FOSSIL MAMMALIA.
BY MR. OWEN.
It may be expected that the description of the osseous remains of extinct Mammalia, which rank amongst the most interesting results of Mr. Darwin’s researches in South America, should be preceded by some account of the fossil mammiferous animals which have been previously discovered in that Continent. The results of such a retrospect are, however, necessarily comprised in a very brief statement; for the South American relics of extinct Mammalia, hitherto described, are limited, so far as I know, to three species of Mastodon, and the gigantic Megatherium.
One of the above species of Mastodon (Mast. Cordillerarum) was established by Cuvier[[1]] on remains discovered by Humboldt, in Quito, near the volcanic mountain, called Imbaburra, at an elevation of 1200 toises above the level of the sea; and likewise at the Cordilleras of Chiquitos, near Santa Cruz de la Sierra, a locality which is near the centre of South America. A second species (Mastodon Humboldtii, Cuv.[[2]]) is indicated by molar teeth, stated to have been discovered by the same philosophic traveller, in Chile, near the city of Concepcion. The third species of Mastodon appears to have once ranged in vast troops over the wide empire of Peru: numerous teeth were brought thence to Paris by Dombey,[[3]] and similar teeth, together with a humerus and tibia from Santa Fé de Bogota were placed by Humboldt at the disposal of Cuvier,[[4]] who considered them to belong to the Mastodon angustidens, a species of which the fossil remains are by no means uncommon in several localities of Europe. Cuvier is also disposed to refer to the same species the teeth of the Mastodon from Brazil and Lima, mentioned by Dr. W. Hunter in his observations on the animal incognitum from the Ohio.[[5]] The Megatherium has been scientifically described and illustrated in the works of Bru, Cuvier, and D’Alton, whose accounts are founded on a nearly complete skeleton of this stupendous quadruped which has existed in the Royal Museum at Madrid for more than half a century. The few deficiencies in its osteography have recently been supplied by the descriptions and figures given by Dr. Buckland[[6]] and Mr. Clift,[[7]] taken from remains of the Megatherium, brought by Sir Woodbine Parish from Buenos Ayres, and which were discovered in the bed of the Rio Salado, a tributary of the Rio Plata. Sir Woodbine Parish’s collection from the same locality, includes also remains of other species of extinct Edentata, which have not yet been described. M. D’Orbigny, in his travels in South America (vol. i. p. 96.), states that, in the banks of the Parana, he found the fossil remains of a large quadruped, of the size of an Ox,—another quadruped of the size of a Cat, apparently of the carnivorous order;—and a third, a Rodent as large as a Rat.
This meagre condition of the historical part of the subject of South American fossils by no means arises from their actual scarcity. The writings of some of the old Spanish authors, for instance, Torrubia, Garcillasso, and others,[[8]] contain frequent allusions to the bones of giants, who in times of old dwelt in Peru. Legentil, also, in 1728, speaks as an eye-witness of these Peruvian remains; and his guides pointed out to him the traces of the thunder-bolts, by which the Anaks of the New World had been exterminated. Bones and teeth of the Mastodon are, according to Humboldt, so abundant in a locality near Santa Fé de Bogota in Columbia, that to this day it bears the name of the “Field of Giants.”
But independently of these indications, the abundance and variety of the osseous remains of extinct Mammalia in South America are amply attested by the materials for the following descriptions, collected by one individual, whose sphere of observation was limited to a comparatively small part of South America; and the future traveller may fairly hope for similar success, if he bring to the search the same zeal and tact which distinguish the gentleman to whom Oryctological Science is indebted for such novel and valuable accessions.
