Please see [Transcriber's Notes] at the end of this document.

THE
ARTISTIC ANATOMY OF ANIMALS

PREFACE

A few lines will suffice to explain why we have compiled the present volume, to what wants it responds, and what its sphere of usefulness may possibly embrace.

In our teaching of plastic anatomy, especially at the École des Beaux-Arts—where, for the past nine years, we have had the very great honour of supplementing the teaching of our distinguished master, Mathias Duval, after having been prosector for his course of lectures since 1881—it is our practice to give, as a complement to the study of human anatomy, a certain number of lessons on the anatomy of those animals which artists might be called on to represent.

Now, we were given to understand that the subject treated in our lectures interested our hearers, so much so that we were not surprised to learn that a certain number repeatedly expressed a desire to see these lectures united in book form.

To us this idea was not new; for many years the work in question had been in course of preparation, and we had collected materials for it, with the object of filling up a void of which the existence was to be regretted. But our many engagements prevented us from executing our project as early as we would have wished. It is this work which we publish to-day.

[vi]

Fig. I.—Reproduction of a Sketch by Barye (Collections of the Anatomical Museum of the École des Beaux-Arts—Huguier Museum).

Putting aside for a moment the wish expressed by our hearers, we feel ourselves in duty bound to inquire whether the utility of this publication is self-evident. Let it be clearly understood that we wish to express here our opinion on this subject, while putting aside every personal sentiment of an author.

No one now disputes the value of anatomical studies made in view of carrying out the artistic representation of man. Nevertheless—for we must provide against all contingencies—the conviction on this subject may be more or less absolute; and yet it must possess this character in an intense degree in order that these studies may be profitable, and permit the attainment of the goal which is proposed in undertaking them. It is in this way that we ever strive to train the students whose studies we direct; not only to admit the value of these studies, but to be materially and deeply convinced of the fact without any restriction. Such is the sentiment which we endeavour to create and vigorously encourage. And we may be permitted to add that we have often been successful in this direction.

Therefore it is that, at the beginning of our lectures, and in anticipation of possible objections, we are accustomed to take up the question of the utility of plastic anatomy. And in so doing, it is in order to combat at the outset the idea—as mischievous as it is false—which is sometimes imprudently enunciated, that the possession of scientific knowledge is likely to tarnish the purity and freshness of the impressions received by the artist, and to place shackles on the emotional sincerity of their representation.

[viii]

Fig. II.—Reproduction of a Sketch of Barye (Collections of the Anatomical Museum of the École des Beaux-Arts—Huguier Museum).

It is chiefly by employment of examples that we approach the subject. These strike the imagination of the student more forcibly, and the presentation of models of a certain choice, although rough in execution, is, in our opinion, preferable to considerations of an order possibly more exalted, but of a character less clearly practical. Let us, then, ask the question: Those artists whose eminence nobody would dare to question, did they study anatomy? If the answer be in the affirmative, we surely cannot permit ourselves to believe that we can dispense with a similar course. And, as proof of the studies of this class which the masters have made, we may cite Raphael, Michelangelo, and, above all, Leonardo da Vinci; and, of the moderns, Géricault. And we may more clearly define these proofs by an examination of the reproductions of their anatomical works, chosen from certain of their special writings.[1]

[1] Mathias Duval and A. Bical, ‘L’anatomie des Maîtres.’ Thirty plates reproduced from the originals of Leonardo da Vinci, Michelangelo, Raphael, Géricault, etc., with letterpress and a history of plastic anatomy, Paris, 1890.

The manuscripts of Leonardo da Vinci of the Royal Library, Windsor, ‘Anatomy, Foliæ A.,’ published by Théodore Sabachnikoff, with a French translation, written and annotated by Giovanni Piumati, with an introduction by Mathias Duval. Édouard Rouveyre, publisher, Paris, 1898.

Mathias Duval and Édouard Cuyer, ‘History of Plastic Anatomy: The Masters, their Books, and Anatomical Figures’ (Library of Instruction of the School of Fine Arts), Paris, 1898.

Accordingly, there is no scope for serious discussion, and it only remains for us to enunciate the opinion that it is necessary that we should imitate those masters, and, with a sense of respectful discipline, follow their example.

Here, with regard to the anatomy of animals, we pursue the same method, and the example chosen shall be that of Barye. His talent is too far above all criticism to allow that this example should be refused. The admiration which the works of this great artist elicit is too wide-spread for us to remain uninfluenced by the lessons furnished by his studies. It is sufficient to see the sketches relating to these studies, and his admirable casts from nature which form part of the anatomical museum of the École des Beaux-Arts, to be convinced that the artistic temperament, of which Barye was one of the most brilliant examples, has nothing to lose by its association with researches the precision of which might seem likely to check its complete expansion.

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Fig. III.—Reproduction of a Sketch of Barye (Collections of the Anatomical Museum of the École des Beaux-Arts—Huguier Museum).

In those sketches we find proofs of observation so scrupulous that we cannot restrain our admiration for the man whose ardent imagination was voluntarily subjected to the toil of study so profound.

If the example of Barye, with whom we associate the names of other great modern painters of animals, can determine the conviction which we seek to produce, we shall be sincerely glad. To contribute to the propagation of useful ideas, and to see them accepted, gives a feeling of satisfaction far too legitimate for us to hesitate to say what we should feel if our hope be realized in this instance.

ÉDOUARD CUYER.

[xii]

Fig. IV.—Reproduction of a Sketch of Barye (Collections of Anatomical Museum of the School of Fine Arts—Huguier Museum).

[Detailed
Table of
Contents.]

CONTENTS

LIST OF ILLUSTRATIONS

THE ARTISTIC ANATOMY OF ANIMALS

INTRODUCTION

GENERALITIES OF COMPARATIVE ANATOMY

Of the animals by which we are surrounded, there are some which, occupying a place in our lives by reason of their natural endowments, are frequently represented in the works of artists—either as accompanying man in his work or in his amusements, or as intended to occupy the whole interest of the composition.

The necessity of knowing, from an artistic point of view, the structure of the human body makes clear the importance we attach, from the same point of view, to the study of the anatomy of animals—that is, the study of comparative anatomy. The name employed to designate this branch of anatomy shows that the object of this science is the study of the relative position and form which each region presents in all organized beings, taking for comparison the corresponding regions in man. The head in animals compared with the human head; the trunk and limbs compared to the trunk and limbs of the human being—this is the analysis we undertake, and the plan of the subject we are about to commence.

Our intention being, as we have just said, the comparison of the structure of animals with that of man, should we describe the anatomy of the human being in the pages which follow? We do not think so. Plastic human anatomy having been previously studied in special works,[2] we take it for granted that these have been studied before undertaking the subject of comparative anatomy. We will therefore not occupy time with the elementary facts relative to the skeleton and the superficial layer of muscles. We will not dilate on the division of the bones into long, short, large, single, paired, etc. All these preliminary elements we shall suppose to have been already studied.

This being granted, it is, nevertheless, necessary to take a rapid bird’s-eye view of organized beings, and to recall the terms used in their classification.

