great amount of motion 
permitted between them. The olecranon is always long 
and strong. 

Order MARSUPIALIA. In the burrowing Wombat (Phasco- 
lomys) the humerus is stout, very broad at the distal end, 

xv.] MARSUPIALIA. 279 

and with strongly developed deltoid and supinator ridges 
(see Fig. 84, p. 242.) These characters prevail generally 
throughout the order, though in a less marked degree. The 
supracondylar foramen is almost always present, some of 
the Dasyures being exceptions. 

The radius and ulna are always distinct and well-developed 
bones. The upper end of the radius is small and rounded, 
and more or less rotation is permitted between the bones, 
even in the carnivorous forms. 

In the animals constituting the order MONOTREMATA, 
the humerus is something like that of the Mole, short and 
extremely broad at both extremities, with greatly produced 
inner and outer condyles, though contracted at the middle 
of the shaft. 

The radius and ulna are stout, and rather flattened at the 
lower end, where they are of about equal size, and closely 
applied together. The upper end of the olecranon is 
widely expanded laterally. 



THE terminal segment of the anterior limb is the hand 
or manus^ Its skeleton consists of three divisions : (i) The 
carpus, a group of small, more or less rounded or angular 
bones, with flattened surfaces applied to one another, and, 
though articulating by synovial joints, having scarcely any 
motion between them ; (2) the metacarpus, a series of 
elongated bones placed side by side, with their proximal 
ends articulating by almost immovable joints with the 
carpus ; (3) the phalanges, or bones of the digits, usually 
three in number to each, articulating with one another by 
freely movable hinge-joints, the first being connected in 
like manner to the distal end of the corresponding meta- 
carpal bone. 

To understand thoroughly the arrangement of the bones 
of the carpus in Mammals, it is necessary to study their 

1 "On account of the ambiguity arising from the as yet unsettled con- 
notation of the terms ' hand ' and ' foot,' I think it better, in a scientific 
treatise, to disuse them altogether, and ... to adopt for the anterior 
extremity (the carpus and all beyond it) the term manus, and for the 
homotypal posterior segment the term pes. The all but necessity for 
distinct homological terms for such parts is obvious." Mivart, " On the 
Appendicular Skeleton of the Primates," Phil. Trans. 1867. 

CH. XVI.] 



condition in some of the lower vertebrates. 1 Fig. 90 repre- 
sents the manus in one of its most complete, and at the 
same time most generalised, forms, as seen in one of the 
Water Tortoises (Chelydra serpentina). 

The carpus consists of two principal rows of bones, an 
upper or proximal row, containing three bones, to which 
Gegenbaur has applied the terms radiah (r), intermedium (/'), 

FIG. 90. Dorsal surface of the right manus of a Water Tortoise (Chelydra ser- 
ftntitta), after Gegenbaur. u ulna ; R radius ; ulnare ; intermedium ; r radiale ; 

c centrale ; 15 the five bones of the distal row of the carpus ; m 1 / 

the five 

and ttlnare (u), the first being on the radial or preaxial side 
of the limb. The lower, or distal row, contains five bones, 
called carpale i, 2, 3, 4, and 5 respectively, commencing on 
the radial side. Between these two rows, in the middle of 
the carpus, is a single bone, the centrale (c). 

In this very symmetrical carpus, it will be observed that 

1 See Gegenbaur, " Untersuchungen zur vergleichenden Anatomic," 
Ites Heft, Carpus und Tarsus. 1864. 


the radiale supports on its distal side two bones, carpale i 
and 2 ; the intermedium is in a line with the ccntrale and 
carpale 3, which together form a median axis of the hand, 
while the ulnare has also two bones articulated with its 
distal end, viz. carpale 4 and 5. Each of the carpals of the 
distal row supports a metacarpal. 

In the carpus of the Mammalia, there are usually two 
additional bones developed in the tendons of the flexor 
muscles, one on each side of the carpus, which may be 
called the radial and ulnar sesamoid bones ; the latter is 
most constant and generally largest, and is commonly known 
as the pisiform bone. The fourth and fifth carpals of the 
distal rows are always united into a single bone, and the 
centrale is often absent. As a general rule all the other 
bones are present and distinct, though it not unfrequently 
happens that one or more may have coalesced to form a 
single bone, or may be altogether suppressed. 

The table bilow shows the principal names in use for the 
various carpal bones. Those in the second column, being 
most generally employed by English anatomists, will be 
adopted in the following pages : 

Jtaiiia'.e --- Scaphoid Naviculare. 

Intermedium Lunar Scmiliinare, Lunatum. 

Ulnare - Cuneiform - Triquclrum, Pyramidal/'. 

Ct nt rale - Central Intermedium (Cuvier). 

Carpale \ - Trapezium - Multangnlum inajns. 

Carpale 2 = Trapezoi.l -- Mnllattguhiin niinu*. 

Carpale 3 Magnum - Capitatum. 

P Unciform Ifamatunt, Uncinatnin. 

Carpale 5 ) 

The metacarpal bones, with the digits which they sup- 
port, never exceed five in number, and are distinguished by 
numerals, counted from the radial towards the ulnar side. 
The digits are also sometimes named (I) fiollex, (II) index. 


(III) medius, (IV) annularis, and (V) minimus. One or more 
may be in a very rudimentary condition, or altogether 
suppressed. If one is absent, it is most commonly the 
first, next follows the fifth ; the third is never lost, although 
either the second or fourth, or both, may be absent. 

Excepting the Cetacea, no Mammals have more than three 
phalanges to each digit, but they may occasionally have 
fewer by suppression or ankylosis. 

The first or radial digit (also called pollex) is an exception 
to the usual rule, one of its parts being constantly absent, 
for while each of the other digits has commonly a meta- 
carpal and three phalanges, it has only three bones altogether. 
Whether the missing one is the metacarpal or one of the 
phalanges, is a subject which has occasioned much dis- 
cussion, but has not yet been satisfactorily decided. In 
accordance with the most usual custom, the proximal bone 
of this digit will here be treated of as a metacarpal. 1 

The terminal phalanges of the digits are often specially 
modified to support the nail, claw or hoof, and are called 
" ungual phalanges." 

Very frequently a pair of small sesamoid bones are 
developed in connection with the tendons passing over the 
palmar surface of the articular heads of the metacarpals and 
phalanges, and occasionally (as in the Armadillos) a larger 
bone of similar nature is met with in the middle of the 
same surface of the carpus and metacarpus. More rarely 
similar bones occur on the dorsal surface of the phalangeal 
articulations. 2 

Each of the carpal bones ossifies from a single nucleus. 

1 See Allen Thomson " On the Ossification of the First Metacarpal 
Bone." ("Journ. Anat. Physiology," vol. iii. p. 131.) 

2 These are almost always lost in prepared skeletons, but they occur 
constantly in the common dog. 



The metacarpals and phalanges have each a main nucleus 
for the greater part of the bone, and usually an epiphysis at 
one end only, this being the distal end of the metacarpals 
(except the first), and the proximal end of the first meta- 
carpal and of all the phalanges. In many of the Cetacea 
epiphyses are found at both ends of the bones, and the 
same takes place regularly in the Elephant Seal (Macro- 
rhinus leoninus), and occasionally in some other Mammals. 1 


FIG. 91. Bones of the right human carpus, J. s scaphoid ; / lunar ; c cuneiform ; 
tin trapezium ; td trapezoid ; in magnum ; it unciform ; / pisiform ; i v the 

In Man the carpus (see Fig. 91) is short and broad. Its 
upper border, by which it articulates with the radius, forms 
a regular curve, with the convexity upwards. It has the 
three bones of the proximal row the scaphoid (s), lunar 
(/), and cuneiform (c) distinct ; also the usual four bones 
of the distal row, but no central. 2 There is a well-developed 

1 The first phalanx of the Horse's toe has a distal epiphysis, which 
unites to the shaft at a much earlier period than that at the proximal 
end. See Struthers "Brit. Assoc. Report, 1885." 

2 The os cenlrale carpi has been found in the cartilaginous stage in 
the embryo, but disappears early, although in rare cases it persists in 
the adult as a separate bone. See E. Rosenberg, " Morphol. Jahrb. '' 
I. p. 172. H. Leboucq, " Acad. Roy. Belg." 3rd ser. torn. iv. 1882. 
W. Gruber, " Beobachtungen aus der menschlichen unct vergleichenden 
Anatomic," IV. Heft. Berlin, 1883. 

xvi.] PRIMATES. 285 

rounded ulnar sesamoid (the pisiform bone, /) which articu- 
lates by a smooth facet with the cuneiform, but no radial 
sesamoid. The trapezium (////) has a saddle-like articular 
surface for the movable first metacarpal. The magnum 
(m\ as its name implies, is the largest bone rather an 
exceptional condition among Mammals ; it has a large 
rounded part or head projecting upwards and fitting into a 
concavity in the distal surface of the bones of the proximal 
row. The unciform (u) has a strong hook-like process 
from its palmar surface curved towards the radial side. 

The first metacarpal is shorter, though somewhat broader 
than any of the others. It is articulated in a different plane 
from them, its palmar surface facing towards the ulnar side 
of the hand, and it is capable of a considerable range or 
movement. The other four metacarpals are nearly equally 
developed, diminishing slightly from the radial to the ulnar 
side ; their shafts are slender and rather compressed, 
especially towards the palmar aspect ; but they enlarge at 
each extremity, particularly at the rounded distal end or 
head. They are so articulated with the carpus as to allow 
of very little motion. The phalanges are of the normal 
number to each digit, all broad, convex on their dorsal, and 
flattened or slightly hollowed on their palmar, side. The 
proximal is the largest, and the ungual the smallest. The 
latter is flattened and slightly expanded or spatulate at its 
terminal portion. Those of the first, digit or thumb are 
stouter than any of the others, those of the fifth digit (little 
finger) are the most slender. The third digit is the longest, 
the second and fourth somewhat shorter and nearly equal , 
the fifth considerably shorter, and the first still more so. 
Sesamoid bones are only developed below the metacarpo 
phalangeal joint of the pollex. 

In the other PRIMATES, the manus is generally longer an 



narrower than in Man, and as a general rule the first digit or 
thumb is less developed and less freely movable. In the 
genera Troglodytes and Simia, as in Man, the proximal 
surface of the carpus articulates with the radius alone, in all 
others it articulates also with the ulna. The scaphoid and 
lunar bones are always distinct. An os centrale, (Gegen- 
baur, Fig. 92, ce) is present in all, except in the Gorilla and 
Chimpanzee, and some of the Lemurs. Where it is wanting 
as an independent bone it probably has become fused with, 
and then forms part of, the scaphoid. The pisiform is 
present in all, and generally of a more elongated form, and 
more salient than in Man : and there is usually a small 

FIG. 92. Bones of the carpus of a Baboon (fyiiocephalus anubis), J. s scaphoid ; 
/ lunar ; c cuneiform ; p pisiform ; ce central ; rs radial sesamoid ; tin trapezium ; 
td trapezoid ; in magnum ; it unciform ; I v the metacarpals. 

rounded radial sesamoid (rs) articulating movably to the 
border of the scaphoid and trapezium, and connected with 
the tendon of the flexor carpi radialis. 

In the Potto (Perodicticus) there is an additional bone on 
the palmar side of the carpus, an ossification in the ligament 
connecting the palmar processes of the trapezium and 
unciform bones, and forming with these processes a com- 
plete bony ring, through which the flexor tendons pass. 1 

1 Mivart, " On the Appenclicular Skeleton of the Primates ; " Phil. 
Trans. 1867. 




The metacarpals and phalanges are of the complete and 
normal number in all, with the following exceptions. In the 
African genus of Long-tailed Monkeys (Cclobus), and also in 
the American Spider Monkeys (Ateles\ the thumb is rudi- 
mentary, having usually but one very minute phalanx, in 
addition to the metacarpal. In the Potto (Perodicticus) and 
some allied Lemurina, the second (or index) digit is very 
much shorter than the others, and has but two rudimentary 

The phalanges are generally more curved than in Man, 
most notably so in the Orang. The hand of the Mada- 
gascar Aye- Aye (Chiromys) is remarkable for the extreme 
attenuation of the bones of the third digit. 

FIG. 93.-l!onesof the carpus of a lk-ar (Ursus anteriranus), }. si scapho-iunar 
bone ; c cuneiform ; / pisiform ; ti unciform ; tit magnum ; td trapezoid ; ttn tra- 
pezium ; rs radial scsamoid ; I v the metacarpals. 

In the CARNIVORA, the scaphoid and lunar bones always 
coalesce into a single scapho-lunar bone (Fig. 93, sl\ with 
which the centrale is united, the latter never appearing 1 as 
a distinct bone, except sometimes in very young animals. 
The radial accessory ossicle or sesamoid (rs) is generally 
present. All have five digits, with the complete complement 
of phalanges, except the Hyaena, in which genus the pollex 

1 See B. G. Wilder, " On the Composition of the Carpus in Dogs." 
Bull. Cornell University, vol. i. p. 301, 1874. 

288 7 HE MAN US. [CHAP. 

is represented only by a rudimentary metacarpal. This digit 
is usually much reduced in size, and often, as in the Dog, 
does not reach the ground in walking. It is best developed 
in the Bears and allied forms. The first metacarpal is never 
more freely movable than any of the others. As a general 
rule the middle digit is somewhat the longest, the second 
and fourth nearly equal to it, the fifth shorter, and the first 
the shortest. 1 

As the toes are nearly always armed with large, strong, 
curved, and sharp claws (see Fig. 94), the ungual phalanges 

i<;. 94. The phalanges of the middle digit of the manus of the Lion (Fells led). I. 
///' proximal phalanx ; //r middle phalanx ; flfi ungual phalanx ; a the central 
portion forming the internal support to the horny claw ; /'the bony lamina reflected 
around the base of the claw. 

are large, strongly compressed, and pointed, and they 
develop from their base a broad thin lamina of bone (/>), 

1 The fissiped Carnivora have been divided into two groups, accord- 
ing to the position of the feet in walking the Plantigrade, or those 
that place the whole of the palmar and plantar surface to the ground ; 
and the Digitigrade, or those that walk only upon the phalanges, the 
metacarpals and metatarsals being vertical and in a line with the fore- 
arm or leg. This distinction, however, is quite an artificial one, and 
every intermediate condition exists between the extreme typical planti- 
grade gait of the bears and the true digitigrade action of the cats and 
dogs. In fact, the greater number of the Carnivora belong to neither 
of these groups, but maybe called "subplantigrade," often, when at 
rest, applying the whole of the sole to the ground, but keeping the heel 
raised to a greater or less extent when walking. 

xvi.] CARNIVOKA. 289 

which is reflected over the root (a) of the horny claw, and 
holds it more firmly in its place. In those genera, as Felts, 
in which the claws are retractile, the middle phalanx (ph'} 
is deeply hollowed on its ulnar side to receive the ungual 
phalanx when folded back upon it, in the quiescent state of 
the foot. 

In the Pinnipedia, the manus is broader and flatter 
than in the terrestrial Carnivora. The scaphoid and lunar 
coalesce. The ulnar side of the carpus is much reduced, 
the unciform being especially small, and consequently the 
fifth metacarpal articulates partly with the cuneiform of the 
proximal row of the carpus. The pisiform is small. The 
first digit is nearly as long as the second ; the remainder 
gradually diminish in length to the fifth. The ungual pha- 
langes in the ordinary Seals are slender, pointed, slightly 
curved, and not much compressed. In the Otariidtc they are 
prolonged beyond the part which bears the very small claw, 
and flattened and truncated at the ends, being continued 
onwards in the living animal as cartilaginous rays, which 
support lobed expansions of the skin. 

Among the INSF.CTIVORA, the scaphoid and lunar coalesce 
in Galfopithecus, Ttipaia, CtnMes, Solenodon, Erinaceus, and 
Gymnura, but in most of the other forms these bones are 
distinct. A distinct os centrale is found in all except 
Galeopithect4S, Potamogale, Chrysochloris, and Sorex. 

There are nearly always five digits, but Rhynchocyon, 
Oryzoryctes tctradactylus and Chrysochloris have but four, 
the pollex being absent. The whole hand is generally 
of moderate size, with pointed, conical, slightly curved, 
ungual phalanges. 

In common with every segment of the anterior extremity, 
the manus of the Mole (Talpd) and its immediate allies is 
extremely modified to suit its fossorial habits (see Fig 95). 




It is extremely broad and strong, its breadth being increased 
by the great development of the radial sesamoid (rs), which, 
being sickle-shaped, has received the special name of os 
faldforme. The ungual phalanges are very large, and cleft 
at their extremities 

FIG. 95. Bones of fore-arm and manus of Mole (Talpa europccd) X 2. R radius 
U ulna ; s scaphoid ; / lunar ; c cuneiform ; p pisiform ; unciform ; m magnum : 
id trapezoid ; tin trapezium ; ce central ; rs radial sesamoid (falciform) ; i v ihc 

In the CHIROPTERA, the hand is especially modified in a 
totally different manner, constituting the organ of flight. 

In the carpus the scaphoid and lunar are united, and in 
some genera (as Ptcropus) the cuneiform is joined with them, 
so that the proximal row contains but a single bone. There 
is no centrale. The pisiform is very small. 




The pollex is short, divaricated from the other digits, not 
enveloped, as they are, in a cutaneous expansion, and armed 
with a curved claw. The other digits are extremely long and 

In the Frugivorous Rats (Fteropida) the second digit has a 
short ungual phalanx and claw, but in each of the other 


Fit;. 96. B^nes of the manus of Beaver (Castor canadensis), j. 

digits the middle phalanx is elongated, and gradually tapers 
to the termination, the ungual phalanx being absent. 

In the Insectivorous Bats, the pollex alone has a claw, 
and the elongation of the other digits is chiefly due to the 
metacarpals, the phalanges being small and very slender, 
and usually only two in number, except in the third digit, 
which occasionally (as in Phyllostomida J ) has three. 

' G. E. Dobson, " Catalogue of Chiroptera," Brit. Mus.'i878. 
U 2 




In the RODENTIA, the scaphoid and lunar are very 
generally united (as in Castor, Dasyprocta, Hydrochcems, 
Capromys, Sciurus, Arctomys, Mus, &c.), but not in all. An 
os centrale is present in many, as Lepus, Dasyprocta, Hydro- 
choerus, Capromys, and Castor, while in other genera it is 
absent. There is very frequently an accessory ossicle on the 
radial side of the carpus, which is particularly large in the 
Beaver (Fig. 96). There are nearly always five digits, with 


FIG. 97. Bones of the manus of Cape Hyrax (Hyra.v capensis), nat. size. 

the normal number of phalanges, though sometimes (as in 
Hydrochcenis} the pollex is rudimentary or suppressed. 

HYRAX and the ELEPHANTS, together with various extinct 
forms (Dinoceras, Coryphodon, Toxodon, &c.), are united 
by Professor Cope under the name of Subungulata, in 
opposition to the Ungulata vera. In all of these the 
bones of the proximal and distal rows of the carpus retain 
their primitive or more typical relation to each other. The 




os magnum of the second row articulates mainly with the 
lunar of the first, or with the cuneiform, but not with the 
scaphoid. The axis of the third toe passes therefore through 
the os magnum and through the os lunare. 

In the Cape Hyrax (Fig. 97) there is an additional carpal 
hone (), which is probably the os centrale, though in 


FIG 98. B^nes of the manus of Indian Elephant (Elefhas indicm\ 

form and situation it looks as if it were a dismemberment 
of the proximal part of the trapezoid. The scaphoid and 
lunar are not united, and there are five digits, of which 
the first is extremely small, and has only one minute 
nodular phalanx. In another species, H. dorsalis, the pollex 


is reduced to a short metacarpal ; the fifth digit has but 
two phalanges, and the centrale is united with the trapezoid. 
The ungual phalanges of the three middle digits are small 
and somewhat conical in form. 

In the Elephant (Fig. 98) the manus is short and broad, 
the carpal bones are massive and square, and articulate by 
very flat surfaces ; they consist of scaphoid, lunar and cunei- 
form, a pisiform and the usual four bones of the distal row ? 
all distinct, without the centrale. There are five digits, with 
short stout phalanges, the terminal ones being very small 
and of irregular form. 

