having at its extremity a minute perforation, through which the vessels and nerves required to maintain the vitality of the tooth enter IV WW FIG. 1.--Diagrammatic Sections of various forms of Teeth. I. Incisor or tusk of Elephant, with pulp cavity persistently open at base. II. Human incisor during development, with root imperfectly formed, and pulp-cavity widely open at base. III. Completely formed human incisor, with pulp-cavity contracted to a small aperture at the end of the root. IV. Human molar, with broad crown and two roots. V. Molar of the Ox, with the enamel covering the crown deeply folded, and the depressions filled up with cement. The surface is worn by use; otherwise the enamel coating would be continuous at the top of the ridges. In all the figures the enamel is black, the pulp white, the dentine represented by horizontal lines, and the cement by dots. the pulp-cavity, which is tooth. The latter is therefore a more primitive condition, the formation of the root being a completion of the process of tooth development. Functionally it is, however, difficult to say that the one is a higher form than the other, since they both serve important and different purposes in the animal economy. As is almost always the case in nature, intermediate conditions between these two forms of teeth are met with. Thus some teeth, as the molars of the Horse, and of many Rodents, are for a time rootless, and have growing pulps producing very long crowns with parallel sides, the summits of which may be in use and beginning to wear away while the bases are still growing; but ultimately the pulp contracts, forms a neck and distinct roots, and ceases to grow. The canine tusks of the Musk Deer and of the Walrus have persistent pulps, and are open at their base until the animal is of advanced age, when they close, and the pulp ceases to be renewed. The same sometimes happens in the tusks of very old Boars. The simplest form of the crown of a tooth is that of a cone; but this may be variously modified. Thus it may be flattened, with its edges sharp and cutting, and pointed at the apex, as in the laterally compressed premolars of most Carnivora; or it may be chisel- or awl-shaped, with a straight truncated edge, as in the human incisors; or it may be broad, with a flat or rounded upper surface. Very often there is a more or less prominent ridge encircling the whole or part of the base of the crown just above the neck, called the cingulum, which serves as a protection to the edge of the gum in masticating, and is most developed in flesh-eating and insectivorous animals, in which the gums are liable to be injured by splinters of bone or other hard fragments of their food. The form of the crown is frequently rendered complex by the development upon its surface of elevations or tubercules called cusps or cones, or by ridges usually transverse, but sometimes variously curved or folded. When the crown is broad and the ridges are greatly developed, as in the molars of the Elephant, Horse, and Ox (Fig. 1, V.), the interspaces between them are filled with cement, which supports them and makes a solid compact mass of the whole tooth. When such a tooth wears away at the surface by friction against the opposed tooth of the other jaw, the different density of the layers of the substances of which it is composed-enamel, dentine, and cement arranged in characteristic patterns, causes them to wear unequally, the hard enamel ridges projecting beyond the others, and thus giving rise to a grinding surface of great mechanical advantage. Succession.-The dentition of all mammals consists of a definite set of teeth, almost always of constant and determinate number, form, and situation, and, with few exceptions, persisting in a functional condition throughout the natural term of the animal's life. In many species these are the only teeth which the animal ever possesses,—the set which is first formed being permanent, or, if accidentally lost, or decaying in extreme old age, not being replaced by others. These animals are called Monophyodont. But in the larger number of mammals, certain of the teeth are preceded by others, which may be only of a very transient, rudimentary, and functionless character (being in the Seals, for example, shed either before or within a few days after birth), or may be considerably developed, and functionally occupy the place of the permanent teeth for a somewhat lengthened period, during the growth and development of the latter and of the jaws. In all cases these teeth disappear (by the absorption of their roots and shedding of the crowns) before the frame of the animal has acquired complete maturity, as evidenced by the coalescence of the epiphyses of the osseous system. As these teeth are, as a general rule, present during the period in which the animal is nourished by the milk of the mother, the name of "milk-teeth" (French dents de lait, German milchzähne) has been commonly accorded to them, although it must be understood that the epoch of their presence is by no means necessarily synchronous with that of lactation. Animals possessing such teeth are called Diphyodont. No mammal is known to have more than two sets of teeth; and the definite and orderly replacement of certain members of the series is a process of quite a different nature from the indefinite succession which takes place in all the teeth continuously throughout the lifetime of the lower vertebrates. When the milk-teeth are well developed, and continue in place during the greater part of the animal's growth, as is especially the case with the Ungulata, and, though to a less degree, with