Chapter II: General Anatomical Characters (2)

The dentition of all the members of the orders Primates, Carnivora, Insectivora, Chiroptera, and Ungulata can clearly be derived from the above-described generalised type. The same may be said of the Rodents, and even the Proboscidea, though at least in the existing members of the order with greater modification. It is also apparent in certain extinct Cetacea, as _Zeuglodon_ and _Squalodon_, but it is difficult to find any traces of it in existing Cetacea, Sirenia, or any of the so-called Edentata. All the Marsupials, different as they are in their general structure and mode of life, and variously modified as is their dentition, present in this system of organs some deep-lying common characters which show their unity of origin. The generalised type to which their dentition can be reduced presents considerable resemblance to that of the placental mammals, yet differing in details. It is markedly heterodont, and susceptible of division into incisors, canines, premolars, and molars upon the same principles. The whole number is, however, not limited to forty-four. The incisors may be as numerous as five on each side above, and they are almost always different in number in the upper and the lower jaw. The premolars and molars are commonly seven, as in the placental mammals, but their arrangement is reversed, as there are four true molars and three premolars.

The larger number of incisive and molar teeth among the Marsupials suggests that their additional teeth have disappeared in the Eutheria,[7] and Mr. O. Thomas has endeavoured to construct a generalised dental formula from which both the Marsupial and Eutherian modifications may have been derived by the suppression of particular teeth. Thus the hypothetical formula _i_ ¹,²,³,⁴,⁵⁄₁,₂,₃,₄,₅, _c_ ¹⁄₁, _p_ ¹,²,³,⁴⁄₁,₂,₃,₄, _m_ ¹,²,³,⁴,⁵⁄₁,₂,₃,₄,₅, by the loss of the fifth lower incisor, and of the second premolars (which we know to be those which disappear in the Marsupials) and the fifth molars, will give _i_ ¹,²,³,⁴,⁵⁄₁,₂,₃,₄,₀, _c_ ¹⁄₁, _p_ ¹,⁰,³,⁴⁄₁,₀,₃,₄, _m_ ¹,²,³,⁴⁄₁,₂,₃,₄; or the formula of the Opossum (_Didelphys_), usually written _i_ ⁵⁄₄, _c_ ¹⁄₁, _p_ ³⁄₃, _m_ ⁴⁄₄. Again, in the same formula the loss of the fourth and fifth incisors in both jaws, and also of the fourth molars, gives us _i_ ¹,²,³,⁰,⁰⁄₁,₂,₃,₀,₀, _c_ ¹⁄₁, _p_ ¹,²,³,⁴⁄₁,₂,₃,₄, _m_ ¹,²,³⁄₁,₂,₃, or the formula of a typical Eutherian, like the Pig, which we generally write as _i_ ³⁄₃, _c_ ¹⁄₁, _p_ ⁴⁄₄, _m_ ³⁄₃. Such a generalised formula will admit of modification into that of all existing, and a large number of fossil Marsupials, but it is possible that some of the Mesozoic types may have had more than four premolars, although there is no absolutely decisive evidence that such was the case. The presence of seven or eight true molars in some Mesozoic forms merely entails the addition of two or three additional figures to the ideal generalised formula.

The milk-dentition of all known Marsupials, existing or extinct, is (if not entirely absent) limited to a single tooth on either side of each jaw, this being the predecessor of the last permanent premolar. And if the view that the milk-dentition is an additional series grafted upon the original permanent series be correct, it is evident that we have in this single replacement the first stage of this additional development.

In very few mammals are teeth entirely absent. Even in the Whalebone Whales their germs are formed in the same manner and at the same period of life as in other mammals, and even become partially calcified, but they never rise above the gums, and completely disappear before the birth of the animal. In some species of the order Edentata, the true Anteaters and the Pangolins, no traces of teeth have been found at any age. The adult Monotremata are likewise devoid of teeth of the same structure as those of ordinary mammals; but well-developed molars occur in the young _Ornithorhynchus_, although no traces of teeth have hitherto been detected in _Echidna_.

_Modifications of the Teeth in Relation to their Functions._—The principal functional modifications noticed in the dentition of mammalia may be roughly grouped as piscivorous, carnivorous, insectivorous, omnivorous, and herbivorous, each having, of course, numerous variations and transitional conditions.

The essential characters of a piscivorous dentition are best exemplified in the Dolphins, and also (as modifications of the carnivorous type) in the Seals. This type consists of an elongated, rather narrow mouth, wide gape, with numerous subequal, conical, sharp-pointed, recurved teeth, adapted simply to rapidly seize, but not to divide or masticate, active, slippery, but not powerful prey. All animals which feed on fish as a rule swallow and digest them entire, a process which the structure of prey of this nature, especially the intimate interblending of delicate, sharp-pointed bones with the muscles, renders very advantageous, and for which the above-described type of dentition is best adapted.