Animals are primarily classed in great divisions, based on the general characters which differentiate them most. These divisions, or branches, allow of their being so grouped that in each of them we find united the individuals whose general structure is uniform; and under the name of vertebrates are included man and the animals with which our studies will be occupied. The vertebrates, as the name indicates, are recognised by the presence of an interior skeleton formed by a central axis, the vertebral column, round which the other parts of the skeleton are arranged.

The vertebrate branch is divided into classes: fishes, amphibians or batrachians, reptiles, birds, and mammals.

The mammals—from the Latin mamma, a breast—are characterized by the presence of breasts designed for the alimentation of their young. Their bodies are covered with hair, hence the name pilifères proposed by Blainville; and, notwithstanding that in some individuals the hairs are few, the character is sufficient to distinguish them from all other vertebrates.

We find united in this class animals which, at first, seem out of place, such as the whale and the bat; and, from their external appearance alone, the former would appear to belong to the fishes, and the latter to birds. Yet, on studying their structure, we find that, not only do these animals merit a place in the class which they occupy, because they possess the distinctive characters of mammals; but, still further, their internal structure is analogous to that of man and of the other individuals of this class.

Notwithstanding this similarity of structure, the whale is not without some points of difference from its neighbours the horse and the dog; therefore, in order to place each of these animals in a position suitable to it, mammals are divided into secondary groups called orders. The first of these orders includes, under the name primates, man and apes. The latter contain animals which approach birds in certain characters of their organism, forming a link between the latter and mammals.

We find, in studying the regions of the body in some of the vertebrates, that, while they present differences from the corresponding regions of the human body, they also offer most striking analogies. We can, for example, recognise the upper limb of man in the anterior one of quadrupeds, in the wing of the bat, in the paddle of the seal, etc. It is, so to speak, those variations of a great plan which give such a charm to the study of comparative anatomy.

The division of classes into orders, which we have just mentioned, being still too general, it was found necessary to establish subdivisions—more and more specialized—to which the names families, genera, species, and varieties were given.

[2] Mathias Duval, ‘Précis of Anatomy for the Use of Artists’: Paris, 1881. ‘Artistic Anatomy of the Human Body,’ third edition, plates by Dr. Fau, text with figures by Édouard Cuyer: Paris, 1896. ‘Artistic Anatomy of Man,’ by J. C. L. Sparkes, second edition, text with 50 plates: Baillière, Tindall and Cox, London, 1900.

CHAPTER I

OSTEOLOGY AND ARTHROLOGY

THE TRUNK

The Vertebral Column

We commence the study of the skeleton with a description of the trunk.

Fig. 1.—A Human Skeleton in the Attitude of a Quadruped. To give a general Idea of the position of the Bones in other Vertebrates.

The trunk being, in quadrupeds, horizontal in direction ([Fig. 1]), the two regions of which it consists occupy, for this reason, the following positions: the thorax occupies the anterior part, the abdomen is placed behind it; the vertebral column is horizontal, and is situated at the superior aspect of the trunk; it projects beyond the latter: anteriorly, to articulate with the skull; and, posteriorly, to form the skeleton of the tail, or caudal appendix.

The number of the vertebræ is not the same in all mammalia. Of the several regions of the vertebral column, the cervical shows the greatest uniformity in the number of the vertebræ of which it consists, with but two exceptions (eight or nine in the three-toed sloth, and six in the manatee); we always find seven cervical vertebræ, whatever the length of the neck of the animal. There are no more than seven vertebræ in the long neck of the giraffe, but they are very long ones; and not less than seven in the very short neck of the dolphin, in which they are reduced to mere plates of bone not thicker than sheets of cardboard. If the cervical region presents uniformity in the number of its bones, it is not so with the other regions of the column.

The following table shows their classification in some animals:

Vertebræ.

It is worthy of notice that in birds the number of the cervical vertebræ is not constant, as in mammals; they are more numerous than the dorsal. These latter are almost always joined to one another by a fusion of their spinous processes; the two or three last vertebræ are similarly united to the iliac bones, between which they are fixed. The dorsal vertebræ thus form one piece, which gives solidity to the trunk, and provides a base of support to the wings, for the movements of flying. There are, so to speak, no lumbar vertebræ, the bones of that region, which cannot be differentiated from the sacrum, having coalesced with the bones of the pelvis.

Vertebræ.

In reptiles, the relation between the number of the cervical vertebræ and that of the dorsal is very variable; some serpents are devoid of cervical vertebræ, having only dorsal ones—that is, vertebræ carrying well-developed ribs.

Vertebræ.

Fig. 2.—Size of the Atlas compared with the Transverse Dimensions of the Corresponding Parts of the Skull in Man.

1, Atlas; 2, mastoid process; 3, external occipital protuberance; 4, inferior maxilla.

Regarding the direction of the vertebral column in animals, in which the trunk is not vertical, it is evident that the spinous processes point upward, and that in comparing them with those of man they must be arranged so that the superior surface of the human vertebra will correspond to the anterior surface of that of the quadruped. Of the cervical vertebræ, the atlas and axis call for special notice. Apropos of the atlas, we find that it, in the human being, is narrower than the corresponding parts of the skull, and is therefore hidden under the base of the cranium ([Fig. 2]); in quadrupeds its width is equal to that of the skull, and sometimes exceeds, because of the great development of its wing-shaped transverse processes, that of the neighbouring parts of the head ([Fig. 3]). On this account those transverse processes often project under the skin of the lateral surfaces of the upper part of the neck.

Fig. 3.—Size of the Atlas compared with the Transverse Dimensions of the corresponding Regions of the Skull in a Dog.

1, Atlas; 2, zygomatic arch; 3, external occipital protuberance; 4, inferior maxilla.

The axis is furnished on its anterior surface with the odontoid process, which articulates with the anterior (or inferior) arch of the atlas, according to the direction of the neck. The spinous process, flattened from without inwards, is more or less pointed; it is elongated from before backwards, so as partly to overlap the atlas and the third cervical vertebra.

We find that this process overlaps less and less the neighbouring vertebræ when we examine in succession the bear, the cat, the dog, the ox, and the horse. With regard to the other vertebræ of this region, they diminish in width from the second to the seventh; and, in some animals, the anterior surface of the body presents a tubercle which articulates with a cavity hollowed in the posterior surface of that of the vertebra before it; this feature dwindles away in the dorsal and lumbar regions.

The spinous process, slightly developed in the third cervical vertebra, gradually increases in size to the seventh, the spinous process of which, long and pointed, well deserves the name of the prominent which is bestowed on it; but it should not be forgotten that the spinous process of the axis is equally developed.

On the inferior surface of the body of each of the vertebræ is found a prominent crest, especially well marked at the posterior part; this crest is but slightly developed in the bear and in the cat tribe, and is not found in swine.

The transverse processes of the cervical vertebræ, from their relation to the trachea, are known as the tracheal processes.

The most marked characteristic of the dorsal vertebræ is furnished by the spinous processes. They are long and narrow. As a rule, the spinous processes of the foremost dorsal vertebræ are the most developed and are directed obliquely upwards and backwards. As we approach the last vertebræ of this region, the processes become shorter and tend to become vertical, and the last ones are even, in some cases, directed upwards and forwards; this disposition is well marked in the dog and the cat. In the cetaceans, on the contrary, the length of the spinous processes increases from the first to the last.