All the existing true Ungulate Mammals agree in the 
complete suppression of the pollex, in the absence of 
an os centrale, and in the complete separation of the 
scaphoid and lunar. The carpus is very compact, the bones 
being generally more or less square, and articulating by flat 
surfaces with each other, and with the radius and ulna 
above. The second or distal row of the carpal bones has, 
in comparison with the Subungulata, been shifted altogether 
towards the radial side of the limb, so that the os magnum 
is brought considerably in relation with the scaphoid, and is 
entirely removed from the cuneiform as in the great majority 
of existing Mammals. The axis of the third toe passes 
therefore through the os magnum but between the lunar 
and scaphoid. The Ungulata are eminently digitigrade, the 
limb being (except in the Tylopoda] entirely supported on 
the ungual phalanges, which are large, and encased in a hoof. 

The digits are arranged according to one or the other of 
two distinct types, each characteristic of, and giving name 
to, one of the sub-orders. 

i. The Perissodactyla, or " odd-toed " Ungulates, have the 
middle or third digit the longest, and symmetrical in itself, 
the free border of the ungual phalanx being evenly rounded. 




The second and fourth toes may be subequally developed, as 
in the Rhinoceros (Fig. 100), or they may be represented 
only by mere splint like rudiments of their metacarpals, as in 
the Horse (Fig. 101). All intermediate conditions are met 
with in various extinct forms, as Palaotherium, Anchitherium, 

F,G. 99. 

FIG. loo. 

FIG. 101. 

FIG. 99. Bone* of the manus of Tapir (Tapirus indicus), {. 
FIG. loo. Bones of the nianus of Rhinoceros (Rhinoceros sHinatrrnsis), \. 
Fi<;. tot. --Bones of the manus of Horse (Equus catalius), J. Hand iv rudimentary 

and Hipparion. In the Tapir (Fig. 99) there are four 
complete toes, in consequence of the fifth being developed, 
though it scarcely reaches the ground in walking. In other 
respects the foot resembles that of the Rhinoceros, the third 
toe being longest, and symmetrical in itself and having on 


each side of it the nearly equal second and fourth. In the 
Rhinoceros there is a rudiment only of the fifth metacarpal. 

In the Horse (Fig. 101), the three bones of the first row 
of the carpus are subequal. The second row consists of a 
very broad and flat magnum (;//), supporting the great third 
metacarpal, having to its radial side the trapezoid (td), and 
to its ulnar side the unciform (?/), which are both small, and 
articulate distally with the rudimentary second and fourth 
metacarpals. The pisiform is large and prominent, flattened 
and curved ; it articulates partly to the cuneiform, and 
partly to the lower end of the radius. The single digit 
consists of a moderate-sized proximal, a very short middle, 
and a wide, semilunar, ungual phalanx. There is a pair of 
large nodular sesamoids behind the metacarpo-phalangeal 
articulation, and a single, transversely extended, " navicular " 
sesamoid behind the joint between the second and third 
phalanx. 1 

2. The Ariiodaetyla have the third and fourth digits 
almost equally developed, and their ungual phalanges 
flattened on their inner or contiguous surfaces, so that each 
is symmetrical in itself, but when the two are placed together 
they form a figure symmetrically disposed to a line drawn 
between them. Or, in other words, the axis or median 
line of the whole manus is a line drawn between the third 
and fourth digits, while in the Perissodactyles it is a line 
drawn down the centre of the third digit. 

In the S/iina, Pigs (Fig. 102), Peccaries, and Hippopotamus, 

1 With reference to the interesting cases of atavistic polydactylism, 
see Gegenbaur's " Critical Remarks on Polydactylyas Atavism," Journ. 
of Anat. and Physiol. xvi. p. 615 ; Boas, " Bemerkungen ueber die 
Folydactylie des Pferdes," Morphol. Jahrb. x. p. 182. By far the 
larger number of cases of polydactyly in the Horse as in other 
Mammals, are merely due to a teratological multiplication of digits, 
and are not reversions to an ancestral condition, as frequently supposed. 




the second and fifth toes are well developed, though always 
considerably smaller than the third and fourth, all four 
metacarpal bones are distinct, and the manus is compa- 
ratively broad. The second row of carpal bones in the 
Pig consists of a small trapezoid, a moderate-sized magnum, 


FIG. 103. 

FIG. 104. 


FIG. 102. Bones of the manus of Pig (Sus scrofa), J. 

f IG. 103. Bones of the manus of Red Deer (Cervus elaphus), 1. 

FIG. 104. Bones of the manus of Camel (Cainelus bactrianus), 

and a large unciform. In the Hippopotamus there is also a 

In the ruminating sections of the sub-order (Figs. 103 and 
104), the third and fourth metacarpals, though originally 
distinct, become more or less conjoined, generally so as to 


form what appears externally to be a single bone, though 
traces of their separate origin always remain ; the two distal 
articular surfaces are quite distinct, each supporting a digit. 
The lateral (second and fifth) metacarpus and digits are 
generally rudimentary, sometimes completely absent. Some- 
times not even the hoofs remain, as in the Giraffe, Prongbuck 
(Antilocapra), and some other Antelopes ; sometimes the 
hoofs alone, as in the Sheep, and Ox, supported, it may be, 
by irregular nodules of bone, rudiments of the ungual 
phalanges. In the Deer the three phalanges are complete, 
sometimes with the lower end of the metacarpal, tapering 
above, and not directly attached to other parts of the 
skeleton of the foot. In other species (as in Fig. 103), 
rudiments of the proximal ends only of the metacarpals are 
present. 1 

In the Tragulina these metacarpals are completely deve- 
loped, and articulate with the carpus. In Ifyomoschus, 
belonging to this section, the third and fourth metacarpals 
commonly remain distinct through life, so that the manus of 
this animal scarcely differs from that of one of the St/ina, 

The Tylopoda, or Camels, differ considerably from the 
true Ruminants in the structure of the fore-foot (see Fig. 
104). In the carpus the trapezoid and magnum are distinct, 
as in the Suina and Perissodactyla, whereas these bones are 
confluent in the Pecora and Traguiina. There are no traces 
of any metacarpals or digits, except the third and fourth. 
The metacarpals of these are very long and, as in the Fecora, 

1 The last-named condition occurs in most of the deer of the Old 
World, the former in all the American deer, with A Ices, Rangifer, 
Hydropotes, and Cafreoliis. See Sir Victor Brooke, " I'roc. Zoological 
Society, 1874," p. 36. 

Regarding the occasional indication of a trapezium in the Cen'idir, 
see Baur : " Der Carpus der Paarhufer," Morphol. Jahrb. ix. pp. 599- 

xvi.] ARTIODACTYLA. 299 

confluent throughout the greater part of their length, though 
separated for a considerable distance at the lower end. The 
distal articular surfaces, instead of being pulley-like, with 
deep ridges and grooves, are simple, rounded, and smooth. 
The proximal phalanges are expanded at their distal ends, 
and the wide and depressed middle phalanges are imbedded 
in a broad cutaneous pad, forming the sole of the foot, on 
which the animal rests in walking, instead of on the hoofs, 
as in other Ruminants. The ungual phalanges are very 
small and nodular, not flattened on their inner or opposed 
surfaces, and not completely encased in hoofs. These 
characters are better marked in the true Camels than in the 

In the animals constituting the order CETACEA, the manus 
has undergone a special modification, being converted into 
a simple, flattened, oval or falciform, usually pointed flipper 
or paddle, showing externally no signs of division into 
separate digits, nor any traces of nails or claws. The skele- 
ton, however, consists, as in other Mammals, of a carpus, 
metacarpus, and either four, or more commonly five, digits, 
the great peculiarity of which is, that the number of pha- 
langes is not limited to three, as in all other animals of 
the class, but may extend even to twelve or thirteen. 

In the Whalebone Wloales, a large portion of the 
skeleton of the hand remains permanently cartilaginous, but 
the cartilages composing the various carpal bones and pha- 
langes are separated from each other by interposed tracts 
of fibrous tissue, and sometimes by synovial cavities. Nodules 
of bone are deposited in the centre of some of these 
cartilaginous masses, and slowly reach the surface as the 
animal attains maturity : there are commonly not more 
than five such ossifications. The phalanges appear like 
cylindrical or slightly flattened bony masses, with roughly 


truncated ends, set in a continuous rod of cartilage. In 
this way a certain amount of flexibility and elasticity is 
secured in the flipper, but beyond this there is no actual 
motion between the various bones of which it is composed. 

The manus of the Right Whale (Baliena mysticetus) is 
comparatively short and very broad, having all five digits 
present, and being also extended on the ulnar side by a 
flattened cartilage projecting from the edge of the carpus, 
representing the pisiform bone. In adult specimens there 
are usually only three distinct ossifications in the carpus. 
The numbers in the digits are respectively I. i, II. 4, 
III. 5, IV. 4, and V. 3. In the Rorquals (Balanoptera) 
the first digit is absent, and the manus is of an elongated 
and narrow form. The carpus has five ossifications, and the 
number of phalanges varies somewhat in different species. 
In Megaptera, the phalanges are both numerous and long, 
producing the remarkable form of manus characteristic of 
the genus. 

In the Odontocetes, the ossification of the skeleton of 
the manus is usually more complete than in the Whalebone 
Whales, the carpal bones generally coming in close contact 
at their edges, and assuming a somewhat polygonal form. 
The phalanges are also better ossified, often having epi- 
physes at each extremity, and they are connected together 
by imperfect synovial joints. They are always very much 
flattened, and their extremities being truncated and their 
sides nearly parallel, they are either square or oblong 
in form. In size they gradually decrease to the end of 
the digit, the last often consisting of minute nodules or 
granules, so irregularly or imperfectly ossified, and so easily 
lost in cleaning, that it is in many cases impossible, when 
describing the skeleton of one of these animals, to give the 
exact number of phalanges to each digit. 

xvi.] CETACEA. 301 

The determination of the homologies of the carpal bones 
of the Cetacea with those of other Mammalia is beset with 
difficulties, and has consequently led to some differences 
of opinion among those anatomists who have attempted it. 
Moreover every species appears liable to certain individual 
variations, and sometimes the different sides of the same 
animal are not precisely alike, either in the arrangement, or 
even the number of the carpal ossifications. 

The pisiform is occasionally represented by a small ossifi- 
cation on the ulnar border of the carpus. Excluding the 
above, the carpus of the Odontocetes appears rarely to con- 
sist of more than six bones, three belonging to the proximal 
and three to the distal row. 1 

The three bones of the proximal row are constant, and 
may easily be identified as corresponding to the scaphoid, 
lunar and cuneiform of human anatomy, or the radiate, in- 
termedium, and ulnare of Gegenbaur. The middle one is 
usually the largest and most thoroughly ossified. 

The three bones of the distal row are generally represented 
by distinct ossifications (corresponding apparently with the 
trapezoid, magnum, and unciform) in the genera Hyperoodon, 
Mesoplodon, Beluga, and Monodon. 

In many cases (see Fig. 105) the bones of the distal row 
of the carpus are reduced to two, which appear to corre- 
spond best with the trapezoid and unciform, the magnum 
being either absent or amalgamated with the trapezoid. 2 

The trapezium appears never to be present as a distinct 

1 At the meeting of the British Association in 1885, Prof. Turner 
exhibited the carpus of a Hyperoodon with five distinct bones in the 
distal row. See " Journ. Anatomy and Physiology," October, 1885. 

8 For the reasons for this determination (about which, however, 
there is considerable doubt), see Flower, " On the Osteology of the 
Sperm Whale " ; Trans. Zoological Society, vol. vi. p. 360. 

3 02 



bone, although the first meta- 
carpal so often assumes the 
characters and position of a 
carpal bone, that it may easily 
be taken for it. 

The cuneiform always di- 
rectly supports the fifth meta- 
carpal, and frequently some 
part of the fourth. Moreover, 
in those species in which the 
ulnar side of the carpus is 
greatly reduced, as Globice- 
phalus, the fifth metacarpal is 
even connected with the ulna. 

In the Cachalot (Physeter} 
many of the carpal bones, in 
addition to the usual central 
nucleus, have epiphysial ossifi- 
cations developed in the peri- 
phery of the cartilage, which 
ultimately unite with the cen- 
tral piece of bone. 

All the Cetacea with teeth 
have five digits, though the 
first is usually rudimentary 
and in close contact with the 
metacarpal of the second. 
G. 105. Dorsal surface of bones of In some forms, as Fhysctcr, 

right anterior limb of Round-headed TT , , f , J, 

Dolphin (Globicephalus me/as), ^. HyperOOdOH, M Oil 0(1 OH, BelUgd, 

The shaded portions of the digits are j . jj, . 1 /n u 

cartilaginous, id trapezoid or magnum. Jnid^ J. l(ltCintSt(l, and C//YY7, the 

manus is short, broad, and 

rounded at its distal extremity ; the digits being nearly 
equally developed, spreading from each other, and without 


xvi.] CETACEA. 303 

any excessive number of phalanges. In the Grampus (Oreo) 
all the phalanges are broader than they are long. 1 In the 
round-headed Dolphins (Globicephalus, Fig. 105), on the 
other hand, the manus is extremely elongated, narrow, and 
pointed. This elongation is mainly due to the- great develop- 
ment of ihe second, and, though to a less extent, of the 
third digit ; the fourth and fifth being quite short, and 
having but few phalanges. The number of phalanges (in- 
cluding the metacarpals) in the different digits are respec- 
tively I. 4, II. 14, III. 9, IV. 3, and V. i. 

In the common Dolphin (Delphiwts) the manus has the 
same essential form, though in a less exaggerated degree, 
the numbers of the phalanges being I. 2, II. 10, III. 7, 
IV. 3, and V. i. The digits are all in close contact. 

Order SIRKNIA. Though in external form, and in being 
inclosed in an undivided integument, the terminal segment of 
the fore limb of the animals constituting this order much 
resembles that of the Cetacea, its skeleton is totally different. 

The carpus is short and broad. In the genus Afanatus 
the seven most usual bones of this region are all distinct, 
though there is no pisiform. The trapezoid is very small 
and placed almost on the dorsal surface of the trapezium. 
The cuneiform is large, and supports the greater part of the 
fifth metacarpal. In Halicore many of the bones of the 
carpus usually coalesce ; thus the first row may consist of 
two bones, a scapho-lunar and a cuneiform, and all the 
bones of the second row may unite together. 

In both genera the digits are five in number, with 
moderately elongated and flattened phalanges, which are 
never increased in number beyond the limit usual in the 

1 This genus is remarkable for the imperfect ossification of the carpal 




Among the animals constituting the order EDENTATA 
there is great diversity in the structure of the fore-foot. 
They agree, however, in wanting an os centrale, and (with 
the exception of Mam's) in .the presence of distinct scaphoid 
and lunar bones. 

In the existing Sloths the whole manus is long, very 
narrow, habitually curved, and terminating 
in two or three pointed, curved claws, in 
close apposition with each other, incap- 
able, in fact, of being divaricated, so that 
it is reduced to the condition of a hook, 
by which the animal suspends itself to 
the boughs of the trees among which it 

The carpus is small, and articulates by 
a smooth rounded surface with the lower 
end of the radius. In the Three-toed 
Sloths (genus Bradypus) it consists of dis- 
tinct scaphoid, lunar, and cuneiform bones 
in the first row, but usually of only two 
bones in the second row, the unciform, 
and a connate magnum and trapezoid, the 
trapezium being generally ankylosed to 
the rudimentary first metacarpal. 1 There 

FIG. 106. Bones of the ,, jj'T i j'i 

right manus of the is a small rounded pisiform, but no radial 

sesamoid. The first and fifth nietacarpals 
are present in a rudimentary condition 
but bear no phalanges. The three middle digits are nearly 
equally developed. The proximal phalanges are extremely 
short, and soon become ankylosed to the ends of the 
metacarpals, so that in adult animals one of the usual bones 
of the digit appears to be entirely wanting. The middle 
1 See "Journal of Anatomy and Physiology," vol. vii. p. 255. 




phalanges are long and compressed. The ungual phalanges 
are also long, much compressed, gently curved, and pointed. 
Bony laminae reflected from their bases encase and support 
the roots of the claws. 

In the Two-toed species (genus Cholaepus, Fig. 106), the 
magnum and trapezoid are distinct. The functional digits 

FIG. 107 Manusof Great Anteater 
(Myrmecophagii jutata), J. 

FIG. 108. Manus of Little Anteater 
(Cyrlatuna didactylus), X 2. 

are the second and third, and there are rudiments of the 
first and fourth metacarpals, though not of the fifth. The 
proximal phalanges (/') are extremely short, as in Bradypus, 
but do not ankylose with the metacarpals. The ungual 
phalanges are not so long as in Bradypus. 

In the Anteaters (Myrmecophaga, Fig. 107), all the usual 
carpal bones are distinct. The unciform supports the fifth 


fourth, and a considerable part of the third, metacarpals. 
The first digit is very slender, the second also slender, 
with compressed phalanges of nearly equal length. The 
third digit is immensely developed ; though its proximal 
phalanx is extremely short, its ungual phalanx is so long 
that the entire length of the digit exceeds that of the second. 
The fourth has a long and rather slender metacarpal, and 
three phalanges gradually diminishing in size, the ungual 
phalanx being very small. The fifth has the metacarpal 
nearly as long, but not so stout as the fourth, and followed 
by two small phalanges, the last rudimentary and conical, 
and bearing no nail. 

The little Tree Anteater (Cydoturus didactylus, Fig. 108), 
has a remarkably modified manus. The third digit is greatl) 
developed at the expense of all the others ; it has a stout, 
short metacarpal, and but two phalanges, of which the most 
distal is large, compressed, pointed, and much curved, 
bearing a very strong hook-like claw. The second digit 
has the usual number of phalanges and bears a claw, but 
is very much more slender than the third. The fourth is 
represented only by a styliform metacarpal, and one small 
phalanx without a claw. The first and fifth metacarpals are 
present but quite rudimentary. The pisiform is very large. 
The trapezoid, magnum and unciform often coalesce. 

In the Armadillos (Dasypodidce), the manus is stout and 
broad, with strongly developed ungual phalanges, adapted 
for digging and scratching. The fifth metacarpal articulates 
with the cuneiform as well as the unciform. There is always 
a very large palmar sesamoid. The digits are almost always 
five in number, but vary much in relative si/.e and structure. 

In the six-banded Armadillos (genus Dasypus), all the 
digits have the normal number of phalanges (see Fig. 109). 
The first digit is rather short and especially slender ; the 




second is the longest, and has all the phalanges, as well as 
the metacarpal, of nearly equal length ; the third has a long 
metacarpal, then two short broad phalanges, the first being 
especially short, and a long, curved, compressed, ungual 
phalanx. The fourth and fifth are shorter, but present 

FIG too. - Bones of the ri?ht manus of FIG. no. Bones of the manus of the 

the Hairy Armadillo (Dasyfus villa- Great Armadillo (Prioiion giganteus\ 

s \ 2 4- a an accessory carpal ossicle in front 

of the pisiform, which is not seen in 
the figure. 

the same general characters, and their metacarpals are also 
reduced in length. 

All the deviations from the normal type of manus ob- 
served in the common Armadillos, when greatly exaggerated 
produce the curiously modified condition seen in the Cabas - 
sou (Xcnurus unitindus). The first and second digits are 
still more slender and elongated, and retain the normal 

x 2 


number of phalanges ; but in the other three the metacarpal 
is short and broad, the proximal phalanx is either sup- 
pressed or incorporated with the metacarpal, as in some of 
the Sloths, the middle phalanx is very short, but the ungual 
phalanx is enormously developed, larger in the third than in 
the fourth and fifth digits. 

A still further modification of the same type is seen in 
the extraordinary mantis of the great Armadillo (Priodon 
giganteus], the largest existing member of the group (Fig. 
no). The metacarpals of the three outer toes are still 
further reduced in length, the ungual phalanx of the third 
is increased in size, while that of the fourth, and especially 
the fifth, are greatly diminished. 