The carnivorous type of dentition is shown in its most specialised development among existing mammals in the _Felidæ_. The function being here to seize and kill struggling animals, often of large size and great muscular power, the canines are immensely developed, trenchant, and piercing, and are situated wide apart, so as to give the firmest hold when fixed in the victim’s body. The jaws are as short as is consistent with the free action of the canines, so that no power may be lost. The incisors are very small, so as not to interfere with the penetrating action of the canines, and the crowns of the molar series are reduced to scissor-like blades, with which to pare off the soft tissues from the large bones, or to divide into small pieces the less dense portions of the bones for the sake of nutriment afforded by the blood and marrow they contain. The gradual modification between this and the two following types will be noticed in their appropriate places.

In the most typical insectivorous animals, as the Hedgehogs and Shrews, the central incisors are elongated, pointed, and project forwards, those of the upper and lower jaw meeting like the blades of a pair of forceps, so as readily to secure small active prey, quick to elude capture, but powerless to resist when once seized. The crowns of the molars are covered with numerous sharp edges and points, which, working against each other, rapidly cut up the hard-cased insects into little pieces fit for swallowing and digestion.

The omnivorous type, especially that adapted for the consumption of soft vegetable substances, such as fruits of various kinds, may be exemplified in the dentition of Man, of most Monkeys, and of the less modified Pigs. The incisors are moderate, subequal, and cutting. If the canines are enlarged, it is usually for other purposes than those connected with food, and only in the male sex. The molars have their crowns broad, flattened, and elevated into rounded tubercles. The name _Bunodont_, or hillock-toothed, has been proposed for molars of this type, and will frequently be found convenient.

In the most typically herbivorous forms of dentition, as seen in the Horse and Kangaroo, the incisors are well developed, trenchant, and adapted for cutting off the herbage on which the animals feed; the canines are rudimentary or suppressed; the molars are large, with broad crowns, which in the simplest forms have strong transverse ridges, but may become variously complicated in the higher degrees of modification which this type of tooth assumes.

Various forms of teeth of this type will be noticed among the Ungulates and Rodents.

The natural groups of mammals, or those which in our present state of knowledge we have reason to believe are truly related to each other, may each contain examples of more than one of these modifications. Thus the Primates have both omnivorous and insectivorous forms. The Carnivora show piscivorous, carnivorous, insectivorous, and omnivorous modifications of their common type of dentition. The Ungulata and the Rodentia have among them the omnivorous and various modifications, both simple and complex, of the herbivorous type. The Marsupialia exhibit examples of all forms, except the purely piscivorous. Other orders, more restricted in number or in habits, as the Proboscidea and Cetacea, naturally do not show so great a variety in the dental structure of their members.

_Taxonomy._—In considering the taxonomic value to be assigned to the modifications of teeth of mammals, two principles, often opposed to each other, which have been at work in producing these modifications, must be held in view:—(1) the type, or ancestral form, as we generally now call it, characteristic of each group, which in most mammals is itself derived from the still more generalised type described above; and (2) variations which have taken place from this type, generally in accordance with special functions which the teeth are called upon to fulfil in particular cases. These variations are sometimes so great as completely to mask the primitive type, and in this way the dentition of many animals of widely different origin has come to present a remarkable superficial resemblance, as in the case of the Wombat (a Marsupial), the Aye-Aye (a Lemur), and the Rodents, or as in the case of the Thylacine and the Dog. In all these examples indications may generally be found of the true nature of the case by examining the earlier conditions of dentition; for the characters of the milk-teeth or the presence of rudimentary or deciduous members of the permanent set will generally indicate the route by which the specialised dentition of the adult has been derived. It is perhaps owing to the importance of the dental armature to the well-being of the animal in procuring its sustenance, and preserving its life from the attacks of enemies, that great changes appear to have taken place so readily, and with such comparative rapidity, in the forms of these organs—changes often accompanied with but little modification in the general structure of the animal. Of this proposition the Aye-Aye (_Chiromys_) among Lemurs, the Walrus among Seals, and the Narwhal among Dolphins form striking examples; since in all these forms the superficial characters of their dentition would entirely separate them from the animals with which all other evidence (even including the mode of development of their teeth) proves their close affinity.

_Trituberculism._—Recent researches, and more especially those of Professors Cope and Osborn, tend to show that almost all of the extremely different forms of tooth-structure found among Mammals may be traced to one common type, in which the crown of each tooth carried three cusps, and hence termed the _tritubercular_ type; these three cusps being arranged in a triangle, with the apex directed inwardly in the upper teeth (Fig. 4, ₆), and outwardly in the lower ones (Fig. 4, ₇). It is further probable that this tritubercular type was itself derived from a type of dentition in which the teeth were in the form of almost a quite simple cone; such a presumably primitive type of dentition—being apparently retained among some existing Edentates, like the Armadillos, while it is possible that we should regard the dentition of the existing Cetacea (Fig. 2) as a reversion to the same primitive type. None of the Mesozoic mammals at present known exhibit this simple conical type of teeth, although we have an approximation to it in the extremely generalised genus _Dromatherium_. Starting then from this presumed simple cone it appears that the teeth of _Dromatherium_ (Fig. 4, ₁) present the first stage towards trituberculism, the crown of each tooth having one main cone, with minute lateral cusps, and the root being grooved. In the next or true Triconodont stage (Fig. 4, ₃₋₅) the crown has become elongated antero-posteriorly, and consists of one central and two lateral cones or cusps, while the root is divided. From this the transition is easy to the tritubercular type, in which the three cusps, instead of being placed in a line, are arranged in a triangle; the upper teeth (Fig. 4, ₆) having one inner and two outer cusps, while the reverse condition obtains in those of the lower jaw (Fig. 4, ₇). These three cusps of the simple tritubercular tooth are collectively designated as the primitive triangle; in the upper tooth the inner cusp is termed the protocone, the antero-external one the paracone, and the postero-external the metacone; the corresponding cusps of the lower tooth being named protoconid, paraconid, and metaconid—the protoconid being here on the outer side of the crown.