In the horse the spinous processes of the first dorsal vertebræ produce the prominence at the anterior limit of the trunk, where the mane ends, which is known as the withers.

Fig. 4.—Lumbar Vertebræ of a Quadruped (the Horse): Superior Surface.

1, Spinous process; 2, anterior articular process and transverse process of the first lumbar vertebra of the left side; 3, costiform process.

The lumbar vertebræ are thicker than the preceding; they are known by their short and latterly-flattened spinous processes, and still more readily by their transverse processes, which, as they are evidently atrophied ribs, it is more accurate to denominate costiform processes ([Fig. 4]). These are long, flattened from above downwards, and directed outwards and forwards.

The true transverse processes are represented by tubercles situated on the superior borders of the articular processes of each of the vertebræ of the lumbar region. Apropos of these different osseous processes, we are reminded that they are also present in the human skeleton.

In the horse the costiform processes of the fifth and sixth lumbar vertebræ articulate, and are sometimes ankylosed, one with the other; the terminal ones articulate with the base of the sacrum. Sometimes the processes of the fourth and fifth are thus related; this is the case in the figure ([4]) given; here the costiform processes of the fourth and fifth vertebræ articulate, and the two terminal ones have coalesced.

In the ox, the same processes are more developed than in the horse; their summits elevating the skin, produce, especially in animals which have not much flesh, prominences which limit the flanks in the superior aspect. The costiform processes of the last lumbar vertebræ are separate from each other; those of the latter are not in contact with the sacrum.

The Sacrum.[3]—This bone, single and median, is formed by the mutual coalescence of several vertebræ, which vary in number according to the species observed.

[3] In human anatomy, the sacrum and the coccyx are studied as part of the pelvis; we, therefore, in the study of the artistic anatomy of man, study these bones with the bones of the lower limbs. Here we do not follow this plan. In animals the sacrum and the coccyx, as a matter of fact, clearly continue the superior border of the skeleton of the trunk; hence we study them with the vertebral column.

Vertebræ Constituting the Sacrum.—Bears, 5; dogs, 3; cats, 3; rabbits, 4; swine, 4; horses, 5; camels, 4; oxen, 5; sheep, 4.

The sacrum is situated between the two iliac bones; with which it articulates, and contributes to the formation of the pelvis. It is obliquely placed, from before backwards, and from below upwards; immediately behind the lumbar section of the vertebral column; and is continued by the coccygeal vertebræ, which form the skeleton of the tail.

It is triangular in outline, and is generally more narrow in proportion than in the human being. All things considered, it is more large and massive, and of greater density, in species which sometimes assume the upright posture, rather than in those which cannot assume that attitude; for example, the sacrum of the ape, of the bear, of the dog, and of the opossum are proportionately larger than those of the horse.[4]

[4] This is particularly striking only in those portions of the sacrum that are not in relation with the other bones of the pelvis. We think that the general form of this bone depends on the mode of its connexions with the iliac bones and the extent of the articular surfaces by which it is in contact with the latter.

Its superior surface presents a crest, formed by the fusion of the spinous processes of the vertebræ which form it. In certain species these processes are attached only by their bases, and are separated from each other superiorly. In the pig they are wholly wanting.

The Coccygeal Vertebræ.—These vertebræ, few in number (and sometimes ankylosed) in the human being, form in the latter a small series, the coccyx; which is inclined forwards, that is to say, towards the interior of the pelvis. In quadrupeds, on the contrary, their number is large; they are not ankylosed, and they form the skeleton of the caudal appendix.

The first coccygeal vertebræ—that is, those which are next the sacrum—present characters which are common to those of other regions: they have a body, a foramen, and processes. As we trace them backwards, these characters become gradually effaced; and they become little more than small osseous cylinders simply expanded at their extremities.

Direction and Form of the Spinal Column

The curves of the vertebral column are, in quadrupeds, slightly different from those which characterize the human spine. First, instead of their being, as in the latter, curves in the antero-posterior aspect, because of the general attitude of the body, they are turned in the supero-inferior direction.

The cervical region is not a single curve, as in the human being. It presents two: one superior, with its convexity looking upwards; the other inferior, the convexity of which is turned downwards. This arrangement reminds one of that of a console.

The dorsal and lumbar regions are placed in a single curved line, more or less concave downwards; so that in the lumbar region there is no curve analogous to that which exists in man; a form which, in the latter, is due to the biped attitude—that is to say, the vertical position of the trunk. Briefly, there is in quadrupeds one dorso-lumbar curve; and not both a dorsal and a lumbar, with convexities in opposite directions.

At the extremity of the dorso-lumbar region is the sacrum and the caudal appendix, which describe a curve of which the concavity is directed downwards and forwards.

It is necessary to point out that it is not the curves of the three anterior portions of the spinal column which determine the form of the superior border of the neck and shoulders, and of the same part of the trunk. For the first portion, there is a ligament which surmounts the cervical region, and substitutes its modelling influence for that of the vertebræ. It is the superior cervical ligament, which arises from the spinous process of the first cervical vertebræ, and is inserted into the external occipital protuberance on the upper part of the posterior surface of the skull. The summits of the spinous processes of the vertebræ alone give form to the superior median border of the trunk. In this connection we here repeat that it is not the general curvature of the vertebral column which produces the withers, but the great length of the spinous process of the first vertebræ of the dorsal region.

The Thorax

The dorsal vertebræ form the posterior limit in man, and superior in quadrupeds, of the region of the trunk known as the thorax. A single bone, the sternum, is situated at the aspect opposite; the ribs bound the thorax on its sides.

In its general outlines the thorax in quadrupeds resembles that of man—that is to say, that, as in the latter, the anterior portion—superior in the human being—is narrower than the part opposite. But the progressive widening takes place in a more regular and continuous fashion, so that it presents a more definitely conical outline. This purely conical form is nevertheless found in the human species, but only during infancy; the inferior portion of the thoracic cage being then widely expanded, because of the development of the abdominal viscera, which at that period are relatively large.

Fig. 5.—A Transverse Section of the Thorax of a Man placed Vertically—that is to say, in the Direction which it would assume in a Man placed in the Attitude of a Quadruped (a Diagrammatic Figure).

1, Dorsal vertebra; 2, sternal region; 3, costal region of one side; 3′, costal region of the other side.

But the proportionate measurements of the thorax are different. Indeed, we may recall that in man the thorax is flattened from before backwards, so that the distance between the sternum and the vertebral column is shorter than the distance from the rib of one side to the corresponding one of the opposite side ([Fig. 5]). In animals, on the contrary, it is flattened laterally. Its vertical diameter—measured from the sternum to the vertebral column—is greater than the transverse measurement ([Fig. 6]).

Fig. 6.—A Vertical Section of the Thorax of a Quadruped (Diagrammatic).

1, Fifth dorsal vertebra; 2, sternal region; 3, costal region of one side; 3′, costal region of the opposite side.

