1911 Encyclopædia Britannica/Mammalia
MAMMALIA (from Lat. mamma, a teat or breast), the name proposed by the Swedish naturalist Linnaeus for one of the classes, or primary divisions, of vertebrated animals, the members of which are collectively characterized by the presence in the females of special glands secreting milk for the nourishment of the young. With the exception of the lowest group, such glands always communicate with the exterior by means of the teats, nipples or mammae, from which the class derives its name. The class-name (modified by the French into Mammifères, and replaced in German by the practically equivalent term Säugethiere) has been anglicized into “Mammals” (mammal, in the singular). Of recent years, and more especially in America, it has become a custom to designate the study of mammals by the term “mammalogy.” Etymologically, however, that designation cannot be justified; for it is of hybrid (Latin and Greek) origin, and is equivalent to “mastology,” the science which deals with the mammary gland (Gr. μαστός, woman’s breast), a totally different signification. As regards existing forms of life, the limitations of the class are perfectly well defined and easy of recognition; for although certain groups (not, by the way, whales, which, although excluded in popular estimation from the class, are in all essential respects typical mammals) are exceedingly aberrant, and present structural features connecting them with the lower vertebrate classes, yet they are by common consent retained in the class to which they are obviously most nearly affiliated by their preponderating characteristics. There is thus at the present day a great interval, unbridged by any connecting links, between mammals and the other classes of vertebrates.
Not so, however, when the extinct forms of vertebrate life are taken into consideration, for there is a group of reptiles from the early part of the Secondary, or Mesozoic period, some of whose members must have been so intimately related to mammals that, were the whole group fully known, it would clearly be impossible to draw a distinction between Mammalia on the one hand and Reptilia on the other. Indeed, as it is, we are already partially acquainted with one of these early intermediate creatures (Tritylodon), which forms a kind of zoological shuttlecock, being, so to speak, hit from one group to another, and back again, by the various zoologists by whom its scanty remains have been studied. Considered collectively, mammals, which did not make their appearance on the earth for some time after reptiles had existed, are certainly the highest group of the whole vertebrate sub-kingdom. This expression must not, however, be considered in too restricted a sense. In mammals, as in other classes, there are low as well as high forms; but by any tests that can be applied, especially those based on the state of development of the central nervous system, it will be seen that the average exceeds that of any other class, that many species of this class far excel those of any other in perfection of structure, and that it contains one form which is unquestionably the culminating point amongst organized beings.
Mammals, then, are vertebrated animals, possessing the normal characteristics of the members of that primary division of the animal kingdom. They are separated from fishes and batrachians (Pisces and Batrachians) on the one hand, and agree with reptiles, and birds (Reptilia and Aves) on the other, in the possession during intra-uterine life of the membranous vascular structures respectively known as the amnion and the allantois, and likewise in the absence at this or any other period of external gills. A four-chambered heart, with a complete double circulation, and warm blood (less markedly so in the lowest group than in the rest of the class), distinguish mammals from existing reptiles, although not from birds. From both birds and reptiles the class is distinguished, so far at any rate as existing forms are concerned, by the following features: the absence of a nucleus in the red corpuscles of the blood, which are nearly always circular in outline; the free suspension of the lungs in a thoracic cavity, separated from the abdominal cavity by a muscular partition, or diaphragm, which is the chief agent in inflating the lungs in respiration; the aorta, or main artery, forming but a single arch after leaving the heart, which curves over the left terminal division of the windpipe, or bronchus; the presence of more or fewer hairs on the skin and the absence of feathers; the greater development of the bridge, or commissure, connecting the two halves of the brain, which usually forms a complete corpus callosum, or displays an unusually large size of its anterior portion; the presence of a fully developed larynx at the upper end of the trachea or windpipe, accompanied by the absence of a syrinx, or expansion, near the lower end of the same; the circumstance that each half of the lower jaw (except perhaps at a very early stage of development) consists of a single piece articulating posteriorly with the squamosal element of the skull without the intervention of a separate quadrate bone; the absence of prefrontal bones in the skull; the presence of a pair of lateral knobs, or condyles (in place of a single median one), on the occipital aspect of the skull for articulation with the first vertebra; and, lastly, the very obvious character of the female being provided with milk-glands, by the secretion of which the young (produced, except in the very lowest group, alive and not by means of externally hatched eggs) are nourished for some time after birth.
In the majority of mammals both pairs of limbs are well developed and adapted for walking or running. The fore-limbs may, however, be modified, as in moles, for burrowing, or, as in bats, for flight, or finally, as in whales and dolphins, for swimming, with the assumption in this latter instance of a flipper-like form and the complete disappearance of the hind-limbs. Special adaptations for climbing are exhibited by both pairs of limbs in opossums, and for hanging to boughs in sloths. In no instance are the fore-limbs wanting.
In the great majority of mammals the hind extremity of the axis of the body is prolonged into a tail. Very generally the tail has distinctly the appearance of an appendage, but in some of the lower mammals, such as the thylacine among marsupials, and the aard-vark or ant-bear among the edentates, it is much thickened at the root, and passes insensibly into the body, after the fashion common among reptiles. As regards function, the tail may be a mere pendent appendage, or may be adapted to grasp boughs in climbing, or even to collect food or materials for a nest or sleeping place, as in the spider-monkeys, opossums and rat-kangaroos. Among jumping animals it may serve as a balance, as in the case of jerboas and kangaroos, while in the latter it is also used as a support when resting; among many hoofed mammals it is used as a fly-whisk; and in whales and dolphins, as well as in the African Potamogale and the North American musquash, it plays an important part in swimming. Its supposed use as a trowel by the beaver is, however, not supported by the actual facts of the case.
As already indicated, the limbs of different mammals are specially modified for various modes of life; and in many cases analogous modifications occur, in greater or less degree, throughout the entire body. Those modifications most noticeable in the case of cursorial types may be briefly mentioned as examples. In this case, as might be expected, the greatest modifications occur in the limbs, but correlated with this is also an elongation of the head and neck in long-legged types. Adaptation for speed is further exhibited in the moulding of the shape of the body so as to present the minimum amount of resistance to the air, as well as in increase in heart and lung capacity to meet the extra expenditure of energy. Finally, in the jumping forms we meet with an increase in the length and weight of the tail, which has to act as a counterpoise. As regards the feet, a reduction in the number of digits from the typical five is a frequent feature, more especially among the hoofed mammals, where the culmination in this respect is attained by the existing members of the horse tribe and certain representatives of the extinct South American Proterotheriidae, both of which are monodactyle. Brief reference may also be made to the morphological importance of extraordinary length or shortness in the skulls of mammals—dolichocephalism and brachycephalism; both these features being apparently characteristic of specialized types, the former condition being (as in the horse) often, although not invariably, connected with length of limb and neck, and adaptation to speed, while brachycephalism may be correlated with short limbs and an abbreviated neck. Exceptions to this rule, as exemplified by the cats, are due to special adaptive causes. In point of bodily size mammals present a greater range of variation than is exhibited by any other living terrestrial animals, the extremes in this respect being displayed by the African elephant on the one hand and certain species of shrew-mice (whose head and body scarcely exceed an inch and a half in length) on the other. When the aquatic members of the class are taken into consideration, the maximum dimensions are vastly greater, Sibbald’s rorqual attaining a length of fully 80 ft., and being probably the bulkiest and heaviest animal that has ever existed. Within the limits of individual groups, it may be accepted as a general rule that increase in bulk or stature implies increased specialization; and, further, that the largest representatives of any particular group are also approximately the latest. The latter dictum must not, however, be pushed to an extreme, since the African elephant, which is the largest living land mammal, attaining in exceptional cases a height approaching 12 ft., was largely exceeded in this respect by an extinct Indian species, whose height has been estimated at between 15 and 16 ft.
In regard to sense-organs, ophthalmoscopic observations on the eyes of living mammals (other than man) have revealed the existence of great variation in the arrangement of the blood-vessels, as well as in the colour of the retina; blue and violet seem to be unknown, while red, yellow and green form the predominating shades. In the main, the various types of minute ocular structure correspond very closely to the different groups into which mammals are divided, this correspondence affording important testimony in the favour of the general correctness of the classification. Among the exceptions are the South American squirrel-monkeys, whose eyes approximate in structure to those of the lemurs. Man and monkeys alone possess parallel and convergent vision of the two eyes, while a divergent, and consequently a very widely extended, vision is a prerogative of the lower mammals; squirrels, for instance, and probably also hares and rabbits, being able to see an object approaching them directly from behind without turning their heads.
An osteological question which has been much discussed is the fate of the reptilian quadrate bone in the mammalian skull. In the opinion of F. W. Thyng, who has carefully reviewed all the other theories, the balance of evidence tends to show that the quadrate has been taken up into the inner ear, where it is represented among the auditory ossicles by the incus.