It is thus apparent that in the first, or haplodont type, as well as in the triconodont type, the upper and lower molars are alike; while in the simple tritubercular type they have a similar pattern, but with the arrangement of the cusps reversed. This simple tritubercular type occurs in the Mesozoic genus _Spalacotherium_ (Fig. 4, ₆ and ₇), and apparently in the existing _Chrysochloris_; but in the majority of tritubercular forms, while this primitive triangle forms the main portion of the crown, other secondary cusps are added, the homologies of which in the upper and lower teeth are somewhat doubtful. At the same time that we have the addition of these secondary cusps we also find trituberculism differentiating into a secodont and a bunodont series, according as to whether the dentition becomes of a cutting or a crushing type.

Thus in the lower molars (Fig. 4, ₈ and ₉) we very frequently find the three cusps of the primitive triangle elevated and connected by cross crests, while there is an additional low posterior heel or talon, which may be termed the hypoconid. This tubercular-sectorial sub-type, as it is termed, is found in the lower molars of many Polyprotodont Marsupials and Insectivores, and it also occurs in the lower carnassial teeth of the true Carnivora. The presence of two cusps (inner and outer) to the talon converts this modification into a quinquetubercular form; while, by the suppression of one of the three primitive cusps, it develops into the quadritubercular type of the bunodont series.

In the upper molars the primitive triangle in the secodont series may remain purely tricuspid; but the addition of intermediate cusps, both in the secodont and bunodont series, may give rise to a quinquetubercular type; these intermediate cusps being respectively designated as the protoconule and metaconule (Fig. 5, _ml_, _pl_). Finally, in the bunodont series, the addition of a postero-internal cusp (Fig. 5, _hy_), termed the hypocone, forms the sextubercular molar.

The following table exhibits, in a collective form, the names and relations of all the above-mentioned cusps, and the letters by which they are indicated in the figures:—

UPPER MOLARS.

Antero-internal cusp = protocone = _pr_.
Postero ” or 6th cusp = hypocone = _hy_.
Antero-external cusp = paracone = _pa_.
Postero ” ” = metacone = _me_.
Anterior intermediate cusp = protoconule = _ml_.
Posterior ” ” = metaconule = _pl_.

LOWER MOLARS.

Antero-external cusp = protoconid = _prd_.
Postero ” ” = hypoconid = _hyd_.
Antero-internal or 5th cusp = paraconid = _pad_.
Intermediate (or in quadritubercular
molars antero-internal) cusp = metaconid = _med_.
Postero-internal cusp = entaconid = _end_.

The common occurrence of trituberculism in the mammals of the earlier geological epochs is, as remarked by Osborn, very significant of the uniformity of molar origin. Thus, among the Mesozoic mammals (with the exception of the group known as Multituberculata, in which the molars are constructed on a different type), trituberculism occurs in the great majority of the genera; while out of 82 species, belonging to five different suborders from the Lowest or Puerco Eocene of the United States, all but four exhibit this feature; and the same holds good for the mammals of the corresponding European horizon. At the present day trituberculism persists in the Lemuroidea, Insectivora, Carnivora, and Marsupialia. In the Carnivora there is a tendency to lose the metaconid, while in the bunodont molars of the Ungulata it is the paraconid that disappears.

III. THE SKELETON.

_Definition._—The skeleton is a system of hard parts, forming a framework which supports and protects the softer organs and tissues of the body. It consists of dense fibrous and cartilaginous tissues, portions of which remain through life in this state, but the greater part is transformed during the growth of the animal into bone or osseous tissue. This is characterised by a peculiar histological structure and chemical composition, being formed mainly of a gelatinous basis, strongly impregnated with salts of calcium, chiefly phosphate, and disposed in a definite manner, containing numerous minute nucleated spaces or cavities called lacunæ, connected together by delicate channels or canaliculi, which radiate in all directions from the sides of the lacunæ. Parts composed of bone are, next to the teeth, the most imperishable of all the organs of the body, often retaining their exact form and internal structure for 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.[8]

In the Armadillos and their extinct allies alone is there an ossified exoskeleton, or bony covering developed in the skin. In all other mammals the skeleton is completely internal. It may be described as consisting of an axial portion belonging to the head and trunk, and an appendicular 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.

It is characteristic of all the larger bones of the mammalia that their ossification takes its origin from several distinct centres. One near the middle of the bone, and spreading throughout its greater portion, constitutes the _diaphysis_, or “shaft,” in the case of the long bones. Others near the extremities, or in projecting parts, form the _epiphyses_, which remain distinct during growth, but ultimately coalesce with the rest of the bone.

_Axial skeleton._—The axial skeleton consists of the skull, the vertebral column (prolonged at the posterior extremity into the tail), the sternum, and the ribs.