In the genus Tolypentes the manus is formed on a some- 
what similar type, except that the first digit is suppressed ; 
but in the Nine banded Armadillos (genus Tatusia) it is 
altogether different, the second and third toes being subequal 
(the third the longest), with moderate, conical, and slightly 
compressed ungual phalanges : and the first and fourth 
also nearly equal and smaller, all with the normal 
number of phalanges. The fifth is absent, or (as in 
T. hybrida) represented by three very rudimentary nodular 

In the Pangolins (Manis) the scaphoid and lunar are 
united but all the other carpal bones are distinct. There 
are five digits with the complete number of phalanges, which, 
except in the pollex, are short and broad. The distal ends 
of the ungual phalanges have deep median clefts. This 
phalanx in the third digit is immensely developed, and con- 
siderably so in the fourth. The first, second, and fifth digits 
are comparatively small. 

In the Cape Anteater (Orycteropus) the pollex is entirely 
suppressed, but all the other digits are well developed, and 

xvi.] EDENTATA. 309 

terminate in subequal, compressed, ungual phalanges of 
moderate size. The second and third digits are nearly 
equal, the fourth and fifth shorter. A sesamoid bone is 
developed on the dorsal side of the metacarpal-phalangeal 

Order MARSUPIALIA. The carpus never has a distinct 
os centrale. It is commonly stated that there is a scapho- 
lunar bone ; but the lunar, though always small, is distinct 
in Didelphys, Perameles, Dasyurus, Thylacinus, Phalangista, 
and Hypsiprymnus (where it is very minute) ; and its ab- 
sence in Macropus appears to be due rather to suppression 
than to coalescence with the scaphoid. In Phascolomys a 
small lunar is present in some individuals, and not in others. 

With the exception of the genus Chceropus, all known 
Marsupials possess the normal number of digits and pha- 
langes, and the manus is short and rather broad, with 
moderately developed, compressed, curved, ungual pha- 

The little " pig- footed " insectivorous Marsupial, Chotropus 
castanotis, belonging to the family Peranulida, has a remark- 
ably modified manus (see Fig. 112) in which only two 
digits are functionally developed ; and as the metacarpals 
are very long, and closely pressed together (though not 
ankylosed), and the phalanges are short, and the nails rather 
hoof-like, the whole manus has much general resemblance 
to that of the Artiodactyle Ungulates. It presents, how- 
ever, the essential difference that the functional digits are 
the second and third of the normal series, instead of the 
third and fourth. This is proved by comparing it with 
the less modified manus of a true Perameles (Fig. in). 
The principal changes from the typical Mammalian manus 
observed in Perameles are the great reduction of the 
first and fifth digits; while the second, third, and fourth 



remain functional, with long ungual phalanges cleft me- 
sially at their extremities. The third is longer than the 
second, and this longer than the fourth. In Chceropus 
the second and third remain, and retain their relative 
size, though comparatively longer, more slender, and 
with smaller ungual phalanges. The fourth digit is 

Fit;, in. Bones of manus of Bandicoot Fro. 112. Hones of manus of Chccrof>u 
(J't-rame/cs), X i$. castanotis, X 2. 

rudimentary, but with the normal number of phalanges ; 
the first and fifth are entirely suppressed. The carpal 
bones have their normal relations, but the trapezium is 
exceedingly reduced. 

Order MONOTREMATA. In the Echidna the carpus is 
short and broad, and has a very complex articulation 
with the distal ends of the radius and ulna. The first row 

xvi.] MONOTREMATA. 311 

consists of a scapho-lunar and a cuneiform. There is no 
central. The dibtal row has the usual four bones. The 
pisiform is large, and articulates with the ulna as well as the 
cuneiform, and there is a small radial sesamoid, articulating 
with the scapho-lunar. There are also two large sesamoids, 
sometimes united, in the palmar tendons. The digits are 
five in number, all with the normal number of phalanges, 
which are short and broad, except those that bear the 
long, slightly curved, broad nails, with which the animal 
scratches and burrows in the ground. The pollex is more 
slender than the other digits ; it is of about the same length 
as the fifth, the second and fourth are nearly equal and 
longer, and the third slightly the longest. 

In the Ornithorh)nchus the manus is comparatively more 
slender and elongated ; but the number and arrangement 
ot the bones are the same as in the Echidna. 



THE posterior limb consists of a pelvic girdle and three 
segments belonging to the limb proper, viz. the thigh, the 
leg, and the foot, or pes. 

The PELVIC GIRDLE is present in some form in all Mam- 
mals, though in the Cetacea and the Sirenia it is in an 
exceedingly rudimentary condition. 

In all Mammals, except those belonging to the two 
orders just named, each lateral half of the pelvic girdle 
consists primitively, like the corresponding part of the an- 
terior limb, of a rod of cartilage crossing the long axis of 
the trunk, having an upper or dorsal, and a lower or ventral, 
end. The upper end diverges from that of the opposite 
side, but the lower end approaches, and, in most cases, 
meets it, forming a symphysis, without the intervention of 
any bone corresponding to the sternum. 

The pelvic girdle differs from the shoulder girdle in being 
articulated to the vertebral column, at a point near to, but 
not at, the upper end of the rod. 

Like the shoulder girdle, it bears on its outer side, near 
the middle, a cup shaped articular cavity (acctabulum or 
cotyloid cavity, Fig. 113, a, p. 313), into which the proximal 
extremity of the first bone of the limb proper is received. 




Like the shoulder girdle it is divided, by its mode of 
ossification, into an upper (dorsal) and a lower (ventral) 
segment, and the point of union between these is near the 
middle of the articular cavity. 

Unlike the shoulder girdle of most Mammals, the lower 
segment is always largely developed, and ossifies from two 
separate centres, which form an anterior and a posterior 

KlG. 113. External surface of right innominate bone of a young Lamb (Ovfs arits), 
J. // ilium (gluteal surface); j/ supra-iliac border ; ah acetahular border ; /^ischial 
border ; /s ischium ; s/> spine ; /;' tuberosity of ischium ; /".pubis ; s symphysis ; 
M_/ thyroid or obturator foramen : acetabulum. 

bar, in contact above and below, but leaving a space 
between them in the middle, filled only by membrane, and 
called the thyroid or obturator foramen (//</). 

The upper segment is named the ilium (//), the anterior 
bar of the lower segment the pubis (/*), and the posterior 
bar the ischium (Is). In the process of growth these three 


osseous pieces always coalesce into a single bone, called the 
os innominatum. 

This is further completed by the addition of epiphyses ; 
one for the upper extremity of the ilium (corresponding to 
the supra-scapular epiphysis of the shoulder), and others for 
the most prominent parts of the lower or free borders of the 
pubis and ischium (symphysis pubis and tuber ischii). 

In most Mammalia (perhaps with the exception of the 
Monotremata, Lemuroidea and Chiroptera) a fourth pelvic 
bone is found, situated between the ilium, pubis, and ischium. 
This, the so-called os acetabuli^- either persists as a separate 
ossification (it is especially large in Talpa, Sorex, Viverra, 
and Lepus) or it fuses with one of the three other pelvic 
bones. Its morphological meaning is as yet unknown, but 
it can scarcely be considered as an epiphysis. 

The three principal bones of the pelvis nearly always 
enter into the formation of the acetabulum, but in a few 
genera, as Lepus, \\\e pubis is excluded. 2 

There is never any secondary osseous bar in the pelvic 
girdle corresponding to the clavicle of the upper extremity. 

The ilium of Mammals is essentially an elongated, three- 
sided, or prismatic bone, though the relative size and posi- 
tion of the various surfaces and angles may differ greatly 
in different species. In the most characteristic form, one 
of the surfaces is internal, or directed towards the ver- 
tebral column, articulating by a flat irregular surface with the 
lateral " pleurapophysial " ossifications of the sacral vertebrae. 

1 See W. Krause, " Ueber den Pfannenknochen," Centralblatt f. 
d. medicinisch. \Vissenschaft. 1876, No. 46 ; W. Leche, " Zur 
Anatomic der Beckenregion bei Insectivora." K. Svenska Vetensk. ; 
Akad.'s Handlingar, xx. 1883 ; and Bronn's " Clas en und Ordnungen 
des Thierreichs, Mammalia." 

2 Gegenbaur, " Ueber den Ausschluss der Schambeines vom Hueft- 
gelenk," Morph. Jahrb. 1876. 


This may be called the sacral surface (see Figs. 114, and 
Ir 5. P- 3 X 7; and 116, ss, p. 325.) Another is directed 
mainly forwards, and may be called anterior or iliac (M), as 
it gives origin to the iliacus muscle. The third is posterior 
or gluteat (gs), as it gives origin to the gluteal muscles. 

Of the borders one is external or acetabular (ab\ as it 
ends below at the margin of the acetabulum ; another is 
antero-internal or pubic ( pb), and the third is postero-internal 
or ischial (ib) t so called because they end below by joining 
the pubis and the ischium respectively. 

The innominate bone is always placed more or less 
obliquely to the vertebral column, the upper or iliac end 
inclining forwards, and the lower or ischio-pubic end turning 
backwards, contrary to the usual direction of the scapular arch. 
In order to give still greater stability and fixity to the pelvic 
girdle, and to incorporate it more completely for mechanical 
purposes with the vertebral column, there is, in addition to 
the articulation between the ilium and true sacral vertebrae, 
a very strong double ligamentous union between the ischium 
and the side of the anterior caudal or pseudo-sacral vertebrae, 
constituting the greater and lesser sacro-sciatic ligaments, 
which are replaced in some Mammals (as most of the Eden- 
tata) by a complete bony union. 1 

The two innominate bones, together with the sacrum, 
constitute the pelvis, a complete circle of bone, or rather a 
short tube. This has two outlets : an anterior (sometimes 
called inlet or brim) bounded by the inferior surface of the 
first sacral vertebra above, by the pubic borders of the 
ilia laterally, and by the anterior borders of the converging 
pubic bones, meeting at the symphysis below ; and a 

1 Practically, though not morphologically, the pelvis is a part of the 
trunk or axial skeleton. The functions of the hind limb in propelling 
and raising the body necessitate that it should be so. 


posterior outlet, bounded by the posterior part of the 
sacrum above, by the great sacro-sciatic ligaments laterally, 
and by the converging posterior borders of the ischia below. 
In consequence of the oblique position of the innominate 
bones, the plane of the anterior outlet (in the horizontal 
position of the body) looks downwards and forwards ; that 
of the posterior outlet upwards and backwards ; but these 
two planes are not exactly parallel, the long axis of the 
cavity being usually more or less curved. 

Modifications of the Pelvic Girdle in the different Groups 
of Mammalia. 1 

Order PRIMATES. The pelvis of man is very consider- 
ably modified from the usual form met with in Mammals. 
The innominate bone (Fig. 114) is remarkably broad in 
proportion to its length. The ilium is flattened and ex- 
panded, and has a greatly extended, almost semicircular 
supra-iliac border (si). The sacral surface (ss) is small, and 
scarcely rises above the vertebral attachment. The iliac 
surface (is) is very broad and hollowed. The gluteal surface 
is likewise much expanded, and, though presenting several 
curves, is, in the main, convex. The acetabular border (a/>) 
is veiy short, and lias a strong, rounded, rough prominence 
for the attachment of the tendon of the rectus (extensor) 
muscle of the leg. The pubic border (pl>) is slightly 
marked, constituting the linea arcuata interna, or linca ilio- 
pectinca of human anatomy. The ischial border (//;) is 
short and deeply hollowed. The acetabulum (a) is large, 
circular, with very prominent borders, incomplete only for a 
small space on the infero-internal aspect. 

1 See Weyhe, " Uebersicht der Saeugethiere nnch ihren Eecken- 
formen, "Zeitschr. f. d. gesammten Natun\iss. xi. iS75. 




The pubis and ischium are short, and widely divergent 
from each other, so that the thyroid foramen (////") is elon- 
gated in the direction across the main axis of the bones. 
The symphysis (s) is rather short, and formed by the 
pubis alone. Each of the apposed surfaces of bone is 


KIG. 114. Ventral surface of right inno- 
minate bone of Man, i 

FIG. 115. Ventral surface of right inno- 
minate bone of Dog, J. 

it supra-iliac border or crest of the ilium ; s.t sacral surface ; is iliac surface ; ab 
acetabular border ; pb pubic border ; ib ischial border of ilium ; a acetabulum ; 
My thyroid or obturator foramen ; /' toberosity of ischium ; * symphysis of pubis. 

capped by a plate of nbro-cartilage ; these are held to- 
gether by strong ligamentous fibres, while between them 
there is usually a more or less perfect synovial cavity. 
Ankylosis at this spot, so common in the lower Mammalia, 
very rarely takes place in Man. 


The posterior and inferior border of the ischium is thick- 
ened and rounded, and distinguished as the tuber ischii (//). 
Above this, on the hinder border of the same bone, is a 
smooth, hollowed surface, called the lesser sciatic notch, 
surmounted by an angular prominence called the spine ; 
above the spine the edge of the ischium passes into the 
great concavity of the posterior or ischial border of the 
ilium, and which is called the great sciatic, or, more pro- 
perly, ilio-ischiatic notch. The strong ligaments (sacro- 
sciatic) which pass from the side of the pseudo-sacral and 
caudal vertebrae, the one to the tuber and the other to the 
spine of the ischium, convert these notches in the living 
body into foramina. 

The anterior or superior (in the vertical position) outlet of 
the pelvis is subcircular, usually rather broader from side to 
side than from the vertebral to the pubic border. Its plane 
is not far from a right angle with the axis of the vertebral 
column. The posterior outlet is also very wide. In con- 
sequence of the great curve of the sacrum, and the short- 
ness of the symphysis, the axis of the whole pelvis is 
strongly curved. 

In all the Simiina the innominate bone, especially the 
iliac portion, is more elongated than in Man ; the anterior 
outlet of the pelvis is longer from above downwards, nar- 
rower, and more oblique; the tuberosities of the schia 
are more everted, and the spine and sciatic notches less 

In the highest forms, such as the Gorilla and Chimpanzee, 
the upper part of the ilium is expanded, flattened, and 
everted, the iliac surface being even wider than in Man, 
though much flatter ; but in the Baboons and Monkeys, 
the whole ilium is long and narrow, the sacral surface rises 
considerably above the sacral articulation, the iliac surface 

xvn.J CARNIVORA. 319 

is narrow, the gluteal surface is very hollow, and the borders 
all approximate to straight and {arallel lines. In the Old- 
World monkeys the tuberosities of the ischia are greatly 
everted, and terminate in broad, triangular, flattened, rough 
surfaces, to which the ischial cutaneous callosities are 

In the true Lemurs the pelvis is very wide ; the ilia are 
long, narrow, and have a sigmoid curve, while the pubes 
approach each other at the symphysis at a very open angle, 
giving an elegant lyre shape to the anterior outline of the 
pelvis. On the other hand, in the genus Loris of the same 
group (and to a less extent in Tarsius, Perodicticus, and 
others) the cavity of the pelvis is remarkably narrow from 
side to side ; the ilia are straight slender rods, from the 
lower end of which the large, flattened, and compressed 
pubes project forward at a right angle, forming a prominent 
keel at the symphysis. 

In the CARNIVORA the pelvis is generally elongated and 
narrow, the ilium and ischium being in a straight line, and 
of nearly equal length. In most species the ilia are straight, 
flattened, and not everted above (see Fig. 115, p. 317), the 
iliac surface (is) is very narrow, and confined to the lower 
part of the bone, as the acetabular and pubic borders meet 
in front above ; the gluteal surface looks directly outwards 
and is concave ; the sacral surface (ss) forms a broad flat 
plane above the attachment to the sacrum, the crest being 
formed by the united edges of the sacral and gluteal sur- 
faces, instead of the iliac and gluteal surfaces, as in Man. 
The symphysis is long ; it includes part of both pubis and 
ischium, and commonly becomes completely osseous in 
adult animals. The thyroid foramen (//{/) is oval, with its 
long axis parallel to that of the whole bone. The ischia are 
wide and divergent posteriorly. 


In the Hyasna the pelvis is shorter and wider than in most 
other Carnivora, both the upper ends of the ilia and lower 
ends of the ischia being considerably everted. 

In the Bears the ilia are short and everted above. 

In the Seals the pelvis is small, and of a different form 
from that of the terrestrial Carnivora. The ilia are exceed- 
ingly short, and with much everted upper borders ; the 
pubes and ischia are very long and slender, inclosing a long 
and narrow obturator .foramen, and meeting at a symphysis 
of very small extent, in which the bones of the two sides 
are very slightly connected, and capable of being widely 
separated during parturition. 

The pelvis of the INSECTIVORA varies considerably in 
form. In Galeopithecus, Rhynchocyon, Macroscelides and 
Tupaia, the symphysis is long, as in the Carnivora, and 
becomes ankylosed ; in Erinaceus, Centetes, Potamogale, and 
Myogale, it is short, though the bones of the two sides are 
in contact, but in many other genera, e.g. in Talpa, Chry- 
sochloris and Sorex, the pubic bones are widely separated in 
the middle line below. 

The Mole has an exceedingly long, narrow, and straight 
pelvis, the innominate bones lying almost parallel with the 
vertebral column. Both ilium and upper end of the 
ischium are firmly ankylosed with the sacrum, leaving a 
small sacro-sciatic foramen between them, but the pubic 
bones do not quite meet in the middle line below. The 
pubes and ischia are very long and straight, and inclose a 
large, but narrow, oval thyroid foramen. 

In the CHIROPTERA, the pelvis is small and narrow ; the 
ilia are rod-like, the pubes and ischia are not in a line with 
them, but project forwards. In most species the pubic 
bones of the opposite sides are very loosely united in front 
in males, and in females they are widely separated ; in the 


Rhinohptuda alone do these bones form a symphysis 
(Dobson). There is usually a strongly developed " pectineal " 
process, near the acetabular end of the anterior border 
of the pubis, and which in some genera (e.g. Phyllorhina, 
triienops), is prolonged so far as to unite with a process from 
the superior extremity of the ilium, forming a pre-acetabular 
foramen, as large as, or larger than, the thyroid foramen, an 
arrangement unique among Mammals. 

In many of the RODENTIA, as the Beaver, the ilia are 
markedly trihedral, with sides of nearly equal extent; but 
in the Hares the outer (acetabular) border is almost obso- 
lete, the gluteal and iliac surfaces are confluent, and both 
face outwards, and the internal surface is largely deve- 
loped above the sacral attachment. 

The pubes and ischia are always largely developed, flat, 
and diverging posteriorly, the obturator foramen is of con- 
siderable si/e, and the symphysis is long, and usually 
becomcs osseous ; in the Guinea Pig (Cavia), however, it 
remains ligamentous, and the bones are widely separated 
during parturition. 

Order UNGUI.ATA. In the Pecora the pelvis generally is 
elongated. The ilium is expanded and everted at the 
upper extremity ; but between the sacral attachment and the 
acctabulum it is much contracted, and its borders rounded, 
so that the divisions of its surfaces are no longer distinct. 
There is usually a deep oval depression above the ace- 
tabulum, just within the attachment of the rectus femoris 
muscle. The anterior outlet forms a regular oval with the 
long diameter between the sacrum and symphysis. The 
latter is very long, including a considerable portion of the 
ischia. The margins of the bones are completed by large 
epiphyses in this region, but ultimately coalesce across 
the middle line. The ischia are much developed ; the 


tuberosities are large, and have on the middle of their outer 
side a well-marked conical process, directed outwards, and 
very characteristic of this group of animals. 

In the Giraffe the pelvis is shorter than in most of the 
other Pecora ; the upper ends of the ilia are more ex- 
panded, the thyroid foramen is nearly circular, and the 
supra-acetabular fossa is almost obsolete. 

These characters are still more strongly marked in the 
Camels ; while, on the other hand, in the Pigs the pelvis is 
elongated, and much resembles that of the Pecora, but the 
supra-acetabular fossa is wanting. 

In the Perissodactyla, the ilia are widely expanded above, 
but much contracted on approaching the acetabulum. The 
ischia are less elongated than in the Pecora, and the thy- 
roid foramen is more circular. 