From this results a peculiar arrangement of the muscles that we are able to bring directly into prominence, which presents points of interest from the point of view of the contraction of the subcutaneous layer. Indeed, in man the region occupied by the pectorals is very broad; it is a wide surface turned directly forward. In quadrupeds, this region of the pectorals is narrowed. It is not spread out, as in the preceding instances; and the appearance it presents is explained by the fact that the thorax is compressed laterally. If we examine the thorax on one of its lateral surfaces, the muscles, on the contrary, are more extended. We see the contour of the vertebral column, and the median part of the abdomen; and, especially in the horse, between the great dorsal and the great oblique of the abdomen, we find a large space, in which the ribs, with the intercostals which join them, are uncovered; the muscles in question separate the one from the other, under the influence, it would seem, of the great dimensions of the lateral wall of the thorax.

The Sternum.—The sternum is, in quadrupeds, directed obliquely downwards and backwards; its form varies in different species. In the carnivora, it consists of eight bones, irregularly cylindrical in form, being slightly flattened from within outwards, and thickened at their extremities. They remain separate, and this contributes elasticity and flexibility to the thorax. The first nine costal cartilages articulate directly with the sternum. The first of these cartilages articulates with a nodule situated a little above the middle of the first bone of the sternum.

In the horse the sternum is flattened laterally in its anterior portion, and from above downwards in its posterior half. The six bones which form the sternum are connected by cartilage. The keel-shaped piece, situated in front of the sternum, is also cartilaginous. This process, but slightly marked posteriorly, becomes more and more prominent in front, and terminates at its anterior extremity by a prolongation, slightly curved backwards, which projects for some centimetres beyond the cavity in which the first costal cartilage is received. This process is known as the tracheal process, or rostral cartilage. The posterior extremity of the sternum, flattened from above downwards, ends in a cartilaginous plate; concave superiorly, and convex inferiorly: this is the abdominal prolongation, or xiphoid appendix.

In the ox, the sternum is formed of two distinct bones, which are united by an articulation. One, the anterior, is short, and forms the first portion of the sternum; it is slightly flattened from side to side, and vertical in direction. The other, the posterior, is longer, and is formed by the fusion of several small bones; it is placed horizontally, and is flattened from above downwards. At the level of articulation of these two portions, and because of their different directions, the bone is bent. This bend occurs at the point of articulation of the second costal cartilage. On the superior border of the anterior segment the cartilage of the first rib is articulated. The xiphoid appendix, which is cartilaginous, is attached to the extremity of a long process of the last bone of the sternum.

The shape of the anterior extremity of the sternum is influenced by the presence or absence of clavicles. We have seen that in some quadrupeds the clavicles are wanting. In the first case, this extremity is large, and approaches in shape to the corresponding part of the human sternum, which is so clearly designed to give a point of support to the anterior bone of the shoulder. In the second, on the contrary, this extremity is narrow.

The sternum in birds is very different from that in mammalia, which we have been studying. It varies greatly in extent and shape, under the influence of certain conditions. To understand the cause of these variations it is necessary to remember that in man (as, indeed, in other animals; but the example of man, for that which follows, will be more striking, on account of the mobility of his upper limbs) the sternum gives origin to the pectoral muscles, and that these muscles are inserted into other parts of the thoracic limbs, designed by their contraction to draw the arms downwards, forwards, and inwards—that is, when these are in a state of abduction and in a horizontal direction, they draw them towards the anterior surface of the thorax and downwards. Now, this movement is similar to that made by birds during flight. It is necessary to add that, in the latter case, the more the displacement of the upper limbs has of force and extent, the more the pectoral muscles are developed.

For these reasons, birds, in which, during flight, the movements of the thoracic limbs—the wings—are necessarily energetic, present a great development of the pectoral muscles; having consequently, because an extent of surface for the origin of the muscles commensurate with their development is necessary, a very large and peculiarly shaped sternum ([Figs. 18], 6; and [21], 6). Indeed, not only is the sternum large, but, further, in order to form a deeper surface, proportionately adapted to the muscles which arise from and cover it, its anterior surface presents, in the median line, a prominent crest known as the keel. This prominence forms two lateral fossæ. We cite as examples, the sternum of the eagle, the vulture, the falcon, and the hawk.

Fig. 7.—Sternum of a Bird (the Cock): Left Side, External Surface.

1, Keel; 2, internal slot; 3, external slot; 4, internal process; 5, external process; 6, inferior ribs; 7, costal process; 8, surface for articulation with the coracoid bone.

All birds are not, however, equally adapted to flight, for in the domestic cock, which flies but a short distance, and badly, the sternum is less developed ([Fig. 7]); it is also diminished by slots, which diminish its surface. These slots, two on each side, are called from their position the internal and external slots. They are bounded by narrow, elongated, bony processes, an internal and an external; the expanded lower extremity of the latter overlaps the last inferior ribs (see [p. 19]). The part of the external border which surmounts this external process is hollowed out into grooves, which receive the inferior ribs, and terminates superiorly in an osseous projection known as the costal prominence.

In the ostrich, the cassowary, and the apteryx, which run, but do not fly, the sternum has the form of a plate of bone slightly convex, but without a keel.

The shape of the sternum, correlated to the faculty of flight (or of swimming; apropos of which we may cite the penguin, of which the rudimentary wings resemble fins, and perform their functions only), or the absence of this faculty, has furnished the division of birds into two groups. In one are included, under the name Carinates (carina, keel), those in which the sternum is provided with a keel; in the other division are those in which the sternum is not furnished with one. These latter, on account of their unique mode of progression, are more nearly allied to the mammals.

The keel is developed in flying mammals (bats).

Ribs and Costal Cartilages.—There are on each side of the thorax as many ribs as there are dorsal vertebræ. In animals, as in man, the ribs which articulate with the sternum by their cartilages are called true, or sternal ribs; those whose cartilages do not articulate with the sternum are called false, or asternal. The longer ribs are those situated in the middle region of the thorax.

The ribs are directed obliquely downwards and backwards, and this obliquity is more marked in the posterior ones than in the anterior. They are, however, less oblique than in the human being; what proves this is that the first rib in man is oblique, while in quadrupeds it is vertical.

The curvature of the ribs is less pronounced in quadrupeds than in the human being, but this is not equal in all animals. The ribs of the bear are more curved than those of the dog; the latter has ribs more curved than those of the horse.

Each rib, at its vertebral extremity, presents, from within outwards, a wedge-shaped head for articulation with two dorsal vertebræ, a neck, and a tuberosity. External to the tubercle are found some rough impressions, for muscular attachments, which correspond to the angle of the human rib.

In the following table, we give the number and classification of the ribs of some animals:

Number of the Ribs on Each Side of the Thorax.

