Part 6

The intermaxillary bones (D, Pls. II. and III.), though large, are relatively of less extent than in the Rodents generally. The nasal processes do not reach the frontal bone, but are limited to the anterior half of the nasal boundary; approaching in this respect to the Herbivorous Cetacea. In the outward expansion of their anterior extremities, the intermaxillaries resemble those of the Hippopotamus, in which, however, this character is more strongly marked. The intermaxillaries in the Hippopotamus are also much less firmly united to the maxillary bones than in the Toxodon, and are consequently commonly lost in the fossil crania. On the palatal surface of the intermaxillary bones there are two grooves which diverge forwards from the line of the suture; and anteriorly to these grooves there are the two large anterior palatine foramina. The maxillo-intermaxillary sutures on the palate converge as they extend backwards to a point; there appears to have been a fissure left between this suture and the mesial suture of the intermaxillaries; in which structure the Toxodon resembles the Hippopotamus.

After summing up the different affinities, or indications of affinity, which are deducible from the cranium of this most curious and interesting fossil mammal, we are led to the conclusion, assuming it to have had extremities cased in hoofs, that it is referrible to the Order Pachydermata. But the structure, form, and kind of teeth in the upper jaw, prove, indisputably, that the gigantic Toxodon was intimately related to the Rodent Order. From the characters of this order, as afforded by the existing species, the Toxodon, however, differs in the relative position of the supernumerary incisors, and in the number, and direction of the curvature, of the molars. If, moreover, the lower jaw, next to be described, belong, as I believe, to the Toxodon, the dental character of the genus will be incisors ⁴⁄₆; pro laniariis diastema; molares ⁷⁄₇ ⁷⁄₇.

The Toxodon again deviates from the true Rodentia, and resembles the Wombat, and the Pachyderms, in the transverse direction of the articular cavity of the lower jaw.

It deviates from the Rodentia, and resembles the Pachydermata in the relative position of the glenoid cavities and zygomatic arches, and in many minor details already alluded to.

In the aspect of the plane of the occipital foramen, and occipital region of the skull; in the form and position of the occipital condyles; in the aspect of the plane of the anterior bony aperture of the nostrils; and in the thickness and texture of the osseous parietes of the skull, the Toxodon deviates both from the Rodentia and existing Pachydermata, and manifests an affinity to the Dinotherium and Cetaceous Order, especially the Herbivorous section.

At present we possess no evidence to determine whether the extremities of the Toxodon were organized on the ungulate or unguiculate type, nor can we be positive, from the characters which the skull affords, that the genus may not be referrible to the Mutica of Linnæus; although the development of the nasal cavity and the presence of large frontal sinuses render it extremely improbable that the habits of this species were so strictly aquatic, as the total absence of hinder extremities would occasion.

Where the dentition of a mammiferous animal is strictly carnivorous, this structure is obviously incompatible with a foot incased in a hoof:—but where the teeth are adapted for triturating vegetable substances the case is different. If animals so characterized are of small size and seek their food in trees, or if they burrow for roots or for shelter, the vegetable type of dentition must co-exist with unguiculate extremities, as in the Edentata and Rodentia generally: but the largest genus (Hydrochærus) of the Rodent Order, whose affinity to the Pachydermata is manifested in its heavy shapeless trunk, thinly scattered bristly hair, and many other particulars, has each of its toes inclosed in a miniature hoof.

The affinity above alluded to, is too obvious to have escaped popular notice, and the Capybara, from its aquatic habits, has obtained the name of Water-hog. It is highly interesting to find that the continent to which this existing aberrant form of Rodent is peculiar, should be found to contain the remains of an extinct genus, characterized by a dentition which closely resembles the Rodent type, but manifesting it on a gigantic scale, and tending to complete the chain of affinities which links the Pachydermatous with the Rodent and Cetaceous Orders.

ADMEASUREMENTS OF THE CRANIUM OF TOXODON. feet inches lines Extreme length 2 4 Extreme breadth 1 4 Extreme height, (exclusive of the lower jaw) 10 Length of zygomatic process 1 1 6 Depth or vertical extent of do. 6 Transverse extent of zygomatic fossa 6 Transverse diameter of cranium between the zygomatic arches 5 Transverse diameter of occipital plane of the cranium 1 From the outside of one condyle to that of the opposite condyle 8 6 Length of the bony palate 1 6 Extreme breadth of ditto 6 Breadth of palate at the intermaxillary suture 2 6 Breadth of palate behind the molar alveoli 3 Longitudinal extent of the molar alveoli 9 6 Longitudinal extent of the diastema 5 6 Transverse diameter of posterior nasal aperture 3 9 Transverse diameter of occipital foramen 3 Transverse diameter of glenoid cavity 4 6 Antero-posterior do. of ditto 1

DESCRIPTION OF FRAGMENTS OF A LOWER JAW AND TEETH OF A TOXODON.