The Elephant has a very peculiar pelvis, the form of the 
ilium and the arrangement of its surfaces somewhat recalling 
those of the human pelvis. The supra-iliac border or crest 
is greatly extended and curves outwards and downwards. 
The sacral surface of the ilium is narrow, and scarcely rises 
above the attachment to the sacrum. The iliac and gluteal 
surfaces are widely expanded, especially at the upper part, 
and, the pelvis being set nearly vertically to the vertebral 
column, they face almost directly forwards and backwards. 
The outer or acetabular border is short and deeply hollowed. 
The pubic and ischial portions are comparatively small, the 
latter being very little produced backwards beyond the 

In the SIRENIA, the pelvis is extremely rudimentary, being 
composed, in the Dugong, of two slender, elongated bones 
on each side, placed end to end, and commonly ankylosing 
together. The upper one, which represents the ilium, is 
connected by a ligament with the end of the transverse 

xvn.] SfKENIA. 323 

process of the sacral vertebra, usually the fourth behind the 
last that bears a free rib ; the lower one is the ischium, or 
ischium and pubis combined, and approaches, though it 
does not meet, its fellow of the opposite side. 

In the adult Manatee, the innominate bone is represented 
by a single irregular triangular bone, connected by rather 
long ligaments with the vertebral column above, and with 
the opposite bone across the middle line. 

There is no trace of an acetabulum, or of any portion of 
the limb proper in any of the existing Sirenia ; but in the 
extinct Halitherium an acetabular depression and rudimentary 
styliform femur have been discovered. 

In the CETACEA the pelvis is reduced to a pair of elon- 
gated slender bones (each of which ossifies from a single 
centre), placed on each side of, and rather below, the ver- 
tebral column, lying nearly parallel to its long axis (though 
they converge somewhat anteriorly), and opposite the spot 
where the chevron bones commence to be developed 
beneath the bodies of the vertebrae. These bones probably 
represent the ischia, and their principal function is to 
give attachment to the crura of the penis or clitoris, as 
the case may be. Hence they are usually more largely 
developed in the male than in the female. 

In the Whalebone Whales they usually have a project- 
ing angle placed about the middle, near which, in some 
species, a second small bone, which probably represents 
the femur, is attached by ligament (see Fig. 120, p. 335). 
In a full-grown male Rorqual {Balanoptera musculus), 
sixty-seven feet in length, each pelvic bone was sixteen 
inches long. In the Greenland Whale (Balczna mysticetus,} 
they are rather shorter and stouter. As might be expected, 
from the rudimentary character of these bones, they vary 
considerably in size and form in different individuals of 

Y 2 


the same species, and often on the two sides of the same 

In the Dolphins, they are generally smaller, and more 
simple than in the Whalebone Whales, and usually quite 
straight, though sometimes arched, or presenting a sigmoid 
curve. In Platanista alone no traces of pelvic bones have 
yet been found. 

Order EDENTATA. In the Sloths, the pelvis is very short 
and wide, with tolerably broad flattened ilia, and slender 
pubes and ischia, inclosing a large oval thyroid foramen, 
the inferior boundary of which, and the extremely narrow 
ossified symphysis, are formed by the pubis alone. The 
spine of the ischium is produced backwards to unite with 
the transverse processes of some of the pseudo-sacral ver- 
tebras, inclosing a sacro-sciatic foramen. The sacro-iliac 
articulation is commonly ankylosed. 

In all the other Edentates the pelvis is more or less elon- 
gated, the ilia trihedral, the ischia largely developed, (he 
pubes slender, the symphysis exceedingly short, but usually 
ossified, and the thyroid foramen large. In all, except Oryc 
teropus, the ischii unite with the vertebral column. This 
union is carried to its greatest extent in the Armadillos, in 
which animals the broad transverse processes of as many as 
five pseudo-sacral vertebrae may coalesce with each other and 
with the side of the ischium, converting the pelvis into a long 
bony tube, the more so as the ilia are also firmly and exten- 
sively united with the true sacrum. There is usually, especially 
in Orycteropus, a strongly developed " pectineal " tubercle. 

Order MARSUPIALIA. In the American Opossums 
(Didelphys), the ilium is a very simple, straight, three-sided 
rod, thicker at its upper end than at its acetabular end, 
each side being nearly equal in extent, hollowed, and sharply 
defined by prominent straight borders. 




In the Kangaroo (Macropus, Fig. 116), the three surfaces 
of the ilium are also well marked and nearly equal : but the 
whole bone is curved outwards at the upper end. 

In the Thylacine and Dasyures the ilia are compressed 
laterally, the acetabular and pubic 
borders meeting above in front, so that 
the iliac surface is (as in the Carnivora) 
very narrow, and disappears in the 
upper half of the bone, the "crest" 
being formed by the united edges of 
the sacral and gluteal surfaces ; where- 
as in the wide, depressed pelvis of the 
Wombat (Phascolomys\ the flattening 
has taken place in the contrary direc- 
tion, and the iliac surface spreads out 

to form with the gluteal surface behind 

a wide, arching, supra-iliac border and 


The ischia and pubes are always 

largely developed, and the symphysis 

is long and generally ossified. In the 

Kangaroos the pectineal tubercle (//) 

of the pubis is strongly developed. 
Nearly all Marsupials have a pair of 

elongated, flattened, slightly curved 

bones (Fig. 116, m\ movably articu- 
lated by one extremity to the anterior 

edge of the pubis, near the symphysis, 

and, passing forwards, diverging from * symphysis: m -mar- 

supial bone. 

each other, within the layers of the 
abdominal parietes. They are, in fact, ossifications in, or 
intimately connected with, the inner tendon or " pillar " of 
the external oblique muscle, and therefore come under the 


FIG. 116 Ventral surface 
of innominate bone of 
Kangaroo (Mncropus 
major), J. si supra- 
iliac border ; ss sacral 
surface ; is iliac sur- 
face ; al> acetabular bor- 
der ; pb pubic border of 
ilium ; // pectineal tu- 
bercle ; a acetabulum ; 
////"thyroid foramen ; //' 
tuberosity of ischium ; 


category of sesamoid bones. They vary in size and shape 
in different species. In Didelphys they are nearly as long 
as the ilia, while in the Kangaroo they are scarcely half the 
length of that bone. Though largely developed in the 
Dasyures, in the allied genus Thylarinus they are repre- 
sented only by small, unossified fibro-cartilages. 

These bones are commonly called " marsupial bones," 
though they have no special function relating to the ventral 
pouch of the female, being nearly equally developed in both 
sexes, and also in those species in which the marsupiurn is 
not present. 1 

In the MONOTREMATA the pelvis is short and broad. 
The ilia are short, distinctly trihedral and everted above. 
The ischia are large, and prolonged into a considerable 
backward-directed tuberosity. The symphysis is long, and 
formed about equally by pubes and ischium. The thyroid 
foramen is round. The acetabulum is perforated in Echidna 
as in Birds, but not in Ornithorhynchus. The pectineal 
tubercle is greatly developed. There are large " marsupial " 
bones in both genera. 

1 See O. Katz, " Zur Kenntniss der Bauchdecke uncl der mit ibr 
verkniipften Organe bei den Beutclthieren. Zeitschr. f. wiss Zoolog." 
36, 1882, p. 6ll. 



THE skeleton of the first segment of the limb proper 
consists of an elongated, more or less cylindrical bone, 
the femur^ which is described as having a shaft and two 

The dorsal, or (in the ordinary position of the limb) 
anterior surface of the shaft, is smooth and rounded from 
side to side, and generally arched somewhat forwards from 
above downwards ; the ventral or posterior surface is more 
or less compressed, and has a rough longitudinal ridge, the 
linta aspcra. 

At the proximal extremity is a hemispherical, smooth, 
articular " head " (Fig. 117, //, p. 328) which fits into the 
aretabulum of the innominate bone, and is generally more 
round and more distinctly separated from the rest of the 
hone by a constriction or "neck" (//) than is the corre- 
sponding part of the humerus. The axis of the head does 
not coincide with that of the shaft of the bone, but crosses 
it at an angle varying in different animals, being directed 
towards the preaxial 1 or (in the ordinnry position of the 
limb) internal side, and slightly also towards the anterior 

1 See note to p. 269. 



aspect. In nearly all Mammals there is a rounded depres- 
sion near the middle of the surface of the head into which 
the Hgamentum feres of the hip-joint is inserted. Both 
ligament and depression are, however, wanting in the (Drang 
Utan, Seals, the Sea-Otter, Elephant, 
Sloths, Wombat, and the Monotremata. 
Immediately below the neck of the 
femur are two tuberosities, called tro- 
chanters. One (//) is a comparatively 
small, conical eminence, situated rather 
to the pre-axial side, and called the 
lesser or inner trochanter. The other 
(gf) is generally very prominent, pro- 
jecting upwards, as high or higher than, 
the top of the head, situated mainly 
on the post-axial border of the bone, 
but curving inwards and backwards at 
its extremity; this is called the great 
or outer trochanter. To the posterior 
side of its base there is usually a deep 
depression, the digital fossa. 

In some Mammals, as the Perisso- 
dactyle Ungulates, some Rodents and 
Edentates, there is a compressed ridge 
for attachment of the great gluteal 
muscle on the post axial side of the 
shaft, a short distance below the great 
trochanter, distinguished as the third trochanter. (See Fig. 
118, /', p. 333.) 

The distal extremity of the femur is thickened, and has a 
large trochlear articular surface for the bones of the leg. 
This surface is narrow in front, and bounded by more or less 
prominent ridges : posteriorly it is divided by a deep median 

FIG. 117. Right human 
femur, d >rsal or ante- 
rior aspect, \. The 
boundary lines of the 
various epiphyses are 
shown. A head ; neck ; 
ft greater trochanter ; 
ill lesser trochanter ; ec 
external condyle ; ic in- 
ternal condyle. 


notch (intercondylar) into two prominent rounded eminences, 
called condyles (Fig. 117, cc and if). The slightly elevated 
roughed portions of bone above the articular condyles are 
termed the tuberosities. 

The femur has a main centre of ossification for the shaft, 
and epiphysial centres for the head, for each trochanter, and 
for the lower extremity. (See Fig. 117.) In most Mammals 
the great trochanter and head coalesce together before they 
join the shaft. The lower epiphysis is the last to become 

The skeleton of the second segment of the limb consists 
of two bones, the tibia and fibula? of which the former is 
the larger in all Mammals. These bones always lie in their 
primitive, unmodified position, parallel to each other, the 
tibia on the pre-axial, and the fibula on the post-axial side 
and are never either permanently crossed or capable of any 
ronsiderable amount of rotation, as in the corresponding 
bones of the fore-limb. In the ordinary walking position the 
tibia is internal, and the fibula external. 

The tibia has an expanded proximal end, with a flattened 
articular surface, divided into two slightly concave facets, by 
a rough median eminence, to which the intra-articular or 
crucial ligaments of the knee-joint are attached. The shaft 
is usually more or less trihedral, with one flat surface directed 
backwards, and one border forwards. The upper end of 
this border is thickened into a rough tuberosity, into which 
the tendon of the great extensor muscles of the leg are 
inserted. The lower end is slightly expanded, and has a 
somewhat square articular surface to receive the proximal 
bone of the tarsus, or astragalus. The inner (pre-axial) side 
of the bone is prolonged beyond this surface, forming a 

1 Also occasionally called perone, whence " peroneal," applied to 
structures in relation with it. 


process called internal malledus, which is applied to the 
side of the astragalus, giving additional strength to the 
articulation called "ankle-joint." 

The fibula has a slender and generally compressed shaft, 
and is somewhat expanded at each extremity. Its upper 
end usually takes no part in the knee-joint, being con- 
nected, by a separate synovial joint, with the tibia just 
below that articulation. The lower end, however, forms the 
outer side of the ankle-joint, under the name of external 

In many Mammals the fibula is in a more or less rudi- 
mentary condition, and it often ankyloses with the tibia at 
one or both ends. 

As a general rule each of these bones has a principal 
centre of ossification for the shaft, and an epiphysis at 
either end. 

In the neighbourhood of the knee-joint, certain sesamoid 
bones are often found in connection with the tendons which 
pass over the various bony prominences. 

The largest and most constant is the patella, placed on 
the anterior surface of the joint, in the conjoined tendon 
of the four great extensor muscles of the leg, and having 
a smooth articular facet, which plays upon the narrow 
anterior part of the inferior articular surface of the femur, 
and forms part of the wall of the cavity of the knee joint. 
This bone varies considerably in form, being in some cases 
broad, flattened, or lozenge-shaped, and in others, laterally 
compressed or oval. It is found in an ossified con- 
dition in all Mammals, with the exception of a few of the 

There are also very frequently smaller ossicles, one 
or two in number, situated behind the femoral condyles, 
called fabclltc ; and occasionally there is a wedge-shaped 

x v 1 1 1 ] rRIMA TES. 33 j 

bone within the joint, lying on the articular surface of the 
tibia, an ossification of the internal inter-articular semilunar 

Special Characters of tht Bones of the Thigh and Leg in the 
various Groups. 

In Man, the femur (see Fig. 117, p. 328) is long and 
rather slender, the shaft is curved forwards, the head is large 
and globular, the neck elongated and narrow. 

In the Gorilla, the femur is much shorter and broader ; 
the head is smaller and less globular, the neck is shorter and 
set on the shaft more at a right angle. In the Chimpanzee 
the femur more resembles that of Man. In the Lemurs it 
is very slender and straight, the head is globular, a.nd the 
neck very short. 

The tibia and fibula are distinct, and well developed in 
all the Primates, and are united with each other only at 
their extremities. Fabellae are wanting in the highest forms, 
but generally present in the others. The patella is usually 
broad and flat, and more or less lozenge-shaped. 

In the terrestrial CARXIVORA, the femur is straight, mode- 
rately slender, and with rather a small head. The fibula is 
slender, and in the Dogs curved towards the tibia, the lower 
half being closely applied to that bone ; but in the Bears, 
and many others, there is a considerable interval between 
the bones throughout, except at their articular extremities. 
Fabellaj are generally present. 

In the Seals, the femur is exceedingly short, broad, and 
flattened, with a globular head and an extremely short neck- 
The fibula is almost as large as the tibia, especially at the 
distal end. These bones are commonly ankylosed together 
at their proximal extremities. 


Among the INSECTIVORA, the Hedgehog has a. strong 
ridge distally from the great trochanter of the femur, and 
several other forms have a similar rudiment of a third 
trochanter. As a general rule the fibula is slender, and in 
its distal half ankylosed with the tibia, but it is complete 
and distinct in the genera Galeopithecus, Tupaia, Centetes, 
Ericulus and Solcnodon. 

In the CHIROPTEKA, the femur is slender and straight, 
with trochanters of nearly equal size, and with a small 
globular head, set on a very short neck, with its axis 
pointing almost directly to the anterior or dorsal surfaces of 
the bone. The fibula varies in condition ; in the Molossiiitf 
alone it is well developed. In all other species it is either 
very slender or ligamentous in its proximal third (Pteropns)^ 
reduced to a small bony process above the heel, as in 
Megaderma, or altogether absent, as in AJycteris (Dobson). 

In the RODENTIA the femur varies much. In the Hares 
and Squirrels it is long and slender, with a third trochanter 
immediately below the great trochanter. In the Beaver it is 
broad and flat, and has a strong ridge about the middle of 
the outer side of the shaft. In many other forms (e.g. 
Hystrix) neither of these accessory prominences exist, but 
the great trochanter is usually much developed. 

In Myopotamus and still more in Capromys there is a 
strong ridge on the inner side of the middle of the shaft, 
indicating the point of insertion of the adductor muscles. 

In some forms, as the Beaver, the fibula is distinct, 
strongly developed, and separated from the tibia, except at 
the extremities, by a wide inter-osseous space. In others, as 
the Hares, it is slender, and in its distal half united with the 
tibia. The patella is generally elongated, fabellie are usually 
developed, and there are often wedge-shaped ossifications in 
the semilunar cartilages of the knee-joint. 

xvin.] UNCULATA. 333 

In the UNC.ULATA, the femur is rather compressed, espe- 
cially at the lower end. There is no distinct constriction 
of the neck, separating the head from the rest of the bone. 
The great trochanter is very large, and usually rises above 
the level of the head. The small trochanter is not very 
salient, and sometimes, as in the Rhinoceros (Fig. 118), is a 

Kir. i:2 - Antcri. r n<rc of K ht '"" ur of Rhinoceros (Rkinoeeroi unicorttfs), J. 
h head ; / great irochanter ; f third trochanler. 

mere rough ridge. The inner edge of the anterior part of 
the inferior articular surface is very prominent. In all the 
Perissodactyles there is a strongly marked third trochanter 
(0, in the form of a compressed ridge curving forwards. 
This is entirely absent in all the known Artiodactyles. 

The fibula is subject to great variations among the different 
members of the order. In the Rhinoceros, Tapir, Pig, and 


Hippopotamus it is complete and distinct, though slender 
in proportion to the tibia. In the horse a mere styliform 
rudiment of the proximal end is present. 

In all Pecora and Tylopoda, a small distinct bone, having 
a very definite form, articulating with the lower end of the 
tibia, and forming the external malleolus, appears to 
represent the distal extremity of the fibula (see Fig. 119). 
There is occasionally in addition a slender styliform rudi- 
ment of the proximal extremity, but the two are never united 
together by bone. 

FIG. 119. Anterior aspect of lower end of the right tibia and fibula of Red Deer 
(Corns claphus), \. t tibia ; f fibula. 

In the Tragnlina the fibula is long and slender, and com- 
plete, but its lower end is indistinguishably blended with 
the tibia. 

The patella is well ossified, and usually somewhat trian- 
gular, with the broad end upwards ; but fabellrc are not 
commonly developed in the Ungulata. 

In the Hyrax there is a slight ridge on the femur in the 
place of a third trochanter. The fibula is complete, and 
thickest at its proximal end, where it generally ankyloses 
with the tibia. The styloid process of the tibia articulates by 
two facets with two corresponding surfaces of the astragalus. 

xvin.] CETACEA. 335 

The femur of the Elephant is long and very straight ; the 
axis of the head is more in a line with that of the shaft than 
usual. The great trochanter is not much developed, and 
the small trochanter is nearly obsolete. The fibula is com- 
plete, distinct, and slender, though considerably enlarged at 
the lower end. 

In the CETACEA, certain small nodular bones or cartilages 
attached by fibrous tissue to the outer side of the pelvic 
bone in some of the Whalebone Whales, are commonly 

Fir;. 120 Side view of bones of posterior extremity of Greenland Fight Whale 
(Raltrna tnyititctus), J, from Eschricht and Keinhardt. / ischium ; f femur ; 
/ accessory ossicle representing the tibia. 

regarded as rudimentary and functionless representatives 
of the skeleton of the hind limb. In the Greenland Whale 
(Fig. 120), there is a proximal, somewhat pear-shaped bone 
(/), about eight inches in length, and a smaller conical distal 
bone (/), which may represent the femur and tibia 
respectively, as suggested by their discoverer, Professor 
Reinhardt. 1 In Megaptera longimana there is but one such 

1 See " Recent Memoirs on the Cetacea," Ray Society, 1866, p. 134. 
The bones, articulations, and muscles of the rudimentary hind limb of 
the Greenland Whale have been fully described by Professor Struthers, 
" Journ. of Anat. and Physiology," vol. xv., i8ii. 


bone, and in an adult Fin Whale (Bal&noptera musculus], 
sixty-seven feet long, this was found to be only repre- 
sented by an oval nodule of cartilage about the size of a 
walnut. 1 Even this is wanting in some species of the group, 
as B. rostra ta and B. boreal is. 

No trace of any structure representing the skeleton of 
the hind limb, beyond the pelvis, has yet been detected in 
any of the Odontocetes. 

In none of the existing SIREXIA are there any rudiments 
of the hind limb proper, but the extinct Ifalit/uniim had an 
ossified femur, articulated to a well-defined acetabulum in 
the pelvis. 