The costal cartilages, by which the first ribs are united to the sternum (sternal ribs), whilst the latter are united one to the other without being directly connected with the sternum (asternal ribs), are, as a rule, in quadrupeds, directed obliquely downwards, forwards, and inwards; each forms, with the rib to which it belongs, an obtuse angle more or less open anteriorly. Their length is proportionate to that of the ribs. The cartilages, which are continued from the asternal ribs, unite and form the borders, directed obliquely downwards and forwards, of the fossa which is found at the inferior and posterior part of the thorax, and which forms the lateral limits of the epigastric region. In the dog and cat the ribs are thick and almost cylindrical; the costal cartilages are thicker at the margin of the sternum than at their costal extremity. In the ox, the ribs are flattened laterally and are very broad, the more so as we examine a portion further from the vertebral column. From the second to the twelfth they are quadrangular in the superior fourth, and thicker than in the rest of their extent. The first costal cartilage is vertical; the following ones are progressively more oblique in a direction downwards and forwards. The four or five cartilages which succeed the first unite with slight obliquity to the sternum; their union with that bone gives the impression of a very strong, well-knit apparatus. The costal cartilages which unite with the sternum are flattened laterally in the portions next the ribs, and flattened from front to back in the rest of their extent.

In the horse the ribs increase in length from the first to the ninth; they are flattened from without inwards, and increase in width from the first to the sixth or seventh, and the following ones become narrower. The costal cartilages, from the second to the eighth, are, as in the ox, at first flattened laterally, near the ribs; while near the sternum they are flattened from front to back.

In birds, the ribs are each furnished with a flat process ([Fig. 18], 10), which springs from the posterior border, is directed backwards, and overlaps the external surface of the succeeding rib. These processes are not found, as a rule, on the first or last ribs.

As for the costal cartilages, they are, as a rule, ossified, and receive the name of inferior ribs ([Fig. 18], 11), united to the preceding (superior ribs; [Fig. 18], 9) by articulation; by the other extremity they are joined to the sternum; the first superior ribs generally want them. Sometimes the last inferior rib becomes connected with the one that precedes it, not articulating with the sternum; and thus recalls the relations of the asternal ribs which we have noticed in our study of the mammals.

In the bat, as in birds, the costal cartilages are ossified.

THE ANTERIOR LIMBS[5]

[5] Consult [Figs. 21], [33], [34], [38], [39], [46].

The anterior limbs, homologous to the upper limbs in man, are formed, as in the latter, of four segments: the shoulder, the arm, the forearm, and the hand. These limbs, considered in the vertebral series, present themselves under very different aspects, which are determined by the functions they are called upon to perform.

Fig. 8.—Anterior Limb of the Bat: Left Side, Anterior Surface.

1, Clavicle; 2, scapula; 3, humerus; 4, radius; 5, cubitus; 6, carpus; 7, thumb; 8, metacarpus; 9, phalanges.

They constitute the forepaw in terrestrial mammals; in aerial vertebrates they form wings; in aqueous mammals they act as paddles. In whatever series we study them, we can readily find the relationship of the different parts; it is very easy to recognise the same bones in the upper limbs of the human being, the wings of the bat ([Fig. 8]) and of birds ([Fig. 21]), and in the anterior paddles of the seal ([Fig. 9]) and of the dolphin.

Fig. 9.—Anterior Limb of the Seal: Left Side, External Surface.

1, Scapula; 2, humerus; 3, radius; 4, ulna; 5, carpus; 6, metacarpus; 7, phalanges of the fingers.

In quadrupeds, the shoulder and arm are hidden, the latter more or less completely, in the muscular mass which binds it to the lateral wall of the trunk; so that the anterior limbs only present; free from the trunk: the elbow, forearm, and hand.

The Shoulder

In some vertebrates, the shoulder is formed of two bones—the scapula and clavicle; in others of only one bone—the scapula; the clavicle in this case does not exist.

The Scapula or Omoplate.—The scapula is situated on the lateral surface of the thorax, and is directed obliquely, from above downwards and from behind forwards.

We must first recall, so as to be able to make a comparison, that in man this bone is placed at the posterior surface of the thoracic cage; so that if we look at the human thorax on one of its lateral aspects we see chiefly the external border of the scapula; it is the external surface (homologous to the posterior surface of the human scapula) which we see in its full extent when we look on the same surface of the thorax in quadrupeds.

Fig. 10.—Situation and Direction of the Scapula in the Human Being, the Trunk being Horizontal, as in Quadrupeds. Vertical and Transverse Section of the Thorax (Diagrammatic Figure).

1, Contour of the thorax; 2, 2, the scapula.

Fig. 11.—Position and Direction of the Scapula in Quadrupeds. Vertical and Transverse Section of the Thorax (Diagrammatic Figure).

1, Contour of the thorax; 2, 2, the scapula.

To sum up, if we fancy the human being in the position of the quadruped, the scapula will have its surfaces almost parallel to the ground ([Fig. 10]); while in quadrupeds, the surfaces are situated in a plane which is almost perpendicular to the ground ([Fig. 11]). This position of the scapula in an almost vertical plane is designed to give the necessary point of support to the osseous columns that form the skeleton of the other portions of the anterior limbs.

Fig. 12.—Left Scapula of the Human Being, Posterior Surface, placed in the Position which it would Occupy in the Skeleton of a Quadruped.

1, Cervical border; 2, spinal border—the scapula here represented, being from a hoofed animal, has a cartilage of extension attached to its spinal border; 3, axillary border; 4, supraspinous fossa; 5, subspinous fossa; 6, spine of the scapula; 7, glenoid cavity; 8, coracoid process. The scapula of the horse has no acromion process, but it is easy, if we compare the human scapula, to judge of the position which this process would occupy if it were present.

Because of this position of the scapula ([Figs. 12] and [13]), the spinal border is superior, the cervical, anterior, and the axillary, posterior. In direct contrast to what obtains in the human scapula, the spinal border is the shortest of the three; except in the bat, and the majority of the cetaceans.

Fig. 13.—Left Scapula of a Horse: External Surface.

1, Cervical border; 2, spinal border; 3, axillary border; 4, supraspinous fossa; 5, subspinous fossa; 6, scapular spine; 7, glenoid cavity; 8, coracoid process; 9, acromion process.

In certain animals (in the ungulates [hoofed[6]]—pigs, oxen, sheep, horses) the superior, or spinal, border of the scapula is surmounted by a cartilage called the cartilage of prolongation.

[6] For the definition of the word hoofed, see [p. 37].

Fig. 14.—Vertical and Transverse Section, at the Site of the Shoulders, of the Thorax of the Horse (Diagrammatic Figure).

1, Outline of the thorax at the level of the third dorsal vertebra; 2, 2, scapula; 3, spinal border of the scapula; 4, cartilage of prolongation; 5, contour of the skin.

This is the cause why the border to which it is fixed is so slightly noticeable under the skin in these animals; indeed, in the upper part, the bone and cartilage are not distinguishable in the contour of the corresponding region of the back; being applied to the lateral surfaces of the spinous processes, the prominence formed by the extremities of which is directly continuous with the plane of the scapula ([Fig. 16]).

Fig. 15.—Vertical and Transverse Section, at the Plane of the Shoulders, of the Thorax of a Dog (Diagrammatic Figure).

1, Outline of the thorax at the level of the third dorsal vertebra; 2, 2, scapula; 3, spinal border of the scapula; 4, contour of the skin.

