CHAPTER V DENTITION

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In the most generalized snakes—those which show the nearest approach to lizards—teeth are present not only on the rami of both jaws, but also on the premaxillary bone, on the palatines, and on the pterygoids. A reduction of the dentition takes place in various genera, in which the teeth of either the upper or the lower jaw, and of the palatines or pterygoids, or both, may be absent, and the premaxillary is devoid of teeth in the great majority, including all European representatives, of the Ophidia.

In the egg-eating snakes of the genera Dasypeltis and Elachistodon the dentition is very much reduced, in accordance with the peculiar rÉgime, and this deficiency is compensated by the development on some of the anterior thoracic vertebrÆ of long, tooth-like processes (hypapophyses) directed forwards, and capped with a remarkably dense, vitreous tissue simulating enamel, the function of these tooth-like processes being to break the shell of the egg within the gullet, where none of its contents are lost, the shell being afterwards rejected through the mouth in the form of a pellet.

With the exception of the worm-like TyphlopidÆ, which are provided with a few teeth in the upper jaw only, European snakes have teeth on the maxillary, palatine, pterygoid, and dentary bones. Unless the maxillary be strongly abbreviated and modified in connexion with the poison apparatus, as in the ViperidÆ, the teeth in the jaws as well as on the palate form single longitudinal series; they are elongate, conical, with or without a sharp posterior edge, more or less recurved, acutely pointed, sometimes needle-like, and directed backwards, as behoves their function, which, in addition to attack and defence, is to prevent the retrogression of the prey in the act of prehension and deglutition. A notable exception occurs in the genus Iguanognathus, from Sumatra, all the teeth having spatulate crowns ribbed along the outer side. Unfortunately, nothing is known as to the food of this remarkable snake. The teeth are coated with a thin layer of enamel. It was held, for a time, that the glossy outer coating was only due to a denser structure of the dentine. As in all living Reptiles with the exception of the Crocodiles, the teeth are not implanted in true sockets, but simply ankylosed to the bone on which, when detached, their slightly enlarged base, or rather the bony tissue on which it rests, leaves a shallow impression, or pseudo-socket. In the process of biting or feeding, some of the teeth are frequently lost, and are readily replaced by others lying in reserve in the gum at the inner side, and becoming fixed to the bone soon after a vacancy occurs. Such replacement teeth, of different grades of development, form several series, so that in a snake like our common Tropidonotus the mouth may contain four times as many teeth as are functional, without reckoning different earlier stages of tooth germs which escape ordinary observation, being placed vertically one above the other.

Three types of teeth, connected by every intermediate step, are distinguished: the solid, the grooved, and the canaliculated or tubular, so-called “perforated”; the third, as we shall explain, being only a further modification of the second. In the grooved tooth, a sulcus runs along the anterior or outer surface, its object being to convey into the wound the secretion of a poison gland. It varies in depth according to the species, and may be so slight as to escape detection without a very strong magnifying glass. In some the sulcus may be very deep and wide, forming a canal round which the tooth folds to the extent of its borders nearly meeting; from this condition the so-called “perforated” fang is derived through the complete fusion of the borders of the tooth, and the obliteration of the line of union except at each extremity. The structure of such a fang may be best understood by imagining a tooth, lined all round with the same layer of dentine and enamel, being flattened out in a vertical plane and then folded over, the outer edges coalescing on the front median line in such a way that the inner wall of the tooth is in reality the anterior surface, and the outer wall the posterior surface, of the ordinary tooth.

Grooved teeth, with open canal, are situated either at the anterior extremity (Proteroglyphs) or at the posterior extremity (Opisthoglyphs) of the maxillary bone, usually followed or preceded by a series of solid teeth, which in some cases may likewise show a more or less distinct groove. Such may also be present on the teeth of the lower jaw, as in the European Coelopeltis, in some specimens of which a faint groove is visible on the outer side with the aid of a strong lens.

The tubular fangs of the ViperidÆ are inserted on the posterior extremity of the much abbreviated and erectile maxillary bone, which bears no other teeth. The Proteroglyphs (Cobras, Coral-snakes, Sea-snakes) and the Solenoglyphs (Vipers, Pit-vipers, Rattlesnakes) may be regarded as the diverging extremes in the development of the poison apparatus, both culminating in forms with tubular fangs, the former as derived directly from the Aglyphs (harmless snakes), the latter from the Opisthoglyphs, likewise evolved out of the Aglyphs. That the insertion of the poison fangs of the ViperidÆ is really on the posterior extremity of the maxillary bone is evident from the condition of the bone in its recumbent position, especially in the African Viper, Causus, which in several respects departs less markedly from the Colubrid type than our European Vipers.

