Chapter II: Part 2

The most important deviation from the Rodent structure presented by the teeth, occurs in the direction of the reflected fold of enamel, and such a deviation might have been inferred, even in the absence of the teeth, from the structure of the articular surface, or glenoid cavity for the reception of the condyle of the lower jaw. As the ridge of enamel runs, as above described, in a direction approaching that of the longitudinal axis of the skull, it is obvious that the grinding motions of the lower jaw should be in a proportionate degree in the transverse direction. The glenoid cavity, therefore, instead of being a longitudinal groove, and open behind, as in the true Rodents, is extended transversely, and is defended behind by a broad descending bony process preventing the retraction of the jaw, and showing marks of the forcible pressure to which it was subject.

It is worthy of observation that, in the Wombat,—which exhibits the Rodent type of dentition, and, like the Toxodon, has remarkably curved molars, but in an opposite direction,—the condyle of the lower jaw is also extended transversely, and adapted to an articular surface, which admits of lateral motion in the trituration of the food. In the outward span of the zygomatic arches, in which Toxodon deviates from the Rodentia, we may trace a relation of subordinacy to the above structure of the grinding teeth and joint of the lower jaw: the widening of the arches giving to the masseter muscles greater power of drawing the jaw from side to side. The depth of the zygoma bespeaks the magnitude of these masticatory muscles, and the included space shews that the temporal muscles were also developed to a degree, which indicates the force with which the great incisors at the extremity of the jaws, were used; probably, like the canines of the Hippopotamus, to divide or tear up by the roots the aquatic plants, growing on the banks of the streams, which the Toxodon may have frequented.

In the Rodentia, the zygoma, though sometimes as deep as in the Toxodon, is generally almost straight, and the space included between it and the cranium is consequently narrow. The zygoma also is placed more forwards in all true Rodents, than in the Toxodon; and, instead of abutting against the posterior alveoli, it terminates opposite the anterior ones. It thus affords such an attachment to the masseter, that this muscle extends obliquely backwards to its insertion in the lower jaw, at an angle which enables it to act with more advantage in drawing forwards the lower jaw,—a motion for which the joint is expressly adapted. In many Rodents, also, there is a distinct muscle, or portion of the masseter, which passes through the ant-orbital foramen, which is on that account of large size. In examining the cranium of Toxodon, with reference to this structure, it was found that the ant-orbital foramen was not larger than might have been expected to give transmission to nerves requisite for supplying with sensibility the large lips, and whiskers with which the expanded muzzle of this remarkable quadruped was probably furnished.

Having thus examined the cranium of the Toxodon in its relation, as a mechanical instrument, subservient to the function of digestion; we next proceed to consider the structure and composition of those cavities of the skull which gave lodgment and protection to the organs of _special_ sense, and endeavour to deduce from their structure conclusions as to the degree in which the organs were developed, and the circumstances under which the senses were exercised.

The orbit of Toxodon forms the anterior boundary of the zygomatic area; it is about as distinctly defined as in the Tapir or Dugong, having its osseous rim less complete than in the Hippopotamus, yet more developed than in the Capybara, Coypus, and many other Rodentia, in which the orbit is scarcely distinguishable in the cranium from the small space occupied by the origin of the temporal muscle.

The lower boundary of the orbit in Toxodon is formed by an excavation in the upper and anterior part of the zygoma; the upper boundary by a strong and rugged overarching process of the frontal bone, the posterior angle of which (_a_, Pl. III.) descends a little way, but leaves a space of three inches and a half between it and the opposite angle of the malar bone below (_b_, Pl. II. and III.), the circumference of the orbit being completed probably by ligament in the recent subject. The cavity thus circumscribed is remarkable for the preponderance of the vertical over the transverse or longitudinal diameter, and indicates great extent of motion of the eyeball in the vertical direction, such as may be supposed to be well adapted to the exigencies of an amphibious quadruped. The orbit of the Capybara, or Water-hog, makes a near approach to the form just described. In the elevation of the supra-orbital boundary, and its outward projection in the Toxodon, we perceive an approximation to the form of the orbit in the Hippopotamus, but the size of the orbit is relatively larger in the Toxodon, which in this respect manifests its affinity to the Rodentia.