In quadrupeds whose scapula, on the contrary, is wanting in the cartilage of prolongation (in the clawed,[7] such as the cat and dog), the superior border of the scapula is visible, especially when the animal is resting on its fore-limbs, particularly when it crouches; at such a time the skin is markedly raised by that border; and the spinous processes of the vertebræ, beyond which it projects, occupy the bottom of a fossa ([Fig. 15]). The internal surface of the scapula is turned towards the ribs; it is known, as in man (in whom this surface is anterior), as the subscapular fossa.

[7] For the definition of this word, see [p. 37].

Fig. 16.—Left Clavicle of the Cat: Superior Surface (Natural Size).

1, Internal extremity; 2, external extremity.

Fig. 17.—Clavicle of the Dog (Natural Size).

Its external surface is divided into two parts by the spine of the scapula; which, in some animals, terminates inferiorly in a flat and clearly distinct process, the homologue of the acromion process of the human scapula. The two regions separated by the spine are known as the supraspinous fossa and the infraspinous fossa. The supraspinous fossa is anterior to the spine, and the infraspinous is posterior to it. The surfaces of the scapula are, in quadrupeds, flatter than in the human being, and in particular the subscapular fossa, which is also less concave. Some authors attribute this to the lesser curvature of the ribs in quadrupeds. A few words will suffice to prove that there must be another reason. The scapula is not in immediate contact with the ribs; the subscapular fossa is not moulded on them. Besides, the form of the scapula is, as in other parts of the skeleton, dependent on the disposition of muscles, and the development of these latter is correlated to the extent and energy of the movements which the individual is able or required to execute. But the movements which those muscles produce (more especially the rotation of the humerus) are, in quadrupeds, less extensive than in the human being; and, consequently, the muscles which produce them are, proportionally, less strongly developed. The inferior angle (superior and external in man), situated at the junction of the cervical and axillary borders, presents the glenoid cavity, which, looking downwards, receives the articular surface of the superior extremity of the bone of the arm—that is to say, the head of the humerus. Above this cavity, on the lower part of the cervical border, is situated a tubercle which reminds us of the coracoid process of the human scapula. The region occupied by the glenoid cavity is separated from the body of the bone by a constriction—the neck of the scapula.

Fig. 18.—Skeleton of the Shoulder of a Bird (Vulture): Antero-External View of the Left Side.

1, Left clavicle; 2, inferior portion of the right clavicle, forming by its ankylosis with that of the other side the fourchette; 3, coracoid bone; 4, scapula; 5, articular surface for humerus; 6, superior half of the sternum; 7, keel of sternum; 8, spinous process of the dorsal vertebræ; 9, superior ribs; 10, process of one of these ribs; 11, inferior ribs.

In birds the scapula is elongated in a direction parallel to the vertebral column, and very narrow in the opposite ([Fig. 18]): it is also flat, and has no spine. Its coracoid process is represented by a peculiar bone—the coracoidean or coracoid bone—which we shall describe later on when we come to the study of the clavicle and of the anterior region of the shoulder (see [p. 26]).

The Clavicle.—The clavicle is found only in the human being, and in animals whose anterior limbs, possessing great freedom of movement in all directions, require that the scapula should possess a point of support which, while affording this, can be displaced with it, or draw it in certain directions. Now, this point of support is furnished by the clavicle.

In animals possessed of hoofs (ungulates), such as the sheep, ox, and horse, the clavicle does not exist. Indeed, in them the freedom of movement of the anterior limbs is limited; they move by projection in the forward and backward directions only; they merely fulfil the functions of giving support to and carrying about the body. The clavicle is rudimentary in the cat and the dog; in the cat it is a small, elongated bone ([Fig. 16]), 2 centimetres in length, thin and curved, connected with the sternum and the scapula by ligamentous bundles. In the dog it is represented by a small osseous plate only ([Fig. 17]), which is not connected with any of the neighbouring bones.

It is on the deep surface of a muscle which passes from the head and neck to the humerus (mastoido-humeral, a muscle common to the arm, neck, and head) in which this rudimentary bone is found to be developed.

The clavicle exists in perfect state in mammals which use their limbs for digging, grasping, or flying; the insectivora (hedgehog, mole) and some rodents (squirrel, woodchuck) are provided with it.

The cheiroptera (bats) possess an extremely well-developed clavicle, on account of the varied movements which their thoracic limbs execute.

This formation of the shoulder which favours flight in the bat is even more remarkable in birds. In these latter ([Fig. 18]) the clavicles, fused together by their lower extremities, form one bone, having the shape of the letter V or U, which is known as the fourchette; this bone, acting as a true spring, keeps the shoulders apart, and prevents their approximation during the energetic movements which flight necessitates.

In birds whose power of flight is strong, the two limbs of this bone are widely separated and thick, and the fourchette is U-shaped. Those whose flight is awkward and but slightly energetic have the limbs of the fourchette slender; they unite at a more acute angle, and the bone is V shaped.

Furthermore, a bone named the coracoid joins the scapula to the sternum; this bone, often fused with the scapula, where it contributes to the formation of the glenoid cavity, represents in birds the coracoid process of the human scapula. If we fancy this process directed inwards, and sufficiently lengthened to join the sternum, we shall have an idea of the disposition of the bone we are now discussing, and the reasons for which the name has been chosen by which it is designated. The coracoid bone, like the fourchette which it reinforces, offers to the wings a degree of support proportionate to the efforts developed by those limbs; for this reason it is thick and solid in birds of powerful flight.

The superior extremity of each branch of the fourchette, at the level of its junction with the coracoid and the scapula, bounds, with these latter, a foramen which gives passage to the tendon of the elevator muscle of the wing, or small pectoral. The importance of the fourchette being, as we have seen, in proportion to the movements of flying, it is easy to understand that the bone is not found in the ostrich.

The Arm

A single bone, the humerus, forms the skeleton of this portion of the thoracic limb.

The Humerus.—The bone of the arm is, in quadrupeds, inclined from above downwards and from before backwards.

It is, with relation to other regions, short in proportion as the metacarpus is elongated, and as the number of digits is lessened. In the horse, for example, whose metacarpus is long, and in which but one digit is apparent, the humerus is very short. The slight development in length of the humerus explains its close application to the side of the animal as far as the elbow.

In animals in which the humerus is longer, the bone is slightly free, as well as the elbow, at its inferior extremity. [Later on] we will return to the consideration of this peculiarity and of the proportions of the humerus, after we have studied the other parts of the fore-limbs.

The humerus in quadrupeds is inflected like the letter S; in man this general form is less accentuated, the humerus being almost straight. On its body, which appears twisted on its own axis, we find the musculo-spiral groove,[8] which crosses the external surface, and is very deep in some animals. Above this groove, and on the external surface, there exists a rough surface which is the impression of the deltoid. In some species this rugosity is very prominent, and is called the tuberosity of the deltoid; it is prolonged downwards by a border which forms the anterior crest of the musculo-spiral groove and limits this latter in front. The external border of the bone, or posterior crest of the groove, limits it behind.