Found at Bahia Blanca, in latitude 39° on the East coast of South America.

In looking over some fragments of jaws and teeth, forming part of Mr. Darwin’s collection of South American mammiferous remains, and which had been set aside with mutilated specimens referrible to species belonging to the family of Edentata, my attention was caught by the appearance of roots of teeth projecting, in a different direction from the grinders, from the fractured anterior extremity of a lower jaw, and I was induced to examine minutely the structure of the teeth in this specimen, and to search the collection for corresponding fragments. The result was the discovery of portions of the two rami, and the commencement of the symphysis of a lower jaw, containing anteriorly the roots of six incisors, and at least six molars on each side; but as the rami had been fractured through the middle of the sixth alveolus, the number of grinders may have corresponded with those in the upper jaw of the Toxodon.

The most perfect of these fragments is figured in Pl. V. figures 1 and 4; figure 2 shows the form of the teeth in transverse section, and the disposition of the enamel upon the grinding surface of the molars on the right side, as restored from a comparison of the fractured teeth in the two rami. From the remains of the symphysis shown at fig. 4, it will be seen that the jaw was remarkably compressed, or narrow from side to side; while the rami (fig. 1.) were of considerable depth, in order to give lodgment to the matrices and bases of grinders enjoying uninterrupted growth.

The pulps of the six incisors in this lower jaw are arranged in a pretty regular semicircle, whose convexity is downwards; the teeth themselves are directed forwards, and curved upwards, like the inferior incisors of the Rodentia. The form and degree of the curvature are shown in the almost perfect incisor (Pl. V. fig. 5) which corresponds with the left inferior incisor of the lower jaw, and was found in the same stratum, but belonged to another individual.

These incisors are nearly equal in size: they are all hollow at their base, and the indurated mineral substance impacted in their basal cavities well exhibits the form of the vascular pulps which formerly occupied them. Sufficient of the tooth itself remains in four of the sockets to show that these incisors, like the nearly perfect one (fig. 5), had only a partial investment of enamel; but though in this respect, as well as in their curvature and perpetual growth, they resemble the dentes scalprarii of the Rodentia, they differ in having a prismatic figure, like the inferior incisors of the Sumatran Rhinoceros, or the tusks of the Boar. Two of the sides, viz., those forming the anterior convex and mesial surfaces of the incisor have a coating of enamel, about half a line in thickness, which terminates at the angles between these and the posterior or concave surface. In plate V. fig. 4, the enamel of the broken incisors is represented by short lines, showing the direction of its crystalline fibres; the white space immediately within the enamel shows the thickness of the ivory at the base of the tooth, the included gray substance represents a section of the formative matrix or pulp of the tooth, which was of the usual conical form: the inferior broken end of the incisor (fig. 5,) appears to have been distant about one-third from the apex of the pulp.

From the relative position of the bases or roots of these incisors, we may infer that they diverged from each other as they advanced forwards, in order to bring their broadest cutting surfaces into line. That they were opposed to teeth of a corresponding structure in the upper jaw is proved by the oblique chisel-like cutting surface of the more perfect incisor: and it is not without interest to find that the presence of dentes scalprarii at the anterior part of the mouth has not been necessarily limited to Mammalia of small size.

The position of the pulps of these incisors, in close proximity with the anterior grinders, corresponds with the position of the pulps of the incisors in the upper jaw of the Toxodon, and indicates, in conjunction with the size of the pulps, that a considerable extent of the inferior incisors was lodged in the substance of the anterior part of the jaw. It is most likely that no vertically directed tooth would be developed in the part of the jaw so occupied by the curved bases of the incisors, and hence a diastema or toothless space would intervene between the molars and incisors of this lower jaw, as in the upper jaw of the Toxodon.

It is interesting, also, to observe, that as the deviations from the Rodent type, which occur in the cranium of the Toxodon, are the same, in some instances, as those which obtain in the Wombat; so we find a corresponding deviation in the size and relative position of the inferior incisors, which, as in the Wombat, terminate anterior to the molar teeth, instead of extending backwards beyond the last grinder, as in most of the true Rodents. The Capybara presents the nearest approach to this structure, the pulps of the inferior incisors being situated opposite the interspace of the first and second grinders.