In the terrestrial and fossorial EDENTATA the femur is 
generally short and broad. There is a third trodianter in 
the Armadillos and Orycteropus, and a sharp ridge along 
the whole external border in Myrmecophaga. The fibula is 
as long as the tibia. In the Armadillos these bones are 
commonly ankylosed together at each extremity, but curve 
away from each other at the middle, leaving a wide inter- 
osseous space. In the Anteaters they are both nearly 
straight and parallel. 

In the Sloths the femur is long, slender, ard flattened 
from before backwards. There is no third trochanter ; the 
head is large and globular, and placed near the middle of 
the proximal end of the shaft, with the axis of which it 
more nearly coincides than in most Mammals. The tibia 
and fibula are complete, and more nearly equal in si/.e than 
in most Mammals. They are both curved, so as to be 
separated considerably in the middle part of the leg. The 
lower end of the fibula has a conical prominence which 
turns inwards, and fits into a depression on the outer side 

1 " Proceedings of the Zoological Society, 1865," p. 704. 




of the articular surface of the astragalus, as a pivot into 
a socket. 

In none of the MARSUPIALIA is a third trochanter pre- 
sent on the femur. The fibula is always well developed, 
and its proximal extremity is often produced into a well- 
marked process, to the top of which a sesamoid bone is not 

Fin. ui. Anterior aspec* of loncs of right leg of Ornithorh^nckut nnatinut, * . 
/femur ; / tibia ; /' fibula ; / patella. 

unfrcquently attached ; but, on the other hand, the patella, 
except in the Pframelidie, is unossified or quite rudimentary. 
In Chatrofus and Ifyf>sif>rymnus the distal end of the fibula 
is united with the tibia. In the climbing Australian 
Phalangers and Koalas, which have broad hind feet, with an 
opposable hallux, there is a greater freedom of movement 
between the fibula and tibia than in other Mammals, 


approaching in some degree to the rotation often permitted 
between the radius and ulna. 

In the MONOTREMATA the femur (Fig. 1 2 1,/) is of very 
remarkable form, being short, flattened from before back- 
wards, narrow in the middle of the shaft, and very broad 
at each end ; the trochanters are both well developed, and 
the head placed between them on a very short neck, and 
with its axis directed quite towards the anterior or dorsal 
aspect of the bone. The fibula (_/') is large and straight : 
it has a broad flattened process, completed by an epiphysis, 
projecting from the upper extremity above the point of articu- 
lation with the tibia, much resembling the olecranon of the 
fore limb. The tibia (/) is strongly curved in Ornithorhynchus. 
but straight in Echidna . In both genera the patella (/) is 
well developed. 



THE terminal segment of the hind limb is the foot or pes. 
Its skeleton presents in many particulars a close re- 
semblance to that of the manus, being divisible into three 
parts : (i) a gro'ip of short, more or less rounded or square- 
shaped bones, constituting the tarsus ; (2) a series of 
long bones placed side by side, forming the metatarsus ; 
and (3) the phalanges of the digits or toes (see Fig. 
122, p. 341). 

The metatarsal bones never exceed five in number, and 
the phalanges follow the same numerical rule as in the 
manus, never exceeding three in each digit. Moreover, the 
first digit (counting from the tibial side), or hallitx, resem- 
bles the pullex of the hand in always having one segment 
less than the other digits. If one toe only is absent, it is 
the first or hallux. Unlike those of the manus, any of the 
five toes of the foot may be absent or at least rudimentary ; 
the fourth and fifth are best developed in Macropus, the 
third and fourth in J'fcora, the fourth alone in Choeropus, and 
the third alone in Eqniis. 

The bones of the tarsus in many of the lower Vertebrata 
closely resemble both in number and arrangement those of 

z 2 




the carpus, as shown in Fig. 90, p. 281. They have been 
described in their most generalised condition by Gegenbaur 
under the names expressed in the first column of the 
following table. 1 The names in the second column are 
those by which they are most generally known in this 
country, and which will be used in the present work, while 
in the third column some synonyms, occasionally employed, 
are added. 2 

Tibiale \ 
Intermedium i 





- Os cahis. 

Tar* ale I 
Tar sal,' 2 

Internal Cuneiform 
Middle Cuneiform 

- Entocuneiforme. 
- Mi'socunei forme. 

Tar sale 3 
TarsaL' 4 ) 
Tar sal,' 5 i 

External Cuneiform 

- Ectocumi/ormc. 

The bones of the tarsus of Mammals present fewer diver- 
sities of number and arrangement than those of the carpus. 

1 " Untersuchungen zur vergleichenden Anatomic ; Carpus und 
Tarsus/' 1864. 

- Recent researches of Baur, "On the Morphology of the Tarsus in 
the Mammals/' American Naturalist, January, 1885, make it probable 
that a certain bone on the tibial side of the tarsus of Ilyrax, many 
Edentates, Ornithorhynchus and Rodents (cf. p. 349) and hitherto 
looked upon as a sesamoid bone is the rudimentary tarsale tibiale, 
whilst the astragalus is the intermedium, representing the lunare of the 
hand. This tibiale is frequently fused with the centrale, the navicular 
bone in such cases containing the elements of the centrale and tibiale. 

The homologies of the proximal row of the Mammalian tarsus would 
therefore, according to Baur, be the following- - 

Fibulare Calcaneum. 
Centrale Navicular. 

JWitilt Sesamoid. 

Intermedium Astragalus. 

See also H.irdelebcn, " Zur Morphogie des Hand- und Fuss-skelets. " 
Sitzungsbcrichte lena. Gesellsch. Med. Nat. 1885. 



The proximal row (see Fig. 122) always consists of two 
bones, the astragalus (a), which according to Gegenbaur's 
view represents the coalesced scaphoid and lunar of the 
hand, and the calcaneum (r). The former is placed more 
to the dorsal side of the foot than the latter, and almost 
exclusively furnishes the tarsal part of the tibio-tarsal or 

Kic ill. Bones of a right human foot, the epiphyses. i- T tarsus; M 
metatarsus ; /'A phalanges : c calcaneum ; a astralagus ; c b cuboid ; naytcular ; 
t ' internal cuneiform : f a middle cuneiform ; c* external cuneiform ; the digits are 
indicated by Roman numerals, counting from the tibial to the fibular side. 

ankle-joint. It has a rounded anterior or distal projection 
called the " head." The calcaneum, placed more to the 
ventral or " plantar " side of the foot, is elongated back- 
wards to form a more or less prominent tuberosity, the tuber 
is, to which the tendon of the great extensor muscles of 

342 THE HIND FOOT OR PES. [ciiAr. 

the foot is attached. The naricular bone (//) is interposed 
between the proximal and distal row on the inner, or tibial, 
side of the foot, but on the outer side the bones of the two 
rows come into contact. The distal row, when complete, 
consists of four bones, which, beginning on the inner side, 
are the three cuneiform bones, internal (^ 1 ), middle (r 2 ), and 
external (^), articulated to the distal surface of the navicular, 
and the cuboid (cl>] articulated with the calcaneum. Of these 
the middle cuneiform is usually the smallest in animals in 
which all five digits are developed; but when the hallux is 
wanting, the internal cuneiform may be rudimentary or 
altogether absent. 

The three cuneiform bones support the first, second, and 
third metatarsals respectively, the cuboid supports the fourth 
and fifth ; they thus exactly correspond with the four bones 
of the distal row of the carpus. 

In addition to these constant tarsal bones, there may 
be supplemental or sesamoid bones; one situated near the 
middle of the tibial side of the tarsus, largely developed in 
many Carnivora and Rodents; 1 another, less frequent, on the 
iibular side ; and a third often developed in the tendons 
of the plantar surface of the tarsus, especially large in the 
Armadillos. There is also usually a pair of sesamoid 
bones opposite each metatarso-phalangeal articulation, on 
its plantar aspect. 

The development of the bones of the foot corresponds in 
the main with that of the bones of the manus. Each tarsal 
bone is ossified from a single centre, but the calcaneum has 
in addition an epiphysis for the most projecting part or 
tuberosity. The four outer metatarsals have each one 
centre from which the shaft and proximal end are ossified, 
and a large epiphysis at the distal end ; the first meta- 
1 See footnote on p. 340. 


tarsal (if it should be so called) and all the phalanges have 
an epiphysis only at the proximal extremity. This rule is 
almost universal, the most notable exception being found 
in the Seals, in which animals (see Fig. 124, p. 347) each 
of the metatarsals and all the bones of the toes, except 
the terminal phalanges, have epiphyses at both ends of 
the shaft. 

Order PRIMATES. In Man (see Fig. 122, p. 341) the foot 
is broad, and in the ordinary standing position the whole 
length of the plantar surface (at least its outer edge) rests 
on the ground, the main axis of the foot being at a right 
angle with that of the leg. The inner or tibial side of the 
foot is arched before backwards, each extremity only resting 
on the ground. 

The tarsus is longer than the metatarsus, and the latter 
is longer than the digits, but the forms and relations of the 
tarsal bones are quite characteristic of the general Mam- 
malian type, and the five digits are present with the complete 
number of phalanges. The hallux is much stouter than any 
of the others, sometimes longer and sometimes shorter than 
the second toe. Its metatarsal is articulated to a nearly flat 
surface on the internal cuneiform, directed distally, so that 
it is placed in the same plane as the other toes, and cannot 
be freely separated from, or opposed to, them. There are 
no supplementary tarsal bones, and sesamoids are deve- 
loped under the metatarso-phalangeal joint of the hallux 
only. The phalanges are much smaller, shorter, and more 
compressed than are those of the manus. The ungual 
phalanges are very small, depressed, and somewhat spatulate. 

The principal distinction of the foot of the Simiina from 
that of Man is that it is more or less modified into a grasping 
organ. The tarsal and metatarsal bones and phalanges are 


the same in number and relative position, but the articular 
surface of the internal cuneiform for the hallux is saddle 
shaped, and is directed obliquely towards the inner or 
tibial side of the foot. The consequence is that the hallux 
is not only somewhat separated from the other digits, but 
is also set in a different plane, so that when it is flexed it 
turns towards the sole of the foot, and becomes opposed to 
the others, much as the thumb does in the human hand. 
It is this peculiarity of the pes which has given rise to the 
term quadrumanous,Qi "four-handed," occasionally applied 
to this group of animals. 

The hallux is usually relatively shorter than it is in Man. 
In the Orang (Simla satynts] it is particularly short, and 
often wants the terminal phalanx, while the metatarsals and 
the phalanges of the other digits are long and curved, the 
proportions of the three segments of the foot being exactly 
the reverse of those of Man, as the tarsal segment is 
shortest, and the phalangeal the longest. 

The form of the articular surface of the astragalus, and 
especially the free mobility of the navicular and cuboid 
bones on the astragalus and calcaneum, cause the foot of 
the Orang to be set very obliquely on the leg, so that when 
placed on a level surface the fibular border only rests on the 
ground, and the sole is directed inwards. This position suits 
well for grasping vertically-placed boughs of trees, but is ill 
adapted for standing or walking on the ground. A similar 
disposition is seen in a varying degree in most of the 
Monkeys, but in none so markedly as the Orang, in which 
animal all the peculiarities by which the simian is distin- 
guished from the human foot are most strikingly displayed. 

There are usually two sesamoid bones below each 
metatarso-phalangeal joint, and a single one below the 
cuboid in the tendon of the peroneus longus muscle. 




The structure of the foot of the Lemunna resembles 
generally that of the Simiina, and is in fact one of the 
principal bonds of union between these groups. The hallux 
is large and opposable, with a flattened ungual phalanx. 

The second digit in Lemur has a narrow, pointed, ungual 
phalanx, while that of the other digits is flat and spatulate 
as in the Sinriina. In Chiromys all the ungual phalanges, 


Kl .. 173. Right pe< of Tarsias spectrum (tint. size), a astragalus; e calcaneum , 
x : f ' internal cuneiform ; S* middle cuneiform ; C* external cuneiform ; 
i A cuboid ; I to V the digits. 

except that of the hallux, are compressed, curved, and 

In Perodicticus there is a supplemental ossicle in the 
transverse ligament of the plantar surface of the tarsus, 
corresponding to that met with in the carpus (see p. 286). 

A remarkable elongation of the tarsal segment of the pes 


occurs in the Galagos and certain other Lemurs, owing to 
the modification of two bones, the calcaneum and the navi- 
cular; the distal portion of the former, and the whole of 
the latter, having the form of nearly cylindrical rods placed 
side by side, while the other bones retain nearly their normal 
form and proportions. A precisely similar modification is 
carried to a still greater extent in the genus Tarsii/s (see 
Fig. 123, p. 345). 

All the terrestrial GARNI VORA have the normal number of 
tarsal bones, with very little deviation from their normal 
form and relations. 

The hallux is present and well developed, though shorter 
than the other toes, in the Ursidic, Procyonida, Mustelidie, 
and most of the Virerridcc. In the Canidcc, Hyanidtf, and 
Felidic, it is only represented by a rudimentary metatarsal. 
The other four metatarsals and digits are always well de- 
veloped and subequal. The ungual phalanges in the Felida: 
present the same characters as those of the fore limb (see 
p. 288). 

In the Bears, the foot is flat, broad, and plantigrade. In 
the Dogs and Cats, it is longer and narrower, and the heel 
is raised from the ground in walking. 

In the Sea Otter (Enhydra), the hind foot approximates 
to that of the Seals. It is very large and flattened, almost 
fin-like, and much everted ; but the hallux is still shorter 
than any of the other digits, and the two outer toes are the 

In the Seals the pes is completely modified for a special 
purpose. It has no longer any function as an organ of 
support or progression on land, and is habitually directed 
backwards, with the dorsal surface outwards, and the 
plantar surface in contact with the corresponding part of 
the opposite limb. The calcaneum is very short, its tube- 




rosily being almost obsolete. The two lateral digits (first 
and fifth) are both longer and much stouter than the others ; 
the middle digit is the shortest. In the Elephant Seal 
( Macrorhinus leoninus\ all the phalanges (except the 
terminal ones) have epiphyses at both ends of the shaft 
(see Fig. 124). 

FIG. I2 Porsal Mirfa.6 of rij{ht pes of young Elephant Seal (Mactorhinus 
ttoniniit), \, showing the epiphyses. The letters as before. 

In the Otariidic, or Eared Seals, and the Walrus, which 
use the hind feet in walking, these modifications from the 
ordinary type are not so marked, the calcaneum having a 
greater backward projection, and all the digits being of 
nearly equal length. In both there is a large sesamoid on 
the tibial side of the tarsus. 

In the greater number of the animals of the order 


INSECTIVORA the tarsus is quite normal, and there are five 
digits, all with curved, pointed, moderately developed 
ungual phalanges, the hallux being the shortest. In the 
Mole, the pes is narrow, having none of the modifications 
of structure observed in the manus, except that there is 
an unusually large slender sesamoid on the tibial side of 
the tarsus, corresponding to the falciform bone of the fore 
limb. In the Water Moles (Myogale), the pes is remark- 
ably large and almost fin-like. 

In the African genera Petrodromus and Rhyncfwcyon, the 
hallux is only represented by a rudimentary metatarsal. 
The last-named animal has a remarkably elongated pes, 
produced partly by the length of the metatarsals, and partly 
by a peculiar elongation of all the bones of the distal 
row of the tarsus, the cuboid and three cuneiform bones. 
Contrary to what occurs in the Galagos, the navicular and 
calcaneum are of normal proportions. 

Order CHIROPTKRA. The tarsus is very short ; the tuber 
calcanei is a slender curved process sometimes of con- 
siderable length, supporting the interfemoral membrane ; 
the metatarsals are equal and rather short ; the phalanges 
elongated and subequal in length, the hallux being rather 
the shortest ; the ungual phalanges are long, curved, com- 
pressed and pointed. In the Leaf-nosed Bats (PhyllorhinintR) 
and the anomalous genera, Thyroptcra and Myxopoda all 
the toes have only two phalanges. 

Order RODKNTIA. The structure of the pes varies much 
in different members of this order. In the Beaver, as in 
most swimming quadrupeds, it is disproportionately large 
and fiat. The five digits are well developed, but the third 
and fourth are considerably longjr and stouter than the 
others. The base of the fifth metatarsal is articulated to 
the outer side of the fourth metatarsal, and not directly to 





the cuboid. The middle cuneiform is very small. There is 
a large sesamoid bone on the tibial side of the tarsus, 
articulating with the astragalus, navicular, and internal 
cuneiform. 1 The tuberosity of the calcaneum is long and 
obliquely compressed. 

The functional digits in other Rodents may be five, as 
in the Rats, Porcupines, and Squirrels ; 
or the hallux may be suppressed, as in 
the Hares ; and occasionally the fifth 
digit is also wanting, reducing the 
number to three, as in the Capybara, 
Viscacha, and Agouti. The last-named 
animal has the three metatarsals elon- 
gated and closely pressed together, 
and all the digits with short subequal 
phalanges. A still further modification 
of the same type leads to the singular 
condition of pes met with in the 
Jerboas (genus Dipus, see Fig. 125), 
which at first sight resembles that of 
a bird. The three metatarsals are 
ankylosed together to form a single 
bone, which supports the three separate 
short digits, each with three phalanges. 
These alone are applied to the ground, 
the tarsus and long metatarsal segment 
being raised almost vertically. The hallux is wanting or 
rudimentary, but in some species (Alactaga) there is a small 
fifth digit. 

All the animals of the order UNGULATA are digitigrade, 

1 In Cercolabts and Erelhizon there is also developed a claw-like bone, 
articulating with the " sesamoid." G. 13aur, " American Naturalist," 

FIG. 125. liones of righ: 
pes of Jerboa (Dipvs 
agyptius), \. 


the heel being raised from the ground, and the metatarsal 
segment usually much elongated. There is no trace of a 
hallux in any existing species. As in the corresponding 
segment of the fore limb, the pes is formed upon one or 
other of two distinct types, each characteristic of one of the 

In the Perissodactyla, the third digit is the largest, in tin- 
centre of the foot, and symmetrical in itself; the second and 
fourth are smaller, and nearly equal in length, though some- 
times quite rudimentary. A line drawn through the centre 
of the foot passes through the axis of the third digit, and 
the middle of the external cuneiform, navicular, and as- 
tragalus. The distal surface of the astragalus has a large 
articular surface for the navicular, and a very small one for 
the cuboid, which bone is of comparatively less importance 
than in the Artiodactyla. The calcaneum does not articu- 
late with the lower end of the fibula. 

The Rhinoceros (Fig. 126) and Tapir have all the usual 
bones of the tarsus well developed. The internal cuneiform 
has a curved process projecting backwards. The middle 
cuneiform (c 2 ) is very small. The whole foot is compara- 
tively short and broad. The second and fourth toes 
are well developed, being nearly as long as the middle toe. 
The phalanges resemble those of the fore limb. In the 
Tapir the pes differs from the manus in wanting the fifth 

In the Horse (Fig. 127), the middle toe is greatly 
enlarged, and the second and fourth reduced to slender 
styliform metatarsals, about three-fourths the length of the 
second, but supporting no phalanges. The navicular (//) 
and the external cuneiform (c*) are very broad and flui. 
The cuboid (cl>) is small, and the internal and middle 
cuneiform bones are small and united together. 

XIX. ] 



Various gradational stages between the complete tridac- 
tyle foot of the Rhinoceros and the monodactyle foot of 
the Horse are met with in extinct species of the Perisso- 

In the Artiodactyla the third and fourth digits are nearly 
equally developed, and their ungual phalanges are flattened 
on their contiguous sides, so that together they constitute a 

Kl<;. 116.- Dorsal surface of right laru 
of khinocero> (A'A/rtiHvnM Sumatra/ 

FH;. 127. Dorsal surface of right tarstifc 
of Horse (Equus cakaJliiS), \. 

symmetrical form. The second and fifth toes, when present 
are also equal, but smaller than the others. A line drawn 
through the centre of the foot has on its tibial side the third 
digit and metatarsal, the external cuneiform, the navicular 
and half the astragalus ; and on its fibular side the fourth 
digit and metatarsal, the cuboid and the other half of the 



astragalus. The distal articular surface of the astragalus is 
divided into two nearly equal facets, one for the navicular 
and one for the cuboid. The calcaneum has an articular 
facet for the lower end of the fibula. 