_Skull._—In the _skull_ of adult mammals, all the bones, except the lower jaw, the auditory ossicles, and the bones of the hyoid arch, are immovably articulated together, their edges being in close contact, and often interlocking by means of fine denticulations projecting from one bone and fitting into corresponding depressions of the other; they are also held together by the investing periosteum, or fibrous membrane, which passes directly from one to the other, and permits no motion, beyond perhaps a slight yielding to external pressure. In old animals there is a great tendency for the different bones to become actually united by the extension of ossification from one to the other, with consequent obliteration of the sutures. The cranium, thus formed of numerous originally independent ossifications, which may retain throughout life more or less of their individuality, or be all fused together, according to the species, the age, or even individual peculiarity, consists of a brain-case, or bony capsule for enclosing and protecting the brain, and a face for the support of the organs of sight, smell, and taste, and of those concerned in seizing and masticating the food. The brain-case articulates directly with the anterior cervical vertebra, by means of a pair of oval eminences, called condyles, placed on each side of the large median foramen which transmits the spinal cord. It consists of a basal axis, continuous serially with the axes or centra of the vertebræ, and of an arch above, roofing over and enclosing the cavity which contains the cephalic portion of the central nervous system (see Fig. 6). The base with its arch is composed of three segments placed one before the other, each of which is comparable to a vertebra with a greatly expanded neural arch. The hinder or occipital segment consists of the basioccipital, exoccipital, and supraoccipital bones; the middle segment of the basisphenoid, alisphenoid, and parietal bones; and the anterior segment of the presphenoid, orbitosphenoid, and frontal bones. The axis is continued forwards into the mesethmoid, or septum of the nose, around which the bones of the face are arranged in a manner so extremely modified for their special purposes that anatomists who have attempted to trace their serial homologies with the more simple portions of the axial skeleton have arrived at very diverse interpretations. The characteristic form and structure of the face of mammals is mainly dependent upon the size and shape of (1) the orbits, a pair of cup-shaped cavities for containing the eyeball and its muscles, which may be directed forwards or laterally, placed near together or wide apart, and may be completely or only partially encircled by bone; (2) the nasal fossæ, or cavities on each side of the median nasal septum, forming the passage for the air to pass between the external and the internal nares, and containing in their upper part the organ of smell; (3) the zygomatic arch, a bridge of bone for the purpose of muscular attachment, which extends from the side of the face to the skull, overarching the temporal fossa; (4) the roof of the mouth, with its alveolar margin for the implantation of the upper teeth. The face is completed by the mandible, or lower jaw, consisting of two lateral rami, articulated by a hinge joint with the squamosal (a cranial bone interposed between the posterior and penultimate segment of the brain-case, where also the bony capsule of the organ of hearing is placed), each being composed of a single solid piece of bone, and the two united together in the middle line in front, at the symphysis,—which union may be permanently ligamentous or become completely ossified. Into the upper border of the mandibular rami the lower teeth are implanted.

]

In addition to the bones already mentioned as entering into the formation of the cranium, there are many others, the most important of which may be briefly noticed. The anterior extremity of the skull is formed by the premaxillæ (Figs. 6, 7, _PMx_), which carry the incisors; behind them are the maxillæ, in which all the remaining upper teeth are implanted. Both the premaxillæ and maxillæ meet in a median suture on the palate, where they form a floor to the nasal passage; this floor being continued backwards by the plate-like palatines, at the hinder extremity of which the posterior nares are usually situated. In a few instances, however, as in certain Edentates and Cetaceans, the small pair of bones forming the posterior continuation of the lateral borders of the palatines, and known as the pterygoids (Fig. 6, _Pt_), likewise meet in the middle line below the nasal passage, and thus cause the aperture of the posterior nares to be situated near the occiput. On the upper, or frontal aspect of the cranium the paired nasals roof over the nasal passage and fill the interval left between the premaxilla and maxilla of either side. Behind the nasals and maxillæ, the anterior part of the brain-case is formed by the large paired frontals (Figs. 6, 7, _Fr_), behind which are the parietals, which may be of still larger size, and form the greater part of the brain-case. A median interparietal ossification (Fig. 6, _IP_) may divide the parietals posteriorly, and is itself articulated with the supraoccipital, to the lateral borders of which the parietals are also joined. The squamosal (Fig. 7, _Sq_) forms the lateral wall of the hinder part of the brain-case, and articulates superiorly with the parietal, and posteriorly with the exoccipital. The glenoid cavity (Fig. 8), for the reception of the articular condyle of the mandible, is formed by the inferior portion of the squamosal, at the point where it gives off the zygomatic process to form the hinder portion of the zygomatic arch. The middle portion of that arch is formed by the jugal, or malar bone (Fig. 7, _Ma_), which articulates posteriorly with the zygomatic process of the squamosal, and anteriorly with the maxilla. The jugal (as in Fig. 7) may also articulate with a small bone situated on the anterior border of the orbit known as the lachrymal. It is important to observe that the zygomatic or temporal arch is a squamoso-maxillary one, and that an arcade thus composed is found elsewhere only among the extinct Anomodont reptiles, which have already been mentioned as showing signs of mammalian affinity. The relative position occupied by the orbito- and alisphenoid is sufficiently indicated in Fig. 7.