Although the present article does not discuss mammalian osteology in general (for which see Vertebrata), it is interesting to notice in this connexion that the primitive condition of the mammalian tympanum apparently consisted merely of a small and incomplete bony ring, with, at most, an imperfect ventral wall to the tympanic cavity, and that a close approximation to this original condition still persists in the monotremes, especially Ornithorhynchus. The tympano-hyal is the characteristic mammalian element in this region; but the entotympanic likewise appears to be peculiar to the class, and to be unrepresented among the lower vertebrates. The tympanum itself has been regarded as representing one of the elements—probably the supra-angular—of the compound reptilian lower jaw. The presence of only seven vertebrae in the neck is a very constant feature among mammals; the exceptions being very few.
Two other points in connexion with mammalian osteology may be noticed. A large number of mammals possess a perforation, or foramen, on the inner side of the lower end of the humerus, and also a projection on the shaft of the femur known as the third trochanter. From its occurrence in so many of the lower vertebrates, the entepicondylar foramen of the humerus, as it is called, is regarded by Dr E. Stromer as a primitive structure, of which the original object was to protect certain nerves and blood-vessels. It is remarkable that it should persist in the spectacled bear of the Andes, although it has disappeared in all other living members of the group. The third trochanter of the femur, on the other hand, can scarcely be regarded as primitive, seeing that it is absent in several of the lower groups of mammals. Neither can its presence be attributed, as Professor A. Gaudry suggests, to the reduction in the number of the toes, as otherwise it should not be found in the rhinoceros. Its general absence in man forbids the idea of its having any connexion with the upright posture.
Hair.—In the greater number of mammals the skin is more or less densely clothed with a peculiarly modified form of epidermis known as hair. This consists of hard, elongated, slender, cylindrical or tapering, thread-like masses of epidermic tissue, each of which grows, without branching, from a short prominence, or papilla, sunk at the bottom of a pit, or follicle, in the true skin, or dermis. Such hairs, either upon different parts of the skin of the same species, or in different species, assume very diverse forms and are of various sizes and degrees of rigidity—as seen in the fur of the mole, the bristles of the pig, and the spines of the hedgehog and porcupine, which are all modifications of the same structures. These differences arise mainly from the different arrangement of the constituent elements into which the epidermal cells are modified. Each hair is composed usually of a cellular pithy internal portion, containing much air, and a denser or more horny external or cortical part. In some mammals, as deer, the substance of the hair is almost entirely composed of the central medullary or cellular substance, and is consequently very easily broken; in others the horny part prevails almost exclusively, as in the bristles of the wild boar. In the three-toed sloth (Bradypus) the hairs have a central horny axis and a pithy exterior. Though generally nearly smooth, or but slightly scaly, the surface of some hairs is imbricated; that is to say, shows projecting scale-like processes, as in some bats, while in the two-toed sloth (Choloepus) they are longitudinally grooved or fluted. Though usually more or less cylindrical or circular in section, hairs are often elliptical or flattened, as in the curly-haired races of men, the terminal portion of the hair of moles and shrews, and conspicuously in the spines of the spiny squirrels of the genus Xerus and those of the mouse-like Platacanthomys. Hair having a property of mutual cohesion or “felting,” which depends upon a roughened scaly surface and a tendency to curl, as in domestic sheep, is called “wool.”
It has been shown by J. C. H. de Meijere that the insertion of the individual hairs in the skin displays a definite arrangement, constant for each species, but varying in different groups. In jerboas, for example, a bunch of twelve or thirteen hairs springs from the same point, while in the polar bear a single stout hair and several slender ones arise together, and in the marmosets three equal-sized hairs form regular groups. These tufts or groups likewise display an orderly and definite grouping in different mammals, which suggests the origin of such groups from the existence in primitive mammals of a scaly coat comparable to that of reptiles, and indeed directly inherited therefrom.
In a large proportion of mammals there exist hairs of two distinct types: the one long, stiff, and alone appearing on the surface, and the other shorter, finer and softer, constituting the under-fur, which may be compared to the down of birds. A well-known example is furnished by the fur-bearing seals, in which the outer fur is removed in the manufacture of commercial “seal-skin,” leaving only the soft and fine under-fur.
Remarkable differences in the direction or slope of the hair are noticeable on different parts of the body and limbs of many mammals, especially in certain apes, where the hair of the fore-limbs is inclined towards the elbow from above and from below. More remarkable still is the fact that the direction of the slope often differs in closely allied groups, as, for instance, in African and Asiatic buffaloes, in which the hair of the middle line of the back has opposite directions. Whorls of hair, as on the face of the horse and the South American deer known as brockets, occur where the different hair-slopes meet. In this connexion reference may be made to patches or lines of long and generally white hairs situated on the back of certain ruminants, which are capable of erection during periods of excitement, and serve, apparently, as “flags” to guide the members of a herd in flight. Such are the white chrysanthemum-like patches on the rump of the Japanese deer and of the American prong-buck (Antilocapra), and the line of hairs situated in a groove on the loins of the African spring-buck. The white underside of the tail of the rabbit and the yellow rump-patch of many deer are analogous.
The eye-lashes, or ciliae, are familiar examples of a special local development of hair. Special tufts of stout stiff hairs, sometimes termed vibrissae, and connected with nerves, and in certain cases with glands, occur in various regions. They are most common on the head, while they constitute the “whiskers,” or “feelers,” of the cats and many rodents. In other instances, notably in the lemurs, but also in certain carnivora, rodents and marsupials, they occupy a position on the fore-arm near the wrist, in connexion with glands, and receive sensory powers from the radial nerve. In some mammals the hairy covering is partial and limited to particular regions; in others, as the hippopotamus and the sea-cows, or Sirenia, though scattered over the whole surface, it is extremely short and scanty; but in none is it reduced to so great an extent as in the Cetacea, in which it is limited to a few small bristles confined to the neighbourhood of the lips and nostrils, and often present only in the young, or even the foetal condition.
Some kinds of hairs, as those of the mane and tail of the horse, persist throughout life, but more generally, as in the case of the body-hair of the same animal, they are shed and renewed periodically, generally annually. Many mammals have a longer hairy coat in winter, which is shed as summer comes on; and some few, which inhabit countries covered in winter with snow, as the Arctic fox, variable hare and ermine, undergo a complete change of colour in the two seasons, being white in winter and grey or brown in summer. There has been much discussion as to whether this winter whitening is due to a change in the colour of the individual hairs or to a change of coat. It has, however, been demonstrated that the senile whitening of human hair is due to the presence of phagocytes, which devour the pigment-bodies; and from microscopic observations recently made by the French naturalist Dr E. Trouessart, it appears that much the same kind of action takes place in the hairs of mammals that turn white in winter. Cold, by some means or other, causes the pigment-bodies to shift from the normal positions, and to transfer themselves to other layers of the hair, where they are attacked and devoured by phagocytes. The winter whitening of mammals is, therefore, precisely similar to the senile bleaching of human hair, no shift of the coat taking place. Under the influence of exposure to intense cold a small mammal has been observed to turn white in a single night, just as the human hair has been known to blanch suddenly under the influence of intense emotion, and in both cases extreme activity of the phagocytes is apparently the inducing cause. The African golden-moles (Chrysochloris), the desmans or water-moles (Myogale), and the West African Potamogale velox, are remarkable as being the only mammals whose hair reflects those iridescent tints so common in the feathers of tropical birds.
The principal and most obvious purpose of the hairy covering is to protect the skin. Its function in the hairless Cetacea is discharged by the specially modified and thickened layer of fatty tissue beneath the skin known as “blubber.”
Scales, &c.—True scales, or flat imbricated plates of horny material, covering the greater part of the body, are found in one family only of mammals, the pangolins or Manidae; but these are also associated with hairs growing from the intervals between the scales or on the parts of the skin not covered by them. Similarly imbricated epidermic productions form the covering of the under-surface of the tail of the African flying rodents of the family Anomaluridae; and flat scutes, with the edges in apposition, and not overlaid, clothe both surfaces of the tail of the beaver, rats and certain other members of the rodent order, and also of some insectivora and marsupials. Armadillos alone possess an external bony skeleton, composed of plates of bony tissue, developed in the skin and covered with scutes of horny epidermis. Other epidermic appendages are the horns of ruminants and rhinoceroses—the former being elongated, tapering, hollow caps of hardened epidermis of fibrous structure, fitting on and growing from conical projections of the frontal bones and always arranged in pairs, while the latter are of similar structure, but without any internal bony support, and situated in the middle line. Callosities, or bare patches covered with hardened and thickened epidermis, are found on the buttocks of many apes, the breast of camels, the inner side of the limbs of Equidae, the grasping under-surface of the tail of prehensile-tailed monkeys, opossums, &c. The greater part of the skin of the one-horned Asiatic rhinoceros is immensely thickened and stiffened by an increase of the tissue of both the skin and epidermis, constituting the well-known jointed “armour-plated” hide of those animals.