It is remarkable that all the fossils, collected by Mr. Darwin, belong to herbivorous species of mammalia, generally of large size. The greater part are referrible to the order which Cuvier has called Edentata, and belong to that subdivision of the order (Dasypodidæ) which is characterized by having perfect and sometimes complex molar teeth, and an external osseous and tesselated coat of mail. The Megatherium is the giant of this tribe; which, at the present day, is exclusively represented by South American species, the largest (Dasypus Gigas, Cuv.) not exceeding the size of a Hog. The hiatus between this living species and the Megatherium, is filled up by a series of Armadillo-like animals, indicated more or less satisfactorily by Mr. Darwin’s fossils, some of which species were as large as an Ox, others about the size of the American Tapir. The rest of the collection belongs, with the exception of some small Rodents, to the extensive and heterogeneous order Pachydermata; it includes the remains of a Mastodon, of a Horse, and of two large and singular aberrant forms, one of which connects the Pachydermatous with the Ruminant Order; the other, with which the descriptions in the following pages commence, manifests a close affinity to the Rodent Order.
A DESCRIPTION OF THE CRANIUM OF
TOXODON PLATENSIS;
A gigantic extinct mammiferous animal, referrible to the Order Pachydermata, but with affinities to the Rodentia, Edentata, and Herbivorous Cetacea.
The cranium, which is the subject of the present description, was found in the Sarandis, a small stream entering the Rio Negro, and about 120 miles to the N. W. of Monte Video: it had been originally embedded in a whitish argillaceous earth, and was discovered lying in the bed of the rivulet, after a sudden flood had washed down part of the bank.
The zoological characters deducible from this cranium, forbid its association, generically, with any known Mammiferous animal, and it must therefore be referred to an extinct genus, which I propose to call Toxodon,[[9]] from the curved or arched form of the teeth, as will afterwards be described. The specific name, in the absence of other means of knowing the peculiarities of the animal than those afforded by the skull, may be most conveniently taken from the district (La Plata), in which its remains were first discovered.
The dimensions of the cranium of the Toxodon Platensis amply attest that the animal to which it belonged was of a magnitude attained by few terrestrial quadrupeds, and only to be compared, in this respect, with the larger Pachyderms, or the extinct Megatherium. The length of the skull (of which a base view of the natural size is given in Plate [I].) is two feet four inches: the extreme breadth one foot four inches. The other requisite admeasurements are given in the table at the conclusion of this description.
The general form of the skull, as seen from above, is pyriform; but viewed sideways, and without the lower jaw, it is semi-ovate; it is depressed, elongate, of considerable breadth, including the span of the zygomatic arches, but becoming rather suddenly contracted anterior to them, the facial part thence growing narrower to near the muzzle, which again slightly expands.
Among the first peculiarities which strike the observer, is the aspect of the plane of the occipital foramen, and of the occipital or posterior region of the cranium, the latter of which inclines from below upwards and forwards at an angle of 50° with the basal line of the skull. This slope of the back part of the skull is one of the characteristics of the Dinotherium; it is common to all the Cetacea, and is met with in a slighter degree in many Rodentia, and in the great Ant-eater and some others of the Edentate order. The corresponding aspect of the foramen magnum presents nearly the opposite extreme to man in the occipital scale, proposed by Daubenton to determine the diversities of the form of the cranium, as a gage of the intelligence of different animals[[10]]; and the indication of the limited capacity of the Toxodon, thus afforded, is strengthened by the very small proportion, which the bony walls of the cerebral cavity bear to the zygomatic and maxillary parts of the skull, and to the size of the vertebral column, as indicated by the condyloid processes, and foramen magnum.
The zygomatic arches are of remarkable size and strength; they commence immediately anterior to the sides of the occipital plane, increase in vertical extent as they pass outwards, forwards and downwards, and are suddenly contracted as they bend inwards to abut against the sides of the sockets of the two posterior molar teeth.
The cranial cavity is remarkably narrow at the space included by the zygomatic arches; being, as it were, excavated on each side to augment the space for the lodgment of the temporal muscles, so that its diameter at this part is less than that of the anterior extremity of the upper jaw. The upper surface of the cranium expands to form the post-orbital processes, and again contracts anterior to these.
The muscular ridges, or other characters, at the top of the skull, cannot be precisely determined, as a great proportion of the outer table of the bone is broken away, exposing a coarse and thick diplöe. There seems, however, to have been a strong ridge separating the occipital from the coronal or upper surface of the cranium. The form of the remaining parts, which are modified in relation to the attachment of the muscles of the jaws, indicates that these were powerfully developed both for the offices of mastication and prehension. The general form of the skull, while it presents certain points of resemblance to that of the aquatic Pachydermata, and even of the Carnivora, has much that is peculiar to itself; but, in the facial part, it approaches the nearest to that of the Rodentia; and the dentition of the Toxodon, as exhibited in the upper jaw, corresponds with that which characterizes the Rodent Order.