Wedged in between the squamosal and the bones of the occipital and basisphenoidal region are the bones connected with the organ of hearing, known as the periotic and tympanic. The position of the periotic, which encloses the labyrinth or essential organ of hearing, is shown in Fig. 6. The periotic is divided into a very dense antero-internal moiety known as the petrosal, and a postero-external or mastoid portion (Fig. 8), which appears on the outer wall of the brain-case. The tympanic is produced horizontally outwards to form the external auditory meatus or tube of the ear, while the inner and under surface is frequently dilated into a shell-like auditory bulla (Fig. 8). The small bones of the internal ear known as the malleus, incus, and stapes are contained in the membranous _tympanic cavity_, which is situated in a space left among this group of bones. Further mention of these bones is made below under the head of the sense organs.

In the Carnivora and some other groups the foramina on the base of the skull for the passage of blood-vessels and nerves are of considerable taxonomic importance. The position of the more important of these foramina is indicated in Fig. 8; but for details the reader may refer to the work on the _Osteology of the Mammalia_ already mentioned. Attention may, however, be particularly directed to the so-called alisphenoid canal, the position of which is shown in Fig. 8, since this is a feature of some importance in the classification of the Carnivora. This canal is a short channel running horizontally forward from near the foramen ovale through the alisphenoid, and opening anteriorly with the foramen rotundum; it is traversed by the external carotid artery.

Only in those species, as Man and the smaller kinds of the Primates and some other orders, in which the brain holds a large relative proportion to the rest of the body, does the external form of the skull receive much impress from the real shape of the cavity containing the brain. The size and form of the mouth, and the modifications of the jaws for the support of teeth of various shape and number, the ridges and crests on the cranium for the attachment of the muscles necessary to put this apparatus in motion, and outgrowths of bone for the enlargement of the external surface required for the support of sense organs or of weapons, such as horns or antlers (which outgrowths, to prevent undue increase of weight, are filled with cells containing air), cause the principal variations in the general configuration of the skull. These variations are, however, only characteristically developed in perfectly adult animals, and are in many cases more strongly marked in the male than the female sex. Throughout all the later stages of growth up to maturity the size and form of the brain-case remain comparatively stationary, while the accessory parts of the skull rapidly increase and assume their distinctive development characteristic of the species.

The hyoidean apparatus in mammals (Fig. 6) supports the tongue and larynx, and consists of an inferior median portion termed the basihyal, from which two pairs of half arches, or cornua, extend upwards and outwards. The anterior is the more important, being connected with the periotic bone of the cranium. It may be almost entirely ligamentous, but more often has several ossifications, the largest of which is usually the stylohyal. The posterior cornu (thyrohyal) is united at its extremity with the thyroid cartilage of the larynx, which it suspends in position. The median portion, or basihyal, is sometimes, as in the Howling Monkeys, enormously enlarged and hollowed, admitting into its cavity an air-sac connected with the organ of voice.

_Vertebral Column._—The _vertebral column_ consists of a series of distinct bones called vertebræ, arranged in close connection with each other along the dorsal side of the neck and trunk, and in the median line.[10] 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. The number of distinct bones composing the vertebral column varies greatly among the Mammalia, the main variation being due to the degree of elongation 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 (_Hyrax_ and _Cholœpus_). The different vertebræ, 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. The exceptions are the following:—(1) near the posterior part of the trunk, in nearly all mammals which possess completely developed hinder limbs, two or more vertebræ become ankylosed together to form the “sacrum,” or portion of the vertebral column to which the pelvic girdle is attached; (2) in some species of Whales and Armadillos there are constant ossific unions of certain vertebræ of the cervical region.

Although the vertebræ of different regions of the column of the same animal or of different animals present great diversities of form, yet there is a certain general resemblance among them, or a common plan on which they are constructed, which is more or less modified by alteration of form or proportions, or by the addition or suppression of parts to fit them to fulfil their special purpose in the economy. An ordinary or typical vertebra consists, in the first place, of a solid piece of bone, termed the body or centrum (Fig. 9, _c_), of the form of a disk or short cylinder. The bodies of contiguous vertebræ are connected together by a very dense, tough, and elastic material called the “intervertebral substance,” of peculiar and complex arrangement. This substance forms the main, and in some cases the only, union between the vertebræ. Its elasticity provides for the vertebræ 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 (_p_) arises on each side from the dorsal surface of the body. These processes, meeting in the middle line above, form an arch, surmounting a space or short canal (_nc_). Since it contains the posterior prolongation of the great cerebro-spinal nervous axis, or spinal cord, this space is called the neural canal, and the arch the neural arch, in contradistinction to another arch on the ventral surface of the body of the vertebræ, called the hæmal arch. The latter is, however, never formed in mammals by any part of the vertebra itself, but by certain distinct bones placed more or less in apposition to it, namely the ribs in the thoracic, and the “chevron bones” in the caudal region. In most cases the arch of one vertebra is articulated with that of the next by distinct surfaces with synovial joints, placed one on each side, called “zygapophyses” (_az_, _pz_), but these 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, chiefly serving for the attachment of the numerous muscles which move the vertebral column. Of these two are single and median, viz. the spinous process, neural spine, or neurapophysis (_s_), arising from the middle of the upper part of the arch, and the hypapophysis from the under surface of the body. The latter, however, is as frequently absent as the former is constant. The other processes are paired and lateral. They are the transverse processes (_t_), of which there may be two, an upper and a lower, in which case the former is called, in the language of Owen (to whom we are indebted for the terminology of the parts of vertebræ in common use), “diapophysis,” and the latter “parapophysis.” Other processes less constantly present are called respectively “metapophyses” (_m_) and “anapophyses” (_a_).