Nails, Claws and Hoofs.—With few exceptions, the terminal extremities of the digits of both limbs of mammals are more or less protected or armed by epidermic plates or sheaths, constituting the various forms of nails, claws or hoofs. These are absent in the Cetacea alone. A perforated spur, with a special secreting gland in connexion with it, is found attached to each hind-leg of the males of the existing species of Monotremata.
Scent-glands, &c.—Besides the universally distributed sweat-glands connected with the hair-system, most mammals have special glands in modified portions of the skin, often involuted to form a shallow recess or a deep sac with a narrow opening, situated in various parts of the surface of the body, and secreting odorous substances, by the aid of which individuals recognize one another. These probably afford the principal means by which wild animals are able to become aware of the presence of other members of the species, even at great distances.
To this group of structures belong the suborbital face-gland, “larmier,” or “crumen,” of antelopes and deer, the frontal gland of the muntjak and of bats of the genus Phyllorhina, the chin-gland of the chevrotains and of Taphozous and certain other bats, the glandular patch behind the ear of the chamois and the reed-buck, the glands on the lower parts of the legs of most deer and a few antelopes (the position of which is indicated by tufts of long and often specially coloured hair), the interdigital foot-glands of goats, sheep, and many other ruminants, the temporal gland of elephants, the lateral glands of the musk-shrew, the gland on the back of the hyrax and the peccary (from the presence of which the latter animal takes the name Dicotyles), the gland on the tails of the members of the dog-tribe, the preputial glands of the musk-deer and beaver (both well known for the use made of their powerfully odorous secretion in perfumery), and also of the swine and hare, the anal glands of Carnivora, the perineal gland of the civet (also of commercial value), the caudal glands of the fox and goat, the gland on the wing-membrane of bats of the genus Saccopteryx, the post-digital gland of the rhinoceros, &c. Very generally these glands are common to both sexes, and it is in such cases that their function as a means of mutual recognition is most evident. It has been suggested that the above-mentioned callosities or “chestnuts” on the limbs of horses are vestigial scent-glands; and it is noteworthy that scrapings or shavings from their surface have a powerful attraction for other horses, and are also used by poachers and burglars to keep dogs silent. The position of such glands on the lower portions of the limbs is plainly favourable to a recognition-taint being left in the tracks of terrestrial animals; and antelopes have been observed deliberately to rub the secretion from their face-glands on tree-trunks. When glands are confined to the male, their function is no doubt sexual; the secretion forming part of the attraction, or stimulus, to the other sex.
Fig. 1.—Upper and Lower Teeth of one side of the Mouth of a Dolphin (Lagenorhynchus), as an example of the homoeodont type of dentition. The bone covering the outer side of the roots of the teeth has been removed to show their simple character.
Dentition.—In the great majority of mammals the teeth form a definite series, of which the hinder elements are of a more or less complex type, while those in front are simpler. With the exception of the marsupials, a set of deciduous, or milk, teeth is developed in most mammals with a complicated type of dentition; these milk-teeth being shed at a comparatively early period (occasionally even in utero), when they are succeeded by the larger permanent series, which is the only other ever developed. This double series of teeth thus forms a very characteristic feature of mammals generally. Both the milk and the permanent dentition display the aforesaid complexity of the hinder teeth as compared with those in front, and since the number of milk-teeth is always considerably less than that of the permanent set, it follows that the hinder milk-teeth are usually more complex than the teeth of which they are the predecessors in the permanent series, and represent functionally, not their immediate successors, but those more posterior permanent teeth which have no direct predecessors. This character is clearly seen in those animals in which the various members of the lateral or cheek series are well differentiated from each other in form, as the Carnivora, and also in man.
In mammals with two sets of teeth the number of those of the permanent series preceded by milk-teeth varies greatly, being sometimes, as in marsupials and some rodents, as few as one on each side of each jaw, and in other cases including the larger portion of the series. As a rule, the teeth of the two sides of the jaws are alike in number and character, except in cases of accidental or abnormal variation, and in the tusks of the narwhal, in which the left is of immense size, and the right rudimentary. In mammals, such as dolphins and some armadillos, which have a large series of similar teeth, not always constant in number in different individuals, there may indeed be differences in the two sides; but, apart from these in describing the dentition of any mammal, it is generally sufficient to give the number and characters of the teeth of one side only. As the teeth of the upper and the lower jaws work against each other in masticating, there is a general correspondence or harmony between them, the projections of one series, when the mouth is closed, fitting into corresponding depressions of the other. There is also a general resemblance in the number, characters and mode of succession of both series; so that, although individual teeth of the upper and lower jaws may not be in the strict sense of the term homologous parts, there is a great convenience in applying the same descriptive terms to the one which are used for the other.
The simplest dentition is that of many species of dolphin (fig. 1), in which the crowns are single-pointed, slightly curved cones, and the roots also single and tapering; so that all the teeth are alike in form from the anterior to the posterior end of the series, though it may be with some slight difference in size, those at the two extremities being rather smaller than the others. Such a dentition is called “homoeodont” (Gr. ὄμοιος, like, ὀδούς, tooth), and in the case cited, as the teeth are never changed, it is also monophyodont (Gr. μόνος, alone, single, φύειν, to generate, ὀδούς, tooth). Such teeth are adapted only for catching slippery living prey, like fish.
In a very large number of mammals the teeth of different parts of the series are more or less differentiated in character; and, accordingly, have different functions to perform. The front teeth are simple and one-rooted, and are adapted for cutting and seizing. They are called “incisors.” The back, lateral or cheek teeth, on the other hand, have broader and more complex crowns, tuberculated or ridged, and supported on two or more roots. They crush or grind the food, and are hence called “molars.” Many mammals have, between these two sets, a tooth at each corner of the mouth, longer and more pointed than the others, adapted for tearing or stabbing, or for fixing struggling prey. From the conspicuous development of such teeth in the Carnivora, especially the dogs, they have received the name of “canines.” A dentition with its component parts so differently formed that these distinctive terms are applicable to them is called heterodont (Gr. ἕτερος, different). In most cases, though by no means invariably, mammals with a heterodont dentition are also diphyodont (Gr. διφυής, of double form).
This general arrangement is obvious in a considerable number of mammals; and examination shows that, under great modifications in detail, there is a remarkable uniformity of essential characters in the dentition of a large number of members of the class belonging to different orders and not otherwise closely allied, so much that it is possible to formulate a common plan of dentition from which the others have been derived by the alteration of some and the suppression of other members of the series, and occasionally, but very rarely, by addition. In this generalized form of mammalian dentition the total number of teeth present is 44, or 11 above and 11 below on each side. Those of each jaw are placed in continuous series without intervals between them; and, although the anterior teeth are simple and single-rooted, and the posterior teeth complex and with several roots, the transition between the two kinds is gradual.
In dividing and grouping such teeth for the purpose of description and comparison more definite characters are required than those derived merely from form or function. The first step towards a classification rests on the fact that the upper jaw is composed of two bones, the premaxilla and the maxilla, and that the division or suture between these bones separates the three front teeth from the rest. These three teeth, which are implanted in the premaxilla, form a distinct group, to which the name of “incisor” is applied. This distinction is, however, not so important as it appears at first sight, for their connexion with the bone is only of a secondary nature, and, although it happens conveniently that in the great majority of cases the division between the bones coincides with the interspace between the third and fourth tooth of the series, still, when it does not, as in the mole, too much weight must not be given to this fact, if it contravenes other reasons for determining the homologies of the teeth. The eight remaining teeth of the upper jaw offer a natural division, inasmuch as the three hindmost never have milk-predecessors; and, although some of the anterior teeth may be in the same case, the particular one preceding these three always has such a predecessor. These three, then, are grouped as the “molars.” Of the five teeth between the incisors and molars the most anterior, or the one usually situated close behind the pre-maxillary suture, very generally assumes a lengthened and pointed form, and constitutes the “canine” of the Carnivora, the tusk of the boar, &c. It is customary, therefore, to call this tooth, whatever its size or form, the “canine.” The remaining four are the “premolars.” This system has been objected to as artificial, and in many cases not descriptive, the distinction between premolars and canine especially being sometimes not obvious; but the terms are now in such general use, and also so convenient, that it is not likely they will be superseded. It is frequently convenient to refer to all the teeth behind the canine as the “cheek-teeth.”