In the Suina (Fig. 128) all the tarsal bones are distinct. 
The four toes are well developed, and the metatarsals are 

FIG. 1 23. Dorsal surface of right tarsus 
of Pig (Sus sere/a)^ J. 

Fir.. 129. Dorsal surface of right tarsu 
of Red Deer (Cen-us tlaphiis), J. 

usually distinct. The foot is relatively shortest and broadest 
in the Hippopotamus, the outer toes being nearly as long as 
the others, and the lingual phalanges very short, broad, 
and rounded in front. The Peccaries show a transition 
towards the ruminating sections of the order in the reduc- 
tion of the size of the outer toes and the confluence of the 

xix.] UNGULATA. 353 

third and fourth metatarsals. On the hind foot the filth 
digit is absent though the second is tolerably well developed. 

In the Tylopoda the cuboid and navicular are distinct. 
There is rio internal cuneiform. The second and fifth digits 
are entirely absent. The metatarsals of the third and fourth 
are united except at their lower extremity. The phalanges 
resemble those of the fore foot. 

In the Tragulina the cuboid, navicular, and two outer 
cuneiforms are united to form a single bone. The third and 
fourth metatarsals are confluent. The second and fifth are 
complete, extending between the small digits and the tarsus, 
but are very slender. 

In all the Ptcora (Fig. 129) the cuboid (cb} and navicular 
(//) are united, as are the second and third cuneiform bones, 
and in some Deer (Ctn'ulus and Pudu) these latter are 
further united with the cubo-navicular. The first cuneiform 
is always distinct, though small. 1 The third and fourth 
metatarsals (m in. and /// iv.) are united in the same manner 
as the mctacarpals, and the phalanges of the digits are very 
similar to those of the manus. The second and fifth 
metatarsals are always wanting, and bones of the corre- 
sponding digits are absent in the Giraffe and most of the 
Oxen, Sheep, and Antelopes. In the Deer there are 
usually three small phalanges to each of these digits, not 
directly articulated with the rest of the skeleton. A large 
oval sesamoid is commonly present in the plantar surface of 
the tarsus. 

The pes of the Hyrax closely resembles that of the 
Rhinoceros, but the ungual phalanx of the second digit is 
cleft almost to its base. The fifth metatarsal is represented 
by a small nodule ; the astragalus articulates with the cuboid. 

1 See Sir Victor Brooke, " Troceedings of the Zoological Society, 
1874," p. 34. 

A A 


In the PROBOSCIDEA the pes is short and broad, but 
smaller and more compressed than the manus, and in the 
more rudimentary condition of the two lateral digits shows 
a greater tendency to approach the Perissodactyle form. 
The astragalus is very flat, and has no articulation with the 
cuboid. The internal cuneiform is produced distally, as is 
the corresponding bone of the manus. 

Order ED KN TAT A. In the Sloths the pes much resembles 
the manus in its general characters, being long, very narrow, 
and curved, terminating in strong, compressed, pointed 
ungual phalanges, supporting hook-like claws. The tarsus 
is short ; the astragalus has a deep, cup-shaped cavity on its 
outer side, into which a conical projection of the lower end 
of the fibula is received. 

The appellation " Two-toed " applied to the genus Cho- 
Lrpus refers only to the anterior limb, for in the pes the 
three middle toes are functionally developed, and of nearly 
equal sixe in both the genera of the family. 

In Bradypus the tuber calcanei is long, compressed, and 
widened at the extremity. The tarsal bones have a great 
tendency to ankylosis. The first and fifth metatarsals are 
very rudimentary, and support no phalanges. The proxi- 
mal phalanges of the three middle digits are very short, and 
coalesce very early with the metatarsals, as in the corre- 
sponding part of the upper extremity. 

In Cholcepus, the tuberosity of the calcaneum is very 
small. The tarsal bones remain distinct from one another. 
The proximal phalanges of the three middle digits are 
extremely short, but not ankylosed with the metatarsals. 
The first and fifth metatarsals are about three-fourths of the 
length of the others, flattened, and gradually diminishing in 
size to their free ends. 

In nearly all the members of the Entomophagous section 


of the Edentata the pes is much more normal in type, and 
adapted for plantigrade progression on the ground. It does 
not even present any modifications corresponding to those 
observed in the manus of the same animals. The normal 
number of tarsal bones, and the complete number of pha- 
langes, are always present in each digit. Of these the second 
and third are usually the longest, the fourth next, and the 
first and fifth shortest. 

The little prehensile-tailed Tree Anteater (Cycloturus 
didactylus) has the pes modified into a climbing organ. 
The hallux is rudimentary, consisting of a metatarsal and 
one phalanx, concealed beneath the skin, but the four other 
toes are subequal and much curved, with long, pointed, 
compressed, ungunl phalanges. The tuber calcanei is di- 
rected towards the plantar surface, and parallel with it, and 
extending to about double its length is the greatly elongated 
sesamoid ossicle of the tibial side of the foot. These 
together support the prominent calcarine cushion to which 
the nails are opposed in climbing. 

In the MARSUPIALIA, the hind foot is subject to great 
modifications, some of the genera presenting very striking 
deviations from the typical condition. 

The seven bones usually found in the Mammalian tarsus 
are always present and distinct from each other, but the 
astragalus is relatively smaller and more flattened than in 
placental Mammals. In the climbing Marsupials espe- 
cially, the articular surface for the fibula, instead of being 
perpendicular to that for the lower end of the tibia, is almost 
in the same plane with it ; and in all, the " head," or portion 
of the astragalus which projects forwards to articulate with 
the navicular, is very slightly developed 

In the American Opossums (Didelphidcz), the foot is short 
and broad ; the hallux is stout, placed at right angles with, 

A A 2 


and opposable to, the other four digits ; it has a short 
rounded terminal phalanx, which bears no nail. The other 
digits are subequal, and have compressed, pointed, curved 
ungual phalanges. 

In the Dasyuridcz, the foot is comparatively narrow ; the 
second,. third, fourth, and fifth toes are subequal; the hallux 
is either very small and placed close to the others, or com- 
pletely suppressed, as in the Thylacine. 

In the Wombats (Phascolornyida), the foot is short and 
broad ; the hallux is very short, with only one rounded 
phalanx, and divaricated from the other toes. These are 
nearly equal in length ; the fourth and fifth are stouter than 
the second and third, thus showing a slight tendency towards 
the condition met with in the next group. 

In all the remaining Marsupials a peculiar condition of 
the pes, called syndactylism, prevails. Whatever the con- 
dition of the other toes, or whatever the general form of 
function of the foot may be, the second and third metar- 
tarsals and digits are very slender, and inclosed nearly to 
their extremities in a common integument, so that they look 
externally like one small toe with two claws. 

In the Kangaroo (Macropiis) the whole foot (Fig. 130) is 
very long and narrow, and rests entirely on the ground in 
the ordinary position of the animal. The tarsal segment is 
short, the metatarsus very long. The cuboid is greatly 
developed, the navicular and the three cuneiform bones 
exceedingly small, in conformity with the condition of the 
digits they respectively support. There is a sesamoid on 
the plantar surface of the tarsus. The fourth metatarsal 
and digit are enormously developed, the fifth moderately 
so ; the second and third are nearly as long as the fifth, but 
excessively attenuated. There is no rudiment of a hallux. 
The ungual phalanges are conical, pointed, slightly curved 

xix.] MARSUPIALIA. 357 

above, and flattened on the under surface. The whole foot 
is much compressed laterally, especially at its hinder part, 
so that the proximal ends of the second and third are thrown 
behind that of the great fourth metatarsal, and entirely 
concealed in a view of the dorsal surface of the foot. 

The Tree-Kangaroos of New Guinea (Dcndro/agJts), 
which habitually live among the boughs of large trees, have 

KH-. i-.o.* of right pe< of K.ati- Fir.. 131. Hones of right foot of Pha- 
it-"'" {Macnifut tuniH-ttiJi, J. lanjjer (PkalaHgista rulftina), [. 

the feet constructed on the same type, but shorter, and more 
laterally extended. Hypsiprymnadon moschatus, a species 
of Rat Kangaroo, lately discovered by Ramsay in North- 
east Australia, differs from all other members of the family 
ida in the possession of a small prehensile hallux> 

7///: H!.\'D I-OOT OK 


without a nail, and presents therefore an approximation to 

the next group. 

In the leaf eating, climbing Australian Opossums (Phalan- 

%:sta, Fig. 131; and Koalas < Phaxolarctos) the second and 

ihird toes are al-o very slender, but the fourth and fifth are 
more equal, especially in length, the 
foot is broad, and there is a strongly- 
d.-veloped prehensile and opposable, 
through nailless, haliu.x. 

The insect- and root eating, ground- 
dwelling Bandicoots (Piramelid*} dif- 
fering in many other respects from the 
Kangaroos, have their hind foot con- 
structed on txact'.y the same type as in 
Macrifus. even to the relative length 
of the different dibits, though there is 
often a rudiment of the metatarsal of 
tiie hallux. In one remarkable genus 
i Ciurrcfus), already mentioned on 
account of the peculiar structure of the 
man us (see Fig. 1 1:. p. 310), the same 
type is carried to a great extreme, the 
fourth toe (>ee Fig. 137) remaining of 
a prodigious size, and the fifth being 
reduced to even smaller dimensions 
than the second or third. 
MOXOTREMATA. In both species the seven usual bones 

of the tarsus are complete and distinct, 1 and the five digits 

have the normal number of phalanges. 

1 It has been stated that the culv 1 1 in the Ornithorhynchus is divided 
into two b >nes, a-< i:i some reptiles, one supporting the fourth and the 
other the fifth metatarsal ; but this U not the cose in any specimen 
which I have examined. 

if,. 132. Fonc< f 

foot of Cf!<rr:'flti .. 

naiis (iiat. >uc). 

xix.] MONOTREMATA. 359 

In the Ornithorhynchus the proximal articular surface of 
the astragalus is divided by a deep groove into two distinct 
heads, one for the tibia, and the other for the ftbula, the 
latter being the larger of the two ; the inner side has a cup- 
shaped socket, into which fits an incurved conical process 
from the lower end of the tibia. The tuberosity of the cal- 
caneum is broad and bifid at the extremity, and directed 
not backwards, but towards the tibial side of the foot. In 
the male there is an additional, large, flat, curved ossicle, on 
the hinder and tibial side of the plantar aspect of the tarsus* 
articulated chiefly to the tibia, which supports the peculiar 
perforated horny spur characteristic of this sex, the function 
of which has not yet been discovered. There is also a small, 
rounded, supplementary ossicle, below the tibial edge of the 
tarsus, near the articulation between the astragalus and 
scaphoid. The metatarsals increase in length from the first 
to the fifth. The phalanges are all rather long and slender- 
The four outer toes are nearly equal ; the hallux is somewhat 
shorter. The ungual phalanges are compressed, slightly 
curved, and very sharp-pointed. 

In the Echidna the astragalus is large, with an irregular^ 
broad, rounded, proximal articular surface, not divided by 
a groove, and with a much less distinct fossa, for the in- 
ternal malleolus. The tuber calcanei is directed forwards, 
it is also bifid, and its external process is much longer than 
the other and curved towards the plantar surface of the 
foot. The spur of the male, and the ossicle which supports 
it, are much smaller than in the Ornithorhynchus: The 
metatarsals are shorter and broader ; they increase in length 
from the first to the fifth. The hallux is very short, and has 
a flattened, conical, ungual phalanx. The proximal and 
middle phalanges are all very short and broad. The ungual 
phalanx of the second digit is extremely long and falcate, 

360 THE HIND FOOT OR PES. [CH. xix. 

the others gradually diminish to the fifth. The ends of the 
toes are turned outwards and backwards in the ordinary 
position of the animal. 

The ungual phalanges of both extremities in the Mono- 
tremata have a deep median groove, near the base of the 
under surface, leading at its distal extremity into a foramen. 



THAT a general correspondence exists in the plan of con- 
struction of the anterior and posterior extremity cannot fail 
to strike the most superficial observer, though to follow out 
this correspondence into all its details has severely exercised 
the ingenuity of many an anatomist 

It would he quite beside the character of the present 
work to give an historical account of the numerous and 
very various views which have been held upon this sub- 
ject, 1 but I propose to lay before the student in a condensed 

1 For the bibliography of this question, see Ch. Martins, " Nouvelle 
c tinparaison cits uieuibres," (Mem. de 1'Academie de Montpellier, 
1^57); Mivart, "On some Points in the Anatomy of Echidna 
hystrix," (I. inn. Soc. Trans, xxv. 1866) ; and Rolleston, " On the 
Homologies of certain Muscles connected with the Shoulder Joint," 
(ibid. xxvi. 1869). See also Humphry's "Observations in Myology," 
1872; A. Cioette, " Ueber Entwickelung und Regeneration des Glied- 
ma-senskclets der Mulche," (Leipzig, 1879) ; H. Strasser, " Entwicke- 
lung der Extremitseten-knorpel bei Salamandern und Tritonen," (Mor- 
phul. Jahrb. 1879); A. Sabatier, " Com para ison des Ceintures et des 


form such portions of the general outcome of these researches 
as appear to be most satisfactorily established, premising, 
however, that all the statements hereinafter to be made 
have not yet met with universal assent. 1 

In the first placj, it is perfectly obvious that the fore and 
hind limbs have each a similar division into four main seg- 
ments ; the shoulder girdle, the arm, the fore-arm, and the 
manus of the one representing respectively the pelvic girdle, 
the thigh, the leg, and the pes of the other. 

To proceed to further details, it is necessary to place the 
limbs (at least in imagination) in an exact corresponding 
position one, in fact, which is often impossible in the adult 
animal on account of the modifications of the articular sur- 
faces to suit the posture best adapted for the habits and 
mode of life of the individual, but which is the position of 
the limbs of many Mammals, when they first appear as bud- 
like processes from the side of the body of the embryo. In 
this position - the limbs are extended at right angles to the 

Membres antericurs et posterieurs dans la serie des Vertebres," (Mont- 
pcllier, 1880); G. Gegenbaur, " Untersuchungen zur vergleich. Ana- 
tomie ; I. Carpus und. Tarsus," (Leipzig, 1864, p. 115 ff. ) ; and "Zur 
Gliedmassenfrage," (Morphol. Jahrl>. v. 1879) ; F. M. BalLmr, " A 
Treatise on Comparative Embryology," (London, iSSi). 

1 For an exposition of the very opposite hypothesis of " Antero- 
posterior Symmetry," see Jeffries Wyuian, " On Symmetry and Homo- 
logy in Limbs," (1'roc. Boston Nat. Hist., June, 1867, p. 277) ; and 
the elaborate series of papers by Dr. Flliott Coues, published in the 
" Medical Record " (New York) for 1870. 

2 This horizontal position is probably not the primary one, and 
wherever it occurs, as in Selachians and many Mammalia, it is a secondary 
modification of the original position of the cheiropterygium, in which 
the long axis of the limb is directed obliquely outwards and backwards 
towards the tail, the ulna and the fifth finger looking dorsally, and the 
palmar and plantar surfaces of hand and foot looking towards the sides 
of the body. This position was that of the limbs of the Enaliusauri, and is 
still repeated by the Cctacea and by Ceratodus and the e-nbryonic limb.t of 


axis of the trunk and parallel to each other, as in Fig. 133, 
A and B. There is then to each limb a superior or dorsal 
surface (turned towards the observer in the figure), an 
inferior or ventral surface, and an anterior and a posterior 
edge. These last are called by Professor Huxley pre-axial 
and post-axial (in reference to the axis of the limb itself) to 
avoid the confusion with anterior and posterior in the modi- 
fied positions they assume in Man and various animals. In 
the figures the pre-axial side is left light, and the post-axial 
side is shaded. 

The dorsal surface of the anterior extremity includes the 
back of the hand and the extensor surface of the fore-arm 
and arm. The dorsal surface of the posterior extremity 
includes the dorsum of the foot, front of the leg, and the 
extensor surface of the thigh. The pre-axial border of the 
anterior extremity has in it the pollex, the radius, the 
condyle commonly called " external," and the greater tuber- 
osity. The pre-axial border of the posterior extremity in- 
cludes the hallux, the tibia, the condyle commonly called 
" internal," and the lesser trochanter. All these parts, then, 
should be regarded as serially homologous. 

Leaving for the present the shoulder and pelvic girdles 
out of consideration, we will next consider the adaptive 
changes which take place in the segments of the limb 
proper in various animals. These will be best understood 
by dividing them into stages (all of which are represented 
in the diagram), though it is not meant to imply that the 
limbs actually go through so many distinct phases in the 
course of development, as all the various modifications from 
the primitive to the most adaptive positions may take place 
gradually and even simultaneously. 

the Amphibia, and is in harmony with Gegenbaur's derivation of the 
limbs from the archipterygium. (H. G.) 


KiG. 133. Diagrammatic representation of the positions of the limbs of Mammalia. 
'1 lie pre-axial border is left light, the post-axial border is shaded, in all the figures. 
Limbs of the right side are represented in all cases. (A) dorsal aspect of the 
anterior extremity in its primitive unmodified position ; gl> glenoid border of the 
scapula; j- spine ; ch coracoid border; ssf sub-scapular fossa : //post-scapular (in- 
fra-spinal)fo:-sa ; c coracoid ; h humerus ; gt greater, radial, or pro-axial tuberosity ; 
// lesser, ulnar, or post-axial tuberosity ; cc external (in the modified position), 
radial, or pre-axial condyle ; ic internal, ulnar, or post-axial condyle ; r radius ; 
H ulna; I polkx ; v fifth digit. (H) dorsal aspect of the posierior extremity in the 
same position; nb acctabular border of the ilium; fb pubic border; it> ischial 
border ; gs gluteal surface ; is iliac s'irf:tce ; i ischiinn ; yJpubis ; yfemur ; // lesser, 
tibial, or pre-axial trochanter ; / greater, fibular, or post-axial trochanter; ic in- 
ternal (in the modified position), tibial, or pre-axial condyle : <r external, fibular, 
or post-axial condyle ; t tibia ; f fibula ; I hallux ; v (ifih digit, (c) th- anterior 
extremity, with the humerus in the same position, bin the elbow- and wrist-joints 
bent. (D) the posterior extremity in the same position. 


In what may be considered the first stage of modification 
(Figs, c and D), each segment of the limb is simply bent 
upon the one above it. The proximal segments (humerus 

FIG. 134. Diagrammatic reprevntation of the positions of the limbs of Mammalia, 
continued. (K) the anterior-extremity with the same flexures as in c, but with the 
whole limb rotated backwards The pre-axial side is external. The letters as 
before, (f) the posterior extremity, with the joints bent, and the whole limb rotated 
forward-, as in the ordinary position of quadruped Mammals. The post-axial side 
is external (<;) the humerus in the same position as in E, but the fore-arm rotr.ted 
as in the ordinary position of quadruped Mammals. Whilst the pre-axial side of 
the humerus remains external, the post-axial side of the inanus is now external. 
(H) the anterior extremity of a Cetacean (H yperoodon), dorsal surface, (t) the 
posterior extremity of a Seal, dorsal surface. 

and femur) remain unchanged in position, the dorsal surface 
still looking upwards, and the ventral surface downwards ; 


the middle segment is bent downwards, so that its ventral 
surface faces inwards and its dorsal surface outwards ; and 
the joints between these segments (elbow and knee) form 
prominent angular projections. The third segment being 
bent to a greater or less degree, in the opposite direction 
to the middle one, retains much of its primitive position, 
the dorsal surface being directed upwards and the ends of 
the digits pointing outwards. The relations of the pre-axial 
and post-axial borders of the limb are unchanged. No 
Mammal habitually carries its limbs in this position, although 
the climbing Galeopitheats and the Sloths are not far from 
it. It is, however, very nearly the normal position of 
some Reptiles, especially the Tortoises, though it is ill 
adapted for anything but a very slow and clumsy mode of 

The next change, and one which takes place at a very 
early period in embryonic life, and which is one of the most 
essential in giving the characteristic conformation of the 
extremities of the higher Vertebrates, is a rotation of the 
whole limb from the proximal end, though in opposite 
directions in each case. 