In that part of the bony structure of the auditory apparatus, which is visible on the exterior of the cranium, the skull of the Toxodon presents a character in which it recedes from the Rodentia. In these, the tympanic portion of the temporal bone is remarkably developed, forming a large bulla ossea between the glenoid cavity and the occiput; and it always remains disunited to the other elements of the temporal bone. In the Toxodon the tympanic bone (_c_, Pl. II.) consists of a rough compressed vertical osseous plate, wedged in transversely between the occiput and the posterior part of the glenoid cavity. The internal extremity of this plate points inwards and forwards, representing the styloid process; behind this is seen the petrous bone, which forms a small angular protuberance at the basis cranii, and is less developed than in the Hippopotamus. Anterior to the petrous bone are the orifices of the Eustachian tube, and carotid canal; external to it is the great foramen lacerum, for the jugular vein and nervus vagus; and behind it is the anterior condyloid foramen. The foramen auditorium externum is only half an inch in diameter, and gives passage to a long and somewhat tortuous meatus, which passes inwards and slightly forwards and downwards; its direction being precisely the same as in the Hippopotamus; it was accompanied, probably, by as small an external auricle.

But the indications of the aquatic habits of the Toxodon, which are presented by the osseous parts relating to the senses of sight and hearing, are of minor import compared with those afforded by the bony boundary of the nostrils. This boundary circumscribes a large ovate aperture, the aspect of whose plane is upwards, and a little forwards, as in the Herbivorous Cetaceans, and especially the Manatee (_Trichecus Manatus_, Cuv.) In one part of the bony structure of the nasal cavity the Toxodon deviates, however, in a marked degree from the Cetaceous structure; I allude to the frontal sinuses, which are exposed by the fracture of the upper part of the skull. (They are shewn in Plate III., and an asterisk is placed on one of the narrow canals of intercommunication between the sinuses and the nasal passages.) The posterior orifice of the nasal cavity is relatively larger and wider than in the Herbivorous Cetaceans, and differs both in form and aspect in consequence of the greater extent of the bony palate. The Toxodon further differs from the Manatee and Dugong, in the firm nature of the connexion of the bones of the head; and it differs from the Hippopotamus in the strong attachment of the intermaxillary bones to the maxillaries.

There next remain to be described, as far as the shattered condition of the skull will permit, the relative position, extent, and connexions of the principal bones composing it.

The _occipital bone_ exhibits a complete confluence of its basilar, condyloid, and supra-occipital elements. The basilar portion, in connexion with the corresponding element of the sphenoid bone, describes a curve whose convexity is downwards. The condyles are large, extended in the transverse direction, completely terminal, and a little inclined downwards below the level of the basilar process. The curve of the articulating surface describes, in the vertical direction, two-thirds of a circle, indicating that the head must have possessed considerable extent of motion upwards and downwards upon the atlas; thus, while the body of the Toxodon was submerged, the head probably could be raised so as to form an angle with the neck, and bring the snout to the surface of the water without the necessity of any corresponding inflection of the spine. Indeed, in the form and position of the condyles, the Toxodon more nearly resembles the true Cetacea than any other existing mammalia; and it is only with these that it can be compared in regard to the aspect of the plane of the occipital foramen, and of the occipital region of the skull. This is inclined forwards from the occipital foramen at such an angle, that on viewing the skull from above, not only the condyles, but the entire circumference of the occipital foramen are visible. (See Pl. III.) The upper part of the supra-occipital plate presents a broad rugous depression, indicative of the insertion of strong cervical muscles, and probably of a _ligamentum Nuchæ_.[13]

The ex-occipital processes advance forwards for about an inch beyond the condyles, and then suddenly extend outwards at right angles to the former line, and terminate in the form of vertically compressed bony plates; the lower rugged margins of which represent or perform the office of the mastoid processes (_d_, _d_, Pls. II. and III.). The breadth of the entire occipital region of the skull (fig. 1, Pl. IV.) appears to have been, allowing for the fractures, about one-third more than the height of the same part.