With regard to the lower teeth the difficulties are greater, owing to the absence of any suture corresponding to that which defines the incisors above; but since the number of the teeth is the same, since the corresponding teeth are preceded by milk-teeth, and since in the large majority of cases it is the fourth tooth of the series which is modified in the same way as the canine (or fourth tooth) of the upper jaw, it is reasonable to adopt the same divisions as with the upper series, and to call the first three, which are implanted in the part of the mandible opposite to the premaxilla, the incisors, the next the canine, the next four the premolars, and the last three the molars.
It may be observed that when the mouth is closed, especially when the opposed surfaces of the teeth present an irregular outline, the corresponding upper and lower teeth are not exactly opposite, otherwise the two series could not fit into one another, but as a rule the points of the lower teeth shut into the interspaces in front of the corresponding teeth of the upper jaw. This is very distinct in the canine teeth of the Carnivora, and is a useful guide in determining the homologies of the teeth of the two jaws.
For the sake of brevity the complete dentition is described by the following formula, the numbers above the line representing the teeth of the upper, those below the line those of the lower jaw: incisors 3—3, canines 1—1, premolars 4—4, molars, 3—3 = 11—11; total 44. As, however, initial letters may be substituted for the names of each group, and it is unnecessary to give more than the numbers of the teeth on one side of the mouth, the formula may be abbreviated into:
i 3, c 1, p 4, m 3; total 44.
The individual teeth of each group are enumerated from before backwards, and by such a formula as the following:—
i 1, i 2, i 3, c, p 1, p 2, p 3, p 4, m 1, m 2, m 3
a special numerical designation is given by which each one can be indicated. In mentioning any single tooth, such a sign as m1 the first lower molar, and so on.will mean the first upper molar,
When, as is the case among nearly all existing mammals with the exception of the members of the genera Sus (pigs), Gymnura (rat-shrew), Talpa (moles) and Myogale (desmans) the number of teeth is reduced below the typical forty-four, it appears to be an almost universal rule that if one of the incisors is missing it is the second, or middle one, while the premolars commence to disappear from the front end of the series and the molars from the hinder end.
The milk-dentition is expressed by a similar formula, d for deciduous, being added before the letter expressive of the nature of the tooth. As the three molars and (almost invariably) the first premolar of the permanent series have no predecessors, the typical milk-dentition would be expressed as follows: di 3, dc 1, dm 3 = 28. The teeth which precede the premolars of the permanent series are called either milk-molar or milk-premolar. When there is a marked difference between the premolars and molars of the permanent dentition, the first milk-molar resembles a premolar, while the last has the characters of the posterior molar. It is sometimes convenient to refer to all the seven cheek-teeth as members of a single continuous series (which they undoubtedly are), and for this purpose the following nomenclature has been proposed:—
|Upper Jaw.||Lower Jaw.|
With the exception of the Cetacea, most of the Edentata, and the Sirenia, in which the teeth, when present, have been specialized in a retrograde or aberrant manner, the placental mammals as a whole have a dentition conforming more or less closely to the foregoing type.
With the marsupials the case is, however, somewhat different; the whole number not being limited to 44, owing largely to the fact that the number of upper incisors may exceed three pairs, reaching indeed in some instances to as many as five. Moreover, with the exception of the wombats, the number of pairs of incisors in the upper always exceeds those in the lower. When fully developed, the number of cheek-teeth is, however, seven; and it is probable that, as in placentals, the first four of these are premolars and the remaining three molars, although it was long held that these numbers should be transposed. The most remarkable feature about the marsupial dentition is that, at most, only a single pair of teeth is replaced in each jaw; this pair, on the assumption that there are four premolars, representing the third of that series. With the exception of this replacing pair of teeth in each jaw, it is considered by many authorities that the marsupial dentition corresponds to the deciduous, or milk, dentition of placentals. If this be really the case, the rudiments of an earlier set of teeth which have been detected in the jaws of some members of the order, represent, not the milk-series, but a prelacteal dentition. On the assumption that these functional teeth correspond to the milk-series of placentals, marsupials in this respect agree exactly with modern elephants, in which the same peculiarity exists.
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, although they never rise above the gums, and completely disappear before birth. In the American anteaters and the pangolins among the Edentata no traces of teeth have been found at any age. Adult monotremes are in like case, although the duck-billed platypus (Ornithorhynchus) has teeth when young on the sides of the jaws. The northern sea-cow (Rhytina), now extinct, appears to have been toothless throughout life.
In different groups of mammals the dentition is variously specialized in accordance with the nature of the food on which the members of these groups subsist. From this point of view the various adaptive modifications of mammalian dentition may be roughly grouped under the headings of piscivorous, carnivorous, insectivorous, omnivorous and herbivorous.
The fish-eating, or piscivorous, type of dentition is exemplified under two phases in the dolphins and in the seals (being in the latter instance a kind of retrograde modification from the carnivorous type). In the dolphins, and in a somewhat less marked degree among the seals, this type of dentition consists of an extensive series of conical, nearly equal-sized, sharp-pointed teeth, implanted in an elongated and rather narrow mouth (fig. 1), and adapted to seize slippery prey without either tearing or masticating. In the dolphins the teeth form simple cones, but in the seals they are often trident-like; while in the otters the dentition differs but little from the ordinary carnivorous type.
This carnivorous adaptation, in which the function is to hold and kill struggling animals, often of large size, attains its highest development in the cats (Felidae). The canines are in consequence greatly developed, of a cutting and piercing type, and from their wide separation in the mouth give a firm hold; the jaws being as short as is consistent with the free action of the canines, or tusks, so that no power is lost. The incisors are small, so as not to interfere with the penetrating action of the tusks; and the crowns of some of the teeth of the cheek-series are modified into scissor-like blades, in order to rasp off the flesh from the bones, or to crack the bones themselves, while the later teeth of this series tend to disappear.
In the insectivorous type, as exemplified in moles and shrew-mice, the middle pair of incisors in each jaw are long and pointed so as to have a forceps-like action for seizing insects, the hard coats of which are broken up by the numerous sharp cusps surmounting the cheek-teeth.
In the omnivorous type, as exemplified in man and monkeys, and to a less specialized degree in swine, the incisors are of moderate and nearly equal size; the canines, if enlarged, serve for other purposes than holding prey, and such enlargement is usually confined to those of the males; while the cheek-teeth have broad flattened crowns surmounted by rounded bosses, or tubercles.
In the herbivorous modification, as seen in three distinct phases in the horse, the kangaroo, and in ruminants, the incisors are generally well developed in one or both jaws, and have a nipping action, either against one another or against a toothless hard pad in the upper jaw; while the canines are usually small or absent, at least in the upper jaw, but in the lower jaw may be approximated and assimilated to the incisors. The cheek-teeth are large, with broad flattened crowns surmounted either by simple transverse ridges, or complicated by elevations and infoldings. In the specialized forms the premolars tend to become more or less completely like the molars; and, contrary to what obtains among the Carnivora, the whole series of cheek-teeth (with the occasional exception of the first) is very strongly developed.
Opinions differ as to the mode in which the more complicated cheek-teeth of mammals have been evolved from a simpler type of tooth. According to one theory, this has been brought about by the fusion of two or more teeth of a simple conical type to form a compound tooth. A more generally accepted view—especially among palaeontologists—is the tritubercular theory, according to which the most generalized type of tooth consists of three cusps arranged in a triangle, with the apex pointing inwards in the teeth of the upper jaw. Additions of extra cusps form teeth of a more complicated type. Each cusp of the primitive triangle has received a separate name, both in the teeth of the upper and of the lower jaw, while names have also been assigned to super-added cusps. Molar teeth of the simple tritubercular type persist in the golden moles (Chrysochloris) among the Insectivora and also in the marsupial mole (Notoryctes) among the marsupials. The type is, moreover, common among the mammals of the early Eocene, and still more so in those of the Jurassic epoch; this forming one of the strongest arguments in favour of the tritubercular theory. (See Professor H. F. Osborn, “Palaeontological Evidence for the Original Tritubercular Theory,” in vol. xvii. (new series) of the American Journal of Science, 1904.)