[8] It would be going outside our province to discuss whether the humerus is really twisted on its axis. This question, often discussed, has been solved in some recent works in the following manner: the humerus has undergone torsion at the level of its superior extremity, and not at the level of its body; this does not authorize us further to accord any definite sense to the denomination ‘groove of torsion’ (musculo-spiral groove). That which we must especially remember in connection with this fact, is, as we shall afterwards see, the difference of direction which the articular head presents according as the torsion has been more or less considerable: because this is established, according to the same order, in man and in quadrupeds.

The superior extremity is enlarged, and remarkable in three portions which it presents; these are: an articular surface and two tuberosities.

The articular surface, or head of the humerus, smooth and round, is in contact with the glenoid cavity of the scapula. This head in the human skeleton is directed upwards and inwards; in quadrupeds its direction is upwards and backwards. The inferior extremity, having in both one and the other its long axis directed transversely, and the point of the elbow looking backwards in all, the result is that the head of the humerus is not situated vertically above the same regions; in the first, it is almost directly above the internal part of this extremity; in the latter, it is situated above its posterior surface, or the point of the elbow in the complete skeleton. This difference of direction is correlated with the position of the scapula, the glenoid cavity of which, as we have already seen, is in man turned outwards, whereas in quadrupeds it looks downwards. In the latter case the scapula consequently rests on the head of the humerus; and this position is most favourable for the performance of the functions which the anterior limbs have to fulfil in these latter.

Of the tuberosities of the head of the humerus, one is situated on the external aspect—it is the great tuberosity, or trochiter; the other is placed internally—it is the small tuberosity, or trochin. The great tuberosity is divided into three parts—summit, convexity, and crest; these different parts give insertion to the muscles of the shoulder. We recollect that the facets (anterior, middle, and posterior) of the great tuberosity of the humerus in man give attachment to the muscles of the same region. The head of the humerus in the human body projects above the tuberosities. We shall see afterwards, when dealing with some special quadrupeds, that in some of these, on the other hand, the tuberosities are on a higher level than the articular head of the bone. Between the two tuberosities is the bicipital groove.

In man, the superior extremity of the humerus, although covered by the deltoid, reveals its presence by elevating the corresponding portion of the latter. In quadrupeds, the anterior part of this extremity, although similarly covered by muscular bundles, produces a prominence under the skin. This prominence is situated at the summit of the angle formed by the opposing directions of the scapula and the bone of the arm, and constitutes what is known by the name of the point of the shoulder, or of the point of the arm.

The inferior extremity, transversely enlarged, presents an undulating articular surface, which reminds us of the trochlea and the condyle of the human humerus; on which, however, the condyle is more sharply defined from the trochlea.

In the human skeleton, the internal lip of the trochlea descends lower than the external; and also lower than the condyle. In the bear, the cat, and the dog, it is the same. In the ox and the sheep, the condyle is lower than the trochlea, but only very little lower. In the horse the arrangement is still the same, but a little more accentuated.

On the lateral parts of this extremity we find: internally, a prominence, the epitrochlea; and, externally, another, the epicondyle. It is from this latter that the crest arises, which, passing upwards, forms the posterior limit of the groove of torsion.

The two prominences, which we have just described from a general point of view, present special arrangements which it is necessary to point out. When we examine the form of the outline of the inferior extremity of the humerus in man, the bear, the cat, the dog, the ox, and the horse, we find in following this order that the extremity tends to become narrow transversely, and that the epicondyle and the epitrochlea are less and less prominent on the external and internal aspects respectively. These two processes, indeed, project backwards; the epitrochlea always remaining more developed than the epicondyle. Because of this projection backwards, the cavity situated on the posterior surface of the inferior extremity, the olecranon fossa, is very deep, more so than in the humerus of man. Its borders being thus formed by the two processes, are very prominent. In front we find the coronoid fossa, which is less deep than that of which we have just spoken.

Fig. 19.—Inferior Extremity of the Left Humerus of a Felide (Lion).

1, Epitrochlea; 2, supra-epitrochlear foramen.

There exists in some mammals an osseous canal, situated above the epitrochlea, and known as the supratrochlear canal ([Fig. 19]). It is bounded by a plate of bone which at its middle portion is detached from the shaft of the humerus, and blends with the latter at both its extremities. The brachial artery and median nerve pass through the foramen.

Fig. 20.—Inferior Extremity of the Left Human Humerus, showing the Presence of a Supratrochlear Process.

1, Epitrochlea; 2, supra-epitrochlear process.

A similar condition is sometimes found, as an abnormality, in man, which presents itself under the following aspect ([Fig. 20]): an osseous prominence more or less long, in the shape of a crochet-needle—supra-epitrochlear process—situated 5 or 6 centimetres above the epitrochlea; the summit of this process gives attachment to a fibrous band, which is inserted by its other end into the epitrochlea and the internal intermuscular aponeurosis. The fibro-osseous ring thus formed gives passage to the brachial artery and the median nerve, or in case of a premature division of this artery to the ulnar branch of the same.[9]

[9] For further details of this anomaly, see Testut, ‘The Epitrochlear Process in Man’ (International Journal of Anatomy and Physiology, 1889); A. Nicolas, ‘New Studies on the Supratrochlear Process in Man’ (Review of Biology of the North of France, t. iii., 1890-1891).

There is also found in some mammals a perforation of the thin plate of bone which, in others, separates the olecranon fossa from the coronoid. This perforation is sometimes found as an abnormality in the human humerus.

Fig. 21.—Skeleton of a Bird (Vulture): Left Surface.

1, Cranium; 2, face; 3, cervical vertebræ; 4, spinous processes of the dorsal vertebræ; 5, coccygeal vertebræ; 6, sternum; 7, keel; 8, superior ribs; 9, inferior ribs; 10, clavicle; 11, coracoid bone (for the details of the skeleton of the shoulder, see [Fig. 18]); 12, humerus; 13, radius; 14, ulna; 15, carpus; 16, hand (for details of the skeleton of this region, see [Fig. 31]); 17, ilium; 18, ischium; 19, pubis (for the details of the pelvis, see [Fig. 46]); 20, femur; 21, tibia; 22, fibula; 23, osseous nodule, which some anatomists think represents the calcaneum; it is the sole vestige of the tarsus; 24, metatarsus; 25, foot; 26, first toe (for the details of the skeleton of the foot, see [Fig. 48]).

As does the sternum and the skeleton of the shoulder, the humerus of birds presents differences correlated to the functions which the thoracic limbs are destined to fulfil. Lying on the side of the thorax, directed obliquely downwards and backwards ([Fig. 21]), it is proportionately longer in individuals of powerful flight than in those which fly less or not at all. In the vulture it projects beyond the posterior part of the pelvis; in the cock it does not even reach the anterior border of the same. To these differences in length are added differences in volume and in the development of the processes which serve for muscular attachment, which are more considerable in birds of powerful flight.

The humerus is so placed that the radial border, external in man and quadrupeds, looks upwards, with the result that the surface of the bone of the arm, which in these latter is anterior, in the former looks outwards. The humeral head, which is turned forwards and a little inwards, is convex and elongated in the vertical direction. Behind and above this head is found a crest for the insertion of muscles. It is the same for the region below, where there is a tuberosity whose inferior surface presents a pretty large opening which looks inwards to a fossa from the floor of which a number of minute openings communicate with the interior of the bone. This is the pneumatic foramen of the humerus.