The great development of the _tympanic_ bones in the Rodentia, occasions the intervention of a considerable space between the occipital bone and the zygomatic process of the temporal; but in the great Toxodon, in which the sense of hearing was doubtless inferior to that enjoyed by the small and timorous Rodents, the tympanic bone is reduced to a thin plate, which is wedged in between the occiput and glenoid cavity. In this structure, and the consequent posterior position of the glenoid cavity, there is a close resemblance between the Toxodon and the Hippopotamus, Tapir, and Rhinoceros.

The _squamous_ element of the temporal bone (N, Pl. II.) forms a small proportion of the lateral walls of the cranium, and also enters into the composition of the lateral and superior parts of the posterior region of the cranium, where two deep fossæ perforated by large vascular foramina, indicate the junction of the squamous bones with the supra-occipital bone. The posterior surface of the skull is thus divided into three broad and shallow depressions, the two lateral facets being slightly over-lapped by the middle one, at their junction with it. In this structure the Toxodon resembles the Hippopotamus, and differs considerably from the Cetacea, in which the occipital region is rendered convex by the extraordinary development of the brain within.

The _zygomatic_ process of the temporal bone projects boldly outwards at its commencement, where it is of great strength, and three-sided; the glenoid cavity extends transversely across the base or inferior surface of this part; the lateral surfaces converge to form the ridge or upper boundary of the zygoma. The depth of the glenoid cavity is increased by a transverse production of bone both before and behind it: the posterior process (_g_, Pl. II.) descends the lowest, and affords the requisite defence against backward dislocation of the lower jaw; the pressure of the condyle against this process is denoted by a well defined, transversely-ovate, flattened and smooth surface, as if the bone had been planed down at that part: the anterior transverse boundary is convex and smooth, and probably formed part of the articulation for the lower jaw. The lower facet of the zygoma anterior to the glenoid cavity gradually contracts in breadth, as it advances forward, and at the distance of three inches from the articular cavity the zygoma changes from a prismatic to a laminar form. It is at this point that the zygomatic suture commences, at the lower margin of the arch; whence it extends directly forwards for more than half its length, and then bends upwards at a right angle. The zygomatic suture has a similar course in the Capybara, and Hippopotamus.

The remainder of the zygoma is formed externally by the _malar_ bone (G Pl. II.), which in its position is intermediate to the Rodent and Pachydermatous structures. It is not suspended in the middle of the zygomatic arch, as in the former order; neither does it extend into the region of the face so far anterior to the orbit as in the Tapir or Hippopotamus. The exterior line of the malo-maxillary suture defines the orbit anteriorly; but from this line the maxillary bone extends backwards, along the inner side of the malar portion of the zygoma, until it almost reaches the temporo-malar suture; thus abutting by an oblique surface against nearly the whole internal facet of the malar bone, and materially contributing to the general strength of the zygomatic arch. The malar bone is of considerable vertical extent, and presents a rugged and thickened inferior margin for the attachment of the masseter. The upper margin of the malar bone is smoothly rounded, and presents a regular semicircular excavation, forming the lower boundary of the orbit. The relative magnitude of the zygomata to the entire cranium far exceeds in the Toxodon that which exists in the Hippopotamus or any other known Pachyderm. This arises from the great vertical development of the malar bone behind the orbit, and the vertical expansion of the temporal portion of the arch. The oblique position of the zygoma, descending as it advances forwards, is deserving of attention, as the Toxodon, in deviating from the Pachyderms in these respects, makes an evident approach to the herbivorous Cetaceans, as the Dugong and Manatee: in the latter Cetacean we observe a similar development of the lower part of the zygomatic process of the malar bone. It is here, also, that we may perceive an indication of a resemblance between the Megatherium and Toxodon.

There is no discernible trace of the _lachrymal_ bone (E, Pl. II.) having extended, as in the Hippopotamus beyond the anterior boundary of the orbit: the lachrymal foramen is situated rather deep in the orbit, and the bone itself appears to have been of very small size.