The teeth of the Toxodon consist of molars and incisors, separated by a long diastema, or toothless space. In the upper jaw the molars are fourteen in number, there being seven on each side; the incisors four, one very large, and one small, in each intermaxillary bone.
The general form and nature of the teeth are indicated by the sockets; and the structure of the grinders is exhibited in a broken molar, the last in the series on the left side of the jaw of the present cranium (See a figure of the grinding surface restored of this tooth, fig. 2, Pl. [I].), and by another perfect molar, the last but one on the right side of the upper jaw, which, though not belonging to the same individual as the skull here described, undoubtedly appertains to the same species. This latter tooth (Fig. 3, Pl. [I].; figs. 2 and 3, Pl. [IV].) was found by itself, embedded in the banks of the Rio Tercero, or Carcarana, near the Parana, at the distance of a hundred and eighty miles from the locality where the head was discovered. Fragments of a molar tooth of a Toxodon, apparently the seventh of the left side, upper jaw, were also found at Bajada de Sta Fé, in the province of Entre Rios, distant forty miles from the mouth of the Rio Tercero.
All the molar teeth are long and curved, and without fangs,[[11]] as in most of the herbivorous species of the Rodent Order: in those, however, with curved grinders, as the Aperea or Guinea-pig, and Cavia Patachonica, the concavity of the upper grinders is directed outward, the fangs of the teeth of the opposite sides diverging as they ascend in the sockets; but, in the Toxodon, the convexity of the grinders is outward, and the fangs converge and almost meet at the middle line of the palate, forming a series of arches, capable of overcoming immense resistance from pressure. (See the upper view of the skull, Plate [III]., in which the fractures expose to view a part of the series of these arched sockets.)
Of the incisors, the two small ones (the sockets of which are indicated at s s, Pl. [III].) are situated in the middle of the front of the upper jaw, close to the suture between the intermaxillaries, and the two large ones in immediate contiguity with the small incisors, which they greatly exceed in size. The sockets of the two large incisors (t t, Pl. [III].) extend backwards, in an arched form, preserving a uniform diameter, as far as the commencement of the alveoli of the molar teeth: the curve which they describe is the segment of a circle; the position, form, and extent of the sockets of these incisors are the same as in those of the corresponding teeth of the Rodentia.
The matrix, or secreting pulp of the large incisors, was lodged, as in the Rodentia, in close proximity with the sockets of the anterior molars; and we are enabled to infer, from the form of the incisive sockets, notwithstanding the absence of the teeth themselves, that the pulp was persistent, and that the growth of these incisors, like those of the Rodentia, continued throughout life.
This condition, joined with the form and curvature of the socket, implies a continual wearing away of the crown of the tooth by attrition against opposing incisors of a corresponding structure in the lower jaw: and as a corollary, it may be inferred that the teeth in question had a partial coating of enamel, to produce a cutting edge, and were, in fact, true dentes scalprarii. The number of incisors in the upper jaw of Toxodon, is not without its parallel in the Rodent Order, the genus Lepus being characterized by four, instead of two superior incisors, which also present a similar relative size but have a different relative position, the small incisors, in the hare and rabbit, being so placed immediately behind the large pair, as to receive the appulse of the single pair of incisors in the lower jaw.
In the Toxodon the position of the incisors, in the same transverse line, might lead to the inference, that they were opposed by a corresponding number in the lower jaw; but the numerous examples of inequality, in the number of incisors, in the upper and lower jaws of existing mammalia, forbid any conclusion on this point.[[12]] The sockets of the small mesial incisors of the Toxodon (s s, Pl. [III].) gradually diminish in size, as they penetrate the intermaxillary bones, and we may, therefore, infer that the pulp was gradually absorbed in the progress of their development; and that, like ordinary incisors, their growth was of limited duration, and their lodgment in the jaw effected by a single conical fang.