The anterior extremity (see Fig. E.) is rotated from the 
shoulder, through nearly a quarter of a circle, backwards, so 
that the humerus, instead of being at a right angle to the 
axis of the trunk, is nearly parallel with it, the elbow points 
backwards, the pre-axial side is outwards, and the post-axial 
side towards the middle line of the body, and as long as the 
radius and ulna retain their primitive parallel position, the 
manus is placed with the ends of the digits directed back- 
wards, the pre-axial side being external. 

The hind limb (see Fig. F.) is, at the same time, rotated 
from the hip to the same extent forwards, so that the femur 
is also nearly parallel to the axis of the body, but with the 


knee projecting forwards ; the pre-axial side is inwards and 
the post-axial side outwards ; the tibia and fibula are parallel, 
the former internal and the latter external ; the foot has the 
ends of the digits directed forwards, the hallux or pre-axial 
digit is on the inner, and the fifth or most post-axial digit 
is on the outer, side. 

In this position the hind limb remains, subject only to 
minor modifications, in nearly all terrestrial Mammals; but 
the Walrus, and to a certain extent the Sea-Lions, alone 
carry the fore limb as described above without further 

The next stage, affecting the fore limb alone, consists in 
the rotation of the lower end of the radius around the ulna 
which brings the distal extremity of the manus round from 
the back to the front of the limb (see Fig. c.). In most 
Mammals the limb is permanently fixed in this position, 
and the bones of the fore-arm become greatly modified in 
consequence, as described in Chapter XV. It will now be 
understood how, though the outer side of the humerus 
corresponds with the inner side of the femur, in ordinary 
quadruped progression, yet the outer side of the manus 
corresponds with the outer and not the inner side of 
the pes. 

To these general conditions there are certain modifica- 
tions met with in some animals, and certain exceptions 
in others. 

The modifications with regard to the anterior extremity 
are that the humerus may be quite horizontal, or its distal 
end may incline upwards, or, as is much more frequently the 
case, it may incline somewhat downwards, so that the dorsal 
surface is posterior and the ventral surface anterior ; the fore- 
arm in the ordinary resting position may be quite vertical, 
or inclined with its upper end backwards ; the whole of the 


manus may rest entirely on the ground, as in the so-called 
" plantigrade " or rather " palmigrade " animals, or the 
proximal part, the tarsus and metatarsus, may be raised and 
placed more or less vertically, the limb resting either 
on all or only on the terminal phalanges, according 
to the completeness of the " digitigrade " mode of 

Similar modifications occur in the hind limb. The 
femur is usually inclined with its distal end downwards, so 
that the dorsal or " extensor " surface is anterior, and the 
ventral or " flexor " surface posterior. In the Elephants it 
is very nearly vertical. In most animals which occasionally 
assume the upright position, as the Kangaroos and some 
Rodents, the femur is ordinarily inclined upwards at its 
distal extremity, so that the knee is above the acetabulum, 
and the pelvis slung as it were between the two hind limbs. 
In Man, on the other hand, in standing or walking the 
femur is nearly vertical with the distal ends downwards, and 
the pelvis is supported on the top of the limbs. 

The positions of the limbs which are quite exceptional 
are those of certain aquatic animals. 

In the Cetacea (Fig. H.) none of the segments of the 
anterior limb undergo any deflection from the primitive 
straight condition, nor is there any rotation of the bones of 
the fore-arm. The only changes which take place are a partial 
rotation backwards from the shoulder, and a slight turning 
downwards of the pre-axial border. In the Sirenia and the 
Seals, there is a slight bend at the elbow and the wrist, 
but little or no rotation of the fore-arm. 

In the hind limb of the Seal (Fig. i) there is very little 
flexure at the joints, and the whole limb is turned backwards 
instead of forwards from the hip, and at the same time 
rotated on its axis, so that the pre-axial border becomes 


turned downwards. The skeleton of this limb, therefore, 
and that of the fore-limb of the Cetacean, being retained 
normally in almost exactly similar positions, are well adapted 
for demonstrating the correspondence between the re- 
spective bones of which they are composed (see Figs. 
H. and i.). 

The necessity of the modifications in the direction of the 
axes of the heads of the humerus and femur spoken of pre- 
viously will easily be understood by a consideration of the 
relative positions that these bones are adapted to assume. 
Thus the axis of the head of the humerus in the majority of 
Mammals is inclined towards the post-axial side of the shaft 
of the bone, while that of the femur is inclined towards 
its pre-axial side. 

Hitherto nothing had been said about the shoulder 
and pelvic girdles, because the correspondence of their 
parts is not so easily explained, nor so generally recog- 
nised, as that of the segments of the limb proper. The 
following view appears to be, of those yet suggested, the 
most probable. 

It has been already shown (Chapters XIV. and XVII.) 
that the lateral half of each girdle consists primarily of a 
bar or rod placed vertically, and divided into an upper and 
a lower segment, the point of attachment of the limb being 
close to the junction of these two segments. The upper 
segment in the fore-limb is the scapula, in the hind-limb the 
ilium ; the lower segment in the fore-limb is the coracoid, 
in the hind-limb the ischium and pubis. 

In every Mammal both scapula and ilium may be re- 
solved into bars or rods of three-sided or prismatic form. 
The two extremities of each bar are placed, as regards the 
general position of the trunk, dorsally and ventrally. The 
dorsal or upper extremity is capped by the supra-scapular 

B B 


epiphysis in the shoulder girdle, and by the corresponding 
supra-iliac epiphysis in the pelvic girdle. The ventral 
or inferior extremity enters into the formation of the 
glenoid or the acetabular articular cavity, as the case 
may be, and joins the coracoid or the ischial element ol 
he girdle. 

The bar, supposed to be in a nearly vertical position, has 
three surfaces and three borders. In what may be, at least 
theoretically, considered one stage of their primary position, 
the surfaces of each bar are (i) Inner or vertebral, turned 
towards the vertebral column ; (2) Pre-axial, corresponding to 
the pre-axial line of the limb (Fig. 125, A ff, B is); (3) 
Post-axial, corresponding to the post-axial line of the limb (A 
ssf, B gs). The borders are (i) External, in a line with the 
middle of the dorsal surface of the limb, and terminating 
below at the upper margin of the glenoid or acetabular cavity 
(A/', i? aft) ; (2) Antero-internal, terminating below in the 
acromion or in the pubis (A s, H //>) ; (3) Postero-internal, 
terminating below in the coracoid, or the ischium, as the 
case may be (A cl>, n il>}. 

The correspondence between these parts of the scapula 
and ilium will be better understood by placing them in 
a tabular form, the middle column showing the names 
expressed in the generalized or ideal condition applicable 
to both in their primitive condition, and the column at each 
side giving the special terms applied to each part in its 
variously modified forms. 




Pre-scapular fossa. I. Vertebral. 
Supra-spinous fossa. 


Sacral surface. 
Inner surface of ilium, 
behind linea arcuata 
interna, including the 
at titular surface for 
the sacrum, and the 
portion of the bone 
above and below this. 

Post-scapular fossa. ;. Pre-axial. Iliac surface. 
Infra-spinous fossa. , Internal iliac fossa. 

Sub-scapular fossa. 3. Post-axial. Gluteal surface. 




Glenoid border. 
Posterior border in 

most animals. 
Axillary border in 



Coracoid bonier. 
Anterior border in 

most animals 
Superior border in 


I. External. 

Acetabular border 
Anterior border. 

2. Antero-internal. Pubic border. 

Linea arcuata interna. 

3. Postero-internal. I schial border. 

Posterior border. 

As the humerus in ordinary quadruped Mammals is 
rotated backwards from its primitive position, and the femur 

B B 2 


is rotated forwards, so that the pre-axial side of the first 
becomes external, and the really corresponding side of the 
other becomes internal, so it is with the scapula and ilium. 
Each has undergone a rotation on its own axis, through 
nearly a quarter of a circle, and in the opposite direction 
(see Fig. 134, E. and F), so that the inner surface of the one 
comes to correspond with the outer surface of the other, 
the anterior border of the one with the posterior border of 
the other. The long axis of each is also differently inclined, 
the upper end of the scapula leaning backwards, while that 
of the ilium is inclined forwards, which makes the resem- 
blance between them seem still more obscure. 

These views are considerably strengthened by a considera- 
tion of the disposition of the muscles connected with the 
various bones in question. 1 

The principal differences between the shoulder and pelvic- 
girdles of the Mammalia are two: (i) The rudimentary 
condition of the inferior or ventral section of the girdle (the 
coracoid) in the former, as compared with the vast deve- 
lopment of the corresponding part of the lower extremity : 
(2) the free condition of the anterior as compared with the 
posterior girdle. It is neither attached to the vertebral 
column above, nor does it (except in the Monotremata) join 
the opposite part in the middle line below. To compensate 
for this, a clavicle is superadded to the anterior girdle in 
many Mammals, for which there is no exact homologue in 
the lower extremity. 

It has been shown in Chapters XVI. and XIX. that 
the terminal segments of each limb present a remarkable 
general correspondence with certain constant differences. 

1 See Flower, "On the Correspondence between the Parts composing 
the Shoulder and the Pelvic Girdles of the Mammalia." Journal of 
Anatomy and Physiology, vol. iv. p. 239, 1870.) 


There can be no question but that the carpus and tarsus, 
the metacarpus and metatarsus, and the various dibits 
beginning, at the pollex in the one and the hallux in the 
other, are serially homologous, or rather homodynamous ; 
the circumstance of the constant absence of one of the 
bones of the pre-axial digit in both fore and hind limbs is 
most significant. 

In the carpus and tarsus, the serial homology of the four 
bones of the distal row in their respective order is generally 
admitted, but with the other bones there is still some differ- 
ence of opinion. Gegenbaur has, however, given good 
reasons, 1 derived chiefly from the results of tracing both 
limbs back to their less modified condition in Reptiles and 
Amphibia, for considering the astragalus as equivalent 
to the scaphoid and lunar united, or the scapho-lunar 
bone of the Carnivora, c. ; the ca'.caneum as repre- 
senting the cuneiform, and not, as often supposed, the 
cuneiform and the pisiform ; and the navicular of the foot 
as representing the os centrale, found only occasionally 
in the manus of Mammals. 

1 " Untcrsuchungcn zur vergleichenden Anatomic," l tcs Heft, 
Carpus und Tarsus, 1864. For a modification of this view see Baur's 
suggestion on p. 340. 


On p. 90, add to footnote: 

Ruge, " Unlersnchungen uehcr Kntwicklungsvorgange am Bnistlieine 
itnd an cler Sternoclavicularverbindung des Menschen." Morpholo- 
gisches Jnhrbuch, vi., ]>. 362, 1880. 

Gegenhaur, " Uehcr ilic episternalen Skelettheilc und ilir Vorkom- 
nien lx?i den Saugethieron and beim Menschen." Jenaischt: Zeitschrift, 
i. p. 175. 1864. 




Ar;un (Arrr/nv/a), 96, 182, 275, 349 
Ailnrm. 8 1 
AUtlara. 85. 349 
AUtt (Klk), 83. 197. 298 

Ano^talktritim. 73 

Anleater. 47. 61. 64. 68. 75. 88, 102, 103, 

111, 114. 229. 230. 333, 333, 359. 305, 

3. 336, 355 

Anlclope, 74. 196, 197, 398, 353 
Antelofiid*. i /' 

Antitixatra (I'rungbuck), 2, 298 

Atmyx. 8 1 

Ape. 29. 30. 54. 72, 372 

Antitli*. So 

Aritomyi (Marmot), 85, 179, 180. 182, 

Anttmyx. 81 

Armadillo, 29. 39. 47. 61. 64. 68. 75. 76. 
104, 107, 113. 114. 231. 232. 259. 260, 
278. 283. 306. 307. 308, 324, 336, 342 

ARTIODACTYLA, n. 29, 41, 57, in, 192, 

29*. Tli 35>- 35 
Arvicpta. 8s 
AteUt (Spider Monkeys). 54, 72, 79, 163, 

273, 287 
Ainkmiit, 81 
Anl^odtit, 86 
Avahit. 79 
Aye-Aye (Ckirvmyt), 79. 273, 287. 345 


. 83 

Baboon (Cynocrfhalus), 72, 160. 161. 162, 

165. 272. 286, 318 
HaLrna (Whale), 43. 44, 84. 09, 100, 112. 

220, 331, 323, 256, 297. 300, 323, 335. 


.'laltrHtip/rra. 43, 58. 84. 100. 101, tit, 330, 

331, 333, 356. 300, 323, 336 

Bandicoot (ftratnelet), 48, 49, 65, 263, 

309. 310. 358 
Kjttaris, 8 1 

Bat, 9, 15. 39. 30. 96, 176. 177. 291, 348 

ffathyrrgvi. 85. 179, 180, 183 

Bear (Ursui). 55, 68. 72. 81, 167, 168. 274, 

287, 388. 330, 331. 346 
Beaver (Castor), 26, 68, 73, 85, 96. 180, 

181, 183, 183. 254, 375, 291, 392, 321, 


Beluga (White Whale), 46, 301. 302 
fifrarrftMS, 84 
fiettemgia, 88 
Bison, 82 

Bot(Ox), 73. 74, 82, in, 195, 298. 353 
Sosf/afAut. 82 
Km'itLr, 196 
flraJypodidff, 259 
Bradypus (Sloth), 48, 6t, 62, 87, 104, 233, 

260, 261, 278, 304, 305. 354 
flitfra/HS, 83 
Buffalo, 57 

CABA;sot.i (Xfnurus), 88, 307 

Cachalot (Physcter), 47, 59. 60. 84, 9?, 99, 

113. 217, 218, 219, 255. 301, 302 
Camel (Camefus), 41, 47, 83, in, 197, 276, 

297, 298, 299. 322 
Canidtr. no. 255, 274, 346 
Cants (Dog), 20. 27, 37, 38, 55. 68. 80. 94, 

117, 120, 136, 139, 143, 144, 148, 173, 192, 

247, 273- 288, 317, 331, 346 
Cape Anteatcr (Otyctfrofius), 47, 6r, 64, 

68, 88, io3, 114. 233, 258, 308, 336 
Cape Buffalo, 57 
Cape Golden Mole {Chrysochloris), 175, 

253. 275, 289 
Cape Jumping Hare (Pedetes), 73, 180, 


Cafra, 83 
Cafrtolus, 83, 298 
Cafronys, 56, 86. 292. 333 
Capybara (Hydroclurtus), 39, 73, 86. 96, 

179, 181, 182. 183, 2,2, 349 
Cartatus, 83 
CARNIVORA. 9, n. 30, 37. 38. 54, 55, 68, 

72. 79, 80, 81, 93. no, 166, 173, 176, 248, 

255, 273, 287, 288, 289, 320, 325. 331, 

342, 346 



Castor (Beaver), 26, 68, 73, 85, 96. 180. 

181, 182, 183, 254. 275. 291, 292, 321, 332 
Cat, 167, 172, 173, 346 
Catarrhina-, 163 

Cavia (Guinea-Pig). 73, 86, 181, 254, 321 
Cavy (Coelogenys), 86, 181 
Cebidce, 160, 161, 163, 164, 273 
Cebus. 79, 161 
Centetes (Tenrec), 38, 56, 72, 6, 174, 274, 

289, 320, 332 
Centetida, 174 
( 'ere o fates, 349 
Cercoleftes. 81 
Cercopitheciis, 78, 165 
Cervidir, 196, 298 
Cervulus, 83. 353 
Cemus (Deer), 40, 83. 97, 257, 273, 237, 

334- 352 
CETACEA, 7, 9. n, 31,43.58,61, 68, 74 78, 

84, 85, 98, III, 2C9, 210, 2l6. 223. 225, 
255, 256, 262, 263, 269, 277, 278, 283.284, 

299. 301, 302, 312, 323, 335, 368 
Cheetah, 173 

Chelydra (Water Tortoise), 281 
Chimpanzee ( Troglodytes), 37, 53, 54, 67, 92, 

no, 160, 161, 251, 286, 318, 331 
Chinchilla, 86 
CJtirogaleus, 79 

( 'hirontys (Aye-Aye), 79, 273, 287, 345 
Ckironectes, 89 
CHIKOHTKKA (Hats), 9, n, 39. 56. 68, 73, 

87. 96, 176, 253, 275, 290, 314, 320, 332, 


Chlamydopkorus, 75, 88, 232 
Cha'ropus, 309, 310, 337, 339, 358 
Cholifpus (Sloth), 15, 30, 48, 61,87, l 4> 

109, 113, 235, 260, 261, 278, 304, 305, 

Chrysochloris (Cape Golden Mole), 56, 86, 

252, 275, 289. 320 
Chrysothrix, 79 
Ctrtofffnyf (Cavy), 86, 181 
Colobus, 78, 287 
Condylura. 251 
Connodurtes, 82 
Coryphodon, 292 
Coypu (Myojotamtis), 86, 179, 253, 275, 


Cricetus, 85 
Crocidura, 86 
Crocodile. 66 
Crossarchus, 80 
Cryptoprocta, 80 
Cyclotitrus (Anteater), 61, 75, 88, 103, 113. 

2 3'. 259> 305, 3 06 ' 355 
Cynocepltalns (Baboons), 79, 160, 161, 162, 

165. 272, 286, 318 
Cynti-litrus. 80 
Cynomys, 85 
Cynonycterts, 96 
Cynoptrnis. 87 
Cystpphora, 55, 82, 173 


Dasypodidte (Armadillos), 47, 61, 64, 68, 

104. 107, 113. 259, 306 
Dasyprocta (Aguti;, 86, 96, 182, 275, 292, 


Dasypus, 62. 88, 231, 306, 307 
Dasyuridtr, 356 
Dasyunts (Dasyure), 89, 239, 279, 309, 

325, 326 
Deer, 40, 74. 97, 196, 257, 273, 297, 298. 

334- 352, 353 
Delphinaptents, 84 
Delphinida, 46, 217 
Ittlphinus (Dolphin), 46, 58, 59, 60, 61, 85. 

98, 112. 256, 303 
Dendrohyrax, 57, 84 
Dendrolaffus ('free Kangaroo), 88, 357 
Desmodus, 73, 87 
Dicotylfs, 83 
Didelphia. 2, 4. n, 48 
Didelphida-, 262, 355 
Didelphys (Opossum). 48, 49. 65, 76, 89. 

239. 39, 324, 326, 355 
Dinoccras, 292 
Dinoccrata, 8 
Dipns (lerb-.a) 39, 85, 349 
1 )og (Cams). 20, 27, 37, 38, 55,68,94, 117. 

120, 136,139, 143, 144, 148, 172, 192,247. 

273. 288, 317, 331. 346 
Dolichotis, 86 
Dolphin. 46, 58. 59, 60, 61, 85. 98, 112, 209, 

211, 214, 217, 219, 221, 226. 255, 256, 302, 

33. 3 2 4 
Dugong (Halicore), 42. 57, 84, 101, 228. 

277. 33. 322 


F.chidna, 3. 25, 50, 65, 68. 77, 89, 104, 114. 
175. 241, 242, 243, 263, 264, 310, 326, 338, 


Ecliidnidtf, 3 
EDENTATA, 6, 11, 31, 47, 61,62. 64,68, j--,, 

87, 88, 102, 113, 22Q. 258, 278, 304, 324. 

328. 336, 340, 354 
Eland, 57 
Elephant (_Elephas), 42, 57. 74. 84, in, 148. 