The surface of the supra-orbital process of the _frontal_ bone(C, Pl. II.) is deserving of attention, as it presents a peculiar ruggedness which is not found in any other part of the skull; the irregularity seems, as it were, to have been produced by the impression of numerous small tortuous and anastomosing vessels. In the skull of a Sumatran two-horned Rhinoceros, in the Museum of the College of Surgeons (No. 816), the circumference of that part of the surface of the skull which supported the posterior horn, and which includes precisely the same part of the os frontis, presents the same character, the surface being broken by numerous vascular impressions. On the supposition that this character of the supra-orbitary arch in the Toxodon might indicate the superincumbency of a bony case, I examined the skulls of two Armadillos, _Dasypus Peba_ and _Das. 6–cinctus_, and found that in the Dasypus 6–cinctus, the supra-orbital ridges, which are slightly elevated, to support the cephalic plate, presented, in a minor degree, a corresponding rugosity. May we venture then to conjecture that the Toxodon was defended by an ossified integument like the Armadillo, or that it was armed with an epidermic production, analogous to the horn of the Rhinoceros; or had the rugous surface in question as little relation with the parts that covered it as the sculptured surface of the malar bones in the Cavy?

After forming the rugged and prominent supra-orbital processes already described, the frontal bone continues to send backwards a slightly elevated ridge or _crista_, circumscribing the origin of the temporal muscles, but the extent of this ridge, and the disposition of the inter-orbital portion of the frontal bones cannot be determined in the present mutilated specimen. The fractures it has sustained are not, however, wholly unattended with advantage; they expose the structure of the diploë, which from its coarseness of texture and thickness, resembles that of the Cetaceous crania; and what is of still more importance, they also demonstrate the existence and form of the frontal sinuses.

The cavity of the nose is extensive, and the remains of the ossa spongiosa superiora testify that the Toxodon enjoyed the sense of smell to a degree equal at least to that of the Hippopotamus.

The _sphenoid bone_ resembles that of the Hippopotamus, but it contributes a larger share to the formation of the internal pterygoid processes (_p_, Pl. II.); these are of a simple form, and more developed than in the _Hippopotamus_; they project outwards to a greater extent, and terminate in a point. The sphenoid also sends off a short and thick pointed process from the posterior part of the base of the internal pterygoid processes. The ala of the sphenoid does not rise so far into the orbit, nor does it articulate with the parietal bone, as in the Hippopotamus; but in this part of its structure, is the same as in the Rhinoceros. The spheno-palatine foramen is relatively larger than in the above-named Pachyderms, and is bounded above by the descending orbital plate of the frontal bone.

The palatal processes of the _palatine_ bones terminate anteriorly between the last molars, and extend backwards for some distance beyond the alveolar processes, increasing the extent of the bony roof of the mouth posteriorly: this is a structure in which the Toxodon deviates both from the Rodents, and Pachyderms, and resembles the Armadillos among the Edentata; excepting that the postdental part of the bony palate in the Toxodon is suddenly contracted in breadth. The palato-maxillary suture is in the form of a chevron, with the angle directed forwards, as in the Hippopotamus and Capybara, but truncated.

The _superior maxillary_ bones (F, Pl. II.) are united posteriorly to the malar, as above described: they ascend and join the frontal and nasal bones: their outer surface is almost vertical, smooth, and slightly undulating; perforated at its posterior part by the ant-orbital foramen, and joined anteriorly to the intermaxillaries by a suture running in the sigmoid direction (as shewn in Pl. II.) from the middle of the nasal cavity, to within four inches of the anterior boundary of the upper jaw. We have, in the position and extent of this suture, and the absence of tusks and their large prominent sockets, a most important difference between the Toxodon and Hippopotamus. The chief peculiarity in the maxillary bones, obtains in the arched form of the alveolar processes, corresponding to the shape and position of the grinders above described, and which are peculiar among known mammalia to the present genus. The palatal surface of the maxillary bones is obliquely perforated by two large foramina, from which two deep longitudinal grooves extend forwards, and are gradually lost; we find the posterior palatine foramina represented by similar grooves and foramina in the Capybara.

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;[14] 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.