I may observe, that the formation of a fang is the necessary consequence of the gradual absorption of the matrix or pulp of a tooth; for the pulp continues, as it diminishes in size, to deposit ivory upon the inner surface of the cavity of the tooth from which it is receding, and the tooth or fang thus likewise progressively diminishes in size. The formation of the socket proceeds uninterruptedly, and the bone encroaching upon the space left by the tooth, closely surrounds the wasting fang, and affords it a firm support; and thus an inference may be drawn from the form of the socket alone, as to whether the tooth it contained had or had not one or more conical fangs, and consequently whether its growth was temporary or uninterrupted.
Applying this reasoning to the molar teeth of the Toxodon, we infer that their growth, like those of most of the Phytiphagous Rodents, of the Megatherium and Armadillo, was perpetual, because their sockets are continued of uniform size from the open to the closed extremity; and the molar tooth which is preserved proves the accuracy of the deduction, inasmuch as its base is excavated by a large conical cavity for the lodgment of the pulp, the continued activity of which was the compensation here designed to meet the effects of attrition on the opposite or grinding surface of the tooth.
The molar tooth discovered by Mr. Darwin in the banks of the Tercero, not only belonged to the same species as the skull under consideration, but to an individual of the same size; it fits exactly into the socket next to the posterior one of the right side. The figures subjoined of this molar tooth (Fig. 3, Pl. [I].; figs. 2 and 3, Pl. [IV].) almost preclude the necessity of a description. The transverse section of the tooth gives an irregular, unequal sided, prism; the two broadest sides of which converge to the anterior angle, which is obtusely rounded. The outer surface of the tooth (fig. 2, Pl. [IV].) is slightly concave in the transverse direction, but undulating, from the presence of two slight convex risings which traverse the tooth lengthwise. The inner surface presents at its anterior part a slightly concave surface, and posteriorly two prominent longitudinal convex ridges, separated by a groove which is flat at the bottom, and from the anterior angle of which the reflected fold of enamel penetrates the substance of the tooth, advancing obliquely forwards, rather more than half-way across the body of the tooth. A longitudinal ridge of bone projects from the internal side of the socket, and fits into the groove above mentioned, and as a corresponding ridge exists in all the sockets of the grinders, save the two anterior small ones, we may infer that the five posterior grinders on each side, had a similar structure to the tooth above described. The external layer of enamel is uniformly about half a line in thickness; it is interrupted for the extent of nearly three lines at the anterior angle, and for more than double that extent at the posterior part of the tooth, which is consequently worn down much below the level of the rest of the grinding surface. Where the ivory is thus unprotected by the enamel, it has a coat of cæmentum, which also fills up the small interval at the origin of the reflected fold of enamel. On the grinding surface of the entire tooth, and on the fractured ends of the mutilated molars, the component fibres, or tubules, of the ivory, are readily perceptible by the naked eye, diverging from the line which indicates the last remains of the cavity of the pulp of the tooth, as it was progressively obliterated during growth.
Although the complication of the grinding surface by the inflection of simple or straight folds of enamel is peculiarly characteristic of the Rodent type, we must regard the number of molar teeth, and their diminution of size as they advance towards the anterior part of the jaw, in the Toxodon, as indicative of a deviation from that order, and an approach to the Pachyderms. The common number of grinders in the upper jaw of Rodent animals is eight, four on each side. In some genera, as Lemmus, Mus, Cricetus, there are only three on each side, and in Hydromys and Aulacodus, only two on each side. In Lepus, however, we find six on each side of the upper, and five on each side of the lower jaw. The Toxodon, like the Tapir and Hippopotamus, has seven on each side of the upper jaw: the first in each of these species being the smallest. It is worthy of notice, however, that the Capybara which adheres to the Rodent type in the number of its molars, presents in the vastly increased size, and additional number of component laminæ of the posterior grinders, an approximation to the pachydermatous character just adduced, and the bony palate at the same time presents an expansion between these molars, offering a resemblance to the Toxodon which I have not found in any other Rodent besides the Capybara.
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 |