the Primates and Carnivora, their use is obvious, since taken all together they form structurally a complete epitome on a small scale of the more numerous and larger permanent set (see Fig. 3), and, consequently, are able to perform the same functions, while time is allowed for the gradual maturation of the latter, and especially while the jaws of the growing animal are acquiring the size and strength sufficient to support the permanent teeth. Those animals, therefore, that have a well-developed and tolerably persistent set of milk-teeth may be considered to be in a higher state of development, as regards their dentition, than those that have the milk-teeth absent or rudimentary. It is a very general rule that individual teeth of the milk and permanent set have a close relationship to one another, being originally formed, as mentioned above, in exceedingly near proximity, and with, at all events so far as the enamel-germ is concerned, a direct connection. Moreover, since the latter ultimately come to occupy the position in the alveolar border temporarily held by the former, they are spoken of respectively as the predecessors or successors of each other. But it must be understood that milk-teeth have no successors in the permanent series, may be present which and, what is far more general, permanent teeth may have no predecessors in the milk series. The complete series of permanent teeth of most mammals forms a complex machine, with its several parts adapted for different functions, the most obvious structural modification for this purpose being an increased complexity of the individual components of the series from the anterior towards the posterior extremity of such series. Since, as has just been said, the complete series of the milk teeth often presents structurally and functionally a similar machine, but composed of fewer individual members, and the anterior of which are as simple, and the posterior as complex as those occupying corresponding positions in the permanent series, and since the milk-teeth are only developed in relation to the anterior or lateral, never to the most posterior of the permanent series,-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 molar series are well differentiated from each other in form, as the Carnivora, and also in Man. In animals which have two sets of teeth the number of those of the permanent series which are 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 sometimes including the larger portion of the series. Although there are difficulties in some cases in arriving at a satisfactory solution of the question, it is, on the whole, safest to assume that when only one set of teeth is present, this corresponds to the permanent teeth of the Diphyodonts. When this one set is completely developed, and remains in use throughout the animal's life, there can be no question on this subject. When, on the other hand, the teeth are rudimentary and transient, as in the Whalebone Whales, it is possible to consider them as representing the milk series; but there are weighty reasons in favour of the opposite conclusion.1 Arrangement, Homologies, and Notation of Teeth.-The teeth of the two sides of the jaws are always alike in number and character, 1 This and other questions concerning the homologies, notation, and succession of the teeth of mammals are more fully developed in two memoirs by one of the present writers :-" Remarks on the Homologies and Notation of the Teeth of the Mammalia," in the Journal of Anatomy and Physiology, vol. iii. p. 262, 1869; and "Notes on the First or Milk Dentition of the Mammalia," in the Trans. Odontological Society of Great Britain, 1871. See also an important memoir by Oldfield Thomas on the "Homologies and Succession of the teeth in the Dasyuride," Phil. Trans. 1887, pp. 443-462. except in cases of accidental or abnormal variation, and in the one remarkable instance of constant deviation from bilateral symmetry among mammals, the tusks of the Narwhal (Monodon), in which the left is of immense size, and the right rudimentary. In certain mammals, such as the Dolphins and some Armadillos, which have a very large series of similar teeth, not always constant in number in different individuals, there may be differences in the two sides; but, apart from these, in describing the dentition of any mammal, it is quite sufficient to give the number and characters of the teeth of one side only. Since 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 any strict sense of the term homologous parts, there is a great convenience in applying the same descriptive terms to the one as are used for the other. The simplest dentition as a whole is that of many species of Dolphin (Fig. 2), in which the crowns are single-pointed, slightly FIG. 2.-Upper and Lower Teeth of one side of the Mouth of a Dolphin (Lagenorhynchus) as an example of the homodont type of dentition. The bone covering the outer side of the roots of the teeth has been removed to show their simple character. curved cones, and the roots also single and tapering, and all 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 of the series being rather smaller than the others. Such a dentition is called Homodont, and in the case cited, as the teeth are never changed, it is also Monophyodont. Such teeth are adapted only for catching slippery living prey, as fish. In a very large number of mammals the teeth of different parts of the series are more or less differentiated in character, and 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- or cheek-teeth have broader and more complex crowns, tuberculated or ridged, and are sup |