Many analogous examples will readily occur to the experienced comparative anatomist. The Horse has been adduced as one to which reference can very readily be made; but I would also cite the Sumatran Rhinoceros, the skull of which, in the Hunterian collection, has already been alluded to. In this species the anterior grinders, in both jaws, are small and simple, and increase in complexity as they recede backwards. The third superior grinder (fig. 8, Pl. V.) presents a single fold of enamel, reflected obliquely forwards from the inner side half-way across the tooth; the outer line of enamel describes a simply undulating line. The opposite grinder of the lower jaw (fig. 9, Pl. V.) has only one-half the breadth of the upper one, but has its grinding surface further complicated by having two inflected folds of enamel from the inner side, and one shorter and broader fold from the outer side. This tooth, therefore, presents a close resemblance to one of the posterior grinders of the lower jaw of the Toxodon, but differs essentially in being of limited growth, and consequently in having fangs.[15]

In speculating upon the nature of the organized substances which the teeth of the Toxodon were destined to grind down, we must not only take the structure of the tooth into consideration, but also the power of perpetual renovation, which will compensate for the defective quantity of enamel in the grinders of the Toxodon, as compared with those of the existing Ruminants and Pachyderms, whose grinders, when once completed, receive no further addition of dental substance at their base. The Toxodon, in this character of its dentition, participated in the same advantages with the Capybara and the Megatherium.

Although we have been enabled to observe the structure of the grinding teeth of the upper jaw of the Toxodon in two examples only; one, an insulated perfect grinder corresponding to the sixth alveolus on the right side, and the other, a portion of the last grinder of the left side remaining in the socket of the head previously described, yet from the relations subsisting between socket and tooth, a very satisfactory opinion may be formed of the structure of those teeth which are wanting, as well as of their size. It thus appears, that the grinders of the upper jaw of the Toxodon, are small and simple at the anterior part of the jaw, and that they increase (chiefly in antero-posterior extent) in size, as well as in complexity, as they recede backwards in the jaw. In this respect, as well as in size, the teeth, in the fragments of the lower jaw just described, exactly correspond. There is, however, a slight difference in the lateral diameter of the two sets of grinders, those of the lower jaw being narrower, as is usually the case, but not in the same degree as in the Horse or Ruminant. A greater difference obtains in the degree of curvature of the two sets of molars, those of the lower jaw, especially the posterior grinders, being much less bent than the corresponding teeth of the upper jaw. It is necessary to observe, also, that the convexity of the curve of the inferior grinders is directed outwards, as in the superior grinders; while in the Guinea Pig and Wombat, which have also curved grinders, the convexity is outwards in the lower jaw, and inwards in the upper jaw.

Nevertheless, if we take into consideration the close similarity which exists between the teeth of the upper jaw of the Toxodon, and those of this lower jaw in more essential points, as in their persistent pulps, their characteristic structure and form, the depth of their sockets, and their relative sizes and complexity; and when we consider how the depth of this lower jaw, and its narrowness in the transverse direction, corresponds with the characteristic form of the upper jaw of the Toxodon, and that to these resemblances is added an apparatus of incisors adequate to oppose the great dentes scalprarii of the upper jaw, the conclusion seems irresistible, that the lower jaw, here described, must be referred, if not to the same, at least to a nearly allied species of Toxodon, as that to which the large cranium belonged.

Further researches in South America, it is hoped, will lead, ere long, to the completion of our knowledge of the osteology of this very remarkable and interesting genus of extinct mammiferous animals.

DESCRIPTION OF PARTS OF THE SKELETON OF
MACRAUCHENIA PATACHONICA;
_A large extinct Mammiferous Animal, referrible to the Order
Pachydermata; but with affinities to the Ruminantia, and especially to
the Camelidæ_.

In the preceding pages the nature and affinities of a large extinct Mammal were attempted to be determined from the cranium and teeth exclusively: we come now to consider the remains of a quadruped consisting of bones of the trunk and extremities, without a fragment of a tooth or of the cranium to serve as a guide to its position in the zoological scale.

It may appear, even to anatomists and naturalists familiar with the kind of evidence afforded by a fossil fragment, that an opinion as to the relation of the present species to a particular family of Ruminants, formed without a knowledge of the important organs of manducation, must be vague and doubtful, but the evidence about to be adduced, will be regarded, it is hoped, as more conclusive than could have been _à priori_ expected.