The vertebral column is divided for convenience of description into five regions—the cervical, thoracic or dorsal, lumbar, sacral, and caudal. This division is useful, especially as it is not entirely arbitrary, and in most cases is capable of ready definition; but at the contiguous extremities of the regions the characters of the vertebræ of one are apt to blend into those of the next region, either normally or as peculiarities of individual skeletons.

_Cervical Vertebræ._—The _cervical_ region constitutes the most anterior portion of the column, or that which joins the cranium. The vertebræ which belong to it are either entirely destitute of movable ribs, or if they have any these are small, and do not join the sternum. As a general rule they have a considerable perforation through the base of the transverse process (the vertebrarterial canal, Fig. 11, _v_); or, as it is sometimes described, they have two transverse processes, superior and inferior, which meet at their extremities to enclose a canal. This, however, rarely applies to the last vertebra of the region, in which only the upper transverse process is usually developed. The transverse process, moreover, very often sends down near its extremity a more or less compressed plate (inferior lamella), which, being considered serially homologous with the ribs of the thoracic vertebræ (though not developed autogenously), is often called the “costal” or “pleurapophysial” plate. This is usually largest on the sixth, and altogether wanting on the seventh vertebra. The first and second cervical vertebræ, 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 the intervertebral substance, 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 vertebræ is the same, viz. seven, with the exception of the Manatee and Hoffman’s Two-toed Sloth (_Cholœpus hoffmanni_), which both have but six, and the Three-toed Sloth (_Bradypus tridactylus_), which has nine, though in this case the last two usually support movable ribs, which are not sufficiently developed to reach the sternum.

According to Parker there may occasionally be eight cervicals in the Pangolins (_Manis_).

_Dorsal Vertebræ._—The _dorsal_ (or, as it would be more correctly termed, _thoracic_) region consists of the vertebræ succeeding those of the neck, which have ribs movably articulated to them. These ribs arch round the thorax—the anterior one, and usually the greater number of those that follow, being attached below to the sternum.

_Lumbar Vertebræ._—The _lumbar_ region consists of those vertebræ of the trunk in front of the sacrum 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 region. As a general rule there is a certain relation between the number of the thoracic and lumbar vertebræ, 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 all known Artiodactyle Ungulata there are 19 dorso-lumbar vertebræ; but these may consist of 12 dorsal and 7 lumbar vertebræ, or 13 dorsal and 6 lumbar, or 14 dorsal and 5 lumbar. The smallest number of dorso-lumbar vertebræ 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 Artiodactyla, nearly all Marsupials, and very many Rodents; 20 or 21 in Carnivora and most Insectivora; and 23 in Perissodactyla. The highest and quite exceptional numbers are in the Two-toed Sloth (_Cholœpus_) 27, and the Hyrax 30. The prevailing number of rib-bearing vertebræ is 12 or 13, any variation being generally in excess of these numbers.

_Sacral Vertebræ._—The _sacral_ region offers more difficulties of definition. Taking the human “os sacrum” as a guide for comparison, it is generally defined as consisting of those vertebræ between the lumbar and caudal regions which are ankylosed together to form a single bone. It happens, however, that the number of such vertebræ varies in different individuals of the same or nearly allied species, especially as age advances, when a certain number of the tail vertebræ generally become incorporated with the true sacrum. Other suggested tests—as those vertebræ which have a distinct additional (pleurapophysial) centre of ossification between the body and the ilium, those to which the ilium is directly articulated, or those in front of the insertion of the ischiosacral ligaments—being equally unsatisfactory or unpractical, the old one of ankylosis, as it is found to prevail in the average condition of adults in each species, is used in the enumeration of the vertebræ in the following pages. The Cetacea, having no iliac bones, have no part of the vertebral column modified into a sacrum.

_Caudal Vertebræ._—The _caudal_ vertebræ are those placed behind the sacrum, and terminating the vertebral column. They vary in number greatly—being reduced to 5, 4, or even 3, in a most rudimentary condition, in Man and in 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 highest known number, 46, is possessed by the African Long-tailed Pangolin. Connected with the under surface of the caudal vertebræ of many mammals which have the tail well developed are certain bones formed more or less like an inverted arch, called chevron bones, or by the French _os en V_. 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.

In some of the Anomodont Reptiles and Labyrinthodont Amphibians these chevrons are attached to the intercentra—or imperfect disks alternating with the true centra—which suggests that they are primarily intercentral elements which have been transferred to the edges of the centra by the disappearance of the intercentra.