The poison fangs of the ViperidÆ appear to be movable, folding in the mouth when at rest, and erected, or even thrust forward, when ready to act. This, however, is simply due to the mobility of the maxillary bone, to which they are ankylosed as in all other snakes. There are normally two equally-developed fangs, close together and side by side, to each maxillary, followed by several replacement fangs loosely attached behind them, usually in two series of four. When the two fangs are in situ, they of course both function in the act of biting, although only one is in relation with the single poison duct; often, however, there is only one fang in position, either the right or the left, the place of the other being indicated by a shallow socket which will soon be filled by one of the posterior reserve fangs moving forward and becoming ankylosed to the bone. Snake-charmers who extract the poison fangs of the snakes they use for their performances have therefore to renew the operation frequently, unless they amputate the bone on which the fangs are inserted, an injury which the creature does not long survive.

The dentition of the snakes in which the maxillary bone is not movable vertically falls under three divisions: the Aglyphs, in which the teeth are all solid; the Opisthoglyphs, in which one or more (usually two) of the hindermost teeth are provided with a groove; and the Proteroglyphs, in which grooved or canaliculated teeth are situated in front, followed or not by solid teeth. Beyond these three principal divisions, the dentition furnishes important characters for the classification, although that importance has sometimes been over-estimated. The maxillary teeth may be equal in length (Isodonts), or the anterior the longer (Lycodonts), or the posterior the longer, increasing gradually in size (Coryphodonts) or abruptly, without (Syncranterians) or with a diastema, or break, in front of them (Diacranterians). These categories are, however, so completely connected as to preclude their use in taxonomy beyond helping to define genera. The number of maxillary teeth and the relative proportions and disposition of the mandibular teeth also afford useful generic characters.

The European genera may be arranged as follows, according to the dentition:

I. Teeth few, disposed in a transverse series in the upper jaw only

Typhlops.

II. Teeth in both jaws and on the palatines and pterygoids.

A. A series of solid teeth along the maxillary; no grooved teeth.

1. Anterior maxillary and mandibular teeth longest; 9 or 10 maxillary teeth

Eryx.

2. Maxillary teeth equal, or increasing in size posteriorly.

a. Mandibular teeth 17 to 30; maxillary teeth 15 to 22.

Posterior maxillary teeth longest; mandibular teeth subequal, more than 20

Tropidonotus.

Posterior maxillary teeth longest; mandibular teeth not more than 20, posterior smallest

Zamenis.

Maxillary teeth subequal; mandibular teeth 20 to 25, posterior smallest

Coluber.

b. Mandibular teeth 14 or 15, subequal; maxillary teeth 12 to 15.

Maxillary teeth increasing in size

Coronella.

Maxillary teeth subequal

Contia.

B. One or two enlarged grooved fangs behind the series of solid maxillary teeth.

14 to 17 subequal solid maxillary teeth, forming a continuous series; 21 to 23 mandibular teeth, anterior strongly enlarged

Coelopeltis.

9 to 11 solid maxillary teeth, fourth and fifth or fifth and sixth enlarged, followed by an interspace; sixth mandibular tooth fang-like, followed by an interspace

Macroprotodon.

9 or 10 solid maxillary teeth, forming a continuous series, decreasing in length posteriorly; anterior mandibular teeth strongly enlarged

Tarbophis.

C. Maxillary with only two large canaliculated fangs side by side, one of which may be missing; anterior mandibular teeth longest

Vipera, Ancistrodon.

In counting the teeth for the purpose of using this key, care must be taken to ascertain the full number, as it frequently happens that one or more are missing; but their place is indicated by the shallow pits in which their base was implanted, the overlooking of which might convey the impression of a hiatus such as is characteristic of certain genera—Macroprotodon, for instance. Needless to say, the loose teeth which are in reserve on the inner side of the jaws or behind the tubular fangs are not taken into consideration, the numbers given being those of functional teeth only. Although as a rule the teeth can be counted easily, on a specimen preserved in spirit, by simply pushing aside the lips and gums with the finger, it is sometimes necessary to remove and clean the bones of the jaws, an operation which does not require much skill.


                                                                                                                                                                                                                                                                                                           

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