The portions of the skeleton of the animal—which, in relation to the affinity above alluded to, as well as from the length of its neck, I propose to call _Macrauchenia_[16]—were discovered by Mr. Darwin in an irregular bed of sandy soil, overlying a horizontal accumulation of gravel on the south side of Port St. Julian: and independently of the circumstances under which they were found, their correspondence with each other in size, colour, texture and general character prove them to have belonged to one and the same individual.

These remains include two cervical vertebræ, seven lumbar vertebræ, all more or less fractured; a portion of the sacrum and ossa innominata; fragments of the right scapula; of the right radius and ulna, and right fore-foot; the right femur nearly entire, the proximal and distal extremities of the right tibia and fibula; and a metatarsal bone of the right hind-foot.

Before entering upon the description of these remains, a few observations may be advantageously premised on some of the distinguishing characters of the Camelidæ. It is well known that the Camels and Llamas deviate in their dentition, viz., in the presence of two incisors in the upper jaw, from the true Ruminants; and we cannot avoid perceiving that in this particular the direction in which they deviate tends towards the conterminous Ungulate Order, in which incisor teeth are rarely absent in the upper jaw. They also further deviate from the Ruminants and approach the Pachyderms in the absence of cotyledons in the uterus and fetal membranes; having, instead thereof, a diffused vascular villosity of the chorion, as in the sow and mare.

But besides these characters, by which, in receding from one type of hoofed mammalia, the Camelidæ claim affinity with another, there are many parts of their organization peculiar to themselves; of some of these peculiarities, the relation to the circumstances under which the animal exists, can be satisfactorily traced; in others, the connection of the structure with the exigencies of the species, is by no means obvious, and in this predicament stands the osteological peculiarity, which is immediately connected with our present subject—a peculiarity in which the Camelidæ differ not only from the other Ruminants, but from all other existing Mammalia, and which consists in the absence of perforations for the vertebral arteries in the transverse processes of the cervical vertebræ, the atlas excepted.

I may observe that what is described as a perforation of a single transverse process in a cervical vertebra is essentially a space intervening between two transverse processes, a rudimental rib, and the body of the vertebra. In the cold-blooded Saurians,—in which the confluence of the separate elements of a vertebra takes place tardily and imperfectly, if at all,—the nature of the so called perforation of the transverse process is very clearly manifested, as in the cervical vertebræ of the Crocodile, in which the interspace of the inferior and superior transverse processes is closed externally by a separate short moveable cervical rib. In the Ornithorhynchus paradoxus the vertebra dentata also preserves throughout life this condition of its lateral appendages: in other Mammalia it is only in the fœtal state that the two transverse processes are manifested on each side with their extremities united by a distinct cartilage, which afterwards becomes ossified and anchylosed to them.

In the Hippopotamus the inferior transverse process sends downwards a broad flat plate extended nearly in the axis of the neck, but so obliquely, that the posterior margins of these processes, in one vertebra, overlap the anterior ones of the succeeding vertebra below, like the cervical ribs in the Crocodile; the same structure obtains in many other mammalia, especially in the Marsupials. In the Giraffe, the inferior transverse processes are represented by relatively smaller compressed laminæ, projecting obliquely downwards and outwards from the anterior and inferior extremity of the body of the vertebra. The superior transverse processes in this animal are very slightly developed in any of the cervical vertebræ, and the perforation for the vertebral artery is above and generally in front of the rudiment of this process, being continued as it were through the side of the substance of the body of the vertebræ.