Digestive System.—As already mentioned, mammals are specially characterized by the division of the body-cavity into two main chambers, by means of the horizontal muscular partition known as the diaphragm, which is perforated by the great blood-vessels and the alimentary tube. The mouth of the great majority of mammals is peculiar for being guarded by thick fleshy lips, which are, however, absent in the Cetacea; their principal function being to seize the food, for which purpose they are endowed, as a rule, with more or less strongly marked prehensile power. The roof of the mouth is formed by the palate, terminating behind by a muscular, contractile arch, having in man and a few other species a median projection called the uvula, beneath which the mouth communicates with the pharynx. The anterior part of the palate is composed of mucous membrane tightly stretched over the flat or slightly concave bony layer which separates the mouth from the nasal passages, and is generally raised into a series of transverse ridges, which sometimes, as in ruminants, attain a considerable development. In the floor of the mouth, between the two branches of the lower jaw, and supported behind by the hyoid apparatus, lies the tongue, an organ the free surface of which, especially in its posterior part, is devoted to the sense of taste, but which by reason of its great mobility (being composed almost entirely of muscular fibres) performs important mechanical functions connected with masticating and procuring food. Its modifications of form in different mammals are numerous. Between the long, extensile, worm-like tongue of the anteaters, essential to the peculiar mode of feeding of those animals, and the short, immovable and almost functionless tongue of the porpoise, every intermediate condition is found. Whatever the form, the upper surface is, however, covered with numerous fine papillae, in which the terminal filaments of the taste-nerve are distributed. In some mammals, notably lemurs, occurs a hard structure known as the sublingua, which may terminate in a free horny tip. If, as has been suggested, this organ represents the tongue of reptiles, the mammalian tongue will obviously be a super-added organ distinctive of the class.
Fig. 3.—Diagrammatic Plan of the general arrangement of the Alimentary Canal in a typical Mammal.
Salivary glands, of which the most constant are the parotid and the submaxillary, are always present in terrestrial mammals. Next in constancy are the “sublingual,” closely associated with the last-named, at all events in the locality in which the secretion is poured out; and the “zygomatic,” found only in some mammals in the cheek, just under cover of the anterior part of the zygomatic arch, the duct entering the mouth-cavity near that of the parotid.
The alimentary, or intestinal, canal varies greatly in relative length and capacity in different mammals, and also offers manifold peculiarities of form, being sometimes a simple cylindrical tube of nearly uniform calibre throughout, but more often subject to alterations of form and capacity in different portions of its course—the most characteristic and constant being the division into an upper and narrower and a lower and wider portion, called respectively the small and the large intestine; the former being arbitrarily divided into duodenum, jejunum and ileum, and the latter into colon and rectum. One of the most striking peculiarities of this part of the canal is the frequent presence of a blind pouch, “caecum,” situated at the junction of the large and the small intestine. Their structure presents an immense variety of development, from the smallest bulging of a portion of the side-wall of the tube to a huge and complex sac, greatly exceeding in capacity the remainder of the alimentary canal. It is only in herbivorous mammals that the caecum is developed to this great extent, and among these there is a complementary relationship between the size and complexity of the organ and that of the stomach. Where the latter is simple the caecum is generally the largest, and vice versa. In vol. xvii. (1905) of the Transactions of the Zoological Society of London, Dr P. Chalmers Mitchell has identified the paired caeca, or blind appendages, of the intestine of birds with the usually single caecum of mammals. These caeca occur in birds (as in mammals) at the junction of the small with the large intestine; and while in ordinary perching-birds they are reduced to small nipple-like buds of no functional importance, in many other birds—owls for instance—they form quite long receptacles. Among mammals, the horse and the dog may be cited as instances where the single caecum is of large size, this being especially the case of the former, where it is of enormous dimensions; in human beings, on the other hand, the caecum is rudimentary, and best known in connexion with “appendicitis.” The existence of paired caeca was previously known in a few armadillos and anteaters, but Dr Mitchell has shown that they are common in these groups, while he has also recorded their occurrence in the hyrax and the manati. With the aid of these instances of paired caeca, coupled with the frequent existence of a rudiment of its missing fellow when only one is functional, the author has been enabled to demonstrate conclusively that these double organs in birds correspond in relations with their normally single representative in mammals.
In mammals both caecum and colon are often sacculated, a disposition caused by the arrangement of the longitudinal bands of muscular tissue in their walls; but the small intestine is always smooth and simple-walled externally, though its lining membrane often exhibits contrivances for increasing the absorbing surface without adding to the general bulk of the organ, such as the numerous small tags, or “villi,” by which it is everywhere beset, and the more obvious transverse, longitudinal, or reticulating folds projecting into the interior, met with in many animals, of which the “valvulae conniventes” of man form well-known examples. Besides the crypts of Lieberkühn found throughout the intestinal canal, and the glands of Brunner confined to the duodenum, there are other structures in the mucous membrane, about the nature of which there is still much uncertainty, called “solitary” and “agminated” glands, the latter more commonly known by the name of “Peyer’s patches.” Of the liver little need be said, except that in all living mammals it has been divided into a number of distinct lobes, which have received separate names. It has, indeed, been suggested that in the earlier mammals the liver was a simple undivided organ. This, however, is denied by G. Ruge (vol. xxix. of Gegenbaur’s Morphologisches Jahrbuch).
Origin of Mammals.—That mammals have become differentiated from a lower type of vertebrates at least as early as the commencement of the Jurassic period is abundantly testified by the occurrence of the remains of small species in strata of that epoch, some of which are mentioned in the articles Marsupialia and Monotremata (q.v.). Possibly mammalian remains also occur in the antecedent Triassic epoch, some palaeontologists regarding the South African Tritylodon as a mammal, while others consider that it was probably a reptile. Whatever may be the true state of the case with regard to that animal probably also holds good in the case of the approximately contemporaneous European Microlestes. Of the European Jurassic (or Oolitic) mammals our knowledge is unfortunately very imperfect; and from the scarcity of their remains it is quite probable that they are merely stragglers from the region (possibly Africa) where the class was first differentiated. It is not till the early Eocene that mammals become a dominant type in the northern hemisphere.
It is now practically certain that mammals are descended from reptiles. Dr H. Gadow, in a paper on the origin of mammals contributed to the Zeitschrift für Morphologie, sums up as follows: “Mammals are descendants of reptiles as surely as they [the latter] have been evolved from Amphibia. This does not mean that any of the living groups of reptiles can claim their honour of ancestry, but it means that the mammals have branched where the principal reptilian groups meet, and that is a long way back. The Theromorpha, especially small Theriodontia, alone show us what these creatures were like.” It may be explained that the Theromorpha, or Anomodontia, are those extinct reptiles so common in the early Secondary (Triassic) deposits of South Africa, some of which present a remarkable resemblance in their dentition and skeleton to mammals, while others come equally near amphibians. A difficulty naturally arises with regard to the fact that in reptiles the occipital condyle by which the skull articulates with the vertebral column is single, although composed of three elements, whereas in amphibians and mammals the articulation is formed by a pair of condyles. Nevertheless, according to Professor H. F. Osborn, the tripartite reptilian condyle, by the loss of its median element, has given rise to the paired mammalian condyles; so that this difficulty disappears. The fate of the reptilian quadrate bone (which is reduced to very small dimensions in the Anomodontia) has been referred to in an earlier section of the present article, where some mention has also been made of the disappearance in mammals of the hinder elements of the reptilian lower jaw, so as to leave the single bone (dentary) of each half of this part of the skeleton in mammals.
Most of the earliest known mammals appear to be related to the Marsupialia and Insectivora. Others however (inclusive of Tritylodon and Microlestes, if they be really mammals), seem nearer to the Monotremata; and the question has yet to be decided whether placentals and marsupials on the one hand, and monotremes on the other are not independently derived from reptilian ancestors.
With regard to the evolution of marsupials and placentals, it has been pointed out that the majority of modern marsupials exhibit in the structure of their feet traces of the former opposability of the thumb and great toe to the other digits; and it has accordingly been argued that all marsupials are descended from arboreal ancestors. This doctrine is now receiving widespread acceptation among anatomical naturalists; and in the American Naturalist for 1904, Dr W. D. Matthew, an American palaeontologist, considers himself provisionally justified in so extending it as to include all mammals. That is to say, he believes that, with the exception of the duckbill and the echidna, the mammalian class as a whole can lay claim to descent from small arboreal forms. This view is, of course, almost entirely based upon palaeontological considerations; and these, in the author’s opinion, admit of the conclusion that all modern placental and marsupial mammals are descended from a common ancestral stock, of which the members were small in bodily size. These ancestral mammals, in addition to their small size, were characterized by the presence of five toes to each foot, of which the first was more or less completely opposable to the other four. The evidence in favour of this primitive opposability is considerable. In all the groups which are at present arboreal, the palaeontological evidence goes to show that their ancestors were likewise so; while since, in the case of modern terrestrial forms, the structure of the wrist and ankle joints tends to approximate to the arboreal type, as we recede in time, the available evidence, so far as it goes, is in favour of Dr Matthew’s contention.