The molar teeth, in this mutilated lower jaw, like those in the upper jaw of Toxodon, had persistent pulps, as is proved by the conical cavity at their base, as represented in fig. 3; they consequently required a deep socket, and a corresponding depth of jaw to form the socket and protect the pulps. In order to economise space, and to increase the power of resistance in the tooth, and perhaps, also, to diminish the effects of direct pressure on the highly vascular and sensible matrix, we find the molars and their sockets are curved, but in a less degree than those of the upper jaw of the Toxodon. They correspond, however, with the superior molars of the Toxodon in the antero-posterior diameter, in being small and simple at the anterior part of the jaw, and by increasing in magnitude and complexity as they are situated more posteriorly. They are, however, narrower from side to side; but supposing them to belong to the Toxodon, it would agree in this respect with most other large herbivorous mammalia;—the fixed surface for attrition in the upper jaw being from obvious principles more extensive than the opposed moveable surface in the lower jaw.

The first grinder, in the lower jaw here described (Pl. V. fig. 2), is of small size and simple structure, being surrounded with a coating of enamel of uniform thickness, and without any fold penetrating the substance of the tooth. It is more curved than any of the other molars, and appears to have differed from the external incisor only in its entire coating of enamel and direction of growth; it is interesting, indeed, to find so gradual a transition, in structure, from molar to incisive teeth, as this jaw presents; for the robust incisors may here be regarded as representing molars simplified by the partial loss of enamel, and with a change in their direction.

In the second molar, we find an increase in the antero-posterior diameter, and in the length of the tooth, and the enamel at the middle of the outer side makes a fold which penetrates a little way into the tooth; the line of enamel, on the inner side, is slightly concave and unbroken.

The third molar presents an increase of dimensions in the same directions as the second; the enamel on the outer side of the tooth presents a similar fold, but it is directed a little more backwards.

In the fourth molar, besides a further increase of size, and a corresponding but deeper fold of enamel in the external side of the tooth, we have the grinding surface rendered more complicated by two folds of enamel entering the substance of the tooth from the inner side: these folds divide the antero-posterior extent of the tooth into three nearly equal parts; they are both directed obliquely forwards, half-way across the substance of the ivory.

The fifth molar presents the same structure as the fourth, which it exceeds only slightly in size.

In the sixth molar we have a proportionally greater increase of size in the antero-posterior diameter, which measures two inches; but the lateral diameter is but slightly augmented; its structure resembles that of the fifth.

As these grinding teeth by no means increase in the lateral diameter in the same proportion as in their antero-posterior diameter, the posterior ones present, but in a greater degree, the compressed form which characterizes the grinders of the upper jaw of the Toxodon.

It will be seen, however, that there is a difference in the structure of the grinders in this fragment of the lower jaw and those of the upper jaw of the Toxodon. In the lower grinders there are two folds of enamel proceeding from the inner side of the tooth into its substance, whilst in the upper grinders there is only one fold continued from the inner side; in the lower grinders there is also a fold of enamel reflected into the substance of the tooth from the outer surface, while in the upper grinders of Toxodon we find the enamel coating on the outer side of the tooth merely bent inwards, so as to describe, in the transverse section, a gently undulating line; fig. 7, Pl. V. is the grinding surface of the sixth molar, right side, upper jaw.

But this difference of structure is by no means incompatible with the co-existence of the two series of teeth in the same animal, since we find the grinders of the upper and lower jaws presenting differences of structure of equal degree in existing herbivorous species. If we examine the jaws of the Horse, for example, we shall find not only an equal amount of difference in the structure of the upper and lower grinders, but that they deviate from one another in a very similar manner to that above described in the Toxodon. In this comparison attention should be confined to the course of the external enveloping layer of enamel, leaving out of consideration the central crescentic islands of enamel which constitute the additional complexity of the Horse’s grinder. Viewing then the course of the external coat of enamel on the worn surface of the tooth, we find it describing on the outer side of the tooth in the upper jaw an undulating line,—a middle convexity being situated between two concavities; on the inner side of the tooth one fold of enamel penetrates to the middle of the tooth, and on each side of this there is a smaller fold. But in the lower jaw the line of enamel on the outer side of the tooth, instead of merely bending outwards midway in its course, is reflected a little way inwards; while on the opposite, or inner side of the tooth, the enamel sends two extensive folds into the substance of the tooth, opposite to the interspace of which the shorter fold projects from the outer side. Now, on the supposition that the fragment of the lower jaw here described belongs to the Toxodon, the kind and degree of difference in the complexity of the grinding surface of the teeth in the upper and lower jaw, are remarkably analogous to those which exist in the Horse. I have only further to remark that in the Horse the inflected folds of enamel, instead of being simple and straight with the two constitutive layers in apposition, as in the Toxodon, are irregular in their course, with cœmentum intervening between the constitutive layers, which also diverge from each other at their angle of reflection, so as to augment the amount of dense material which enters into the composition of the tooth.