In the long cervical vertebræ of the Camel and Llama, the upper and lower transverse processes are not developed in the same perpendicular plane on the sides of the vertebræ, but at some distance from each other; the lower transverse processes (_a_, fig. 1, Pl. VI.; _a_, fig. 1, 3, 4, Pl. VII.) being given off from the lower part of the anterior extremity of the body of the vertebra; the upper ones (_b_, fig. 1, Pl. VI.; _a_, fig. 1, 3, 4, Pl. VII.) from the base of the superior arch near the posterior part of the vertebra, or from the sides of the posterior part of the body of the vertebræ. The extremities of these transverse processes do not become united together, but they either pass into each other at their base, or continue throughout life separated by an oblique groove (as in fig. 1, Pl. VI.) This groove would not, however, afford sufficient defence for the important arteries supplying those parts of the brain which are most essential to life; and, accordingly the vertebral arteries here deviate from their usual course, in order that adequate protection may be afforded to them in their course along the neck. From the sixth to the second cervical vertebræ inclusive in the _Aucheniæ_, and from the fifth to the second inclusive in the _Cameli_,[17] the vertebral arteries enter the vertebral canal itself, along with the spinal chord, at the posterior aperture in each vertebra, run forwards on the outside of the dura mater of the chord between it and the vertebral arch, and when they have thus traversed about two-thirds of the spinal canal, they perforate respectively the superior vertebral laminæ, and emerge directly beneath the anterior oblique or articulating processes, whence they are continued along with the spinal chord into the vertebral canal of the succeeding vertebra, and perforate the sides of the anterior part of the superior arch in like manner; and so on through all the cervical vertebræ until they reach the atlas, in which their disposition, and consequently the structure of the arterial canals, resemble those in other Ruminants.

The two cervical vertebræ of the Macrauchenia present precisely the structure and disposition of the bony canals for the vertebral arteries which are peculiarly characteristic of the Camelidæ among existing Mammalia. In Plate VI. fig. 2, the groove and orifices of the canal for the vertebral artery are shown in a section exposing the spinal canal: in Plate VII. figures 1 and 3 exhibit the orifices at the commencement of the arterial canals, as seen in a posterior view of the vertebræ; in figs. 2 and 4, the terminations of the same canals are shown, in the anterior view of the same vertebræ; the smaller figures (3 and 4) are taken from the fourth cervical vertebra of a Llama. The vertebræ of the Macrauchenia also closely resemble the middle cervical vertebræ of the Vicugna and Llama in their elongated form; approaching the Auchenial division of the Camelidæ, and deviating from the true Camels in the relations of the length of the body of the vertebra to its breadth and depth, and in the much smaller size of the inferior processes. Excepting the Giraffe, there is no existing mammal which possesses cervical vertebræ so long as the Macrauchenia; but the cervical vertebræ of the Giraffe, differ in the situation of the perforations for the vertebral arteries, and in the form of the terminal articular surfaces, as will be presently noticed.

Both of the cervical vertebræ of the _Macrauchenia_ here described, are of the same size, each measures six inches and a half in extreme length, two inches, ten lines in breadth, and two inches, four lines in depth. In the Giraffe and the Camelidæ, the spinous processes are thin laminæ of considerable extent in the axis of the vertebra, but rising to a very short distance above the level of the vertebral arch: the spinous processes have the same form in the corresponding vertebræ of the Macrauchenia, but present a still greater longitudinal extent; they commence at the interspace of the anterior oblique processes, and extend to opposite the base of the posterior oblique processes; the upper margin describing a gentle curve, as shown in fig. 1, Pl. VI. The transverse processes also present the form of slightly produced, but longitudinally extended, laminæ: their disposition is essentially the same as in the Camelidæ, but more nearly corresponds with the modifications presented by the Aucheniæ. The inferior transverse processes,—those which are alone developed in fish, but which are not present in any other vertebræ save the cervical, in mammalia,—these processes in the Macrauchenia are continued from the sides of the under surface of the anterior part of the body of the vertebra; their extremities being broken off, it cannot be determined how far they extended from the body of the vertebræ, but they gradually subside as they pass backwards: the superior transverse processes are continued outwards from the sides of the posterior part of the body of the vertebra, and gradually subside as they advance forwards along three-fourths of the body of the vertebra: they are not continued into the anterior and inferior transverse processes, as in the Vicugna, but are separated therefrom by a narrow and shallow groove. The articular, or oblique processes, closely resemble those of the Auchenia in form, and in the direction of the articular surfaces; those of the anterior processes looking inwards and a little upwards; those of the posterior, outwards and a little downwards.

In the Macrauchenia a small longitudinal process (_c_, fig. 2, Pl. VII.) is given off immediately below the base of the anterior oblique process; this structure is not observable in any of the cervical vertebræ of the Giraffe or Camelidæ.