The same author also discusses the proposition from another standpoint, namely, the condition of the earth’s surface in Cretaceous times. His theory is that in the early Cretaceous epoch the animals of the world were mostly aerial, amphibious, aquatic or arboreal; the flora of the land being undeveloped as compared with its present state. On the other hand, towards the close of the Cretaceous epoch (when the Chalk was in course of deposition), the spread of a great upland flora vastly extended the territory available for mammalian life. Accordingly, it was at this epoch that the small ancestral insectivorous mammals first forsook their arboreal habitat to try a life on the open plains, where their descendants developed on the one hand into the carnivorous and other groups, in which the toes are armed with nails or claws, and on the other into the hoofed group, inclusive of such monsters as the elephant and the giraffe. The hypothesis is not free from certain difficulties, one of which will be noticed later.
Classification.—Existing mammals may be primarily divided into three main groups, or subclasses, of which the second and third are much more closely related to one another than is either of them to the first. These three classes are the Monotremata (or Prototheria), the Marsupialia (Didelphia, or Metatheria), and the Placentalia (Monodelphia, or Eutheria); the distinctive characters of each being given in separate articles (see Monotremata, Marsupialia and Monodelphia.)
The existing monotremes and marsupials are each represented only by a single order; but the placentals are divided into the following ordinal and subordinal groups, those which are extinct being marked with an asterisk (*):—
|1.||Insectivora (Moles, Hedgehogs, &c.).|
|3.||Dermoptera (Colugo, or Flying Lemur).|
|a.||Xenarthra (Anteaters, Sloths and Armadillos).|
|c.||Tubulidentata (Ant-bears, or Aarvarks). |
|5.||Rodentia (Gnawing Mammals):—|
|a.||Duplicidentata (Hares and Picas).|
|b.||Simplicidentata (Rats, Beavers, &c.).|
|a.||Fissipedia (Cats, Dogs, Bears, &c.).|
|b.||Pinnipedia (Seals and Walruses).|
|c.||*Creodonta (Hyaenodon, &c.).|
|8.||Cetacea (Whales and Dolphins):—|
|a.||*Archaeoceti (Zeuglodon, &c.).|
|b.||Odontoceti (Spermwhales and Dolphins).|
|c.||Mystacoceti (Whalebone Whales).|
|9.||Sirenia (Dugongs and Manatis).|
|10.||Ungulata (Hoofed Mammals):—|
|a.||Proboscidea (Elephants and Mastodons).|
|d.||*Toxodontia (Toxodon, &c.).|
|e.||*Amblypoda (Uintatherium, &c.).|
|f.||*Litopterna (Macrauchenia, &c.).|
|g.||*Ancylopoda (Chalicotherium, &c.).|
|h.||*Condylarthra (Phenacodus, &c.).|
|i.||Perissodactyla (Tapirs, Horses, &c.).|
|j.||Artiodactyla (Ruminants, Swine, &c.).|
|a.||Prosimiae (Lemurs and Galagos).|
|b.||Anthropoidea (Monkeys, Apes and Man).|
Separate articles are devoted to each of these orders, where references will be found to other articles dealing with some of the minor groups and a number of the more representative species.
Relationships of the Groups.—As we recede in time we find the extinct representatives of many of these orders approximating more and more closely to a common generalized type, so that in a large number of early Eocene forms it is often difficult to decide to which group they should be assigned.
The Insectivora are certainly the lowest group of existing placental mammals, and exhibit many signs of affinity with marsupials; they may even be a more generalized group than the latter. From the Insectivora the bats, or Chiroptera, are evidently a specialized lateral offshoot; while the Dermoptera may be another branch from the same stock. As to the Edentata, it is still a matter of uncertainty whether the pangolins (Pholidota) and the ant-bears (Tubulidentata) are rightly referred to an order typically represented by the sloths, anteaters, and armadillos of South and Central America, or whether the two first-named groups have any close relationship with one another. Much uncertainty prevails with regard to the ancestry of the group as a whole, although some of the earlier South American forms have a comparatively full series of teeth, which are also of a less degenerate type than those of their modern representatives.
An almost equal degree of doubt obtains with regard to the ancestry of that very compact and well-defined group the Rodentia. If, however, the so-called Proglires of the lower Eocene are really ancestral rodents, the order is brought into comparatively close connexion with the early generalized types of clawed, or unguiculate mammals. Whether the extinct Tillodontia are most nearly allied to the Rodentia, the Carnivore or the Ungulata, and whether they are really entitled to constitute an ordinal group by themselves, must remain for the present open questions.
The Carnivora, as represented by the (mainly) Eocene Creodonta, are evidently an ancient and generalized type. As regards the number and form of their permanent teeth, at any rate, creodonts present such a marked similarity to carnivorous marsupials, that it is difficult to believe the two groups are not allied, although the nature of the relationship is not yet understood, and the minute internal structure of the teeth is unlike that of marsupials and similar to that of modern Carnivora. There is the further possibility that creodonts may be directly descended from the carnivorous reptiles; a descent which if proved might introduce some difficulty with regard to the above-mentioned theory as to the arboreal ancestry of mammals generally. Be this as it may, there can be little doubt that the creodonts are related to the Insectivora, which, as stated above, show decided signs of kinship with the marsupials.
A much more interesting relationship of the creodont carnivora has, however, been established on the evidence of recent discoveries in Egypt. From remains of Eocene age in that country Dr E. Fraas, of Stuttgart, has demonstrated the derivation of the whale-like Zeuglodon from the creodonts. Dr C. E. Andrews has, moreover, not only brought forward additional evidence in favour of this most remarkable line of descent, but is confident—which Professor Fraas was not—that Zeuglodon itself is an ancestral cetacean, and consequently that whales are the highly modified descendants of creodonts. It must be admitted, however, that the links between Zeuglodon and typical cetaceans are at present unknown; but it may be hoped that these will be eventually brought to light from the deposits of the Mokattam Range, near Cairo. Whales and dolphins being thus demonstrated to be nothing more than highly modified Carnivora, might almost be included in the same ordinal group.
An analogous statement may be made with regard to the sea-cows, or Sirenia, which appear to be derivates from the great herbivorous order of Ungulata, and might consequently be included in that group, as indeed has been already done in Dr Max Weber’s classification. It is with the proboscidean suborder of the Ungulata to which the Sirenia are most nearly related; the nature of this relationship being described by Dr Andrews as follows:—
“In the first place, the occurrence of the most primitive Sirenians with which we are acquainted in the same region as the most generalized proboscidean, Moeritherium, is in favour of such a view, and this is further supported by the similarity of the brain-structure and, to some extent, of the pelvis in the earliest-known members of the two groups. Moreover, in the anatomy of the soft-parts of the recent forms there are a number of remarkable points of resemblance. Among the common characters may be noted the possession of: (1) pectoral mammae; (2) abdominal testes; (3) a bifid apex of the heart; (4) bilophodont molars with a tendency to the formation of an additional lobe from the posterior part of the cingulum. The peculiar mode of displacement of the teeth from behind forwards in some members of both groups may perhaps indicate a relationship, although in the case of the Sirenia the replacement takes place by means of a succession of similar molars, while in the Proboscidea the molars remain the same numerically, but increase greatly in size and number of transverse ridges.”
These and certain other facts referred to by the same author point to the conclusion that not only are the Sirenia and the Proboscidea derived from a single ancestral stock, but that the Hyracoidea—and so Arsinöitherium—are also derivatives from the same stock, which must necessarily have been Ethiopian.
Of the other suborders of ungulates, the Toxodontia and Litopterna are exclusively South American, and while the former may possibly be related to the Hyracoidea and Barypoda, the latter is perhaps more nearly akin to the Perissodactyla. The Amblypoda, on the other hand, are perhaps not far removed from the ancestral Proboscidea, which depart comparatively little from the generalized ungulate type. The latter is represented by the Eocene Condylarthra, which undoubtedly gave rise to the Perissodactyla and Artiodactyla, and probably to most, if not all, of the other groups. The Condylarthra, in their turn, approximate closely to the ancestral Carnivora, as they also do in some degree to the ancestral Primates. As regards the latter order, although we are at present unacquainted with all the connecting links between the lemurs and the monkeys, there is little doubt that the ancestors of the former represent the stock from which the latter have originated. C. D. Earle, in the American Naturalist for 1897, observes that “so far as the palaeontological evidence goes it is decidedly in favour of the view that apes and lemurs are closely related. Beginning with the earliest known lemur, Anaptomorphus, this genus shows tendencies towards the anthropoids, and, when we pass up into the Oligocene of the Old World, Adapis is a decidedly mixed type, and probably not far from the common stem-form which gave origin to both suborders of the Primates. In regard to Tarsius, it is evidently a type nearly between the lemurs and apes, but with many essential characters belonging to the former group.”