_Sternum._—The _sternum_ of mammals is a bone, or generally a series of bones, placed longitudinally in the mesial 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.” It is present in all mammals, but varies much in character in the different groups. It usually consists of a series of distinct segments placed one before the other, the anterior being called the presternum or “manubrium sterni” of human anatomy, and the posterior the xiphisternum, or xiphoid or ensiform process, while the intermediate segments, whatever their number, constitute the mesosternum or “body.” In the Whalebone Whales the presternum alone is developed, and but a single pair of ribs is attached to it.

_Ribs._—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. The posterior ribs, however, do not directly articulate with that bone, but are either attached by their extremities to the edges of each rib in front of them, and thus only 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” 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. The portion of each rib nearest the vertebral column and that nearest the sternum differ in their characters, the latter being usually but imperfectly ossified, or remaining permanently cartilaginous. These are called “costal cartilages,” or when ossified “sternal ribs.”

In the anterior part of the thorax the vertebral extremity of each rib is divided into two parts, “head” or “capitulum,” and “tubercle”; the former is attached to the side of the body of the vertebra, the latter to its transverse process; the former attachment corresponds to the interspace between the vertebræ, 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. The number of pairs of ribs is of course the same as that of the thoracic vertebræ.

The circumstance that in some of the Anomodont reptiles and Labyrinthodonts the capitula of the ribs articulate with the intercentral elements of the vertebral column has suggested, as in the instance of the chevron bones, that the intercentral capitular articulation of the ribs of mammals is a feature directly inherited from those extinct types by the gradual disappearance of the intercentra.

_Appendicular Skeleton._—The appendicular portion of the framework consists, when completely developed, of two pairs of limbs, anterior and posterior (Fig. 15).

_Anterior Limb._—The anterior limb is present and fully developed in all mammals, being composed of a shoulder girdle and three segments belonging to the limb proper; viz. the upper arm or brachium, the forearm or antebrachium, and the hand or manus.

_Shoulder-girdle._—The _shoulder_ or _pectoral girdle_ in the large majority of mammals is in a rudimentary or rather modified condition, compared with that in which it exists in other vertebrates. In the Monotremata (_Ornithorhynchus_ and _Echidna_) alone is the ventral portion, or coracoid, complete and articulated with the sternum below, as in the Sauropsida; and in this group alone do we find an anterior ventral element, apparently corresponding with the pre-coracoid of the Anomodont reptiles, although generally known as the epi-coracoid. In all other mammals the coracoid, though ossified from a distinct centre, forms only a process, sometimes a scarcely distinct tubercle, projecting from the anterior border of the glenoid cavity of the scapula. The last-named cavity, which in the Monotremes is formed jointly by the scapula and coracoid, receives the head of the humerus, or arm-bone. The scapula is always well developed, and generally broad and flat (whence its vernacular name “blade bone”), with a ridge called the “spine” on its outer surface, which usually ends in a free curved process, the “acromion.” As the scapula affords attachment to many of the muscles which act upon the anterior limb, its form and the development of its processes are greatly modified according to the uses to which the member is put. Thus it is most reduced and simple in character in those animals whose limbs are mere organs of support, as the Ungulates; and most complex when the limbs are also used for grasping, climbing, or digging. The development or absence of the clavicle or “collar-bone,” an accessory bar which connects the sternum with the scapula and steadies the shoulder-joint, has a somewhat similar relation, though its complete absence in the Bears shows that this is not an invariable rule. A complete clavicle is found in Man and all the Primates, in Chiroptera, all Insectivora (except _Potamogale_), in many Rodents, in most Edentates, and in all Marsupials, except _Perameles_. More or less rudimentary clavicles (generally suspended freely in the muscles) are found in the Cat, Dog, and most Carnivora, _Myrmecophaga_, and some Rodents. Clavicles are altogether absent in most of the _Ursidæ_, all the Pinnipedia, _Manis_ among Edentates, the Cetacea, Sirenia, Ungulates, and some Rodents.

The Monotremes are peculiar in possessing a T-shaped interclavicle like that of many reptiles, lying upon the sternum, and articulating superiorly with the clavicles.

_Brachium and Antebrachium._—The proximal segment of the anterior or pectoral limb proper contains a single bone, the humerus, and the second segment two bones, the radius and the ulna, placed side by side, and articulating with the humerus at their proximal, and with the carpus at their distal extremity (Fig. 15). In their primitive and unmodified condition these bones may be considered as placed one on each border of the limb, the radius being preaxial or anterior, and the ulna postaxial or posterior, when the distal or free end of the limb is directed outwards, or away from the trunk. This is their position in the earliest embryonic condition, and is best illustrated among adult mammals in the Cetacea, where the two bones are fixed side by side and parallel to each other. In the greater number of mammals the bones assume a very modified and adaptive position, usually crossing each other in the forearm, 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 preaxial 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 supination, 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.

The humerus frequently has a foramen piercing the inner border of the distal extremity, known as the entepicondylar foramen, which corresponds with a similar one found in the Anomodont reptiles. The hollow in the head of the ulna for the reception of the head of the humerus is known as the greater sigmoid cavity, and that for the head of the radius as the lesser sigmoid cavity (Fig. 16). The term olecranon is applied to that process of the ulna which forms the prominence of the elbow.