201, 203, 205, 227, 258, 292, 293, 294. 322, 

328, 335, 368 

Elephant seal (Macrorhinns), 82, 284, 347 
Elk (Alecs), 83, 197, 298 
Ellobius, 85 

En/tydraCf>c^ Ouer), Si, 328, 346 
Epotnophorus, 87 
Equus (Horse), 41, 57, 83.97, IIT > '85. 187, 

188, 258, 276, 284, 295, 256, 339, 350, 

Erelhizon, 349 

Ericitliis. 86, 332 

Erinaceidip (Hedgehogs), 174 

Erinaceus (Hedgehog), 56, 72, 6,94, 174, 

175. 274. 289, 320, 332 
F.nplercs, 80 
EuTHERtA, 2, 4, 5, 6 




Felittft. 55. no. 169, 172, 255, 274, 346 
/<// 55. 68. 69, 70, 79, 80, i6>, 170, 171, 

173. 388. 289 
Fiber. 85 
Kin whale (Hatifnoftera\ 44, 45, 46, 58, 

100, 101, 112, 336 


o, 54. j). 346. 348 

t, 39. 56, 72, 86, 87, 173, 174, 

253. 374, 330, 332, 366 
GaJictit, 8 1 
CaliJia, So 
l,azf(la. 8 j 
tifHftta, 80 
Iftamyt. 85 
Georytkui* 85 
Gibbon \Hylsbates), 54, 78, 92, no, 160, 


C.iralTe. 41. 83, 96. 258. 2 (8, 332. 35* 
GtoHirfikalMt, 46, 5>, 84. 209,211, 214. 389. 

30*. y>3 
Glypt xlon, 64 
(ionlla (Trvglodylfi}. 37. 53. 54, 67. 78, 

93, 93. no. 160, 161, 162, 251. 273. 286, 

318. 331 

( irampu* (O na\ 46. 84. 302. 103 
Guinea Pig (Cavia), 73. 86. 181. 354, 321 
Gulo, 8 1 
dymnnrn, 73, 86. 174. 289 


Halickarnt. 82 

HmlUort (Dngonc). 43, 57, 84, 101, 228, 

77- 33> 3" 

Halitktrium. 47. 323, 336 
///a/r. 79. 273 
Hafalfmur, 79 

ftit. 85 
Hare i /.</). IT, 39- 5$ 73- W. 96. 178. 

179. 180, 181. i22, 183, 184. 254. *75, 321, 

332, 319 
Hedgehog (Eri*a(em\ 56. 72, 85, 94, 174, 

175, 2:4. 289. 320. 332 
Hetnigalta, 80 
Hrrfiesln, 80 

Hippopoumus. 41. 83. 97, 199* 2 5i 2 7, 

296. 297- 334- 352 
HOIMP (Man). 78, iff Man 
Horse (A><"M)- 4'. 57, 97- "'- l8 S- t8 7- 

1 88, 358, 376, 284, 295, 296, 339, 350 

Howling monkeys (Myceles\ 36. 37, 55, 93. 

160. 161, 163. 164. 165 
Jlytma, 55,68, 80, 172, 287, 320 

Hyimida, no, 274, 346 

Hydrochtrrus (Ofcpybara), 39 73, 86, 96, 

179. 181, 183, 183, 292, 349 
Ilydrontys, 85 
Hydrofiotcs, 83. 298 
Hylnba.tei, 54, 78, 93, 93, no, 160, 161 
Hyoiiiosckus, 298 
Hyfxroodon, 47, 58, 59, 84, 109, 113, 217, 

218. 301, 302. 365 
Uypsiprymnodon, 357 
Hypfipryntnus (Rat Kangaroo), 88, 240, 

309. 337 

Hyrax, 8. 15, 30, 57, 74. 84, in, 201, 202, 

258, 277, 292. 293. 334, 340, 3?3 
Hystrix ( Porcupine). 73, 86, 178, 179, 182, 

'83. 254. 33*i 349 


Icticyon, 80 

Indris, 79 

Inia, 46. 58, 302 

IKSECTIVORA, 9, 11, 30, 31, 38, 55. 56. 72, 

86. 87, 94, 173. 176. 248, 251, 253, 274. 

389, 320, 332, 348 

JERBOA (/?/>), 39, 85. 349 


KANGAROO, 48, 49, 65. 76, 88, 235,240,325, 

3* 6 - 35*. 357. 358, 368 
Koala (/'fiascolarcttu), 48, 65, 76, 77, 88, 

_ii4, 240,337. 358 
Kagia, 113. 219 
Koodoo antelope, 74, 196 


Lagostomits (Viscacha), 86. 180, 182, 349 
Lagotkrix, 36. 79, 165 
Lamb (Ovis), 313 
Lemur, 9. 79, no, 166, 373. 386, 319, 331, 

3*5- 346 
LEMURINA. 37. 54. 68. 72. no, 166, 173, 

287. 345 

LEMUROIUEA, ii. 314 
Leopard, 55. 69, 70. 173 
Ltfiits, 19,39, 56. 73. 86. 96. 178, 184. 254, 

375. 93. 3'4- 3*1, 332, 349 
Lion, 173. 288 
Lizard. 249. 263 
Llama. 41, 197, 299 
Lontkerts, 56 
LopJtiotttfs, 173 
Loris, 54, 79. 319 

380 INDEX. 

Lutra, 81 Mylodon. 87 

Lycaon, 80 My odes, 85 

Lynx (Felis), 72, 173 Myogale (Water Moles), 38, 86, 175, 320, 


Myopolamus. (Coypu), 86, 179, 253, 275, 

M. 332 

Myrmecobws, 89 

Macacus, 78 Myrniecophaga (Anteater), 47, 61, 6?, 63, 

Macraitchenia, 41 87, -102, 103, 113, 229, 230, 259, 305, 336 

Macropidie. 357 Mymtecophagida, 259 

Macrofns (Kangaroo), 48, 49. 65, 76, 88, Mvsi ACOCETI. 43, 99, 219 

239, 240, 309. 325. 326, 339, 356, 357, Myxopoda, 348 

Macrorhinus (Elephant Seal), 82, 284, ,^ T 


Macroscelid* (Elephant Shrew). 56. 173. Narwha | e (Monodon), 46, 84, 301, 302 

M 174 ;/7 4 ' 3 f^ Nasalis, 78 

Man (Homo), 26. 30. 36, 37, 51, 67. 72, 02- Xasita s=; 81 

108, 120, 132, 150, 163, 240, 270, 271, 272, Weotrueu* 1-7 

273,284,317,331,343.368 Nvcteri* s' 

Manatee (,)///), 26, 42, 43, 57- 64, A%c//"j s 

IO2, HI, 223. 224. 228, 278, 323 fj f.-fi 

Mam's (Pangohn or Scaly Anteater), 31, WttflU ins, 79 

47, 61, 64, 75, 76, 88, 102, 232, 259, 304, 

308 O. 
Marmot (Arctoinys), 85. 179, 180, 182, 292 

MAKSUPIAI.IA, 2. 4, n, 23, 29, 31. 48, 65, Octndon. 86 

68, 76, 77, 88, 89, 104, 114, 235,241, 261, ODON rocirn, 46, 98, 107, 112, 209, 255, 

278, 309, 324. 325, 337. 355 3<. 3'- 33 6 

Megaderma. 87. i;8. 332 Opossum (Diaflfhyt) 48, 49, 65, 76, 239, 

Megaptera, 46, 112, 256, 277, 300. 335 309, 324. 326, 355, 358 

Megatherium, 62, 75, 87, 260 Orans; (Simla), 37. 53. 54. 67, 92, 93, no, 

Meles. 55. 81 160, 161, 162. 286. 287, 328. 344 

Jlfcllivpra, 55, 81 Ore a ((irampus), 46, 84. 302, 303 

MellirstlS.Sl OKMTHOUKLI'HIA, 2, 3, II, 50,246, 248, 

Menopoma, 66 249 

Mephitis, 55. 8 1 Ornithorhynchus, 3. 20. 50, 65, 68, 77, 89, 

Mrsp/>l<>i/<y>i. 84. 301 U'4. 10;, 114, 243, 262, 264,311,326, 337, 

Mesothetium, y 338, 340, 358. 359 

Metat/u-ria, -2, 4 Orycteropus (Cnpc Ante.-iter), 47, 64. 68, 

Microgale, 31, 73, 86 76, S8. 102, 114. 233, 258, 308, 324, 336 

Midas, 79 Oryzoryftcs. 289 

Mimofitcrus, 87 Otaria (Eared Seal), 55, i, 82. 255 

Mole (Talpa\ 38, 56, 86, 95. 175. 246, 251, Otariittif, (Eare.1 Seal-), 173, 289, 347 

2 74, 2 75t 289. 290, 314, 320, 348 Oft'cyti, 80 

Molossimr. 332 Ovifus, 82 

Molossus, 87 O~'is (Sheep), 82, 192, 19;, 298, 313, 353 

Monkeys (Siii'iina), 54, 55, 72 93, no. Ox (/>'<>$), 73, 74, 82, in, 195, 298, 353 

159, 160, 161, 162, 163, 164, 165. 273, 287, 


Monodon (Narwliale), 46, 84, 301, 302 

MONOTREMATA, 2, 4, II, 23, 25, 50, 65, PACA, 182, 183 

68, 77, 89, 104, 107. 114, 241, 262, 279, Paltrotherhtm, 215 

310, 314, 32^, 328, 338, 358, 360 J'aiiH' lin (Ji~ finis), 47, Ci, 64, 75, 76. 10.-, 

Norinops, 178 232, 259, 308 

Moschns (Deer), 74, 83 Paradoxurus, 72. 80 

Mns i Rat), 85. 96, 180, 182, 2^2, 349 Pecari, 258. 296, 352 

Mtiscardirtns. ^-5 PECOKA, n, 39, 199, 257, 298, 321, 322, 

Mnstela (U easel), 55, 80, 81, 169 334, 339, 353 

Musteliihr, no, 169, 274, 346 Pt-detes (Cape Jumping Hare), 73, 85, 180. 

Mycetes (Howling Monkey), 36, 37, 55, 79, 182 

93, 160, 161, 163, 164, 105, 273 Ferameles (Randicoot), 48, 49, 89, 262, 

My dans, 81 309, 310, 358 


rtrameliJa, a6a, 309, 337, 358 
PERISSODACTYLA, n, 30, 41, 57, m, 185, 

aoi, 758, 294, 296. 298, 322. 328, 333, 350, 

Ptrodicticia (Potto), 55, 79, 286, 287, 319, 


Petaurus, 88 
Pttrodramus, 774, 348 
Pkottxkftrtts, 83 
1'kala.Hgist* (I'halanger). 48, 76, 88, 309, 

337. 357, 358 
rkascogaU, 89 
J'katcotarclot (Koala), 48, 65, 76, 77. 88, 

"4. 240. 337, 358 
Pk*stolomys (Wombat), 48, 49. 65, 68. 76, 

77, 88, 114, 240, v6j, 267, 278, 309, 325, 

328, 355 
I'tuxa. (Seal), 55, 68, 72, 8j, 173, 255, a<;6, 

274- 284. 289, 319, 328, 331, 343, 346, 365, 

367, 368 

Pkixtrna (Porpoise), 46. 71, 84, 98, 112 
1'kyllorkina, 87, 321 
1'kyllorkinina (l^af-nosed Bat), 348 
i'ltyilaiti'mit&r, 291 
Pkystttr (Cachalot or Sperm Whale). 47, 

59, 60, 84, 98, 99, 113, 217, 218, 219, 255. 

301, 502 

Pkytttrridir, 59, 113. 217 
Pig (Sat), 26, 41, 83, 97, 146, 147, 175, 197, 

199, 244. 2jB, 296, 297, ^22, 3J3, 352 
Pike Whale (ll.ila-H^fttra). too 
I'l.ssiPEUlA, it. Si, 82, 94, no, 255, 274, 


Piptstrtllits, 253 
Pitkecia, 79 
f'latamitta (Sousou), 43, 46, 84, 217, 256, 

263. 302, 324 
Pliityrrki*<r, 163 
Polar Bear ( Urtui), 167 
Pontapcritt, 46, 217 
Porcupine {Hyttrijc). 73, 178, 182. 183, 254, 


Porpoise (Phofcrna), 46, 71, 84, 98. it 2 
J'otami'fa/f. 38. 73, 253. 789, 320 
Potto (PeraJicticHs), 55, 79, 286, 187, 319, 

PKIMATKS. 9, n. 32, 36. 53. 78, 79. 92. 

109, 150. 159. 249, ^7'. ' 8 5. 3' 6 , 33'. 343 
Priodan (Armadillo), 75, 76, 88, 104, 114, 

260. 307, 308 
PKOSOSCIDEA, 8, ti, 42, 203, 223, 2;6, 


Ptvcjftm. 55, 81 

Prtxytntidte, no, 169, 274, 346 
Prongbuck {AntiLKapra), 298 
Prattles. 68, 80 
I'KOTOTHKKIA. 2, 4, 5 
/'sfui/arca, 46, 84 
Pttropidtt, 291 
Pteropus, 68, 73, 87, 96, 176, 177, 253, 230, 


Pudu, 353 
Pittiua, 83 
Puma, 68, 173 


Rabbit (Letus), 254 

Rangi/er (Reindeer). 83, 298 

Rat (Mus), 96, 180, 182, 292, 349 

Red Deer (Cervus), 40, 97, 257, 273, 297, 

334. 352 
Rhinoceros, 41, 57, 83, 84, 98, in, 189, 

190, 191, 201, 276, 295, 296, 333,350,351, 

Rninolopkida, y\ 

Rhizoniys, 85 

Rhyntkocyoit, 56, 73, 94, 173, 274. 289, 

3*9. 348 
Knytina, 42 
RODENTIA, 8, n. 29, 39, 56, 57, 68, 73, 85, 

86. 96, 178. 179, 180. 181, 201, 248, 253, 

275. 293. 321, 328, 332, 340, 342, 348, 349, 

Rorqual ( Rakmoptera), 44, 100,112,220, 

221, 277, 300, 323 

RfMINANTIA, 96, 196, 276, 298 

Saiga. 82. 196 

Saimtrii, 160 

SACKOPMDA, 3. 4, 50 

Sfa/afs. 251 

Scaly Anteater (Manis), 31, 47, 61, 64, 75, 

76, 88, 102, 232. 259, 308 
Sciurus (Squirrel), 85, 179, 180, 292, 332, 


Sciurofterus, 85 

Sea Bear or Sea Lion (Seal). 173, 367 
Seal (P/ioca). 55, 68, 72, 82, 173, 255, 256, 

274. 284, 289, 320, 328, 331, 343, 346, 365, 

367. 68 
Sea Otter ( 

(Enhydra\ 81, 328, 346 
Stmnofithecus, 72, 78 
Sheep (Ovis), 82, 192, 193, 298, 353 
Shrew (Sorex), 38, 39, 56, 94, 95, 175, 176, 

252, 289, 314, 320 
Siamang ( Hylobates), 93, 160 
Simia. (Orang), 37, 53. '54. 67, 78, 92, 93, 

no, 160. 161, 162, 286, 287, 328, 344 
SIMIINA (Monkeys), 72,110,159, 160,161, 

162, 163, 164, 165, 273, 287, 318, 343, 

SIKENIA, 7, 9, n, 23, 42, 57, 69, 75, 84, 

lot, in, 223, 256, 277, 303, 312, 322, 336, 

Sloth,/*' .3, 61, 62, 68, 75, 104. 107, 109, 

113, -,.33, '235, 248, 259, 260, 261, 278, 304, 

324, 328. 336, 354, 366 
Solenodon, 73. 86, 289, 332 
Sorex (Shrew), 38, 39, 56, 86, 94, 95, 175, 

176, 252, 289, 314, 320 
Soricida. 56, 175, 252 
So!al a, 84 
Sousou (Platanisla), 43. 46, 84, 217, 256, 

263, 32. 324 
Spalax, 85 

382 INDEX. 

Spermophilns, 85 Tupaia, 56, 73, 173, 174, 289, 320, 332 

Sperm Whale (I'/iyseter), 47, 59,60, 84, 98, 7'tirsiops, 84 

99, 113, 217, 218, 219, 255, 301, 302 TYLOPODA, n, 41, 197, 294, 298, 334, 353 
Spider Monkey {A teles), 54, 72, 79, 163, 

273, 287 ., 

Squirrel (Scinnts), 85, 179, 180, 292, 332, 

Stotorkynch**, 55, 82 U OU '' AT R A ' 8 '"' 20 > 24 ' 39, 57- 68, 73, 
SuBUNGULATA.292 82 ' 8 3- 84, 96, no. i8 S , iy8, 199, "|. 

SUINA. it, 41, 296, 298, 352 "5. 24-, 257, 276, 29,, 29.,. 32,, 3 <8, 

Suricata.Zo 333-334-349 

Sus (Pig), 26, 41, 83, 97, 146, 147, 175, 197, 77 ,' R " \ l69 ' i S5 ' 27 t 346 , , 

* ~ - ' 7 " '" l6 " l68 ' "" 

}'csf-crngo. 39. 87 
Talpa (Mole), 38,56, 86, 95, 175, 24^, 251, Viscacha (La^ostoinus), 86, 180, 182, 349 

274, 275, 289, 290, 314, 320, 348 ViTerra, 55, 80, 314 

Tamandua, 87, 231, 259 I'ircrritite, no, 171, 172, 274, 356 

Taimas. 85 
Tap.r ( Tapiriis), 41, 57, 83. 97, 98, in, 191, ^. 

225. 258, 276, 295, 333, 350 

7^2::'79?.66, 3.9, 345, 346 * M -F, (T"'?,"?*' ' ***> 3 " 7 ' * 7 

T t r/^ X QQ / " Weastl (Alnstela) t ss, 80, 81, 160 

JlltltSHl, 47, 00, *Oj M'U i > -.- , 

Ta vitieti 81 V h:lle> 43> 44> 45> 4(S> *?' s8 ' 5< >- 6o ' 8 '- 9 R - 

Ttnrec (Ctntetes), 38, 56, 72, 36, .74, 274, 59, >, ^, m. "2, 2,7, 220, 256. 2yQ , 

7V a S^. 3 i ^^ & 3 M^>f <6. 3d. 302 

Theropitkcms, 79 Womh ^ t (I'***'?*), 48. 4 .,, 65. 68, 76. 

Thylacine (Thyladnnt), -48, 49. 77, 89, 77-8 8. 114, 240, 162, 267, 278, 309, 325, 

235, 236, 237, 238, 309, 325, 326, 356 
Thyroptcra. 348 

Tiger, 68, 16;, 170, 171, 173 X. 

7 olypeutes, 88. 308 

Tortoise. 179, 281, 366 .\\-nurtis (Cabassou), 88, y.j 

Tojcodon, 9, 192 

TKAGUI.INA. n, 41, 197, 298, 334, 353 

Tragulus, 83 Z. 

Tree Anteatcr (f'yflotunis\, 103. 3r6, 355 
Trichechus (Walrus), 55, 82, 347, 367 7.apns, 85 

Troglodytes, 37, 53. 54, 67, 78, 92, 93, no, /.iphiimr. 218 

160, 161, 251, 286, 318, 331 Ziphius, 59, 113. 217 


Alhrtcht, 134, 155, 344, 171 
lialfour, 145. 147, j6i 
Uardeicbtn, 340 
Baur, 298, 340, 349 

IJUmvillr, 2 

Boo.*. 296 

Bnndl, 201 

Brooke, Sir V.ctor. 298, 353 

Callcruler, 155 

Coue*, 362 

DotMon, 176. 291 

Iollo, 134 

Doran. 133 

Kschrichl, 335 

Flower, 77, 190, 301 

Kraser. 134 

Ciarrod, ac8 

Gcgenbaur, 133, 245, 371, 281, 2^6, 314, 

. 3*'. 374 
(roctie, 361 
(irubcr, 284 
Humphry, 61 
Huxley, 2, i ji, 133 
KAU, 326 

Koelliker, 133 

Krause, 314 

l.eboucq, 284 

Lcche, 314 

M.icuns, 361 

Milne-Kdwards, 179 

Mivart, 37,280, 286, 361 

Parker, 90, 93, 98, 102, 133, 145, 245 

Peters, 133 

Keichert, 133 

Keinhardt, 335 

kolleston, 361 

Rosenberg, 284 

Huge. 374 

Sabatier, 361 

Salensky, 133 

Strasser, 361 

Struthers, 44, 284, 335 

Thomson. Allen. 283 

Turner, W. , 45, too, 112, 244, 301 

Turner, H. N., 141 

Weyhe, 316 

Wilder. 287 

Wyman, 362 

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