Distribution.—For an account of the “realms” and “regions” into which the surface of the globe has been divided by those who have made a special study of the geographical distribution of animals, see Zoological Distribution. For the purposes of such zoo-geographical divisions, mammals are much better adapted than birds, owing to their much more limited powers of dispersal; most of them (exclusive of the purely aquatic forms, such as seals, whales, dolphins and sea-cows) being unable to cross anything more than a very narrow arm of the sea. Consequently, the presence of nearly allied groups of mammals in areas now separated by considerable stretches of sea proves that at no very distant date such tracts must have had a land-connexion. In the case of the southern continents the difficulty is, however, to determine whether allied groups of mammals (and other animals) have reached their present isolated habitats by dispersal from the north along widely sundered longitudinal lines, or whether such a distribution implies the former existence of equatorial land-connexions. It may be added that even bats are unable to cross large tracts of sea; and the fact that fruit-bats of the genus Pteropus are found in Madagascar and the Seychelles, as well as in India, while they are absent from Africa, is held to be an important link in the chain of evidence demonstrating a former land-connexion between Madagascar and India.
There is another point of view from which mammals are of especial importance in regard to geographical distribution, namely their comparatively late rise and dispersal, or “radiation,” as compared with reptiles.
As regards terrestrial mammals (with which alone we are at present concerned), one of the most striking features in their distribution is their practical absence from oceanic islands; the only species found in such localities being either small forms which might have been carried on floating timber, or such as have been introduced by human agency. This absence of mammalian life in oceanic islands extends even to New Zealand, where the indigenous mammals comprise only two peculiar species of bats, the so-called Maori rat having been introduced by man.
One of the leading features in mammalian distribution is the fact that the Monotremata, or egg-laying mammals, are exclusively confined to Australia and Papua, with the adjacent islands. The marsupials also attain their maximum development in Australia (“Notogaea” of the distributionists), extending, however, as far west as Celebes and the Moluccas, although in these islands they form an insignificant minority among an extensive placental fauna, being represented only by the cuscuses (Phalanger), a group unknown in either Papua or Australia. Very different, on the other hand, is the condition of things in Australia and Papua, where marsupials (and monotremes) are the dominant forms of mammalian life, the placentals being represented (apart from bats, which are mainly of an Asiatic type) only by a number of more or less aberrant rodents belonging to the mouse-tribe, and in Australia by the dingo, or native dog, and in New Guinea by a wild pig. The dingo was, however, almost certainly brought from Asia by the ancestors of the modern natives; while the Papuan pig is also in all probability a human introduction, very likely of much later date. The origin of the Australasian fauna is a question pertaining to the article Zoological Distribution. The remaining marsupials (namely the families Didelphyidae and Epanorthidae) are American, and mainly South and Central American at the present day; although during the early part of the Tertiary period representatives of the first-named family ranged all over the northern hemisphere.
The Insectivora (except a few shrews which have entered from the north) are absent from South America, and appear to have been mainly an Old World group, the only forms which have entered North America being the shrew-mice (Soricidae) and moles (Talpidae). The occurrence of one aberrant group (Solenodon) in the West Indies is, however, noteworthy. The family with the widest distribution is the Soricidae, the Talpidae being unknown in Africa. The tree-shrews (Tupaiidae) are exclusively Asiatic, whereas the jumping-shrews (Macroscelididae) are equally characteristic of the African continent. Madagascar is the sole habitat of the tenrecs (Centetidae), as is Southern Africa of the golden moles (Chrysochloridae). It is, however, important to mention that an extinct South American insectivore, Necrolestes, has been referred to the family last mentioned; and even if this reference should not be confirmed in the future, the occurrence of a representative of the order in Patagonia is a fact of considerable importance in distribution.
The Rodentia have a wider geographical range than any other order of terrestrial mammals, being, as already mentioned, represented by numerous members of the mouse-tribe (Muridae) even in Australasia. With the remarkable exception of Madagascar, where it is represented by the Nesomyidae, that family has thus a cosmopolitan distribution. Very noteworthy is the fact that, with the exception of Madagascar (and of course Australia) the squirrel family (Sciuridae) is also found in all parts of the world. Precisely the same may be said of the hares, which, however, become scarce in South America. On the other hand, the scaly-tailed squirrels (Anomaluridae), the jumping-hares (Pedetidae), and the strand-moles (Bathyergidae) are exclusively African; while the sewellels (Haplodontidae) and the pocket-gophers (Geomyidae) are as characteristically North American, although a few members of the latter have reached Central America. The beavers (Castoridae) are restricted to the northern hemisphere, whereas the dormice (Gliridae) and the mole-rats (Spalacidae) are exclusively Old World forms, the latter only entering the north of Africa, in which continent the former are largely developed. The jerboa group (Dipodidae, or Jaculidae) is also mainly an Old World type, although its aberrant representatives the jumping-mice (Zapus) have effected an entrance into Arctic North America. Porcupines enjoy a very wide range, being represented throughout the warmer parts of the Old World, with the exception of Madagascar (and of course Australasia), by the Hystricidae, and in the New World by the Erethizontidae. Of the remaining families of the Simplicidentata, all are southern, the cavies (Caviidae), chinchillas (Chinchillidae), and degus (Octodontidae) being Central and South American, while the Capromyidae are common to southern America and Africa, and the Ctenodactylidae are exclusively African. The near alliance of all these southern families, and the absence of so many Old World families from Madagascar form two of the most striking features in the distribution of the order. Lastly, among the Duplicidentata, the picas (Ochotonidae or Lagomyidae) form a group confined to the colder or mountainous regions of the northern hemisphere.
Among the existing land Carnivora (of which no representatives except the introduced dingo are found in Australasia) the cat-tribe (Felidae) has now an almost cosmopolitan range, although it only reached South America at a comparatively recent date. Its original home was probably in the northern hemisphere; and it has no representatives in Madagascar. The civet-tribe (Viverridae), on the other hand, which is exclusively an Old World group, is abundant in Madagascar, where it is represented by peculiar and aberrant types. The hyenas (Hyaenidae), at any rate at the present day, to which consideration is mainly limited, are likewise Old World. The dog-tribe (Canidae), on the other hand, are, with the exception of Madagascar, an almost cosmopolitan group. Their place of origin was, however, almost entirely in the northern hemisphere, and not improbably in some part of the Old World, where they gave rise to the bears (Ursidae). The latter are abundant throughout the northern hemisphere, and have even succeeded in penetrating into South America, but, with the exception of the Mediterranean zone, have never succeeded in entering Africa, and are therefore of course unknown in Madagascar. The raccoon group (Procyonidae) is mainly American, being represented in the Old World only by the pandas (Aelurus and Aeluropus), of which the latter apparently exhibits some affinity to the bears. The birthplace of the group was evidently in the northern hemisphere—possibly in east Central Asia. The weasel-tribe (Mustelidae) is clearly a northern group, which has, however, succeeded in penetrating into South America and Africa, although it has never reached Madagascar.
The extinct creodonts, especially if they be the direct descendants of the anomodont reptiles, may have originated in Africa, although they are at present known in that continent only from the Fayum district. Elsewhere they occur in South America and throughout a large part of the northern hemisphere, where they appear to have survived in India to the later Oligocene or Miocene.
In the case of the great order, or assemblage, of Ungulata it is necessary to pay somewhat more attention to fossil forms, since a considerable number of groups are either altogether extinct or largely on the wane.
So far as is at present known, the earliest and most primitive group, the Condylarthra, is a northern one, but whether first developed in the eastern or the western hemisphere there is no sufficient evidence. The more or less specialized Litopterna and Toxodontia, as severally typified by the macrauchenia and the toxodon, are, on the other hand, exclusively South American. With the primitive five-toed Amblypoda, as represented by the coryphodon, we again reach a northern group, common to the two hemispheres; but there is not improbably some connexion between this group and the much more specialized Barypoda, as represented by Arsinöitherium, of Africa. The Ancylopoda, again, typified by Chalicotherium, and characterized by the claw-like character of the digits, are probably another northern group, common to the eastern and western hemispheres.
Recent discoveries have demonstrated the African origin of the elephants (Proboscidea) and hyraxes (Hyracoidea), the latter group being still indeed mainly African, and in past times also limited to Africa and the Mediterranean countries. As regards the elephants (now restricted to Africa and tropical Asia), there appears to be evidence that the ancestral mastodons, after having developed from African forms probably not very far removed from the Amblypoda, migrated into Asia, where they gave rise to the true elephants. Thence both elephants and mastodons reached North America by the Bering Sea route; while the former, which arrived earlier than the latter, eventually penetrated into South America.