In most mammals walking on four limbs, in which the hand is permanently prone, the ulna is much reduced in size, and the radius increased, especially at the upper end; so that the articular surface of the latter, instead of being confined 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. In many hoofed or Ungulate mammals, and in Bats, the ulna is reduced to little more than its upper articular extremity, and firmly ankylosed to the radius—stability of these parts being more essential than mobility.

_Manus._—The terminal segment of the anterior limb is the hand or manus. Its skeleton consists of three divisions: (1) 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 to one another by freely movable hinge-joints, the first being connected in like manner to the distal end of the metacarpal bone to which it corresponds.

_Carpus._—To understand thoroughly the arrangement of the bones of the carpus in mammals, it is necessary to study their condition in some of the lower vertebrates. Fig. 17 represents the manus in one of its fullest 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. The upper or proximal row contains three bones, to which Gegenbaur has applied the terms _radiale_ (_r_), _intermedium_ (_i_), and _ulnare_ (_u_), the first being on the radial or preaxial side of the limb.[11] The lower or distal row contains five bones, called _carpale_ 1, 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 the _radiale_ supports on its distal side two bones, _carpale_ 1 and 2; the _intermedium_ is in a line with the _centrale_ and _carpale_ 3, which together form a median axis of the hand, while the _ulnare_ has also two bones articulating 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, which is the more constant and generally larger, is commonly known as the pisiform bone. The fourth and fifth carpals of the distal row are always united into a single bone, and the centrale is very often absent. As a general rule all the other bones are present and distinct, though it not unfrequently happens that two may have coalesced to form a single bone, or one or more may be altogether suppressed.

The following table shows the principal names in use for the various carpal bones,—those in the second column being the terms generally employed by English anatomists:—

_Radiale_ = Scaphoid = _Naviculare_.
_Intermedium_ = Lunar = _Semilunare_, _Lunatum_.
_Ulnare_ = Cuneiform = _Triquetrum_, _Pyramidale_.
_Centrale_ = Central = _Intermedium_ (Cuvier).
_Carpale_ 1 = Trapezium = _Multangulum majus_.
_Carpale_ 2 = Trapezoid = _Multangulum minus_.
_Carpale_ 3 = Magnum = _Capitatum_.
_Carpale_ 4 } = Uneiform = _Hamatum_, _Uncinatum_.
_Carpale_ 5 }

The radial and ulnar sesamoids are regarded by Bardeleben[12] as the rudiments of a prepollex and a postminimus digit; the primitive number of digits being thus supposed to have been seven. These bones have been observed in all orders of mammals having five complete digits. Occasionally, as in _Pedetes caffer_, the so-called prepollex consists of two bones, of which the distal one bears a distinct nail-like horny covering. In _Bathyergus maritimus_ the pisiform, or postminimus, is likewise double; the two elements being regarded by their describer as representing the carpal and metacarpal of the presumed seventh digit.

Similarly in the posterior limb the tibial sesamoid, and a fibular ossification corresponding to the pisiform, are regarded as representing a prehallux and a postminimus.

_Metacarpus and Phalanges._—The metacarpal bones, with the digits which they support, are never more than five in number, and are described numerically—first, second, etc., counting from the radial towards the ulnar side. The digits are also sometimes named (1) the pollex, (2) index, (3) medius, (4) annularis, (5) minimus. One or more may be in a rudimentary condition, or altogether suppressed. If one is absent, it is most commonly the first. 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 is an exception to the usual rule, one of its parts being constantly absent, since, while each of the other digits has commonly a metacarpal and three phalanges, it has only three bones altogether; whether the missing one is a metacarpal or one of the phalanges is a subject which has occasioned much discussion, and has not yet been satisfactorily decided. The terminal phalanges of the digits are usually specially modified to support the nail, claw, or hoof, and are called “ungual phalanges.” In walking, some mammals (as the Bears) apply the whole of the lower surface of the carpus, metacarpus, and phalanges to the ground; to these the term “plantigrade” is applied. Many others (as nearly all the existing Ungulata) only rest on the last one or two phalanges of the toes, the first phalanx and the metacarpals being vertical and in a line with the forearm. These are called “digitigrade.” Intermediate conditions exist between these two forms, to which the terms “phalangigrade” (as the Camel) and “subplantigrade” (as in most Carnivora), are applied. When the weight is borne entirely on the distal surface of the ungual phalanx, and the horny structures growing around it, as in the Horse, the mode of progression is called “unguligrade.”

In the Chiroptera the digits are enormously elongated, and support a cutaneous expansion constituting the organ of flight. In the Cetacea the manus is formed into a paddle, being covered by continuous integument, which conceals all trace of division into separate digits, and shows no sign of nails or claws. In the Sloths the manus is long and very narrow, habitually curved, and terminating in two or three pointed curved claws in close apposition with each other, and incapable, 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 lives. These are only examples of the endless modifications to which the distal extremity of the limb is subjected in adaptation to the various purposes to which it is applied.

_Posterior Limb._—The posterior limb is constructed upon a plan very similar to that of the anterior extremity. It consists of a pelvic girdle and three segments belonging to the limb proper, viz. the thigh, the leg, and the foot or pes (Fig. 15).