It is of interest to remember in connection with this subject that in birds, in keeping with the conditions of flight, every system of organs is adapted to diminish the weight of the body. We particularly draw attention to the osseous framework, the structure of which is such that the weight of the animal is greatly lessened. This condition is secured by the pneumaticity. The bone consists of a cover of compact tissue, which, instead of enclosing marrow, is hollowed out by cavities which contain air, and communicate with special pouches, the air-sacs, which are appendages of the lungs.[10]

[10] The presence of air in the bones does not seem to be always associated with the power of flight; as a matter of fact, we find air spaces in the bones of some birds which do not fly (E. J. Marey, ‘The Flight of Birds,’ Paris, 1890, p. 51).

The antibrachial extremity of the humerus is flattened from without inwards. It terminates in two articular surfaces, which articulate with the radius and ulna.

The olecranon process of the ulna being slightly developed, it follows that the olecranon fossa is not large; neither is the coronoid.

General View of the Form of the Forearm and Hand

We now proceed to the study of the two regions of the fore-limbs which present the greatest variety in regard to the number of bones and also in regard to form and proportions. These two regions are the forearm and the hand.

It is first of all necessary to say that in man, when the fore-limb hangs beside the body, and the dorsum of the hand looks backwards, the two bones of the forearm are parallel, and that this position is known by the name of supination. It is also necessary to remember that there is another attitude, in which the radius, crossing the ulna, and carrying the hand with it, displaces the latter in such a way that the palmar surface looks backwards. This second position is known as pronation.

Let us now suppose that a man wishes to walk in the attitude of a quadruped. It will be necessary, in order that his upper limbs, being for the moment anterior ones, may act as members of support, to place the forearm in pronation, in order that, as is more normal, the hands may rest on the ground by their palmar surfaces. In this position the radius, being rotated on its own axis at its upper extremity and around the ulna in the rest of its extent, shall have its inferior extremity situated on the inner side of the corresponding extremity of the latter.

Such is the situation of the bones of the forearm and the attitude of the hand in quadrupeds. In short, quadrupeds have their anterior members in the position of pronation.

Fig. 22.—The Human Hand resting for its Whole Extent on its Palmar Surface: Left Side, External Surface.

The individual whom we have just supposed placed in the attitude of a quadruped would be able to maintain this position by pressing on the ground more or less extensive portions of his hands; the whole palm of the hand may be applied to the ground ([Fig. 22]); or the fingers only—that is to say, the phalanges ([Fig. 23]); or the extremities of the fingers only—that is to say, the third phalanges ([Fig. 24]). This last position, which is certainly difficult to maintain, should here be regarded rather as theoretical.

Fig. 23.—The Human Hand resting on its Phalanges: Left Side, External Surface.

We shall meet with each of these modes of support in certain groups of animals. Thus, the bear, badger, and the majority of rodents, have the paws applied to the ground by the whole extent of the palmar surface of the hand, from the wrist to the tips of the fingers. They are therefore called plantigrade, from the analogy, in this case, of the palm of the hand to the plantar surface, or sole of the foot.

Fig. 24.—The Human Hand resting on the Tips of some of its Third Phalanges: Left Side, External View.

In others, such as the lion, tiger, panther, cat, wolf, and fox, the support is made no longer on the whole extent of the palmar surface, but on the corresponding surface of the fingers only—the metacarpus is turned back, and, consequently, the wrist—that is to say, the carpus—is removed from the ground. These are the digitigrades.

Lastly, the ruminants (sheep, oxen, deer, etc.), and also the pig, ass, and horse, rest on the third phalanx only. In them not only is the metacarpus turned back, but also the two first phalanges. The wrist is very far removed from the ground. In these animals, the third phalanx is enclosed in a case of horn, a nail (the hoof), and because the support of the limb is on that nail, the name of unguligrades has been given them. Nevertheless, as the point of support is on the third phalanx, which is also known by the name of phalangette, we are of opinion that, in order to specify definitely, although they walk on their fingers, as do the digitigrades, the support is provided not by the whole extent of those appendages, they might receive the name of phalangettigrades.

It is necessary among the ruminants to make an exception of the camel and the llama, which are digitigrades.

Just in proportion as the hand is raised from the ground, as we have just seen in passing from the plantigrades to the digitigrades and unguligrades, the number of bones of that region diminishes, the bones of the forearm coalesce, and the ulna tends to disappear; the hand becomes less and less suitable for grasping, climbing, or digging, so as to form an organ exclusively adapted for walking and supporting the body.

Thus, the bear (plantigrade) has five digits, and the power of performing the movements of supination and pronation. Indeed, we know with what facility this animal is able to move his paws in every direction, and climb a tree by grasping it with his fore-limbs. It is well known, however, that no animal except the ape can perform the movements of rotation of the radius around the ulna with the same facility as man; and that none possesses the same degree of suppleness, extent, and variety of movements of the forearm and hand.

In the digitigrades there is one finger which is but slightly developed, and which is always removed from the ground—that is, the thumb: there is also a little less mobility of the radius around the ulna.

In the ungulates the limbs are simply required to perform the movements of walking, and form veritable columns of support, which become the more solid as they are less divided. The bones of the forearm are fused together; there is therefore no possibility of rotation of the radius around the ulna. The metacarpus is reduced to a single piece, which in the horse constitutes what is known as the canon. The number of digits becomes diminished, so that in ruminants there are not more than two, and in the horse but one. We should, however, add that, up to the present, we have taken into account only perfect digits, those that rest on the ground. We shall see further on that there exist supplementary digits, but that they are only slightly developed, and are represented in some cases by mere osseous spurs; it is this fact that has permitted us to ignore them in the general study which we have just made.

Because, as we have already said, the unguligrades have the inferior extremity of the digit encased in a horny sheath, which forms the hoof of the horse and the corresponding structures (onglons) in the ox, those animals have been placed in a special group, which is based on that peculiarity—that is, the group of ungulate mammals.

The plantigrades and digitigrades, of which the paws have their surfaces of support strengthened by an epidermic sole and fatty pads, have the free extremities of the third phalanges covered on their dorsal surface by nails or claws; hence they are named unguiculate mammals.

The bat and birds have the bones of the forearm so arranged that the radius cannot rotate around the ulna. This is necessary in order that during flight, when the wing is being lowered, the radius and hand shall not be able to turn; for, if such rotation took place, each stroke of the wing would place it in a vertical position, which would occasion a loss of resistance incompatible with the effect to be obtained.

The Forearm

The skeleton of the forearm in quadrupeds is vertical in direction; consequently, it forms with the arm an angle open anteriorly; this is well seen on examining the lateral surface. If we examine it on its anterior surface, we find a slight obliquity directed downwards and inwards. In animals in which the bones of the forearm are separate—that is to say, susceptible of supination and pronation—we find a more close resemblance to those of the human skeleton. The ulna, the superior extremity of which always projects beyond that of the radius, has a shaft which gradually narrows from above downwards. Its inferior extremity is terminated by a round head in those animals in which the ulna is fully developed; in others, as it is atrophied, it ends in a thin, long process.