The now waning group of Perissodactyla would appear to have originally been a northern one, as all the three existing families, rhinoceroses (Rhinocerotidae), tapirs (Tapiridae), and horses (Equidae), are well represented in the Tertiaries of both halves of the northern hemisphere. If eastern Central Asia were tentatively given as the centre of radiation of the group, this might perhaps best accord with the nature of the case. Rhinoceroses disappeared comparatively early from the New World, and never reached South America. In Siberia and northern Europe species of an African type survived till a comparatively late epoch, so that the present relegation of the group to tropical Asia and Africa may be regarded as a modern feature in distribution. Horses, now unknown in a wild state in the New World, although still widely spread in the Old, attained a more extensive range in past times, having successfully invaded South America. On the other hand, in common with the rest of the Perissodactyla, they never reached Madagascar. In addition to the occurrence of their fossil remains almost throughout the world, the former wide range of the tapirs is attested by the fact of their living representatives being confined to such widely sundered areas as Malaysia and tropical America.
The Artiodactyla are the only group of ungulates known to have been represented in Madagascar; but since both these Malagasy forms—namely two hippopotamuses (now extinct) and a river-hog—are capable of swimming, it is most probable that they reached the island by crossing the Mozambique Channel. As regards the deer-family (Cervidae), which is unknown in Africa south of the Sahara, it is quite evident that it originated in the northern half of the Old World, whence it reached North America by the Bering Sea route, and eventually travelled into South America. More light is required with regard to the past history of the giraffe-family (Giraffidae), which includes the African okapi and the extinct Indian Sivatherium, and is unknown in the New World. Possibly, however, its birthplace may prove to be Africa; if so, we shall have a case analogous to that of the African elephant, namely that while giraffes flourished during the Pliocene in Asia (where they may have originated), they survive only in Africa. An African origin has also been suggested for the hollow-horned ruminants (Bovidae); and if this were substantiated it would explain the abundance of that family in Africa and the absence from the heart of that continent of the deer-tribe. Some confirmation of this theory is afforded by the fact that whereas we can recognize ancestral deer in the Tertiaries of Europe we cannot point with certainty to the forerunners of the Bovidae. Whether its birthplace was in Africa or to the north, it is, however, clear that the hollow-horned ruminants are essentially an Old World group, which only effected an entrance into North America at a comparatively recent date, and never succeeded in reaching South America. So far as it goes, this fact is also in favour of the African ancestry of the group.
The Antilocapridae (prongbuck), whose relationships appear to be rather with the Cervidae than with the Bovidae, are on the other hand apparently a North American group. The chevrotains (Tragulidae), now surviving only in West and Central Africa and tropical Asia, are conversely a purely Old World group.
The camels (Tylopoda) certainly originated in the northern hemisphere, but although their birthplace has been confidently claimed for North America, an equal, if not stronger, claim may be made on the part of Central Asia. From the latter area, where wild camels still exist, the group may be assumed to have made its way at an early period into North America; whence, at a much later date, it finally penetrated into South America. In the Old World it seems to have reached the fringe of the African continent, where its wanderings in a wild state were stayed.
The pigs (Suidae) and the hippopotamuses (Hippopotamidae) are essentially Old World groups, the former of which has alone succeeded in reaching America, where it is represented by the collateral branch of the peccaries (Dicotylinae). An African origin would well explain the present distribution of both groups, but further evidence on this point is required before anything decisive can be affirmed, although it is noteworthy that the earliest known pig (Geniohyus) is African. The Suinae are at present spread all over the Old World, although the African forms (other than the one from the north) are markedly distinct from those inhabiting Europe and Asia. Hippopotamuses, on the contrary, are now exclusively African, although they were represented in tropical Asia during the Pliocene and over the greater part of Europe at a later epoch.
A brief notice with regard to the distribution of the Primates must suffice, as their past history is too imperfectly known to admit of generalizations being drawn. The main facts at the present day are, firstly, the restriction of the Prosimiae, or lemurs, to the warmer parts of the Old World, and their special abundance in Madagascar (where other Primates are wanting); and, secondly, the wide structural distinction between the monkeys of tropical America (Platyrrhina), and the Old World monkeys and apes, or Catarrhina. It is, however, noteworthy that extinct lemurs occur in the Tertiary deposits of both halves of the northern hemisphere—a fact which has induced Dr J. L. Wortman to suggest a polar origin for the entire group—a view we are not yet prepared to endorse. For the distribution of the various families and genera the reader may be referred to the article Primates; and it will suffice to mention here that while chimpanzees and baboons are now restricted to Africa and (in the case of the latter group) Arabia, they formerly occurred in India.
As regards aquatic mammals, the greater number of the Cetacea, or whales and dolphins, have, as might be expected, a very wide distribution in the ocean. A few, on the other hand, have a very restricted range, the Greenland right whale (Balaena mysticetus) being, for instance, limited to the zone of the northern circumpolar ice, while no corresponding species occurs in the southern hemisphere. In this case, not only temperature, but also the peculiar mode of feeding, may be the cause. The narwhal and the beluga have a very similar distribution, though the latter occasionally ranges farther south. The bottle-noses (Hyperöodon) are restricted to the North Atlantic, never entering, so far as known, the tropical seas. Other species are exclusively tropical or austral in their range. The pigmy whale (Neobalaena marginata), for instance, has only been met with in the seas round Australia, New Zealand and South America, while a beaked whale (Berardius arnouxi) appears to be confined to the New Zealand seas.
The Cetacea, however, are by no means limited to the ocean, or even to salt water, some entering large rivers for considerable distances, and others being exclusively fluviatile. The susu (Platanista) is, for instance, extensively distributed throughout nearly the whole of the river systems of the Ganges, Brahmaputra and Indus, ascending as high as there is water enough to swim in, but apparently never passing out to sea. The individuals inhabiting the Indus and the Ganges must therefore have been for long ages isolated without developing any distinctive anatomical characters, those by which P. indi was separated from P. gangetica having been shown to be of no constant value. Orcella fluminalis, again, appears to be limited to the Irrawaddy; and at least two distinct species of dolphin, belonging to different genera, are found in the Amazon. It is remarkable that none of the great lakes or inland seas of the world is inhabited by cetaceans.
The great difference in the manner of life of the sea-cows, or Sirenia, as compared with that of the Cetacea, causes a corresponding difference in their geographical distribution. Slow in their movements, and feeding on vegetable substances, they are confined to the neighbourhood of rivers, estuaries or coasts, although there is a possibility of accidental transport by currents across considerable distances. Of the three genera existing within historic times, one (Manatus) is exclusively confined to the shores of the tropical Atlantic and the rivers entering into it, individuals scarcely specifically distinguishable being found both on the American and the African. The dugong (Halicore) is distributed in different colonies, at present isolated, throughout the Indian Ocean from Arabia to North Australia; while the Rhytina or northern sea-cow was, for some time before its extinction, limited to a single island in the extreme north of the Pacific Ocean.
The seals (Pinnipedia) although capable of traversing long reaches of ocean, are less truly aquatic than the last two groups, always resorting to the land or to ice-floes for breeding. The geographical range of each species is generally more or less restricted, usually according to climate, as they are mostly inhabitants either of the Arctic or Antarctic seas and adjacent temperate regions, few being found within the tropics. For this reason the northern and the southern species are for the most part quite distinct. In fact, the only known exception is the case of a colony of elephant-seals (Macrorhinus leoninus), whose general range is in the southern hemisphere, inhabiting the coast of California. In this case a different specific name has been given to the northern form, but the characters by which it is distinguished are of little importance, and probably, except for the abnormal geographical distribution, would never have been discovered. The most remarkable circumstance connected with the distribution of seals is the presence of members of the order in the three isolated great lakes or inland seas of Central Asia—the Caspian, Aral and Baikal—which, notwithstanding their long isolation, have varied but slightly from species now inhabiting the Polar Ocean.
Authorities.—The above article is partly based on that of Sir W. H. Flower in the 9th edition of this work. The literature connected with mammals is so extensive that all that can be attempted here is to refer the reader to a few textbooks, with the aid of which, combined with that of the annual volumes of the Zoological Record, he may obtain such information on the subject as he may require: F. E. Beddard, “Mammals,” The Cambridge Natural History, vol. x. (1902); W. H. Flower and R. Lydekker, The Study of Mammals (London, 1891); Max Weber, Die Säugethiere (Jena, 1904); W. T. Blanford, The Fauna of British India—Mammalia (1888-1891); D. G. Elliot, Synopsis of the Mammals of North America (Chicago, 1901) and The Mammals of Middle America and the West Indies (Chicago, 1904); W. L. Sclater, The Fauna of South Africa—Mammals (Cape Town, 1901-1902); W. K. Parker, Mammalian Descent (London, 1885); E. Trouessart, Catalogus mammalium, tam viventium quam fossilium (Paris, 1898-1899); and supplement, 1904-1905; T. S. Palmer, Index generum mammalium (Washington, 1904); W. L. and P. L. Sclater, The Geography of Mammals (London, 1899); R. Lydekker, A Geographical History of Mammals (Cambridge, 1896).